Changing perspectives on early hominin diets.

It is axiomatic that knowledge of the diets of extinct hominin species is central to any understanding of their ecology and our evolution. The importance of diet in the paleontological realm has led to the employment of multiple approaches in its elucidation. Some of these have deep historical roots, while others are dependent upon more recent technical and methodological advances. Historically, studies of tooth size, shape, and structure have been the gold standard for reconstructing diet. They focus on species-level adaptations, and as such, they can set theoretical brackets for dietary capabilities within the context of specific evolutionary moments. Other methods (e.g., analyses of dental calculus, biogeochemistry, and dental microwear) have only been developed within the past few decades, but are now beginning to yield evidence of the actual foods consumed by individuals represented by fossil remains. Here we begin by looking at these more "direct" forms of evidence of diet before showing that, when used in conjunction with other techniques, these "multi-proxy" approaches can raise questions about traditional interpretations of early hominin diets and change the nature of paleobiological interpretations.

Stubby versus stabby: A preliminary analysis of canine microwear in primates: Implication for inferring ingestive behaviors.

Molar and incisor microwear reflect aspects of food choice and ingestive behaviors in living primates and have both been used to infer the same for fossil samples. Canine microwear, however, has received less attention, perhaps because of the prominent role canines play in social display and because they are used as weapons-while outside of a few specialized cases, their involvement in diet related behaviors has not been obvious. Here, we posit that microwear can also provide glimpses into canine tooth use in ingestion. Canines of Sumatran long-tailed macaques (Macaca fascicularis), agile gibbons (Hylobates agilis), lar gibbons (Hylobates lar), Thomas' leaf monkeys (Presbytis thomasi), and orangutans (Pongo abelii), and two African great apes, bonobos (Pan paniscus) and common chimpanzees (Pan troglodytes schweinfurthii), were considered. The labial tips of maxillary canine replicas were scanned using a white-light confocal profiler, and both feature and texture analyses were used to characterize microwear surface patterning. The taxa exhibited significant differences in canine microwear. In some cases, these were consistent with variation in reported anterior tooth use such that, for example, the orangutans, known to use their front teeth extensively in ingestion, had the highest median number of microwear features on their canines, whereas the gibbons, reported to use their front teeth infrequently in food acquisition, had the lowest.

Grit your teeth and chew your food: Implications of food material properties and abrasives for rates of dental microwear formation in laboratory Sapajus apella (Primates).

Dental microwear analysis has been employed in studies of a wide range of modern and fossil animals, yielding insights into the biology/ecology of those taxa. Some researchers have suggested that dental microwear patterns ultimately relate back to the material properties of the foods being consumed, whereas others have suggested that, because exogenous grit is harder than organic materials in food, grit should have an overwhelming impact on dental microwear patterns. To shed light on this issue, laboratory-based feeding experiments were conducted on tufted capuchin monkeys [Sapajus apella] with dental impressions taken before and after consumption of different artificial foods. The foods were (1) brittle custom-made biscuits laced with either of two differently-sized aluminum silicate abrasives, and (2) ductile custom-made "gummies" laced with either of the two same abrasives. In both cases, animals were allowed to feed on the foods for 36 hours before follow-up dental impressions were taken. Resultant casts were analyzed using a scanning electron microscope. We asked five questions: (1) would the animals consume different amounts of each food item, (2) what types of dental microwear would be formed, (3) would rates of dental microwear differ between the consumption of biscuits (i.e., brittle) versus gummies (i.e., ductile), (4) would rates of dental microwear differ between foods including larger- versus smaller-grained abrasives, and (5) would rates of dental microwear differ between molar shearing and crushing facets in the animals in these experiments? Results indicated that (1) fewer biscuits were consumed when laced with larger-grained abrasives (as opposed to smaller-grained abrasives), but no such difference was observed in the consumption of gummies, (2) in all cases, a variety of dental microwear features was formed, (3) rates of dental microwear were higher when biscuits versus gummies were consumed, (4) biscuits laced with larger-grained abrasives caused a higher percentage of new features per item consumed, and (5) the only difference between facets occurred with the processing of biscuits, where crushing facets showed a faster rate of wear than shearing facets. These findings suggest that the impact of exogenous grit on dental microwear is the result of a dynamic, complex interaction between (at the very least) grit size, food material properties, and time spent feeding - which is further evidence of the multifactorial nature of dental microwear formation.

