A new psittacosaur from Inner Mongolia and the parrot-like structure and function of the psittacosaur skull

We describe a new species of psittacosaur, Psittacosaurus gobiensis, from the Lower Cretaceous of Inner Mongolia and outline a hypothesis of chewing function in psittacosaurs that in many respects parallels that in psittaciform birds. Cranial features that accommodate increased bite force in psittacosaurs include an akinetic skull (both cranium and lower jaws) and differentiation of adductor muscle attachments comparable to that in psittaciform birds. These and other features, along with the presence of numerous large gastroliths, suggest that psittacosaurs may have had a high-fibre, nucivorous (nut-eating) diet.Psittacosaurs, alone among ornithischians, generate oblique wear facets from tooth-to-tooth occlusion without kinesis in either the upper or lower jaws. This is accomplished with a novel isognathous jaw mechanism that combines aspects of arcilineal (vertical) and propalinal (horizontal) jaw movement. Here termed clinolineal (inclined) jaw movement, the mechanism uses posteriorly divergent tooth rows, rather than kinesis, to gain the added width for oblique occlusion. As the lower tooth rows are drawn posterodorsally into occlusion, the increasing width between the upper tooth rows accommodates oblique shear. With this jaw mechanism, psittacosaurs were able to maintain oblique shearing occlusion in an akinetic skull designed to resist high bite forces.     

The bite force of the largest fossil rodent (Hystricognathi,Caviomorpha, Dinomyidae)

Blanco R.E., Rinderknecht, A. & Lecuona, G. 2011: The bite force of the largest fossil rodent (Hystricognathi, Caviomorpha, Dinomyidae). Lethaia, Vol. 45, pp. 157–163. An exceptionally well‐preserved skull of the largest fossil rodent Josephoartigasia monesi allows the first analysis of the bite mechanics of this group of South American giant rodents. In this study, we reconstructed the main anatomical features of the skull of this Pliocene rodent, relating them to the bite force at incisors. Bite force was estimated using three different techniques. Two methods suggest that bite forces at incisors of around 1000 N were possible for these mammals. However, the incisors seem to be stronger than expected for this bite force implying that the bite forces may have been greater than 3000 N. We consider three hypotheses: allometric effects, teeth digging or defence against predators, to explain our results. □Bite force, Dinomyidae, incisors, largest rodent, Pliocene.     

Comparison between in vivo and theoretical bite performance: Using multi-body modelling to predict muscle and bite forces in a reptile skull

In biomechanical investigations, geometrically accurate computer models of anatomical structures can be created readily using computed-tomography scan images. However, representation of soft tissue structures is more challenging, relying on approximations to predict the muscle loading conditions that are essential in detailed functional analyses. Here, using a sophisticated multi-body computer model of a reptile skull (the rhynchocephalian Sphenodon), we assess the accuracy of muscle force predictions by comparing predicted bite forces against in vivo data. The model predicts a bite force almost three times lower than that measured experimentally. Peak muscle force estimates are highly sensitive to fibre length, muscle stress, and pennation where the angle is large, and variation in these parameters can generate substantial differences in predicted bite forces. A review of theoretical bite predictions amongst lizards reveals that bite forces are consistently underestimated, possibly because of high levels of muscle pennation in these animals. To generate realistic bites during theoretical analyses in Sphenodon, lizards, and related groups we suggest that standard muscle force calculations should be multiplied by a factor of up to three. We show that bite forces increase and joint forces decrease as the bite point shifts posteriorly within the jaw, with the most posterior bite location generating a bite force almost double that of the most anterior bite. Unilateral and bilateral bites produced similar total bite forces; however, the pressure exerted by the teeth is double during unilateral biting as the tooth contact area is reduced by half.     

