Dental caries in living and extinct strepsirrhines with insights into diet.

Dental caries is one of the most common diseases afflicting modern humans and occurs in both living and extinct non-human primates, as well as other mammalian species. Compared to other primates, less is known about the etiology or frequency of caries among the Strepsirrhini. Given the link between caries and diet, caries frequency may be informative about the dietary ecology of a given animal. Understanding rates of caries in wild populations is also critical to assessing dental health in captive populations. Here, we examine caries frequency in a sample of 36 extant strepsirrhine species (n = 316 individuals) using odontological collections of wild-, non-captive animals housed at the American Museum of Natural History by counting the number of specimens characterized by the disease. Additionally, in the context of studying caries lesions in strepsirrhines, case studies were also conducted to test if similar lesions were found in their fossil relatives. In particular, two fossil strepsirrhine species were analyzed: the earliest Late Eocene Karanisia clarki, and the subfossil lemur Megaladapis madagascariensis. Our results suggest that caries affects 13.92% of the extant individuals we examined. The frugivorous and folivorous taxa were characterized by the highest overall frequency of caries, whereas the insectivores, gummivores, and omnivores had much lower caries frequencies. Our results suggest that caries may be common among wild populations of strepsirrhines, and in fact is more prevalent than in many catarrhines and platyrrhines. These findings have important implications for understanding caries, diet, and health in living and fossil taxa.

Hypoconulid loss in cercopithecins: Functional and developmental considerations.

Cercopithecins differ from papionins in lacking a M3 hypoconulid. Although this loss may be related to dietary differences, the functional and developmental ramifications of hypoconulid loss are currently unclear. The following makes use of dental topographic analysis to quantify shape variation in a sample of cercopithecin M3s, as well as in a sample of Macaca, which has a hypoconulid. To help understand the consequences of hypoconulid loss, Macaca M3s were virtually cropped to remove the hypoconulid and were also subjected to dental topographic analysis. The patterning cascade model and the inhibitory cascade model attempt to explain variation in cusp pattern and molar proportions, respectively. These models have both previously been used to explain patterns of variation in cercopithecines, but have not been examined in the context of hypoconulid loss. For example, previous work suggests that earlier developing cusps impact the development of later developing cusps (i.e., the hypoconulid) and that cercopithecines do not conform to the predictions of the inhibitory cascade model in that the size of the molars is not linear moving distally. Results of the current study suggest that the loss of the hypoconulid is associated with a reduction in dental topography among cercopithecins, which is potentially related to diet, although the connection to diet is not necessarily clear. Results also suggest that the loss of the hypoconulid can be explained by the patterning cascade model, and that hypoconulid loss explains the apparent lack of support for the inhibitory cascade model among cercopithecines. These findings highlight the importance of a holistic approach to studying variation in molar proportions and developmental models.

The largest and earliest known sample of dental caries in an extinct mammal (Mammalia, Euarchonta, Microsyops latidens) and its ecological implications.

Dental cavities or caries is a common disease among modern humans, affecting almost every adult. Caries frequency has been used to study dietary change in humans over time, based on an inferred tie between the incidence of caries and a carbohydrate-rich diet. However, the disease is not unique to our species. Among non-human primates, there is also variation in caries frequency associated with diet, suggesting that this metric may provide a mechanism for studying diet in broader contexts, and across geological time. To date, very few studies have examined caries among fossil mammals, and none have done so among Eocene mammals. Here, we present our analysis of the largest sample to date of fossil caries in a single extinct mammal species, Microsyops latidens, a stem primate from the early Eocene, which is known from over a thousand specimens from the Southern Bighorn Basin of Wyoming (n = 1030). Our results show that Microsyops latidens is characterized by a high prevalence of dental caries (7.48% of individuals), with notable variation through time, reaching 17.24% of individuals from a particular interval. This interval is also associated with a change in overall dental form, as quantified by dental topographic analysis, which measures functional aspects of the chewing surface of teeth. These observations suggest that this species experienced a shift in their diet to include more fruit or other sugar rich-foods for a short period. Our analysis, therefore, suggests that the diet of M. latidens fluctuated over time, as well as providing a framework for assessing caries in other fossil taxa.

The effect of high wear diets on the relative pulp volume of the lower molars.

