Articular scaling and body mass estimation in platyrrhines and catarrhines: Modern variation and application to fossil anthropoids.

Body mass is an important component of any paleobiological reconstruction. Reliable skeletal dimensions for making estimates are desirable but extant primate reference samples with known body masses are rare. We estimated body mass in a sample of extinct platyrrhines and Fayum anthropoids based on four measurements of the articular surfaces of the humerus and femur. Estimates were based on a large extant reference sample of wild-collected individuals with associated body masses, including previously published and new data from extant platyrrhines, cercopithecoids, and hominoids. In general, scaling of joint dimensions is positively allometric relative to expectations of geometric isometry, but negatively allometric relative to expectations of maintaining equivalent joint surface areas. Body mass prediction equations based on articular breadths are reasonably precise, with %SEEs of 17-25%. The breadth of the distal femoral articulation yields the most reliable estimates of body mass because it scales similarly in all major anthropoid taxa. Other joints scale differently in different taxa; therefore, locomotor style and phylogenetic affinity must be considered when calculating body mass estimates from the proximal femur, proximal humerus, and distal humerus. The body mass prediction equations were applied to 36 Old World and New World fossil anthropoid specimens representing 11 taxa, plus two Haitian specimens of uncertain taxonomic affinity. Among the extinct platyrrhines studied, only Cebupithecia is similar to large, extant platyrrhines in having large humeral (especially distal) joints. Our body mass estimates differ from each other and from published estimates based on teeth in ways that reflect known differences in relative sizes of the joints and teeth. We prefer body mass estimators that are biomechanically linked to weight-bearing, and especially those that are relatively insensitive to differences in locomotor style and phylogenetic history. Whenever possible, extant reference samples should be chosen to match target fossils in joint proportionality.

New fossils from Tadkeshwar Mine (Gujarat, India) increase primate diversity from the early Eocene Cambay Shale.

Several new fossil specimens from the Cambay Shale Formation at Tadkeshwar Lignite Mine in Gujarat document the presence of two previously unknown early Eocene primate species from India. A new species of Asiadapis is named based on a jaw fragment preserving premolars similar in morphology to those of A. cambayensis but substantially larger. Also described is an exceptionally preserved edentulous dentary (designated cf. Asiadapis, unnamed sp. nov.) that is slightly larger and much more robust than previously known Cambay Shale primates. Its anatomy most closely resembles that of Eocene adapoids, and the dental formula is the same as in A. cambayensis. A femur and calcaneus are tentatively allocated to the same taxon. Although the dentition is unknown, exquisite preservation of the dentary of cf. Asiadapis sp. nov. enables an assessment of masticatory musculature, function, and gape adaptations, as well as comparison with an equally well-preserved dentary of the asiadapid Marcgodinotius indicus, also from Tadkeshwar. The new M. indicus specimen shows significant gape adaptations but was probably capable of only weak bite force, whereas cf. Asiadapis sp. nov. probably used relatively smaller gapes but could generate relatively greater bite forces.

Convergent evolution of anthropoid-like adaptations in Eocene adapiform primates.

