Craniofacial variation and dietary adaptations of African colobines

African colobine monkeys show considerable craniofacial variation among species, although the evolutionary causes of this diversity are unclear. In light of growing evidence that diet varies considerably among colobine species, we investigated whether colobine craniofacial morphology varies as a function of their diet. We compared craniofacial morphology among five African species: Colobus angolensis, C. guereza, C. polykomos, Piliocolobus badius, and P. verus. Matrix correlation analysis indicated a significant correlation between species-specific morphological distance and dietary distance matrices. The mechanical advantage of the masseter muscle was higher in seed-eaters (C. angolensis and C. polykomos) and lower in those that eat mainly young leaves (C. guereza, P. badius, and P. verus). Canonical correspondence analysis revealed that the durophagous colobines possess relatively wider bigonial breadths, anteroposteriorly shorter faces, shorter postcanine tooth rows, more medially positioned dental batteries, wider bizygomatic arches, and anteroposteriorly longer zygomatic arches. Under the constrained lever model, these morphological features suggest that durophagous colobines have the capacity to generate relatively greater maximum bite forces. However, no consistent relationship was observed between diet and variation in the mandibular corpus and symphysis, implying that robust mandibles are not necessarily adaptations for stress resistance. Factors that may influence mandibular robusticity include allometry of symphyseal curvature and canine tooth support. Finally, linear measures of mandibular robusticity may suffer from error.     

A study of the taxonomic significance of the dental arch.

Sixteen dimensions were measured from the maxillary and mandibular dental arches of different ethnic groups of man, apes and monkeys. Multivariate analysis showed that discrimination was possible among the ethnic groups of man on the one hand and between the ape and monkey samples on the other. Nevertheless, the actual degree of discrimination between the primate samples depended upon whether the maxillary or mandibular arch dimensions were analysed. Furthermore, subsequent inclusion of the dental arch dimensions of fossil hominoid samples into the analysis confirmed the taxonomic significance of the dental arch, although its importance must await the acquisition of the more non-human primate data.     

Dental arch form in the cercopithecidae

The dental arches of the major genera of Old World monkeys (superfamily Cercopithecoidea) were studied by morphometric techniques. Bicanine and bimolar breadths and arch lengths were ascertained for maxillary and mandibular arches. This data was then subjected to a variety of statistical tests. Corresponding arch dimensions of the upper and lower dentition showed the highest correlations, while the lowest correlations were generally observed between comparisons of arch dimensions and body size. A new expression was developed, relative male palate size (RMPS), which quantified the degree of sexual dimorphism while correcting for body size. The four hierarchies examined using RMPS values were sexual dimorphism, modes of sexual selection and predator defense and diet. Maxillary bicanine breadth was the only parameter that exhibited sexual dimorphism consistently in each of the four hierarchies, although differences in arch size were identified for diet and predator defense. Species grouped by predator defense showed the most sexual dimorphism in arch parameters.     

An analysis into age changes of the human dental arch by a multivariate technique

The dimensions of the maxillary and mandibular dental arches were measured from 280 subjects aged between 3 and 15 years in a cross-sectional study. The growth changes were examined by means of canonical analysis, a multivariate technique. This technique enabled the dental arches to be treated as biological units, rather than a series of discrete parameters, as is possible using univariate techniques. The results showed that the size and shape of the dental arches changed maximally between the periods 5–7 and 11–13 years of age. These periods correspond to the major phases of permanent tooth eruption.     

The British dental arch

Thirty-two dimensions of the maxillary and mandibular dental arches were measured on 320 adult skulls derived from Anglo-saxon to modern times. Multivariate analysis showed varying patterns of contrast between the samples, depending upon which arch dimensions were considered. The results suggest that dental arch changes occur independently of jaw changes.     

