Many ways to build an angler: diversity of feeding morphologies in a deep-sea evolutionary radiation.

Almost nothing is known about the diets of bathypelagic fishes, but functional morphology can provide useful tools to infer ecology. Here we quantify variation in jaw and tooth morphologies across anglerfishes (Lophiiformes), a clade spanning shallow and deep-sea habitats. Deep-sea ceratioid anglerfishes are considered dietary generalists due to the necessity of opportunistic feeding in the food-limited bathypelagic zone. We found unexpected diversity in the trophic morphologies of ceratioid anglerfishes. Ceratioid jaws span a functional continuum ranging from species with numerous stout teeth, a relatively slow but forceful bite, and high jaw protrusibility at one end (characteristics shared with benthic anglerfishes) to species with long fang-like teeth, a fast but weak bite and low jaw protrusibility at the other end (including a unique 'wolftrap' phenotype). Our finding of high morphological diversity seems to be at odds with ecological generality, reminiscent of Liem's paradox (morphological specialization allowing organisms to have broader niches). Another possible explanation is that diverse ceratioid functional morphologies may yield similar trophic success (many-to-one mapping of morphology to diet), allowing diversity to arise through neutral evolutionary processes. Our results highlight that there are many ways to be a successful predator in the deep sea.

Heterodonty and double occlusion in Manidens condorensis: a unique adaptation in an Early Jurassic ornithischian improving masticatory efficiency.

New materials of the ornithischian dinosaur Manidens condorensis highlight a strong heterodonty between the upper and lower dentitions and reveal a novel occlusion type previously unreported in herbivorous dinosaurs. The diamond-shaped maxillary teeth have prominent cingular entolophs in a V- to Z-shaped configuration that are absent in dentary teeth. These cingular entolophs bear denticles and serrations with vertical wear that is coplanar with the apical wear facets, supporting their involvement in chewing. The separated apical and basal wear in dentary teeth is consistent with the apical and cingular wear in maxillary teeth, indicating an alternate occlusion, an orthal jaw motion, and shearing interactions between marginal and cingular edges in a double occlusion. Measurements of the length and wear area along the marginal and cingular edges indicate that the latter are functionally equivalent to adding eight teeth to a maxillary toothrow of ten, almost doubling the lengths of cutting edges and the degree of intraoral processing, while maintaining a plesiomorphic skull anatomy, an adaptation to herbivory unique in Ornithischia.

Controlled feeding trials with ungulates: a new application of in vivo dental molding to assess the abrasive factors of microwear.

Microwear, the quantification of microscopic scratches and pits on the occlusal surfaces of tooth enamel, is commonly used as a paleodietary proxy. For ungulates (hoofed mammals), scratch-dominant microwear distinguishes modern grazers from browsers, presumably as a result of abrasion from grass phytoliths (biogenic silica). However, it is also likely that exogenous grit (i.e. soil, dust) is a contributing factor to these scratch-dominant patterns, which may reflect soil ingestion that varies with feeding height and/or environmental conditions (e.g. dust production in open and/or arid habitats). This study assessed the contribution of exogenous grit to tooth wear by measuring the effects of fine- and medium-grained silica sand on tooth enamel using a novel live-animal tooth-molding technique. It therefore constitutes the first controlled feeding experiment using ungulates and the first in vivo experiment using abrasives of different sizes. Four sheep were fed three diet treatments: (1) a mixture of Garrison and Brome hay (control), (2) hay treated with fine-grained silica sand (180-250 µm) and (3) hay treated with medium-grained silica sand (250-425 µm). We found a significant increase in pit features that was correlated with an increase in grain size of grit, corroborating earlier chewing simulation experiments that produced pits through grit-induced abrasion (i.e. the 'grit effect'). Our results support an interpretation of large silica grains fracturing to create smaller, more abundant angular particles capable of abrasion, with jaw movement defining feature shape (i.e. scratch or pit).

A switch in jaw form-function coupling during the evolution of mammals.