Adapiform or 'adapoid' primates first appear in the fossil record in the earliest Eocene epoch ( approximately 55 million years (Myr) ago), and were common components of Palaeogene primate communities in Europe, Asia and North America. Adapiforms are commonly referred to as the 'lemur-like' primates of the Eocene epoch, and recent phylogenetic analyses have placed adapiforms as stem members of Strepsirrhini, a primate suborder whose crown clade includes lemurs, lorises and galagos. An alternative view is that adapiforms are stem anthropoids. This debate has recently been rekindled by the description of a largely complete skeleton of the adapiform Darwinius, from the middle Eocene of Europe, which has been widely publicised as an important 'link' in the early evolution of Anthropoidea. Here we describe the complete dentition and jaw of a large-bodied adapiform (Afradapis gen. nov.) from the earliest late Eocene of Egypt ( approximately 37 Myr ago) that exhibits a striking series of derived dental and gnathic features that also occur in younger anthropoid primates-notably the earliest catarrhine ancestors of Old World monkeys and apes. Phylogenetic analysis of 360 morphological features scored across 117 living and extinct primates (including all candidate stem anthropoids) does not place adapiforms as haplorhines (that is, members of a Tarsius-Anthropoidea clade) or as stem anthropoids, but rather as sister taxa of crown Strepsirrhini; Afradapis and Darwinius are placed in a geographically widespread clade of caenopithecine adapiforms that left no known descendants. The specialized morphological features that these adapiforms share with anthropoids are therefore most parsimoniously interpreted as evolutionary convergences. As the largest non-anthropoid primate ever documented in Afro-Arabia, Afradapis nevertheless provides surprising new evidence for prosimian diversity in the Eocene of Africa, and raises the possibility that ecological competition between adapiforms and higher primates might have played an important role during the early evolution of stem and crown Anthropoidea in Afro-Arabia.

Craniomandibular signals of diet in adapids.

OBJECTIVES: The craniomandibular morphology of the adapid primates of Europe, especially Adapis and Leptadapis (sensu lato), suggests that they possessed enormous jaw adductor muscles. The goal of this study is to estimate jaw adductor muscle mass, physiological cross-sectional area (PCSA), and fiber length in adapid primates from the Eocene of Europe. We also estimated muscle leverage, bite force, and gape parameters. MATERIALS AND METHODS: We use bony morphology and osteological correlates of soft tissues in a sample of extant strepsirrhines to estimate these soft-tissue and performance variables in Adapis and Leptadapis. RESULTS: Our results suggest that, compared to a broad sample of extant strepsirrhines, Adapis and Leptadapis had relatively great jaw adductor muscle mass, PCSA, and bite force. They had moderately great jaw adductor leverage but no sign of adaptation for wide gapes. There is no support for the hypothesis that either adapid was a gouger. DISCUSSION: Our estimates support the inference that Adapis and Leptadapis were primarily folivorous, perhaps also consuming small to medium-sized tough fruits, nuts, and seeds. Explanations for the likely extreme development of the jaw adductor muscles in adapids remain speculative. These include (1) foods that were generally tougher and/or of higher yield strength than those eaten by strepsirrhines today, (2) using the muscles "in shifts" to avoid muscle fatigue in the context of an obdurate diet, and (3) potential constraints on reshaping of the skull for more efficient food processing.

Maximum ingested food size in captive anthropoids.

OBJECTIVES: Maximum ingested food size (Vb ) is an empirically tested performance variable that can shed light on feeding energetics and adaptation in the masticatory system. Until now, this variable had been tested in strepsirrhines alone among primates. Here, we present the first data on Vb in a broad sample of anthropoid primates and describe scaling patterns. MATERIALS AND METHODS: Vb data on anthropoids were collected under captive conditions at the Philadelphia Zoo and compared with published data on strepsirrhines. Data on Vb were scaled against individual body mass and were compared with experimentally determined toughness and stiffness values for the test foods. RESULTS: Unlike in strepsirrhines, where essentially Vb scales isometrically with body mass, Vb in anthropoids scales with negative allometry. There is a significant effect of food material properties on Vb , although bite size in anthropoids varies less based on food properties than in strepsirrhines. Large folivorous strepsirrhines follow the anthropoid trend in bite size scaling, but large frugivorous ones take especially large bites. DISCUSSION: Negative scaling of bite size in the anthropoids sampled could be due to reduced adaptation for gape. Some early anthropoids likely evolved adaptations for maximizing mechanical advantage and fatigue resistance in the chewing muscles, resulting in reduced gape. This might have channeled them toward smaller bites of more-resistant foods and away from taking large bites. This might also be the case for some folivorous strepsirrhines.

Comparing apples and oranges-the influence of food mechanical properties on ingestive bite sizes in lemurs.