A functional analysis of carnassial biting

The jaw mechanism of carnivores is studied using an idealized model (Greaves, 1978). The model assumes: (i) muscle activity on both sides of the head, and (ii) that the jaw joints and the carnassial teeth are single points of contact between the skull and the lower jaw during carnassial biting. The model makes the following predictions: (i) in carnivores with carnassial teeth the resultant force of the jaw muscles will be positioned approximately 60% of the way from the jaw joint to the tooth—this arrangement delivers the maximum bite force possible together with a reasonably wide gape (remembering that bite force and gape cannot both be maximized); (ii) in an evolutionary sense, if greater bite force is required at the carnassial tooth, either the animal will get larger so as to deliver an absolutely larger bite force or the architecture of the muscles may change, becoming more pinnate, for example, but jaw geometry (i.e. the relative positions of the jaw joints, the carnassial tooth, and the muscle resultant force) will not change; (iii) if greater gape is required, the animal will get larger so as to have longer jaws and therefore an absolutely wider gape or change its muscle architecture allowing for greater stretch while the geometry remains unchanged; and (iv) in animals with a longer shearing region (e.g. the extinct hyaenodonts) the shearing region will be approximately 20% of jaw length and the muscle resultant force will be positioned approximately 60% of the way from the jaw joint to the most anterior shearing tooth.     

Feedback control from the jaw joints during biting: An investigation of the reptile Sphenodon using multibody modelling

Sphenodon, a lizard-like reptile, is the only living representative of a group that was once widespread at the time of the dinosaurs. Unique jaw mechanics incorporate crushing and shearing motions to breakdown food, but during this process excessive loading could cause damage to the jaw joints and teeth. In mammals like ourselves, feedback from mechanoreceptors within the periodontal ligament surrounding the teeth is thought to modulate muscle activity and thereby minimise such damage. However, Sphenodon and many other tetrapods lack the periodontal ligament and must rely on alternative control mechanisms during biting. Here we assess whether mechanoreceptors in the jaw joints could provide feedback to control muscle activity levels during biting. We investigate the relationship between joint, bite, and muscle forces using a multibody computer model of the skull and neck of Sphenodon. When feedback from the jaw joints is included in the model, predictions agree well with experimental studies, where the activity of the balancing side muscles reduces to maintain equal and minimal joint forces. When necessary, higher, but asymmetric, joint forces associated with higher bite forces were achievable, but these are likely to occur infrequently during normal food processing. Under maximum bite forces associated with symmetric maximal muscle activation, peak balancing side joint forces were more than double those of the working side. These findings are consistent with the hypothesis that feedback similar to that used in the simulation is present in Sphenodon.     

Correlation between the size,shape and position of the teeth on the jaws and the bite force in Theropoda

Theropods have fascinated both paleontologists and the general public due to their large diversity of sizes and morphologies. They also present a large variation in tooth morphologies. Previous studies have estimated the bite force of several specimens. The goal of this study is to determine if there is a correlation between the tooth size, shape and position on the skull and mandible and the bite force of these dinosaurs. Measurements were made on several theropods, including the bending strength of the teeth on the anterior-posterior and the mediolateral axes of the jaws, as well as the bending strength of the mandible, and were compared to fossil and modern Crocodylia. We observed that several bending strength maxima of the teeth trends were aligned with key areas of the mandible, and that the size, shape as well as the position of the teeth on the jaws were correlated with the bite force of both Crocodylia and theropods, which can be related to their diet and feeding habits.     

Biomechanical properties of the jaws of two species of Clevosaurus and a reanalysis of rhynchocephalian dentary morphospace

Rhynchocephalians were a successful, globally distributed group of diapsid reptiles that thrived in the Mesozoic. Multiple species of Clevosaurus existed worldwide in the Late Triassic and Early Jurassic, characterized by shearing bladelike teeth perhaps functionally analogous to the carnassial teeth of mammals. Morphometric analysis shows that the dentary morphospace of clevosaurs differs significantly from that of other rhynchocephalians. Five Clevosaurus species occupied islands in the Bristol Channel archipelago of the UK, but generally not those occupied by mammaliaforms, suggesting dietary character displacement. Identifying the diet of such ancient, small tetrapods has been difficult. To identify the nature of their feeding mechanics and ecology, we apply finite element analysis to two near complete three-dimensional skulls of the species Clevosaurus hudsoni and Clevosaurus cambrica to estimate bite force, resistance to bending and torsion and the distribution of stresses in the jaws during biting. Both species had bite forces and tooth pressures sufficient to break apart chitin indicating that, like early Mesozoic mammaliaforms, clevosaurs could feed on tough-shelled beetles and possibly small vertebrates. In addition, the mechanical advantage of the jaws falls within the range of early mammaliaforms, so though we cannot demonstrate niche partitioning between members of the two clades, it raises the prospect that they may have been functionally similar.     