The evolutionary shift from a single-element ear, multi-element jaw to a multi-element ear, single-element jaw during the transition to crown mammals marks one of the most dramatic structural transformations in vertebrates. Research on this transformation has focused on mammalian middle-ear evolution, but a mandible comprising only the dentary is equally emblematic of this evolutionary radiation. Here, we show that the remarkably diverse jaw shapes of crown mammals are coupled with surprisingly stereotyped jaw stiffness. This strength-based morphofunctional regime has a genetic basis and allowed mammalian jaws to effectively resist deformation as they radiated into highly disparate forms with markedly distinct diets. The main functional consequences for the mandible of decoupling hearing and mastication were a trade-off between higher jaw stiffness versus decreased mechanical efficiency and speed compared with non-mammals. This fundamental and consequential shift in jaw form-function underpins the ecological and taxonomic diversification of crown mammals. This article is part of the theme issue 'The mammalian skull: development, structure and function'.

New Reconstruction of Cranial Musculature in Ornithischian Dinosaurs: Implications for Feeding Mechanisms and Buccal Anatomy.

The charismatic and diverse ornithischian dinosaurs exhibited some of the most extreme examples of cranial anatomy, inspiring decades of investigation into their muscular anatomy. Current ornithischian jaw muscle reconstructions, although parsimonious, pose concerns of small adductor muscles and caudally displaced insertions relative to mandibular proportions. Here, craniomandibular material of ornithischian genera spanning all subclades is reexamined for osteological correlates indicative of intracranial and oral soft tissues. M. adductor mandibulae externus (mAME) has traditionally been reconstructed as solely inserting along the caudal margin of the coronoid process for jaw closure. Here, a new mAME reconstruction is proposed in derived ornithischians, with the superficial-most mAME layer reconstructed as a rostrolabial expansion of muscle, exiting the cranium rostroventrally beneath a unique, laterally flaring jugal and inserting along the lateral surface of the coronoid process and its rostrally extending, shelf-like labial dentary ridge (LDR). Through previous dental microwear and morphological studies, ceratopsians, hadrosaurids, and ankylosaurs are known to have implemented a major palinal feeding component in their jaw motions, unlike other primarily basal ornithischians. This rostral fan-like extension of muscle in these derived clades would create a greater mandibular support system and mechanical advantage along the labial margin of the jaw, cradling the entire mandible while lifting it up into occlusion and retracting it. In hadrosaurids and ankylosaurs, this rostrolabially expanding muscle also acts in medial rotation of the dentaries about their long axes. With these new reconstructions, the notion of a novel, unparsimonious "cheek" muscle is rejected, with further discussion of plausible buccal soft tissues. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 303:347-362, 2020. © 2018 American Association for Anatomy.

Mechanobehavior and mandibular ramus length in different facial phenotypes.

OBJECTIVES: To test the hypotheses that mechanobehavior scores (MBS) were correlated with mandibular ramus lengths (Co-Go) and differed between facial phenotypes. MATERIALS AND METHODS: Subjects gave informed consent to participate. Co-Go (mm), mandibular plane angles (SN-GoGn, °), and three-dimensional anatomy were derived from cephalometric radiography or cone beam computed tomography. Temporomandibular joint (TMJ) energy densities (ED) (mJ/mm3) were measured using dynamic stereometry and duty factors (DF) (%) were measured from electromyography, to calculate MBS (= ED2 × DF,) for each TMJ. Polynomial regressions, K-means cluster analysis, and analysis of variance (ANOVA) with Tukey post-hoc tests were employed. RESULTS: Fifty females and 23 males produced replete data. Polynomial regressions showed MBS were correlated with Co-Go (females, R2 = 0.57; males, R2 = 0.81). Cluster analysis identified three groups (P < .001). Dolichofacial subjects, with shorter normalized Co-Go, clustered into two subgroups with low and high MBS compared to brachyfacial subjects with longer Co-Go. SN-GoGn was significantly larger (P < .03) in the dolichofacial subgroups combined (33.0 ± 5.9°) compared to the brachyfacial group (29.8 ± 5.5°). CONCLUSIONS: MBS correlated with Co-Go within sexes and differed significantly between brachyfacial and dolichofacial subjects.

Evolutionary Trends in the Jaw Adductor Mechanics of Ornithischian Dinosaurs.