Previously we found that Maximum Ingested Bite Size (Vb )-the largest piece of food that an animal will ingest whole without biting first-scales isometrically with body size in 17 species of strepsirrhines at the Duke Lemur Center (DLC). However, because this earlier study focused on only three food types (two with similar mechanical properties), it did not yield results that were easily applied to describing the broad diets of these taxa. Expressing Vb in terms of food mechanical properties allows us to compare data across food types, including foods of wild lemurs, to better understand dietary adaptations in lemurs. To this end, we quantified Vb in five species of lemurs at the DLC representing large and small frugivores and folivores using ten types of food that vary widely in stiffness and toughness to determine how these properties relate to bite sizes. We found that although most species take smaller bites of stiffer foods, this negative relationship was not statistically significant across the whole sample. However, there is a significant relationship between bite size and toughness. All three of the more frugivorous taxa in our sample take significantly smaller bites of tougher foods. However, the two more folivorous lemurs do not. They take small bites for all foods. This suggests that the species most adapted to the consumption of tough foods do not modulate their ingestive sizes to accommodate larger pieces of weak foods.

Oldest known cranium of a juvenile New World monkey (Early Miocene, Patagonia, Argentina): implications for the taxonomy and the molar eruption pattern of early platyrrhines.

A juvenile cranium of Homunculus patagonicus Ameghino, 1891a from the late Early Miocene of Santa Cruz Province (Argentina) provides the first evidence of developing cranial anatomy for any fossil platyrrhine. The specimen preserves the rostral part of the cranium with deciduous and permanent alveoli and teeth. The dental eruption sequence in the new specimen and a reassessment of eruption patterns in living and fossil platyrrhines suggest that the ancestral platyrrhine pattern of tooth replacement was for the permanent incisors to erupt before M(1), not an accelerated molar eruption (before the incisors) as recently proposed. Two genera and species of Santacrucian monkeys are now generally recognized: H. patagonicus Ameghino, 1891a and Killikaike blakei Tejedor et al., 2006. Taxonomic allocation of Santacrucian monkeys to these species encounters two obstacles: 1) the (now lost) holotype and a recently proposed neotype of H. patagonicus are mandibles from different localities and different geologic members of the Santa Cruz Formation, separated by approximately 0.7 million years, whereas the holotype of K. blakei is a rostral part of a cranium without a mandible; 2) no Santacrucian monkey with associated cranium and mandible has ever been found. Bearing in mind these uncertainties, our examination of the new specimen as well as other cranial specimens of Santacrucian monkeys establishes the overall dental and cranial similarity between the holotype of Killikaike blakei, adult cranial material previously referred to H. patagonicus, and the new juvenile specimen. This leads us to conclude that Killikaike blakei is a junior subjective synonym of H. patagonicus.

Body size and premolar evolution in the early-middle eocene euprimates of Wyoming.

The earliest euprimates to arrive in North America were larger-bodied notharctids and smaller-bodied omomyids. Through the Eocene, notharctids generally continued to increase in body size, whereas omomyids generally radiated within small- and increasingly mid-sized niches in the middle Eocene. This study examines the influence of changing body size and diet on the evolution of the lower fourth premolar in Eocene euprimates. The P4 displays considerable morphological variability in these taxa. Despite the fact that most studies of primate dental morphology have focused on the molars, P4 can also provide important paleoecological insights. We analyzed the P4 from 177 euprimate specimens, representing 35 species (11 notharctids and 24 omomyids), in three time bins of approximately equal duration: early Wasatchian, late Wasatchian, and Bridgerian. Two-dimensional surface landmarks were collected from lingual photographs, capturing important variation in cusp position and tooth shape. Disparity metrics were calculated and compared for the three time bins. In the early Eocene, notharctids have a more molarized P4 than omomyids. During the Bridgerian, expanding body size range of omomyids was accompanied by a significant increase in P4 disparity and convergent evolution of the semimolariform condition in the largest omomyines. P4 morphology relates to diet in early euprimates, although patterns vary between families.