Molar scaling in strepsirrhine primates

We examined how maxillary molar dimensions change with body and skull size estimates among 54 species of living and subfossil strepsirrhine primates. Strepsirrhine maxillary molar areas tend to scale with negative allometry, or possibly isometry, relative to body mass. This observation supports several previous scaling analyses showing that primate molar areas scale at or slightly below geometric similarity relative to body mass. Strepsirrhine molar areas do not change relative to body mass(0.75), as predicted by the metabolic scaling hypothesis. Relative to basicranial length, maxillary molar areas tend to scale with positive allometry. Previous claims that primate molar areas scale with positive allometry relative to body mass appear to rest on the incorrect assumption that skull dimensions scale isometrically with body mass. We identified specific factors that help us to better understand these observed scaling patterns. Lorisiform and lemuriform maxillary molar scaling patterns did not differ significantly, suggesting that the two infraorders had little independent influence on strepsirrhine scaling patterns. Contrary to many previous studies of primate dental allometry, we found little evidence for significant differences in molar area scaling patterns among frugivorous, folivorous, and insectivorous groups. We were able to distinguish folivorous species from frugivorous and insectivorous taxa by comparing M1 lengths and widths. Folivores tend to have a mesiodistally elongated M1 for a given buccolingual M1 width when compared to the other two dietary groups. It has recently been shown that brain mass has a strong influence on primate dental eruption rates. We extended this comparison to relative maxillary molar sizes, but found that brain mass appears to have little influence on the size of strepsirrhine molars. Alternatively, we observed a strong correlation between the relative size of the facial skull and relative molar areas among strepsirrhines. We hypothesize that this association may be underlain by a partial sharing of the patterning of development between molar and facial skull elements.     

Arch form,tooth size,and occlusomandibular kinesis in the Ceboidea

Correlations between dental morphology, arch configuration, and jaw movement patterns were quantitatively investigated in 23 ceboid species to elucidate integrative aspects of occlusal functional anatomy in an adaptive and evolutionary context. Differential maxillary-mandibular arch widths are primary in guiding lateral jaw movements. These movements are characterized according to their associated condylar shifts as either predominantly translatory or rotational. Predominantly translatory movements result from peripheral contact relationships between maxillary arches which are considerably wider posteriorly than their opposing mandibular arches. The greatest degree of mandibular movement is in the molar region in functional association with wide “primitive” maxillary molars, narrow mandibular molars, constricted maxillary intercanine widths, and narrow maxillary incisors. In contrast, predominantly rotational masticatory jaw movements result from differential arch widths which are greatest in the maxillary canine and incisor regions. Here most jaw movement is in the anterior segment and this is reflected in small maxillary-mandibular molar width differences, a high degree of premolarization, wide-set maxillary canine teeth, and wide maxillary incisors. Possible selectional factors in the putative evolution of rotational predominance in mastication from the more primitive translatory pattern are discussed.     

Jaw morphology and function in living and fossil old world monkeys

This allometric investigation on a sample of 29 cercopithecine and 22 colobine taxa augments the data and implications of prior work on subfamilial variation in mandibular form and function in recent Cercopithecidae. To increase the size range encompassed by living cercopithecines and colobines, I included many of the larger-bodied fossil specimens. These analyses serve to fill a gap in our understanding of size-related changes in masticatory function and symphyseal morphology and curvature in extant and extinct Old World monkeys. Results of subfamilial scaling comparisons indicate that for a given jaw length, colobines possess significantly more robust corpora and symphyses than those of cercopithecines, especially at smaller sizes. Following from previous work, the most plausible explanation for why the subfamilies differ in relative corporeal and symphyseal dimensions is that colobine mandibles experience elevated loads and greater repetitive loading during mastication due, on average, to processing a diet of tough leaves and/or seeds. Although colobines have relatively larger symphyses, subfamilial analyses of symphyseal curvature demonstrate that they evince less symphyseal curvature vis-à-vis cercopithecines of a common size. Moreover, both subfamilies exhibit similar allometric changes in the degree of curvature, such that larger-bodied Old World monkeys have more curved symphyses than those of smaller taxa. Subfamilial scaling analyses also indicate that colobines possess a shorter M2 bite-point length relative to masseter lever-arm length, but not versus temporalis lever-arm length. Thus, as compared to cercopithecines, colobines can recruit less masseter-muscle force to produce similar bite forces during mastication. In both clades, M² bite-point length scales with positive allometry relative to masseter lever-arm length, such that larger species are less efficient at generating molar bite forces. This seems especially important due to the lack of subfamily difference in M² bite-point:temporalis lever-arm scaling (which is isometric across cercopithecids). A consideration of extinct cercopithecids indicates that many of the large-bodied papionins have more robust corpora, due perhaps to a diet which was of similar toughness to that of extant and extinct colobines. However, the biomechanical arrangements of the masseter and temporalis in all but one fossil cercopithecine and all of the fossil colobine specimens are much as predicted for a subfamilial member of its skull size. That most large-bodied papionins with tougher diets nonetheless maintain a less efficient jaw-muscle configuration may be due to stronger offsetting selection for increased relative canine size and gape.     