The dental microwear of hard-object feeding in laboratory Sapajus apella and its implications for dental microwear formation.

OBJECTIVES: This study seeks to determine if (a) consumption of hard food items or a mixture of food items leads to the formation of premolar or molar microwear in laboratory capuchin monkeys (Sapajus apella) in one feeding session and (b) rates of microwear formation are associated with the number of food items consumed. MATERIALS AND METHODS: Five adult male capuchins were used in two experiments, one where they were fed unshelled Brazil nuts, and the other where they were fed a mixture of food items. Dental impressions were taken before and after each feeding session. Epoxy casts made from those impressions then were used in SEM analyses of rates of microwear formation. Upper and lower premolars and molars were analyzed. Qualitative comparisons were made and Spearman's rank-order correlations used to examine the relationship between rates of microwear formation and number of Brazil nuts consumed. RESULTS: Premolars and molars generally showed new microwear in the form of pits and scratches. However, the incidence of those features was low (0-6%). Rates of microwear formation were highest during the consumption of Brazil nuts. DISCUSSION: Variations in the rate of microwear formation on the premolars likely reflected patterns of ingestion whereas consistency in the rate of microwear on the molars likely reflected patterns of chewing. While dental microwear formation seemed to be correlated with the number of hard objects consumed, rates did differ between individuals. Differences in results between the two experiments demonstrate some of the limitations in our knowledge of dental microwear formation.

Microwear textures of Australopithecus africanus and Paranthropus robustus molars in relation to paleoenvironment and diet.

The importance of diet in primate ecology has motivated the use of a variety of methods to reconstruct dietary habits of extinct hominin taxa. Dental microwear is one such approach that preserves evidence from consumed food items. This study is based on 44 specimens of Australopithecus africanus from Makapansgat and Sterkfontein, and 66 specimens of Paranthropus robustus from Swartkrans, Kromdraai and Drimolen. These samples enable examination of potential differences between the two assemblages of A. africanus, and among the various assemblages of P. robustus in relation to the paleoenvironmental reconstructions that have been proffered for each fossil site. Sixteen microwear texture variables were recorded for each specimen from digital elevation models generated using a white-light confocal profiler. Only two of these differ significantly between the Makapansgat and Sterkfontein samples of A. africanus. None of the microwear texture variables differs significantly among the samples of P. robustus. On the other hand, P. robustus has significantly higher values than A. africanus for 11 variables related to feature complexity, size, and depth; P. robustus exhibits rougher surfaces that comprise larger, deeper features. In contrast, A. africanus has smoother, simpler wear surfaces with smaller, shallower and more anisotropic features. As for possible habitat differences among the various sites, only a relatively small number of subtle differences are evident between the specimens of A. africanus from Makapansgat and Sterkfontein, and there are none among the specimens of P. robustus from various deposits. As such, it is reasonable to conclude that, while subtle differences in microwear textures may reflect differences in background habitats, the wear fabric differences between P. robustus and A. africanus are most reasonably interpreted as having been driven by dietary differences.

A molar microwear texture analysis of pitheciid primates.