A fossil primate of uncertain affinities from the earliest late Eocene of Egypt.

Paleontological work carried out over the last 3 decades has established that three major primate groups were present in the Eocene of Africa-anthropoids, adapiforms, and advanced strepsirrhines. Here we describe isolated teeth of a previously undocumented primate from the earliest late Eocene ( approximately 37 Ma) of northern Egypt, Nosmips aenigmaticus, whose phylogenetic placement within Primates is unclear. Nosmips is smaller than the sympatric adapiform Afradapis but is considerably larger than other primate taxa known from the same paleocommunity. The species bears an odd mosaic of dental features, combining enlarged, elongate, and molariform premolars with simple upper molars that lack hypocones. Phylogenetic analysis across a series of different assumption sets variously places Nosmips as a stem anthropoid, a nonadapiform stem strepsirrhine, or even among adapiforms. This phylogenetic instability suggests to us that Nosmips likely represents a highly specialized member of a previously undocumented, and presumably quite ancient, endemic African primate lineage, the subordinal affinities of which have been obscured by its striking dental autapomorphies. Discriminant functions based on measurements of lower molar size and topography reliably classify extant prosimian primates into their correct dietary groups and identify Nosmips and Afradapis as omnivores and folivores, respectively. Although Nosmips currently defies classification, this strange and unexpected fossil primate nevertheless provides additional evidence for high primate diversity in northern Africa approximately 37 million years ago and further underscores the fact that our understanding of early primate evolution on that continent remains highly incomplete.

Primate body mass and dietary correlates of tooth root surface area.

OBJECTIVES: This study aims to examine primate postcanine tooth root surface area (TRSA) in the context of two ecological variables (diet and bite force). We also assess scaling relationships within distinct taxonomic groups and across the order as a whole. MATERIALS AND METHODS: Mandibular postcanine TRSA was measured using a three-dimensional computed tomography (CT) method for catarrhine (N = 27), platyrrhine (N = 21), and strepsirrhine (N = 24) taxa; this represents the first sample of strepsirrhines. Two different body size proxies were used: cranial geometric mean (GM) using nine linear measurements, and literature-derived body mass (BM). RESULTS: TRSA correlated strongly with body size, scaling with positive allometry or isometry across the order as a whole; however, scaling differed significantly between taxa for some teeth. Among Strepsirrhini, molar TRSA relative to GM differed significantly between folivores and pliant-object feeders. Additionally, P4 TRSA relative to BM differentiated folivores from both hard- and pliant-object feeders. Among Cercopithecoidea, P4 TRSA adjusted by GM differed between hard- and pliant-object feeders. DISCUSSION: Dietary signals in TRSA appear primarily driven by high frequency loading experienced by folivores. Stronger and more frequent dietary signals were observed within Strepsirrhini relative to Haplorhini. This may reflect the constraints of orthognathism within the latter, constraining the adaptability of their postcanine teeth. Finally, because of the strong correlation between TRSA and BM for each tooth locus (mean r2  = 0.82), TRSA can be used to predict BM in fossil primates using provided equations.

Maximum ingested food size in captive strepsirrhine primates: scaling and the effects of diet.

Little is known about ingested food size (V(b)) in primates, even though this variable has potentially important effects on food intake and processing. This study provides the first data on V(b) in strepsirrhine primates using a captive sample of 17 species. These data can be used for generating and testing models of feeding energetics. Strepsirrhines are of interest because they are hypometabolic and chewing rate and daily feeding time do not show a significant scaling relationship with body size. Using melon, carrot, and sweet potato we found that maximum V(b) scales isometrically with body mass and mandible length. Low dietary quality in larger strepsirrhines might explain why V(b) increases with body size at a greater rate than does resting metabolic rate. Relative to body size, V(b) is large in frugivores but small in folivores; furthermore scaling slopes are higher in frugivores than in folivores. A gross estimate of dietary quality explains much of the variation in V(b) that is not explained by body size. Gape adaptations might favor habitually large bites for frugivores and small ones for folivores. More data are required for several feeding variables and for wild populations.