The craniomandibular mechanics of being human

Diminished bite force has been considered a defining feature of modern Homo sapiens, an interpretation inferred from the application of two-dimensional lever mechanics and the relative gracility of the human masticatory musculature and skull. This conclusion has various implications with regard to the evolution of human feeding behaviour. However, human dental anatomy suggests a capacity to withstand high loads and two-dimensional lever models greatly simplify muscle architecture, yielding less accurate results than three-dimensional modelling using multiple lines of action. Here, to our knowledge, in the most comprehensive three-dimensional finite element analysis performed to date for any taxon, we ask whether the traditional view that the bite of H. sapiens is weak and the skull too gracile to sustain high bite forces is supported. We further introduce a new method for reconstructing incomplete fossil material. Our findings show that the human masticatory apparatus is highly efficient, capable of producing a relatively powerful bite using low muscle forces. Thus, relative to other members of the superfamily Hominoidea, humans can achieve relatively high bite forces, while overall stresses are reduced. Our findings resolve apparently discordant lines of evidence, i.e. the presence of teeth well adapted to sustain high loads within a lightweight cranium and mandible.     

Bite force and body mass of the fossil rodent Telicomys giganteus (Caviomorpha,Dinomyidae)

An exceptionally well-preserved skull of the Pliocene rodent Telicomys giganteus allowed the first estimation of body mass and analysis of the bite mechanics of this species of South American giant rodent. In this study, we reconstructed the main anatomical features of the skull of this Pliocene rodent and related them to the bite force at the incisors. The average of an estimation body mass gives 100 kg. We also estimated the bite force using three different techniques. Two methods suggest that bite forces at the incisors have a range of 500–1000 N. However, the incisors seem to be stronger than expected for this bite force, implying that the bite forces may have been greater than 2000 N. We consider the hypothesis of defense against predators or other agonistic behavior to explain our results.     

Tooth Root Morphology in the Early Pliocene African Bear Agriotherium africanum (Mammalia, Carnivora, Ursidae) and its Implications for Feeding Ecology

Tooth root surface areas serve as proxies for bite force potentials, and by extension, dietary specialization in extant carnivorans. Here, we investigate the feeding ecology of the extinct large-bodied ursid Agriotherium africanum, by comparing its root surface areas (reconstructed with the aid of computed tomography and three-dimensional image processing) and bite force estimates, with those of extant carnivorans. Results show that in absolute terms, canine and carnassial bite forces, as well as root surface areas were highest in A. africanum. However, when adjusted for skull size, A. africanum’s canine roots were smaller than those of extant solitary predators. With teeth being the limiting factor in the masticatory system, low canine root surface areas suggest that A. africanum would have struggled to bring down large vertebrate prey. Its adjusted carnassial root sizes were found to be smaller than those of extant hard object feeders and the most carnivorous tough object feeders, but larger than those of extant omnivorous ursids and Ursus maritimus. This and the fact that it displayed its highest postcanine root surface areas in the carnassial region (rather than the most distal tooth in the tooth row) suggest that A. africanum consumed more vertebrate tissue than extant omnivorous ursids. With an apparent inability to routinely bring down large prey or to consume mechanically demanding skeletal elements, its focus was most likely on tough tissue, which it acquired by actively scavenging the carcasses of freshly dead/freshly killed animals. Mechanically less demanding skeletal elements would have been a secondary food source, ingested and processed mainly in association with muscle and connective tissue.     

Functional morphology of durophagy in black carp,Mylopharyngodon piceus

The black carp, Mylopharyngodon piceus (Osteichthyes: Cyprinidae), crushes its snail and other molluscan prey with robust pharyngeal jaws and strong bite forces. Using gross morphology, histological sectioning, and X‐ray reconstruction of moving morphology (XROMM), we investigated structural, behavioral, and mechanical aspects of pharyngeal jaw function in black carp. Strut‐like trabeculae in their pharyngeal jaws support large, molariform teeth. The teeth occlude with a hypertrophied basioccipital process that is also reinforced with stout trabeculae. A keratinous chewing pad is firmly connected to the basioccipital process by a series of small bony projections from the base of the pedestal. The pharyngeal jaws have no bony articulations with the skull, and their position is controlled by five paired muscles and one unpaired median muscle. Black carp can crush large molluscs, so we used XROMM to compare pharyngeal jaw postures as fish crushed ceramic tubes of increasing sizes. We found that black carp increase pharyngeal jaw gape primarily by ventral translation of the jaws, with ventral rotation and lateral flaring of the jaws also increasing the space available to accommodate large prey items. A stout, robust ligament connects left and right jaws together firmly, but allows some rotation of the jaws relative to each other. Contrasting with the pharyngeal jaw mechanism of durophagous perciforms with fused left and right lower pharyngeal jaws, we hypothesize that this ligamentous connection may serve to decouple tensile and compressive forces, with the tensile forces borne by the ligament and the compressive forces transferred to the prey. J. Morphol. 276:1422–1432, 2015. © 2015 Wiley Periodicals, Inc.     