Dental microwear textures have been examined for a broad range of extant primates to assess their efficacy for reconstructing diets of fossil species. To date though, no dental microwear texture data have been published for pitheciid molars, despite reported variation in degree of sclerocarpy and, by extension, the fracture properties of foods these platyrrhines eat. While all pitheciids eat hard or tough seeds, Chiropotes and Pithecia have been documented to consume more than Callicebus. In this study, we explored whether measures of molar microwear texture complexity discriminate taxa following variation in reliance upon seeds, and whether dispersion among variables is greatest in Callicebus, which has the most variable diet. Here we report results for a study of microwear textures on M2 "Phase II" facets of Ch. satanas (N = 14), P. irrorata (N = 8), and Ca. moloch (N = 24) from the Brazilian Amazon (Oriximina, UHE Samuel, and Taperinha, respectively). Textures examined using a scanning confocal profiler showed significant differences in central tendencies for three measures: mean dale area (Sda), anisotropy (Str), and heterogeneity (HAsfc9 ). Ten measures showed significant differences in dispersion, with Callicebus being significantly more variable in eight of those ten. These results demonstrate that the pitheciids with different morphological adaptations and dietary reliance on seeds differ in their dental microwear textures, though less than initially hypothesized. Measures of dispersion, especially, show potential for identifying dietary variability.

Canine and incisor microwear in pitheciids and Ateles reflects documented patterns of tooth use.

OBJECTIVES: Platyrrhine species differ in the extent to and the manner in which they use their incisors and canines during food ingestion. For example, Ateles uses its anterior teeth to process mechanically nondemanding soft fruits, while the sclerocarp-harvesting pitheciids rely extensively on these teeth to acquire and process more demanding foods. Pitheciids themselves vary in anterior tooth use, with the pitheciines (Cacajao, Chiropotes, and Pithecia) noted to use their robust canines in a variety of ways to predate seeds, while Callicebus, which rarely predates seeds, uses its incisors and exceptionally short canines to scrape tough mesocarp from fruits. To investigate the relationship between tooth use and dental wear, microwear textures were investigated for the anterior teeth of these five genera of platyrrhine primates. METHODS: Using a white light confocal microscope, 12 microwear texture attributes that reflect feature size, anisotropy, density, and complexity were recorded from high-resolution epoxy casts of the incisors and canines of adult wild-collected Brazilian specimens of Ateles, Callicebus, Cacajao, Chiropotes, and Pithecia. RESULTS: Pitheciine canines tend to have deep microwear features and complex, anisotropic microwear textures, while Ateles anterior teeth tend to have very small features, low feature density, and less complex and anisotropic surfaces. Callicebus incisor and canine microwear is generally intermediate in size and complexity between those extremes. CONCLUSIONS: These findings align with expectations from reported field observations of tooth use and illustrate the potential for using microwear texture analysis to infer patterns of anterior tooth use in extinct primates. Am J Phys Anthropol 161:6-25, 2016. © 2016 Wiley Periodicals, Inc.

Dental microwear profilometry of African non-cercopithecoid catarrhines of the Early Miocene.

The Early Miocene of Kenya has yielded the remains of many important stem catarrhine species that provide a glimpse of the East African primate radiation at a time of major faunal turnover. These taxa have been subject to innumerable studies, yet there is still no consensus on their dietary niches. Here we report results of an analysis of dental microwear textures of non-cercopithecoid catarrhines from the Early Miocene of Kenya. Scanning confocal profilometry of all available molar specimens with undamaged occlusal surfaces revealed 82 individuals with unobscured antemortem microwear, representing Dendropithecus, Micropithecus, Limnopithecus, Proconsul, and Rangwapithecus. Scale-sensitive fractal analysis was used to generate microwear texture attributes for each individual, and the fossil taxa were compared with each other using conservative non-parametric statistical tests. This study revealed no discernible variation in microwear texture among the fossil taxa, which is consistent with results from a previous feature-based microwear study using smaller samples. Our results suggest that, despite their morphological differences, these taxa likely often consumed foods with similar abrasive and fracture properties. However, statistical analyses of microwear texture data indicate differences between the Miocene fossil sample and several extant anthropoid primate genera. This suggests that the African non-cercopithecoid catarrhines included in our study, despite variations in tooth form, had generalist diets that were not yet specialized to the degree of many modern taxa.