Maximum Bony Gape in Primates.

Maximum jaw gape has important functional implications for behavior and feeding habits in primates. It has been suggested that gape is correlated to canine height and ingested food size. Extending these correlations to the fossil record would provide insights about the diets and/or social behavior of extinct primates. However, this can be problematic due to uncertainty about size and location of musculature, and it depends on reliability and repeatability of maximum gape estimation using only skeletal elements. In this study, maximum bony gape (MBG) was estimated using reliable landmarks and repeatable methods. The cranium was fixed in position and then the mandible was rotated and translated to the point immediately prior to loss of condyle-glenoid contact. Then it was photographed in a steady position using an adjustable wooden frame. This protocol allowed for photographs and linear measurements to be obtained for many museum specimens in a short time. The sample included 203 individuals, representing 42 species of primates. When scaled for body size, linear MBG correlates with maximum anesthetized gape (Hylander: Am J Phys Anthropol 150 (2013) 247-259), ingested food size (Perry and Hartstone-Rose: Am J Phys Anthropol 142 (2010) 625-635), and canine length but not condylar height. Anat Rec, 302:215-225, 2019. © 2018 Wiley Periodicals, Inc.

Inferring the Diets of Extinct Giant Lemurs from Osteological Correlates of Muscle Dimensions.

The jaw adductor muscles of extinct mammals are often reconstructed to elucidate paleoecological relationships and to make broad comparisons among taxa. Muscle lever arms, bite load arms, muscle dimensions, and gape are often also reconstructed to better understand feeding. Several different approaches to these and related goals are discussed here. A protocol for reconstructing muscle dimensions and bite force using biomechanically informative skull measurements and osteological proxies of muscle dimensions is described and applied to a case study of subfossil Malagasy lemurs. The results of this case study show that most subfossil lemurs emphasized the masseter and medial pterygoid muscles over the temporalis. This supports the inference that these extinct lemurs depended heavily on tough food like leaves. Exceptions include signals of hard-object feeding in Archaeolemur that vary between A. majori and A. edwardsi. Reconstructions of soft-tissue and function are important for understanding past ecological relationships. Even those based on well-supported osteological proxies from extant analogues have limitations for making precise inferences. Anat Rec, 301:343-362, 2018. © 2018 Wiley Periodicals, Inc.

Muscle Functional Morphology in Paleobiology: The Past, Present, and Future of "Paleomyology".

Our knowledge of muscle anatomy and physiology in vertebrates has increased dramatically over the last two-hundred years. Today, much is understood about how muscles contract and about the functional meaning of muscular variation at multiple scales. Progress in muscle anatomy has profited from the availability of broad comparative samples, advances in microscopy have permitted comparisons at increasingly finer scales, and progress in muscle physiology has profited from many carefully designed and executed experiments. Several avenues of future work are promising. In particular, muscle ontogeny (growth and development) is poorly understood for many vertebrate groups. We consider which types of advances in muscle functional morphology are of use to paleobiologists. These are only a modest subset for muscle anatomy and a very small subset for muscle physiology. The relationship between muscle and bone - spatially and mechanically-is critical to any future advances in "paleomyology". Anat Rec, 301:538-555, 2018. © 2018 Wiley Periodicals, Inc.

Craniomandibular Variation in Phalangeriform Marsupials: Functional Comparisons with Primates.