New marine mammal faunas (Cetacea and Sirenia) and sea level change in the Samlat Formation,Upper Eocene,near Ad-Dakhla in southwestern Morocco

The Samlat Formation is well exposed in coastal sections bordering the Atlantic Ocean south of Ad-Dakhla in southwestern Morocco. Here some 22 m of rhythmically-bedded, chert-rich, marine siltstones and marls are overlain by 1–1.5 m of vertebrate-bearing microconglomeratic sandstone, another 4–8 m of rhythmically-bedded siltstone and marl, and finally a second 3–6 m unit of vertebrate-bearing muddy sandstone. The microconglomeratic and muddy sandstones represent low sea stands in what is otherwise a deeper water sequence. Cetacean skeletons are rare but cetacean vertebrae are common in the lower sandstone (bed B1), where many show the effects of reworking. The cetaceans in bed B1represent a minimum of five species, from smallest to largest: cf. Saghacetus sp., cf. Stromerius sp., Dorudon atrox, cf. Dorudon sp., and Basilosaurus isis. Bed B1 yields rib fragments that may represent sirenians, but sirenians, if present, are rare. The only identifiable cetacean found in the upper sandstone (bed B2) is Basilosaurus sp. Dugongid sirenians identified as cf. Eosiren sp. are the most common mammal in bed B2. We interpret co-occurrence of the typically Early Priabonian species Dorudon atrox and Basilosaurus isis with smaller species more like Middle Priabonian genera Saghacetus osiris and Stromerius nidensis to indicate that bed B1 was deposited during low sea stand Pr-2 between the Early and Middle Priabonian (between the early and middle Late Eocene). Bed B2 is separated from B1 by an interval of deeper water sediment accumulation. Bed B2 could represent a later phase of Pr-2 or a subsequent Priabonian low sea stand (possibly Pr-3).     

Implications of predatory specialization for cranial form and function in canids

The shape of the cranium varies widely among members of the order Carnivora, but the factors that drive the evolution of differences in shape remain unclear. Selection for increased bite force, bite speed or skull strength may all affect cranial morphology. We investigated the relationship between cranial form and function in the trophically diverse dog family, Canidae, using linear morphometrics and finite element (FE) analyses that simulated the internal and external forces that act on the skull during the act of prey capture and killing. In contrast to previous FE-based studies, we compared models using a newly developed method that removes the effects of size and highlights the relationship between shape and performance. Cranial shape varies among canids based on diet, and different selective forces presumably drove evolution of these phenotypes. The long, narrow jaws of small prey specialists appear to reflect selection for fast jaw closure at the expense of bite force. Generalists have intermediate jaw dimensions and produce moderate bite forces, but their crania are comparable in strength to those of small prey specialists. Canids that take large prey have short, broad jaws, produce the largest bite forces and possess very strong crania. Our FE simulations suggest that the remarkable strength of skulls of large prey specialists reflect the additional ability to resist extrinsic loads that may be encountered while struggling with large prey items.     

Mechanoreceptor distribution in stag beetle jaws corresponds to the material stress in fights

Male stag beetles (Lucanidae) use their extremely elongated jaws to pinch their rivals forcefully in male–male battles. The morphology of these jaws has to be a compromise between robustness (to withstand the bite forces), length and weight. Cyclommatus metallifer stag beetles circumvent this trade-off by reducing their bite force when biting with their slender jaw tips. Here we describe the functional mechanism behind the force modulation behaviour. Scanning Electron Microscopy and micro CT imaging show large numbers of small sensors in the jaw cuticle. We find a strong correlation between the distribution of these sensors and that of the material stress in the same jaw region during biting. The jaw sensors are mechanoreceptors with a small protrusion that barely protrudes above the undulating jaw surface. The sensors stimulate dendrites that extend from the neuronal cell body through the entire thickness of the jaw exoskeleton towards the sensors at the external surface. They form a sensory field that functions in a feedback mechanism to control the bite muscle force. This negative feedback mechanism enabled the stag beetles to evolve massive bite muscles without risking overloading their valuable jaws.     