OBJECTIVES: One role of dental pulp is in the upkeep and maintenance of dentine. Under wear, odontoblasts in the pulp deposit tertiary dentine to ensure the sensitive internal dental tissues are not exposed and vulnerable to infection. It follows that there may be an adaptive advantage for increasing molar pulp volume in anthropoid primate taxa that are prone to high levels of wear. The relative volume of dental pulp is therefore predicted to covary with dietary abrasiveness (in the sense of including foods that cause high degrees of wear). MATERIALS AND METHODS: We examined relatively unworn lower second molars in pairs of species of extant hominoids, cebids, and pitheciids that vary in the abrasiveness of their diet (n = 36). Using micro-CT scans, we measured the percent of tooth that is pulp (PTP) as the ratio of pulp volume to that of the total volume of the tooth. RESULTS: We found that in each pair of species, the taxa that consume a more abrasive diet had a significantly higher PTP than the closely related taxa that consume a softer diet. CONCLUSIONS: Our results point to an adaptive mechanism in the molars of taxa that consume abrasive diets and are thus subject to higher levels of wear. Our results provide additional understanding of the relationship between dental pulp and diet and may offer insight into the diet of extinct taxa such as Paranthropus boisei or into the adaptive context of the taurodont molars of Neanderthals.

Cladogenesis and replacement in the fossil record of Microsyopidae (?Primates) from the southern Bighorn Basin, Wyoming.

The early Eocene of the southern Bighorn Basin, Wyoming, is notable for its nearly continuous record of mammalian fossils. Microsyopinae (?Primates) is one of several lineages that shows evidence of evolutionary change associated with an interval referred to as Biohorizon A. Arctodontomys wilsoni is replaced by a larger species, Arctodontomys nuptus, during the biohorizon interval in what is likely an immigration/emigration or immigration/local extinction event. The latter is then superseded by Microsyops angustidens after the end of the Biohorizon A interval. Although this pattern has been understood for some time, denser sampling has led to the identification of a specimen intermediate in morphology between A. nuptus and M. angustidens, located stratigraphically as the latter is appearing. Because specimens of A. nuptus have been recovered approximately 60 m above the appearance of M. angustidens, it is clear that A. nuptus did not suffer pseudoextinction. Instead, evidence suggests that M. angustidens branched off from a population of A. nuptus, but the latter species persisted. This represents possible evidence of cladogenesis, which has rarely been directly documented in the fossil record. The improved understanding of both evolutionary transitions with better sampling highlights the problem of interpreting gaps in the fossil record as punctuations.

Mammalian molar complexity follows simple, predictable patterns.

Identifying developmental explanations for the evolution of complex structures like mammalian molars is fundamental to studying phenotypic variation. Previous study showed that a "morphogenetic gradient" of molar proportions was explained by a balance between inhibiting/activating activity from earlier developing molars, termed the inhibitory cascade model (ICM). Although this model provides an explanation for variation in molar proportions, what remains poorly understood is if molar shape, or specifically complexity (i.e., the number of cusps, crests), can be explained by the same developmental model. Here, we show that molar complexity conforms to the ICM, following a linear, morphogenetic gradient along the molar row. Moreover, differing levels of inhibiting/activating activity produce contrasting patterns of molar complexity depending on diet. This study corroborates a model for the evolution of molar complexity that is developmentally simple, where only small-scale developmental changes need to occur to produce change across the entire molar row, with this process being mediated by an animal's ecology. The ICM therefore provides a developmental framework for explaining variation in molar complexity and a means for testing developmental hypotheses in the broader context of mammalian evolution.

Dental Signatures for Exudativory in Living Primates, with Comparisons to Other Gouging Mammals.