Masticatory stress and the mechanics of "wishboning" in colobine jaws

Cercopithecoid monkeys experience relatively high strains along the lingual aspect of the mandibular symphysis because of lateral transverse bending of the mandibular corpora ("wishboning") during mastication. Hylander (Am J Phys Anthropol 64 (1984) 1-46; Am Zool 25 (1985) 315-330) demonstrated that the distribution of strains arising from wishboning loads is comprehensible with reference to the mechanics of curved beams. Theory of curved beams suggests that lingual tensile strains are some multiple of labial compressive strains, yet limitations of experimental methods and uncertainty in estimating parameters needed for theoretical calculations have confounded attempts to characterize the magnitude of this disparity of normal strains. We evaluate the theoretical disparity of normal strains in wishboning in comparison to in vitro strains collected under controlled loads for a sample of mandibles representing two colobine species (N = 6). These data suggest that in colobine monkeys, maximum normal lingual strains should be at least twice maximum labial strains. In addition, we reexamine the distribution of symphyseal stress under an assumption of asymmetric bending, a general approach for calculation of stress appropriate for members that lack a plane of symmetry and are bent along an axis that is not coincident with the member's principal axes. Under asymmetric bending in colobine mandibles, the effect of symphyseal inclination on lingual strain is mitigating at the superior transverse torus and exacerbating at the inferior transverse torus. Relative compliance of colobine mandibular bone further supports the hypothesis that the structural and material properties of the colobine mandibular symphysis do not represent a morphological strategy for minimizing masticatory strain.     

Individual tooth macrowear pattern guides the reconstruction of Sts 52 (Australopithecus africanus) dental arches

The functional restoration of the occlusal relationship between maxillary and mandibular tooth rows is a major challenge in modern dentistry and maxillofacial surgery. Similar technical challenges are present in paleoanthropology when considering fragmented and deformed mandibular and maxillary fossils. Sts 52, an Australopithecus africanus specimen from Sterkfontein Member 4, represents a typical case where the original shape of the dental arches is no longer preserved. It includes a partial lower face (Sts 52a) and a fragmented mandible (Sts 52b), both incomplete and damaged to such an extent to thwart attempts at matching upper and lower dentitions. We show how the preserved macrowear pattern of the tooth crowns can be used to functionally reconstruct Sts 52's dental arches. High‐resolutiondental stone casts of Sts 52 maxillary and mandibular dentition were mounted and repositioned in a dental articulator. The occlusal relationship between antagonists was restored based on the analysis of the occlusal wear pattern of each preserved tooth, considering all dental contact movements represented in the occlusal compass. The reconstructed dental arches were three‐dimensional surface scanned and their occlusal kinematics tested in a simulation. The outcome of this contribution is the first functional restoration of A. africanus dental arches providing new morphometric data for specimen Sts 52. It is noteworthy that the method described in this case study might be applied to several other fossilspecimens. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.     