Jaw mechanics in ornithischian dinosaurs have been widely studied for well over a century. Most of these studies, however, use only one or few taxa within a given ornithischian clade as a model for feeding mechanics across the entire clade. In this study, mandibular mechanical advantages among 52 ornithischian genera spanning all subclades are calculated using 2D lever arm methods. These lever arm calculations estimate the effect of jaw shape and difference in adductor muscle line of action on relative bite forces along the jaw. Results show major instances of overlap between taxa in tooth positions at which there was highest mechanical advantage. A relatively low bite force is seen across the tooth row among thyreophorans (e.g., stegosaurs and ankylosaurs), with variation among taxa. A convergent transition occurs from a more evenly distributed bite force along the jaw in basal ornithopods and basal marginocephalians to a strong distal bite force in hadrosaurids and ceratopsids, respectively. Accordingly, adductor muscle vector angles show repeated trends from a mid-range caudodorsal orientation in basal ornithischians to a decrease in vector angles indicating more caudally oriented jaw movements in derived taxa (e.g., derived thyreophorans, basal ornithopods, lambeosaurines, pachycephalosaurs, and derived ceratopsids). Analyses of hypothetical jaw morphologies were also performed, indicating that both the coronoid process and lowered jaw joint increase moment arm length therefore increasing mechanical advantage of the jaw apparatus. Adaptive trends in craniomandibular anatomy show that ornithischians evolved more complex feeding apparatuses within different clades as well as morphological convergences between clades.

The Predentary Bone and Its Significance in the Evolution of Feeding Mechanisms in Ornithischian Dinosaurs.

The characteristic predentary bone in ornithischian dinosaurs is a unique, unpaired element located at the midline of the mandibular symphysis. Although traditionally thought to only be a plant "nipping" bone, the true functional significance of this bone among feeding mechanisms of ornithischian dinosaurs is poorly known. Recent studies of a select few ornithischian genera have suggested rotation of the mandibular corpora around their long axes relative to their midline joint articulation with the predentary bone. This study aims to re-evaluate these hypotheses as well as provide in-depth qualitative comparative descriptions of predentary bone morphology in ornithischian genera throughout all subclades, including heterodontosaurids, thyreophorans, ornithopods, and marginocephalians. Descriptions evaluate overall shape of the predentary, its articular surfaces contacting the rostral ends of the dentaries, and the morphology of the rostral extent of the dentaries and their midline symphysis. Functionally relevant morphologies in each predentary morphotype are accentuated for further speculation of feeding mechanisms. Three predentary morphotypes are described throughout ornithischian subclades and each plays a unique role in feeding adaptations. Most notably, the predentary likely evolved as a midline axial point of the mandibular symphysis for simultaneous variable movement or rotation of the mandibular corpora in many, but not all, taxa. This simultaneous movement of the hemimandibles would have aided in feeding on both sides of the jaw at once. The function of the predentary as well as other jaw adaptations is discussed for genera throughout all subclades, focusing on both general shape and joint morphology. Anat Rec, 299:1358-1388, 2016. © 2016 Wiley Periodicals, Inc.

Dental microwear reveals mammal-like chewing in the neoceratopsian dinosaur Leptoceratops gracilis.

Extensive oral processing of food through dental occlusion and orbital mandibular movement is often cited as a uniquely mammalian trait that contributed to their evolutionary success. Save for mandibular translation, these adaptations are not seen in extant archosaurs or lepidosaurs. In contrast, some ornithischian dinosaurs show evidence of precise dental occlusion, habitual intraoral trituration and complex jaw motion. To date, however, a robust understanding of the diversity of jaw mechanics within non-avian dinosaurs, and its comparison with other vertebrates, remains unrealized. Large dental batteries, well-developed dental wear facets, and robust jaws suggests that neoceratopsian (horned) dinosaurs were capable chewers. But, biomechanical analyses have assumed a relatively simple, scissor-like (orthal) jaw mechanism for these animals. New analyses of dental microwear, presented here, show curvilinear striations on the teeth of Leptoceratops. These features indicate a rostral to caudal orbital motion of the mandible during chewing. A rostrocaudal mandibular orbit is seen in multituberculates, haramiyid allotherians, and some rodents, and its identification in Leptoceratops gracilis is the first evidence of complex, mammal-like chewing in a ceratopsian dinosaur. The term circumpalinal is here proposed to distinguish this new style of chewing from other models of ceratopsian mastication that also involve a palinal component. This previously unrecognized complexity in dinosaurian jaw mechanics indicates that some neoceratopsian dinosaurs achieved a mammalian level of masticatory efficiency through novel adaptive solutions.