Environmental perturbations can be detected through microwear texture analysis in two platyrrhine species from Brazilian Amazonia.

Recent dental microwear studies have shown that fossil species differ from one another in texture attributes-both in terms of central tendency and dispersion. Most comparative studies used to interpret these results have relied on poorly provenienced museum samples that are not well-suited to consideration of within species variation in diet. Here we present a study of two species of platyrrhine monkeys, Alouatta belzebul (n = 60) and Sapajus apella (n = 28) from Pará State in the Brazilian Amazon in order to assess effects of habitat variation on microwear (each species was sampled from forests that differ in the degree of disturbance from highly disturbed to minimally disturbed). Results indicate that microwear texture values vary between habitats-more for the capuchins than the howler monkeys. This is consistent with the notion that diets of the more folivorous A. belzebul are less affected by habitat disturbance than those of the more frugivorous S. apella. It also suggests that microwear holds the potential to reflect comparatively subtle differences in within-species variation in fossil taxa if sample size and control over paleohabitat allow.

Premolar microwear and tooth use in Australopithecus afarensis.

The mandibular third premolar (P3) of Australopithecus afarensis is notable for extensive morphological variability (e.g., metaconid presence/absence, closure of the anterior fovea, root number) and temporal trends in crown length and shape change over its 700 Ka time range. Hominins preceding A. afarensis have unicuspid, mesiodistally elongated P3s with smaller talonids, and subsequent australopiths have bicuspid, more symmetrically-shaped P3 crowns with expanded talonids. For these features, A. afarensis is intermediate and, thus, evinces the incipient stages of P3 molarization. Here, we examine A. afarensis P3 Phase II microwear and compare it with that of Australopithecus africanus and Cercocebus atys, an extant hard-object specialist, to assess whether the role of the P3 in food processing changed over time in A. afarensis. Premolar Phase II microwear textures are also compared with those of the molars to look for evidence of functional differentiation along the tooth row (i.e., that foods with different mechanical properties were processed by separate regions of the postcanine battery). Microwear textures were also examined along the mesial protoconid crest, the site of occlusion with the maxillary canine, of the A. afarensis P3 and compared with the same region in Pan troglodytes to determine whether microwear can be useful for identifying changes in the occlusal relationship between the P3 and maxillary canine in early Australopithecus. Finally, temporal trends in P3 Phase II and mesial microwear are considered. Results indicate that 1) both the P3 and molar Phase II facets of A. afarensis have less complex microwear textures than in A. africanus or C. atys; 2) A. afarensis P3 and molar Phase II textures differ, though not to the extent seen in taxa that eat hard and tough items; 3) microwear along the A. afarensis mesial protoconid crest is clearly distinct from that of the P. troglodytes, indicating that there is no honing equivalent in A. afarensis; and 4) there is little evidence of change over time in A. afarensis P3 microwear on either the mesial or Phase II facet. In sum, these results provide no evidence that A. afarensis routinely loaded either its premolars or molars to process hard objects or that A. afarensis P3 function changed over time.

Viewpoints: feeding mechanics, diet, and dietary adaptations in early hominins.

Inference of feeding adaptation in extinct species is challenging, and reconstructions of the paleobiology of our ancestors have utilized an array of analytical approaches. Comparative anatomy and finite element analysis assist in bracketing the range of capabilities in taxa, while microwear and isotopic analyses give glimpses of individual behavior in the past. These myriad approaches have limitations, but each contributes incrementally toward the recognition of adaptation in the hominin fossil record. Microwear and stable isotope analysis together suggest that australopiths are not united by a single, increasingly specialized dietary adaptation. Their traditional (i.e., morphological) characterization as "nutcrackers" may only apply to a single taxon, Paranthropus robustus. These inferences can be rejected if interpretation of microwear and isotopic data can be shown to be misguided or altogether erroneous. Alternatively, if these sources of inference are valid, it merely indicates that there are phylogenetic and developmental constraints on morphology. Inherently, finite element analysis is limited in its ability to identify adaptation in paleobiological contexts. Its application to the hominin fossil record to date demonstrates only that under similar loading conditions, the form of the stress field in the australopith facial skeleton differs from that in living primates. This observation, by itself, does not reveal feeding adaptation. Ontogenetic studies indicate that functional and evolutionary adaptation need not be conceptually isolated phenomena. Such a perspective helps to inject consideration of mechanobiological principles of bone formation into paleontological inferences. Finite element analysis must employ such principles to become an effective research tool in this context.