Modified technique of presurgical infant maxillary orthopedics for complete unilateral cleft lip and palate

This article introduces a modified device fabrication and facial taping method that increases the efficiency and efficacy of presurgical infant maxillary orthopedic therapy for babies born with complete cleft lip and palate. Interarch and intra-arch relationships of the maxillary and mandibular dental arches were evaluated on mounted stone models before and after treatment. The palatal plate device was custom-fabricated in a manner that bypassed the need for periodic acrylic addition and removal, thereby eliminating the risk of natural maxillary growth restriction during therapy. Elastic labial tapes were fabricated and applied in a configuration that mimicked normal function of the orbicularis oris muscle. A nasal stent wire was utilized from the initiation of therapy to enhance intraoral retention and stability of the device. Examples of infants undergoing a unilateral complete cleft lip and palate treatment protocol are presented. Treatment objectives were achieved within 7 to 8 weeks of therapy for patients who had an initial alveolar cleft size more than 10 to 12 mm. The modified protocol of presurgical infant maxillary orthopedic therapy is an effective and efficient treatment modality in reversing the pre-existing orofacial dysmorphism by redirecting the infant's natural growth.     

Size and shape of the mandibular condyle in primates

The relationships between the size of the articular surface of the mandibular condyle and masticatory muscle size, tooth size, diet, and biomechanical variables associated with mastication were studied by taking 12 measurements on skulls of 253 adult female anthropoid primates, including three to ten specimens from each of 32 species. In regressions of condylar length, width, or area against body weight, logarithmic transformations substantially improve the fit of the equations compared with untransformed data. There is a strong relationship between condylar measurements and body weight, with all correlations being .94 or higher. The slopes of the allometric regressions of length, width, and area of the condylar head indicate slight positive allometry with body size. Folivorous primates have smaller condyles than frugivorous primates, and colobines have smaller condyles than cebids, cercopithecines, or hominoids. When colobines are eliminated, the differences between frugivores and folivores are not significant. However, the two species with the relatively largest condyles are Pongo pygmaeus and Cercocebus torquatus, suggesting that there may be a relationship between unusually large condylar dimensions and the ability to crack hard nuts between the teeth. Cranial features having strong positive correlations with condylar dimensions include facial prognathism, maxillary incisor size, maxillary postcanine area, mandibular ramus breadth, and temporal fossa area. These data are interpreted as indicating that relatively large condyles are associated with relatively large masticatory muscles, relatively inefficient mandibular biomechanics, and a large dentition. These relationships support the growing evidence that the temporomandibular joint is a stress-bearing joint in normal function.     

Development of the helicoidal plane

In this study of thebelicoidal occlusal plane the relationships between tooth wear, the transverse slopes of mandibular molars and dental arch breadths were examined in 74 pre-contemporary Australian Aboriginal skulls. With increasing age and tooth wear the orientation of the mandibular occlusal surfaces increased towards the buccal. The differential occlusal orientation from first to third molars, present at eruption, tended to increase progressively with tooth wear. These functionally induced changes, together with regional differences in relative breadths of the maxillary and mandibular dental arches, are important in the development of abelicoidal occlusal plane.     

Ancestral loss of the maxillary sinus in Old World monkeys and independent acquisition in Macaca

Cercopithecoid monkeys are unique among primates in that all species (except macaques) lack a maxillary sinus, an unusual condition among eutherian mammals. Although this uncommon distribution of cranial pneumatization was noted previously, the phylogenetic ramifications have not been investigated fully. Recently, character state optimization analysis of computed tomography (CT) data from extant Old World monkeys suggested that the loss of the sinus may have occurred at the origin of the group, unlike previous hypotheses positing only a reduction in size of the structure. To critically evaluate the "early loss" hypothesis, a recently recovered complete cranium of Victoriapithecus macinnesi from Maboko Island, Kenya, was examined by CT to determine the extent of its cranial pneumatization. This taxon is crucial for evaluating character state evolution in Old World monkeys, due to its phylogenetic position, preceding the cercopithecine/colobine split. CT analysis reveals only cancellous bone lateral of the nasal cavity, indicating that Victoriapithecus does not possess a maxillary sinus. Phylogenetic evaluation of the fossil with extant catarrhine taxa strongly supports the early loss of the sinus in cercopithecoids. The results suggest that the maxillary sinus found in the genus Macaca is not homologous with that of other eutherians, which may provide insights into the origin and function (if any) of the paranasal pneumatizations.     