Exudativory, the consumption of gums, is an obligate or a facultative dietary niche for some primates and marsupials. Exudativory has been cited as a dietary niche that may have been present in early primates, so finding a dental signature for exudativory is highly desirable. The present study combines exudativorous lorisoids (galagos and lorises) into one sample to compare to closely related, non-exudativorous lorisoids to search for a consistent dental signature of exudativory. Linear measurements were taken from the toothcomb, P2 , M3 , upper canine, and P2 from skulls of 295 adult galagids and lorisids. Also, differential distribution of enamel on the anterior teeth was qualitatively investigated as a dental signature for gouging (a behavior that facilitates some exudativory) by micro-CT scanning one specimen each from two gougers, Nycticebus coucang and Callithrix jacchus, and two non-gougers, Perodicticus potto, and Saguinus fuscicollis. Non-primate gouging mammals, the vampire bat Desmodus rotundus and the sugar glider Petaurus breviceps, were compared to non-gouging relatives. Statistical analysis revealed that exudativorous galagos and lorises had significantly (P < 0.05) reduced M3 relative to non-exudativorous galagos and lorises. While the sample sizes for assessing enamel thickness were small, preliminary results show that gouging primates and non-primate mammals have reduced lingual enamel thickness on the anterior dentition compared to non-gouging relatives. We suggest that reduction of mastication, and, therefore, M3 dimensions are a likely dental signature for exudativory in Primates. While broader samples are needed to statistically confirm, differential distribution of enamel in the anterior dentition may also be a signature of exudativory. Anat Rec, 2019. © 2018 Wiley Periodicals, Inc. Anat Rec, 303:265-281, 2020. © 2018 American Association for Anatomy.

A Novel Method for Assessing Enamel Thickness Distribution in the Anterior Dentition as a Signal for Gouging and Other Extractive Foraging Behaviors in Gummivorous Mammals.

Gummivory poses unique challenges to the dentition as gum acquisition may often require that the anterior teeth be adapted to retain a sharp edge and to resist loading because they sometimes must penetrate a highly obdurate substrate during gum extraction by means of gouging or scraping. It has been observed previously that the enamel on the labial surface of the teeth used for extraction is thicker relative to that on the lingual surface in taxa that extract gums, while enamel is more evenly distributed in the anterior teeth of taxa that do not regularly engage in extractive behaviors. This study presents a quantitative methodology for measuring the distribution of labial versus lingual enamel thickness among primate and marsupial taxa in the context of gummivory. Computed microtomography scans of 15 specimens representing 14 taxa were analyzed. Ten measurements were taken at 20% intervals starting from the base of the crown of the extractive tooth to the tip of the cutting edge across the lingual and labial enamel. A method for including worn or broken teeth is also presented. Mann-Whitney U tests, canonical variates analysis, and between-group principal components analysis were used to examine variation in enamel thickness across taxa. Our results suggest that the differential distribution of enamel thickness in the anterior dentition can serve as a signal for gouging behavior; this methodology distinguishes between gougers, scrapers, and nonextractive gummivores. Gouging taxa are characterized by significantly thicker labial enamel relative to the lingual enamel, particularly towards the crown tip. Examination of enamel thickness patterning in these taxa permits a better understanding of the adaptations for the extraction of gums in extant taxa and offers the potential to test hypotheses concerning the dietary adaptations of fossil taxa.

Three-Dimensional Geometric Morphometric Analysis of Treeshrew (Scandentia) Lower Molars: Insight into Dental Variation and Systematics.

Scandentia (treeshrews) is an order of small-bodied Indomalayan mammals generally agreed to be a member of Euarchonta with Primates and Dermoptera (colugos). However, intraordinal relationships among treeshrews are less well understood. Although recent studies have begun to clarify treeshrew taxonomy using morphological and molecular datasets, previous analysis of treeshrew dentition has yielded little clarity in terms of species-level relationships within the order. However, these studies made use of character-based methods, scoring traits across the dental arcade, which depend on there being clear differences among taxa that can be encapsulated in coding schemes. Geometric morphometrics has the potential to capture subtler shape variation, so it may be better for examining similarities among closely related taxa whose teeth have a similar bauplan. We used three-dimensional geometric morphometrics on a sample of treeshrew lower second molars and compared the patterns of variation to the results of previous studies. We captured 19 landmarks on a sample of 43 specimens representing 15 species. Using specimen-based principal components analysis and between-group principal component analysis, the two treeshrew families (Tupaiidae and Ptilocercidae) were well separated in morphospace. Moreover, several treeshrew species plot in morphospace according to the clades established in previous molecular work, with closely related species plotting closer to one another than to more distantly related species, suggesting that dental morphology can be useful when studying relationships among treeshrews. As most extinct treeshrews are known only from teeth, understanding morphological patterns in treeshrew molars is important for future work on the evolutionary history of Scandentia. Anat Rec, 302:1154-1168, 2019. © 2019 Wiley Periodicals, Inc.