Dental microwear texture and anthropoid diets.

Dental microwear has long been used as evidence concerning the diets of extinct species. Here, we present a comparative baseline series of dental microwear textures for a sample of 21 anthropoid primate species displaying interspecific and intraspecific dietary variability. Four dental microwear texture variables (complexity, anisotropy, textural fill volume, and heterogeneity) were computed based on scale-sensitive fractal analysis and high-resolution three-dimensional renderings of microwear surfaces collected using a white-light confocal profiler. The purpose of this analysis was to assess the extent to which these variables reflect variation in diet. Significant contrasts between species with diets known to include foods with differing material properties are clearly evident for all four microwear texture variables. In particular, species that consume more tough foods, such as leaves, tended to have high levels of anisotropy and low texture complexity. The converse was true for species including hard and brittle items in their diets either as staples or as fallback foods. These results reaffirm the utility of dental microwear texture analysis as an important tool in making dietary inferences based on fossil primate samples.

A dental topographic analysis of chimpanzees.

Molar tooth morphology is generally said to reflect a compromise between phylogenetic and functional influences. Chimpanzee subspecies have been reported to exhibit differences in molar dimensions and nonmetric traits, but these have not been related to differences in their diets. And in fact, observations to date of the diets of chimpanzees have not revealed consistent differences among subspecies. This study uses dental topographic analyses shown to reflect diet-related differences in occlusal morphology among primate species, to assess within-species variation among chimpanzee subspecies. High-resolution casts from museum collections were examined by laser scanning, and resulting data were analyzed using GIS algorithms and a two-factor ANOVA model. Although differences were noted between wear stages within subspecies in surface slope, relief, and angularity, none were found to distinguish the subspecies from one another in these attributes. This might reflect limitations in the ability of this method to detect diet-related differences, but is also consistent with a lack of differences in functionally relevant aspects of occlusal morphology among chimpanzee subspecies.

Dental microwear and stable isotopes inform the paleoecology of extinct hominins.

Determining the diet of an extinct species is paramount in any attempt to reconstruct its paleoecology. Because the distribution and mechanical properties of food items may impact postcranial, cranial, mandibular, and dental morphologies related to their procurement, ingestion, and mastication, these anatomical attributes have been studied intensively. However, while mechanical environments influence skeletal and dental features, it is not clear to what extent they dictate particular morphologies. Although biomechanical explanations have been widely applied to extinct hominins in attempts to retrodict dietary proclivities, morphology may say as much about what they were capable of eating, and perhaps more about phylogenetic history, than about the nature of the diet. Anatomical attributes may establish boundary limits, but direct evidence left by the foods that were actually (rather than hypothetically) consumed is required to reconstruct diet. Dental microwear and the stable light isotope chemistry of tooth enamel provide such evidence, and are especially powerful when used in tandem. We review the foundations for microwear and biogeochemistry in diet reconstruction, and discuss this evidence for six early hominin species (Ardipithecus ramidus, Australopithecus anamensis, Au. afarensis, Au. africanus, Paranthropus robustus, and P. boisei). The dietary signals derived from microwear and isotope chemistry are sometimes at odds with inferences from biomechanical approaches, a potentially disquieting conundrum that is particularly evident for several species.

Dental microwear texture analysis shows within-species diet variability in fossil hominins.

