Dental topography of prosimian premolars predicts diet: A comparison in premolar and molar dietary classification accuracies.

OBJECTIVES: This study tests whether (1) premolar topography of extant "prosimians" (strepsirrhines and tarsiers) successfully predicts diet and (2) whether the combination of molar and premolar topography yields higher classification accuracy than using either tooth position in isolation. MATERIALS AND METHODS: Dental topographic metrics (ariaDNE, relief index, and orientation patch count rotated) were calculated for 118 individual matched-pairs of mandibular fourth premolars (P4) and second molars (M2). The sample represents 7 families and 22 genera. Tooth variables were analyzed in isolation (P4 only; M2 only), together (P4 and M2), and combined (PC1 scores of bivariate principal component analyses of P4 and M2 for each metric). Discriminant function analyses were conducted with and without a measure of size (two-dimensional surface area). RESULTS: When using topography only, "prosimian" P4 shape predicts diet with a success rate that is slightly higher than that of M2 shape. When absolute size is included, premolars and molars perform comparably well. Including both premolar and molar topography (separately or combined) improves classification accuracy for every analysis beyond considering either in isolation. Classification accuracy is highest when premolar and molar topography and size are included. DISCUSSION: Our findings indicate that molar teeth incompletely summarize the functional requirements of oral food breakdown for a given diet, and that the mechanism selecting for premolar form is more varied than what is expressed by molar teeth. Finally, our findings suggest that fossil P4s (in isolation or with the M2) can be used for meaningful dietary reconstruction of extinct primates.

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.

Lower molar shape and size in prosimian and platyrrhine primates.

The goal of this research is to evaluate the relative strength of the influences of diet, size, and phylogenetic signal on dental geometric shape. Accurate comprehension of these factors and their interaction is important for reconstructing diet and deriving characters for a cladistic analysis in fossil primates. Geometric morphometric analysis is used to identify axes of shape variation in the lower second molars of (a) prosimian primates and (b) platyrrhines. Landmarks were placed on µCT-generated surface renderings. Landmark configurations were aligned using generalized Procrustes analysis. Principal components analysis and phylogenetic principal components analysis (pPCA) were performed on species average landmark co-ordinates. pPCs were examined with phylogenetic generalized least squares analysis for association with size and with diet. PCs from both phylogenetic and non-phylogenetic analyses were sufficient to separate species by broad dietary categories, including insectivores and folivores. In neither analysis was pPC1 correlated with tooth size, but some other pPCs were significantly correlated with size. The pattern of association between pPCs and size altered when centroid size and dietary variables were combined in the model; effects of diet factors typically exceeded effects of size. These results indicate a dominant phylogenetic and dietary signal in molar shape but also show some shape change correlated with size in the absence of obvious dietary associations. Geometric morphometric analysis appears to be useful for tracking functional traits in molars, particularly in tracking differences between folivorous and insectivorous species.

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.

Genetic comparisons yield insight into the evolution of enamel thickness during human evolution.

Enamel thickness varies substantially among extant hominoids and is a key trait with significance for interpreting dietary adaptation, life history trajectory, and phylogenetic relationships. There is a strong link in humans between enamel formation and mutations in the exons of the four genes that code for the enamel matrix proteins and the associated protease. The evolution of thick enamel in humans may have included changes in the regulation of these genes during tooth development. The cis-regulatory region in the 5' flank (upstream non-coding region) of MMP20, which codes for enamelysin, the predominant protease active during enamel secretion, has previously been shown to be under strong positive selection in the lineages leading to both humans and chimpanzees. Here we examine evidence for positive selection in the 5' flank and 3' flank of AMELX, AMBN, ENAM, and MMP20. We contrast the human sequence changes with other hominoids (chimpanzees, gorillas, orangutans, gibbons) and rhesus macaques (outgroup), a sample comprising a range of enamel thickness. We find no evidence for positive selection in the protein-coding regions of any of these genes. In contrast, we find strong evidence for positive selection in the 5' flank region of MMP20 and ENAM along the lineage leading to humans, and in both the 5' flank and 3' flank regions of MMP20 along the lineage leading to chimpanzees. We also identify putative transcription factor binding sites overlapping some of the species-specific nucleotide sites and we refine which sections of the up- and downstream putative regulatory regions are most likely to harbor important changes. These non-coding changes and their potential for differential regulation by transcription factors known to regulate tooth development may offer insight into the mechanisms that allow for rapid evolutionary changes in enamel thickness across closely-related species, and contribute to our understanding of the enamel phenotype in hominoids.

Dental topography of platyrrhines and prosimians: convergence and contrasts.

Dental topographic analysis is the quantitative assessment of shape of three-dimensional models of tooth crowns and component features. Molar topographic curvature, relief, and complexity correlate with aspects of feeding behavior in certain living primates, and have been employed to investigate dietary ecology in extant and extinct primate species. This study investigates whether dental topography correlates with diet among a diverse sample of living platyrrhines, and compares platyrrhine topography with that of prosimians. We sampled 111 lower second molars of 11 platyrrhine genera and 121 of 20 prosimian genera. For each tooth we calculated Dirichlet normal energy (DNE), relief index (RFI), and orientation patch count (OPCR), quantifying surface curvature, relief, and complexity respectively. Shearing ratios and quotients were also measured. Statistical analyses partitioned effects of diet and taxon on topography in platyrrhines alone and relative to prosimians. Discriminant function analyses assessed predictive diet models. Results indicate that platyrrhine dental topography correlates to dietary preference, and platyrrhine-only predictive models yield high rates of accuracy. The same is true for prosimians. Topographic variance is broadly similar among platyrrhines and prosimians. One exception is that platyrrhines display higher average relief and lower relief variance, possibly related to lower relative molar size and functional links between relief and tooth longevity distinct from curvature or complexity. Explicitly incorporating phylogenetic distance matrices into statistical analyses of the combined platyrrhine-prosimian sample results in loss of significance of dietary effects for OPCR and SQ, while greatly increasing dietary significance of RFI.