Nonmetric tooth crown traits in a Sri Lankan aboriginal Vedda population

This study was conducted to determine the frequencies of non-metric tooth crown traits of Vedda of Sri Lanka and to investigate the affinities of these morphological variations with those of other world populations.Fifty dental plaster casts were observed. The Arizona State University dental anthropology system was adopted for classification of the 16 traits observed. We used 13 traits to compare the Vedda and other world populations. Using the frequencies of 13 traits, Smith Mean Measure of Divergence was calculated to determine inter-population distances. Affinities among the Vedda and other world populations were expressed in two dimensions of the principal coordinate analysis.Cusp number in mandibular second molar and hypocone absence in maxillary second molar had the highest frequency at 95.9% and 93.8%, respectively. Shovelling, double shovelling in the maxillary central incisor and deflecting wrinkle in the mandibular first molar had the lowest frequency at 0%. The principal coordinate analysis showed that Sino American and Western Eurasian populations were separated in negative and positive directions in the first principal coordinate axis. Vedda located with the Western Eurasian population groups. Sahul and Sunda Pacific populations located in the intermediate position between Sino American and Western Eurasian populations.The dental phenotype of Vedda has close affinities with those of early south Asian populations. They are far different from Sino American and Sunda pacific populations. Vedda shows closer affinities to Sahul Pacific and South African (Bantu) populations.     

Metric and morphological variability in the dentition of Colobine monkeys

A morphological study was performed using 97 Colobus polykomos and 41 Nasalis larvatus. The premolars and molars of these two taxa of leaf-eating monkeys were similar in that they possessed deeply grooved and highly pointed cusps. Of 7 dental traits investigated, 4 were significantly different between the 2 forms. Indeed, with 2 of these, the position of the lingual cusp on PM₄ and the number of cusps on PM³ displayed highly contrasting structures.Metric analysis was conducted using 47 C. polykomos and 37 N. larvatus specimens. Mesiodistal and buccolingual measurements were taken on the upper canine and all postcanine teeth. Sexual dimorphism was detected in most maxillary dental measurements and all mandibular estimates of both species. In all cases, males exceeded females in tooth diameter. Testing between species by sex revealed that C. polykomos was significantly larger in several canine and premolar dimensions while N. larvatus had significantly larger molar dimensions. An additional 73 measurements were selected to represent occluding dental structures and a morphological integration analysis was conducted. Results of this analysis indicated that C. polykomos possessed a C¹-PM₃ complex which was more highly intercorrelated whereas the broad lophed N. larvatus molars constituted a more highly correlated functional unit.It was suggested that the above differences between these two groups of leaf-eating monkeys probably resulted from inbreeding, isolation and drift.     

Duplication and divergence of 2 distinct pancreatic ribonuclease genes in leaf-eating African and Asian colobine monkeys

Unique among primates, the colobine monkeys have adapted to a predominantly leaf-eating diet by evolving a foregut that utilizes bacterial fermentation to breakdown and absorb nutrients from such a food source. It has been hypothesized that pancreatic ribonuclease (pRNase) has been recruited to perform a role as a digestive enzyme in foregut fermenters, such as artiodactyl ruminants and the colobines. We present molecular analyses of 23 pRNase gene sequences generated from 8 primate taxa, including 2 African and 2 Asian colobine species. The pRNase gene is single copy in all noncolobine primate species assayed but has duplicated more than once in both the African and Asian colobine monkeys. Phylogenetic reconstructions show that the pRNase-coding and noncoding regions are under different evolutionary constraints, with high levels of concerted evolution among gene duplicates occurring predominantly in the noncoding regions. Our data suggest that 2 functionally distinct pRNases have been selected for in the colobine monkeys, with one group adapting to the role of a digestive enzyme by evolving at an increased rate with loss of positive charge, namely arginine residues. Conclusions relating our data to general hypotheses of evolution following gene duplication are discussed.