Reconstructing the diets of extinct hominins is essential to understanding the paleobiology and evolutionary history of our lineage. Dental microwear, the study of microscopic tooth-wear resulting from use, provides direct evidence of what an individual ate in the past. Unfortunately, established methods of studying microwear are plagued with low repeatability and high observer error. Here we apply an objective, repeatable approach for studying three-dimensional microwear surface texture to extinct South African hominins. Scanning confocal microscopy together with scale-sensitive fractal analysis are used to characterize the complexity and anisotropy of microwear. Results for living primates show that this approach can distinguish among diets characterized by different fracture properties. When applied to hominins, microwear texture analysis indicates that Australopithecus africanus microwear is more anisotropic, but also more variable in anisotropy than Paranthropus robustus. This latter species has more complex microwear textures, but is also more variable in complexity than A. africanus. This suggests that A. africanus ate more tough foods and P. robustus consumed more hard and brittle items, but that both had variable and overlapping diets.

Craniofacial biomechanics and functional and dietary inferences in hominin paleontology.

Finite element analysis (FEA) is a potentially powerful tool by which the mechanical behaviors of different skeletal and dental designs can be investigated, and, as such, has become increasingly popular for biomechanical modeling and inferring the behavior of extinct organisms. However, the use of FEA to extrapolate from characterization of the mechanical environment to questions of trophic or ecological adaptation in a fossil taxon is both challenging and perilous. Here, we consider the problems and prospects of FEA applications in paleoanthropology, and provide a critical examination of one such study of the trophic adaptations of Australopithecus africanus. This particular FEA is evaluated with regard to 1) the nature of the A. africanus cranial composite, 2) model validation, 3) decisions made with respect to model parameters, 4) adequacy of data presentation, and 5) interpretation of the results. Each suggests that the results reflect methodological decisions as much as any underlying biological significance. Notwithstanding these issues, this model yields predictions that follow from the posited emphasis on premolar use by A. africanus. These predictions are tested with data from the paleontological record, including a phylogenetically-informed consideration of relative premolar size, and postcanine microwear fabrics and antemortem enamel chipping. In each instance, the data fail to conform to predictions from the model. This model thus serves to emphasize the need for caution in the application of FEA in paleoanthropological enquiry. Theoretical models can be instrumental in the construction of testable hypotheses; but ultimately, the studies that serve to test these hypotheses - rather than data from the models - should remain the source of information pertaining to hominin paleobiology and evolution.

Ecological stress and linear enamel hypoplasia in Cebus.

Although it is assumed that monkeys in some environments experience more nutritional or physiological stress than others, little research has been conducted on this topic. This study examines the relationship between linear enamel hypoplasia (LEH) frequency, a physiological indicator of stress, and environmental stressors. To test this relationship, LEH frequencies were calculated for 144 Cebus from 54 locations in Brazil. Habitat, temperature range, and annual rainfall were compared between individuals with and without LEH. The LEH frequency for Cebus from semideciduous forests was significantly higher than that for monkeys from coastal areas, the rainforest, and the savanna (chi(2) = 9.97, df = 1; P = 0.0016). A significantly higher LEH frequency was also found for monkeys living in environments with the mean annual temperature between 15 and 18 degrees C than for those in environments greater than 18 degrees C (chi(2) = 7.74, df = 1, P = 0.0054). However, no significant difference was found between LEH frequency and annual rainfall (t = 1.22, P = 0.23) or the average difference in rainfall between the driest and wettest months (t = 0.77, P = 0.44). These results indicate that levels of physiological stress can differ among environments and that habitat and temperature, but not precipitation, may be driving the difference in stress levels among environments.

Diet reconstruction in antebellum Baltimore: Insights from dental microwear analysis.