Feeding biomechanics in Acanthostega and across the fish–tetrapod transition

Acanthostega is one of the earliest and most primitive limbed vertebrates. Its numerous fish-like features indicate a primarily aquatic lifestyle, yet cranial suture morphology suggests that its skull is more similar to those of terrestrial taxa. Here, we apply geometric morphometrics and two-dimensional finite-element analysis to the lower jaws of Acanthostega and 22 other tetrapodomorph taxa in order to quantify morphological and functional changes across the fish–tetrapod transition. The jaw of Acanthostega is similar to that of certain tetrapodomorph fish and transitional Devonian taxa both morphologically (as indicated by its proximity to those taxa in morphospace) and functionally (as indicated by the distribution of stress values and relative magnitude of bite force). Our results suggest a slow tempo of morphological and biomechanical changes in the transition from Devonian tetrapod jaws to aquatic/semi-aquatic Carboniferous tetrapod jaws. We conclude that Acanthostega retained a primitively aquatic lifestyle and did not possess cranial adaptations for terrestrial feeding.     

Dry versus wet and gross: Comparisons between the dry skull method and gross dissection in estimations of jaw muscle cross-sectional area and bite forces in sea otters

Bite force is a measure of feeding performance used to elucidate links between animal morphology, ecology, and fitness. Obtaining live individuals for in vivo bite-force measurements or freshly deceased specimens for bite force modeling is challenging for many species. Thomason's dry skull method for mammals relies solely on osteological specimens and, therefore, presents an advantageous approach that enables researchers to estimate and compare bite forces across extant and even extinct species. However, how accurately the dry skull method estimates physiological cross-sectional area (PCSA) of the jaw adductor muscles and theoretical bite force has rarely been tested. Here, we use an ontogenetic series of southern sea otters (Enhydra lutris nereis) to test the hypothesis that skeletomuscular traits estimated from the dry skull method accurately predicts test traits derived from dissection-based biomechanical modeling. Although variables from these two methods exhibited strong positive relationships across ontogeny, we found that the dry skull method overestimates PCSA of the masseter and underestimates PCSA of the temporalis. Jaw adductor in-levers for both jaw muscles and overall bite force are overestimated. Surprisingly, we reveal that sexual dimorphism in craniomandibular shape affects temporalis PCSA estimations; the dry skull method predicted female temporalis PCSA well but underestimates male temporalis PCSA across ontogeny. These results highlight the importance of accounting for sexual dimorphism and other intraspecific variation when using the dry skull method. Together, we found the dry skull method provides an underestimation of bite force over ontogeny and that the underlying anatomical components driving bite force may be misrepresented.     

NEW GENUS OF DORUDONTINE ARCHAEOCETE (CETACEA) FROM THE MIDDLE-TO-LATE EOCENE OF SOUTH CAROLINA

A new archaeocete whale from the late middle or early late Eocene of South Carolina, Chrysocetus healyorum gen. et sp. nov., is described on the basis of a single subadult specimen. This individual includes: a partial skull; hyoid apparatus; lower jaws; teeth; all cervical, some thoracic and some lumbar vertebrae; ribs and sternum; left forelimb elements; and pelves. The specimen includes portions of much of the body, but while some of the bones are fairly complete, others are damaged, particularly the skull. The pelves resemble those of Basilosaurus , documenting a similar stage of hind limb reduction in dorudontines and suggesting that Chrysocetus was not able to support its body on land. The acetabulum for articulation of the femur is well formed and indicates that the hip joint was functional. Chrysocetus is distinguishable from other described dorudontines based on body size, characteristics of the teeth, and forelimb elements. Absence of deciduous teeth in a subadult individual of Chrysocetus may be indicative of an early stage of the evolution of monophyodonty.     