Phalangeriform marsupials have often been compared with primates because of similarity in the range of external morphology, ecological niches, and body size between the two radiations. We explore morphological convergence in the masticatory anatomy of strepsirrhine primates and phalangeriforms, through osteological measurements of the mandible and facial skeleton, and through dissection of the masticatory musculature, presenting new data on the arrangement and proportions of jaw adductors in phalangeriforms. Phalangeriforms and primates have a large number of shape differences in mandibular morphology. Despite these differences in shape on phylogenetic lines, dietary groups used to pool species of phalangeriforms and strepsirrhines also differed from each other in a range of shape variables. Notably, the striped possum (Dactylopsila), previously described as convergent with the aye-aye (Daubentonia), shares a number of features of mandibular shape with Daubentonia, and the exudate-feeding sugar-glider, Petaurus, shares shape features with gummivorous strepsirrhines. Petaurus also has long-fibered jaw adductors for its body mass, as would be expected for a species with a requirement for large gape. Phalangeriform species on the frugivore-folivore continuum were less clearly comparable to strepsirrhine species with similar diets. There are a number of significant dietary contrasts in osteological measurements, but in the masticatory muscles phalangeriforms did not meet all expectations based on available dietary data, highlighting the possible complexity of dietary adaptation in phalangeriform folivores. Anat Rec, 301:227-255, 2018. © 2018 Wiley Periodicals, Inc.

Do Muscles Constrain Skull Shape Evolution in Strepsirrhines?

Despite great interest and decades of research, the musculoskeletal relationships of the masticatory system in primates are still not fully understood. However, without a clear understanding of the interplay between muscles and bones it remains difficult to understand the functional significance of morphological traits of the skeleton. Here, we aim to study the impacts of the masticatory muscles on the shape of the cranium and the mandible as well as their co-variation in strepsirrhine primates. To do so, we use 3D geometric morphometric approaches to assess the shape of each bone of the skull of 20 species for which muscle data are available in the literature. Impacts of the masticatory muscles on the skull shape were assessed using non-phylogenetic regressions and phylogenetic regressions whereas co-variations were assessed using two-blocks partial least square (2B-PLS) and phylogenetic 2B-PLS. Our results show that there is a phylogenetic signal for skull shape and masticatory muscles. They also show that there is a significant impact of the masticatory muscles on cranial shape but not as much as on the mandible. The co-variations are also stronger between the masticatory muscles and cranial shape even when taking into account phylogeny. Interestingly, the results of co-variation between the masticatory muscles and mandibular shape show a more complex pattern in two different directions to get strong muscles associated with mandibular shape: a folivore way (with the bamboo lemurs and sifakas) and a hard-object eater one (with the aye-aye). Anat Rec, 301:291-310, 2018. © 2018 Wiley Periodicals, Inc.

Biting mechanics and niche separation in a specialized clade of primate seed predators.

We analyzed feeding biomechanics in pitheciine monkeys (Pithecia, Chiropotes, Cacajao), a clade that specializes on hard-husked unripe fruit (sclerocarpy) and resistant seeds (seed predation). We tested the hypothesis that pitheciine crania are well-suited to generate and withstand forceful canine and molar biting, with the prediction that they generate bite forces more efficiently and better resist masticatory strains than the closely-related Callicebus, which does not specialize on unripe fruits and/or seeds. We also tested the hypothesis that Callicebus-Pithecia-Chiropotes-Cacajao represent a morphocline of increasing sclerocarpic specialization with respect to biting leverage and craniofacial strength, consistent with anterior dental morphology. We found that pitheciines have higher biting leverage than Callicebus and are generally more resistant to masticatory strain. However, Cacajao was found to experience high strain magnitudes in some facial regions. We therefore found limited support for the morphocline hypothesis, at least with respect to the mechanical performance metrics examined here. Biting leverage in Cacajao was nearly identical (or slightly less than) in Chiropotes and strain magnitudes during canine biting were more likely to follow a Cacajao-Chiropotes-Pithecia trend of increasing strength, in contrast to the proposed morphocline. These results could indicate that bite force efficiency and derived anterior teeth were selected for in pitheciines at the expense of increased strain magnitudes. However, our results for Cacajao potentially reflect reduced feeding competition offered by allopatry with other pitheciines, which allows Cacajao species to choose from a wider variety of fruits at various stages of ripeness, leading to reduction in the selection for robust facial features. We also found that feeding biomechanics in sympatric Pithecia and Chiropotes are consistent with data on food structural properties and observations of dietary niche separation, with the former being well-suited for the regular molar crushing of hard seeds and the latter better adapted for breaching hard fruits.