Construction in the City of Baltimore during 1996 led to the recovery of human skeletal remains dating from 1792 to 1856. Historical research indicates that the skeletal remains come from two adjacent graveyards: Christ's Church Episcopalian Cemetery and the Potters Field East. The different socioeconomic status of the internees in each cemetery suggests the possibility of marked contrasts in lifestyle, health, and diet. To shed further light on these possibilities, analyses of microscopic wear patterns on teeth, or dental microwear analyses, were undertaken. A sample from Spanish Florida was used to help interpret the results. Epoxy casts of incisor and molar teeth were placed in an SEM and photomicrographs of clean wear facets were taken. The photomicrographs were digitized using the software package Microware 4.02. Statistical analyses of rank transformed data consisted of single-factor ANOVA, followed by post hoc tests. No significant differences were found between Christ's Church and Potters Field East samples for any of the variables examined in either molar or incisor teeth. However, differences between each Baltimore sample and the La Florida samples give suggestions of possible diet differences in antebellum Baltimore. The mosaic of differences between the Baltimore and La Florida samples probably reflects the wide variety of foods available to antebellum Baltimoreans as well as the relative lack of abrasives in their diet.

Variations in the mechanical properties of Alouatta palliata molar enamel.

Teeth have provided insights into many topics including primate diet, paleobiology, and evolution, due to the fact that they are largely composed of inorganic materials and may remain intact long after an animal is deceased. Previous studies have reported that the mechanical properties, chemistry, and microstructure of human enamel vary with location. This study uses nanoindentation to map out the mechanical properties of Alouatta palliata molar enamel on an axial cross-section of an unworn permanent third molar, a worn permanent first molar, and a worn deciduous first molar. Variations were then correlated with changes in microstructure and chemistry using scanning electron microscopy and electron microprobe techniques. The hardness and Young's modulus varied with location throughout the cross-sections from the occlusal surface to the dentin-enamel junction (DEJ), from the buccal to lingual sides, and also from one tooth to another. These changes in mechanical properties correlated with changes in the organic content of the tooth, which was shown to increase from approximately 6% near the occlusal surface to approximately 20% just before the DEJ. Compared to human enamel, the Alouatta enamel showed similar microstructures, chemical constituents, and magnitudes of mechanical properties, but showed less variation in hardness and Young's modulus, despite the very different diet of this species.

Getting Humans Off Monkeys' Backs: Using Primate Acclimation as a Guide for Habitat Management Efforts.

Wild primates face grave conservation challenges, with habitat loss and climate change projected to cause mass extinctions in the coming decades. As large-bodied Neotropical primates, mantled howling monkeys (Alouatta palliata) are predicted to fare poorly under climate change, yet are also known for their resilience in a variety of environments, including highly disturbed habitats. We utilized ecophysiology research on this species to determine the morphological, physiological, and behavioral mechanisms howlers employ to overcome ecological challenges. Our data show that howlers at La Pacifica, Costa Rica are capable of modifying body size. Howlers displayed reduced mass in warmer, drier habitats, seasonal weight changes, frequent within-lifetime weight fluctuations, and gradual increases in body mass over the past four decades. These within-lifetime changes indicate a capacity to modify morphology in a way that can impact animals' energetics and thermodynamics. Howlers are also able to consume foods with a wide variety of food material properties by altering oral processing during feeding. While this capability suggests some capacity to cope with the phenological shifts expected from climate change and increased habitat fragmentation, data on rates of dental microwear warn that these acclimations may also cost dental longevity. Lastly, we found that howlers are able to acclimate to changing thermal pressures. On shorter-term daily scales, howlers use behavioral mechanisms to thermoregulate, including timing activities to avoid heat stress and utilizing cool microhabitats. At the seasonal scale, animals employ hormonal pathways to influence heat production. These lines of evidence cumulatively indicate that howlers possess morphological, physiological, and behavioral mechanisms to acclimate to environmental challenges. As such, howlers' plasticity may facilitate their resilience to climate change and habitat loss. While habitat loss in the tropics is unlikely to abate, our results point to a potential benefit of active management and selective cultivation to yield large, interconnected forest fragments with targeted phenology that provides both a complex physical structure and a diversity of food sources. These steps could assist howlers in using their natural acclimation potential to survive future conservation threats.