Diet and dental topography in pitheciine seed predators.

Pitheciines (Pithecia, Chiropotes, and Cacajao) are a specialized clade of Neotropical seed predators that exhibit postcanine teeth with low and rounded cusps and highly crenulated occlusal surface enamel. Data on feeding ecology show that Pithecia consumes proportionally more leaves than other pitheciine species, and comparative studies demonstrate its greater molar relief and relative shearing potential. However, data on pitheciine food mechanics show that Pithecia masticates seeds with greater crushing resistance than those preferred by Chiropotes. This variation predicts an opposing morphology characterized by low and more rounded occlusal surfaces in Pithecia. We build on previous research using new methods for molar surface shape quantification by examining pitheciine second molar shearing crest length, occlusal relief, surface complexity, and surface curvature relative to nonseed specializing platyrrhines and within the context of the observed interspecific variation in pitheciine feeding ecology. Consistent with the previous analyses, our findings demonstrate that pitheciine molars exhibit low shearing, relief, and curvature compared with nonseed predators, independent of phylogeny. Pitheciines also exhibit highly "complex" occlusal topography that promotes the efficient breakdown of tough seed tissues. Overall, Pithecia, Chiropotes, and Cacajao share a similar topographic pattern, suggesting adaptation to foods with similar structural and/or mechanical properties. However, Cacajao differs in surface complexity, which reflects some variation in its feeding ecology. Contrary to the predictions, Pithecia and Chiropotes do not differ in any of the topographic variables examined. The range of demands imposed on the postcanine teeth of Pithecia might therefore select for an average topography, one that converges on that of Chiropotes.

New primate first metatarsals from the Paleogene of Egypt and the origin of the anthropoid big toe.

The specialized grasping feet of primates, and in particular the nature of the hallucal grasping capabilities of living strepsirrhines and tarsiers (i.e., 'prosimians'), have played central roles in the study of primate origins. Prior comparative studies of first metatarsal (Mt1) morphology have documented specialized characters in living prosimians that are indicative of a more abducted hallux, which in turn is often inferred to be related to an increased ability for powerful grasping. These include a well-developed peroneal process and a greater angle of the proximal articular surface relative to the long axis of the diaphysis. Although known Mt1s of fossil prosimians share these characters with living non-anthropoid primates, Mt1 morphology in the earliest crown group anthropoids is not well known. Here we describe two Mt1s from the Fayum Depression of Egypt - one from the latest Eocene (from the ∼34 Ma Quarry L-41), and one from the later early Oligocene (from the ∼29-30 Ma Quarry M) - and compare them with a sample of extant and fossil primate Mt1s. Multivariate analyses of Mt1 shape variables indicate that the Fayum specimens are most similar to those of crown group anthropoids, and likely belong to the stem catarrhines Catopithecus and Aegyptopithecus specifically, based on analyses of size. Also, phylogenetic analyses with 16 newly defined Mt1 characters support the hypotheses that "prosimian"-like Mt1 features evolved along the primate stem lineage, while crown anthropoid Mt1 morphology and function is derived among primates, and likely differed from that of basal stem anthropoids. The derived loss of powerful hallucal grasping as reflected in the Mt1 morphology of crown anthropoids may reflect long-term selection for improved navigation of large-diameter, more horizontal branches at the expense of movement in smaller, more variably inclined branches in the arboreal environment.

Comparing Dirichlet normal surface energy of tooth crowns, a new technique of molar shape quantification for dietary inference, with previous methods in isolation and in combination.

Inferred dietary preference is a major component of paleoecologies of extinct primates. Molar occlusal shape correlates with diet in living mammals, so teeth are a potentially useful structure from which to reconstruct diet in extinct taxa. We assess the efficacy of Dirichlet normal energy (DNE) calculated for molar tooth surfaces for reflecting diet. We evaluate DNE, which uses changes in normal vectors to characterize curvature, by directly comparing this metric to metrics previously used in dietary inference. We also test whether combining methods improves diet reconstructions. The study sample consisted of 146 lower (mandibular) second molars belonging to 24 euarchontan taxa. Five shape quantification metrics were calculated on each molar: DNE, shearing quotient, shearing ratio, relief index, and orientation patch count rotated (OPCR). Statistical analyses were completed for each variable to assess effects of taxon and diet. Discriminant function analysis was used to assess ability of combinations of variables to predict diet. Values differ significantly by diets for all variables, although shearing ratios and OPCR do not distinguish statistically between insectivores and folivores or omnivores and frugivores. Combined analyses were much more effective at predicting diet than any metric alone. Alone, relief index and DNE were most effective at predicting diet. OPCR was the least effective alone but is still valuable as the only quantitative measure of surface complexity. Of all methods considered, DNE was the least methodologically sensitive, and its effectiveness suggests it will be a valuable tool for dietary reconstruction.