Anatomy and adaptations of the chewing muscles in Daubentonia (Lemuriformes).

The extractive foraging behavior in aye-ayes (Daubentonia madagascariensis) is unique among primates and likely has led to selection for a specialized jaw adductor musculature. Although this musculature has previously been examined in a subadult, until now, no one has reported the fascicle length, weight, and physiological cross-sectional area (PCSA) for these muscles in an adult aye-aye specimen. For the present study, we dissected an adult wild-born aye-aye from the Tsimbazaza Botanical and Zoological Park, Antananarivo, Madagascar. The aye-aye follows the general strepsirrhine pattern in its overall jaw adductor muscle anatomy, but has very large muscles and PCSA relative to body size. Fascicle length is also relatively great, but not nearly as much as in the juvenile aye-aye previously dissected. Perhaps chewing muscle fascicles begin relatively long, but shorten through use and growth as connective tissue sheets expand and allow for pinnation and increased PCSA. Alternately, it may be that aye-ayes develop fascicular adaptation to wide gapes early in ontogeny, only to increase PCSA through later development into adulthood. The functional demands related to their distinctive manner of extractive foraging are likely responsible for the great PCSA in the jaw adductor muscles of the adult aye-aye. It may be that great jaw adductor PCSA in the adult, as compared to the juvenile, is a means of increasing foraging efficiency in the absence of parental assistance. Anat Rec, 297:308-316, 2014. © 2013 Wiley Periodicals, Inc.

Phylogenetic, ecological, and allometric correlates of cranial shape in Malagasy lemuriforms.

Adaptive radiations provide important insights into many aspects of evolution, including the relationship between ecology and morphological diversification as well as between ecology and speciation. Many such radiations include divergence along a dietary axis, although other ecological variables may also drive diversification, including differences in diel activity patterns. This study examines the role of two key ecological variables, diet and activity patterns, in shaping the radiation of a diverse clade of primates, the Malagasy lemurs. When phylogeny was ignored, activity pattern and several dietary variables predicted a significant proportion of cranial shape variation. However, when phylogeny was taken into account, only typical diet accounted for a significant proportion of shape variation. One possible explanation for this discrepancy is that this radiation was characterized by a relatively small number of dietary shifts (and possibly changes in body size) that occurred in conjunction with the divergence of major clades. This pattern may be difficult to detect with the phylogenetic comparative methods used here, but may characterize not just lemurs but other mammals.

Bite force estimation and the fiber architecture of felid masticatory muscles.

Increasingly, analyses of craniodental dietary adaptations take into account mechanical properties of foods. However, masticatory muscle fiber architecture has been described for relatively few lineages, even though an understanding of the scaling of this anatomy can yield important information about adaptations for stretch and strength in the masticatory system. Data on the mandibular adductors of 28 specimens from nine species of felids representing nearly the entire body size range of the family allow us to evaluate the influence of body size and diet on the masticatory apparatus within this lineage. Masticatory muscle masses scale isometrically, tending toward positive allometry, with body mass and jaw length. This allometry becomes significant when the independent variable is a geometric mean of cranial variables. For all three body size proxies, the physiological cross-sectional area and predicted bite forces scale with significant positive allometry. Average fiber lengths (FL) tend toward negative allometry though with wide confidence intervals resulting from substantial scatter. We believe that these FL residuals are affected by dietary signals within the sample; though the mechanical properties of felid diets are relatively similar across species, the most durophagous species in our sample (the jaguar) appears to have relatively higher force production capabilities. The more notable dietary trend in our sample is the relationship between FL and relative prey size: felid species that predominantly consume relatively small prey have short masticatory muscle fibers, and species that regularly consume relatively large prey have relatively long fibers. This suggests an adaptive signal related to gape.