Mechanics of biting in great white and sandtiger sharks

Although a strong correlation between jaw mechanics and prey selection has been demonstrated in bony fishes (Osteichthyes), how jaw mechanics influence feeding performance in cartilaginous fishes (Chondrichthyes) remains unknown. Hence, tooth shape has been regarded as a primary predictor of feeding behavior in sharks. Here we apply Finite Element Analysis (FEA) to examine form and function in the jaws of two threatened shark species, the great white (Carcharodon carcharias) and the sandtiger (Carcharias taurus). These species possess characteristic tooth shapes believed to reflect dietary preferences. We show that the jaws of sandtigers and great whites are adapted for rapid closure and generation of maximum bite force, respectively, and that these functional differences are consistent with diet and dentition. Our results suggest that in both taxa, insertion of jaw adductor muscles on a central tendon functions to straighten and sustain muscle fibers to nearly orthogonal insertion angles as the mouth opens. We argue that this jaw muscle arrangement allows high bite forces to be maintained across a wider range of gape angles than observed in mammalian models. Finally, our data suggest that the jaws of sub-adult great whites are mechanically vulnerable when handling large prey. In addition to ontogenetic changes in dentition, further mineralization of the jaws may be required to effectively feed on marine mammals. Our study is the first comparative FEA of the jaws for any fish species. Results highlight the potential of FEA for testing previously intractable questions regarding feeding mechanisms in sharks and other vertebrates.     

Feeding mechanisms of the sauropod dinosaurs Brachiosaurus,Camarasaurus, Diplodocus and Dicraeosaurus

Skull morphologies and dental wear patterns have been examined in four sauropod genera to evaluate their probable feeding mechanisms. Wear facets on teeth are generally confined to their apices in Brachiosaurus and Dicraeosaurus and they are sometimes also present on the mesial and distal carinae. Skull morphology and dental wear patterns in Diplodocus and Dicraeosaurus are consistent with a raking motion of the jaws during feeding. Diplodocus became mechanically adapted to feed in this way by evolving anteriorly directed teeth in the premaxilla and mesial parts of the maxilla, and by changing the direction of jaw adduction relative to the long axis of the skull. Similar features are present in the few known skulls of Apatosaurus and they may also have been present in Dicraeosaurus. In Brachiosaurus dental wear patterns also imply a raking motion of the jaws, although the more robust skull and teeth and the more vertically directed action of the jaw adductor muscles have led some to suggest the possibility of isognathous occlusion. Camarasaurus employed a powerful bite in its feeding, possibly with slight propaliny of the lower jaw, and its skull was modified to cope with increased stresses arising from mastication. Archaic sauropods appear largely to have employed isognathic occlusion in chopping off vegetation. The raking motion employed by diplodocids and dicraeosaurids was an advanced mode of cropping and stripping, linked evolutionarily to their highly apomorphic cranial morphology.     

Insights into the ecology and evolutionary success of crocodilians revealed through bite-force and tooth-pressure experimentation

Background

Crocodilians have dominated predatory niches at the water-land interface for over 85 million years. Like their ancestors, living species show substantial variation in their jaw proportions, dental form and body size. These differences are often assumed to reflect anatomical specialization related to feeding and niche occupation, but quantified data are scant. How these factors relate to biomechanical performance during feeding and their relevance to crocodilian evolutionary success are not known.

Methodology/Principal Findings

We measured adult bite forces and tooth pressures in all 23 extant crocodilian species and analyzed the results in ecological and phylogenetic contexts. We demonstrate that these reptiles generate the highest bite forces and tooth pressures known for any living animals. Bite forces strongly correlate with body size, and size changes are a major mechanism of feeding evolution in this group. Jaw shape demonstrates surprisingly little correlation to bite force and pressures. Bite forces can now be predicted in fossil crocodilians using the regression equations generated in this research.

Conclusions/Significance

Critical to crocodilian long-term success was the evolution of a high bite-force generating musculo-skeletal architecture. Once achieved, the relative force capacities of this system went essentially unmodified throughout subsequent diversification. Rampant changes in body size and concurrent changes in bite force served as a mechanism to allow access to differing prey types and sizes. Further access to the diversity of near-shore prey was gained primarily through changes in tooth pressure via the evolution of dental form and distributions of the teeth within the jaws. Rostral proportions changed substantially throughout crocodilian evolution, but not in correspondence with bite forces. The biomechanical and ecological ramifications of such changes need further examination.