Ontogeny and phylogeny in papionin primates

This study investigates the developmental bases of size and shape variation in papionin primates (Macaca, Cercocebus, Mandrillus, Lophocebus, and Papio). The analysis tests hypotheses predicting that heterochronic changes in ontogeny, particularly in the degree of overall size growth, can account for cranial diversity and "allometric scaling" in this clade. Large developmental samples of extant papionin crania are examined to test heterochronic hypotheses using bivariate allometric methods. Analyses indicate that the crania of larger papionins (Mandrillus and Papio) are generally peramorphic, surpassing size and shape ranges of smaller, and probably less-derived, macaques and mangabeys. At least two heterochronic processes, including acceleration and hypermorphosis, can account for this pattern. Ontogenetic changes include decoupling of growth and development among cranial regions, along with simple shifts in size. Allometric scaling has complex developmental bases. Size change itself is not sufficient to explain all developmental differences among papionins, but these changes are extremely important in comparisons within cranial regions such as the face. Results imply that Papio exhibits strongly derived patterns of brain growth that impact postnatal patterns of size and shape transformation. Consideration of these results in the context of recent socioecological analyses suggests that derived patterns of cranial growth in Papio may be a response to selection during the early periods of ontogeny, resulting in a distinctive life history pattern.     

Allometry, sexual dimorphism, and phylogeny: A cladistic analysis of extant African papionins using craniodental data

This study conducts a phylogenetic analysis of extant African papionin craniodental morphology, including both quantitative and qualitative characters. We use two different methods to control for allometry: the previously described narrow allometric coding method, and the general allometric coding method, introduced herein. The results of this study strongly suggest that African papionin phylogeny based on molecular systematics, and that based on morphology, are congruent and support a Cercocebus/Mandrillus clade as well as a Papio/Lophocebus/Theropithecus clade. In contrast to previous claims regarding papionin and, more broadly, primate craniodental data, this study finds that such data are a source of valuable phylogenetic information and removes the basis for considering hard tissue anatomy “unreliable” in phylogeny reconstruction. Among highly sexually dimorphic primates such as papionins, male morphologies appear to be particularly good sources of phylogenetic information. In addition, we argue that the male and female morphotypes should be analyzed separately and then added together in a concatenated matrix in future studies of sexually dimorphic taxa. Character transformation analyses identify a series of synapomorphies uniting the various papionin clades that, given a sufficient sample size, should potentially be useful in future morphological analyses, especially those involving fossil taxa.     

Homoplasy and the evolution of ontogeny in papionin primates

Recent advances in developmental biology reveal that patterns of morphological development, even during early phases, may be highly susceptible to evolutionary change. Consequently, developmental data may be uninformative with regard to distinguishing homology and homoplasy. The present analysis evaluates postnatal ontogeny in papionin primates to test hypotheses about homology and homoplasy during later periods of development. Specifically, the analysis studies the allometric bases of craniometric resemblances among four papionin genera to test the hypothesis that homoplasy in adult cranial form, particularly of baboons (Papio) and mandrills (Mandrillus), is underwritten by divergent patterns of development. Bivariate and multivariate allometric analyses demonstrate that the developmental patterns in Papio baboons diverge markedly from ontogenetic allometric trajectories in other papionin species. The resemblances between Papio and Mandrillus (assuming that patterns of development in smaller papionins are ancestral) are largely consequences of perinatal increases in relative brain size in juvenile Papio. Postnatal growth to large size and strong negative allometry of neurocranial form results in shape similarities because developmental pathways for large papionin genera intersect. Analyses show that allometric data may not be particularly informative in revealing homoplasy. However, placed into proper phylogenetic context, such data illustrate derived patterns of development that may reflect critically important life-history or ontogenetic adaptations.     

Brief communication: Enamel thickness and durophagy in mangabeys revisited

The documentation of enamel thickness variation across primates is important because enamel thickness has both taxonomic and functional relevance. The Old World monkeys commonly referred to as mangabeys have figured prominently in investigations of feeding ecology and enamel thickness. In this article, we report enamel thickness values for four mangabey taxa (Cercocebus atys, Cercocebus torquatus, Lophocebus aterrimus, and Lophocebus albigena), offer revised interpretation of the significance of thick enamel in papionin evolution, and place our new data in a broader comparative framework. Our data indicate that all mangabeys have thick enamel and that the values obtained for Cercocebus and Lophocebus equal or exceed those published for most extant non-human primates. In addition, new field data combined with a current reading of the dietary literature indicate that hard foods make up a portion of the diet of every mangabey species sampled to date. Clarification on the relationship between diet and enamel thickness among mangabeys is important not only because of recognition that mangabeys are not a natural group but also because of recent arguments that explain thick enamel as an evolved response to the seasonal consumption of hard foods.     

The phenetic affinities of Rungwecebus kipunji

The kipunji, a recently discovered primate endemic to Tanzania's Southern Highlands and Udzungwa Mountains, was initially referred to the mangabey genus Lophocebus (Cercopithecinae: Papionini), but subsequent molecular analyses showed it to be more closely related to Papio. Its consequent referral to a new genus, Rungwecebus, has met with skepticism among papionin researchers, who have questioned both the robustness of the phylogenetic results and the kipunji's morphological distinctiveness. This circumstance has been exacerbated by the immaturity of the single available specimen (FMNH 187122), an M1-stage juvenile. Therefore, a geometric morphometric analysis of juvenile papionin cranial shape was used to explore the kipunji's phenetic affinities and evaluate morphological support for its separation from Lophocebus. Three-dimensional craniometric landmarks and semi-landmarks were collected on a sample of 124 subadult (dp4-M2 stage) cercopithecid crania. Traditional interlandmark distances were compared and a variety of multivariate statistical shape analyses were performed for the zygomaxillary region (diagnostic in mangabeys) and the cranium as a whole. Raw and size-adjusted interlandmark distances show the kipunji to have a relatively taller, shorter neurocranium and broader face and cranial base than is seen in M1-stage Lophocebus. Principal components and cluster analyses consistently unite the two Lophocebus species but group the kipunji with Cercocebus and/or Macaca. Morphological distances (Mahalanobis D²) between the kipunji and Lophocebus species are comparable to distances between recognized papionin genera. Discriminant function analyses suggest phenetic affinities between the kipunji and Cercocebus/Macaca and do not support the kipunji's classification to Lophocebus or to any other papionin taxon. In canonical plots, the kipunji occupies a region intermediate between macaques and African papionins or groups with Cercocebus, suggesting that it retains basal papionin shape characteristics. In shape comparisons among M1-stage papionins, the kipunji cranium is distinguished from Lophocebus by its relatively unrestricted suborbital fossa, more parasagittally oriented zygomatic arches, and longer auditory tube and from all papionins by its relatively tall, short neurocranium, broad face and cranial base, short nasals, dished nasal profile, and dorsally oriented rostrum. The kipunji is thus a cranially diagnosable phenon with a unique combination of cranial traits that cannot be accommodated within Lophocebus as currently defined. Based upon these results, Rungwecebus appears to be a valid and useful nomen that accurately reflects the morphological diversity of African papionins.     

Functional morphology of the mangabey mandibular corpus: relationship to dental specializations and feeding behavior

Recent molecular and morphological surveys suggest that mangabeys do not represent a monophyletic group. Specifically, Cercocebus is the sister taxon of Mandrillus, whereas Lophocebus forms an unresolved trichotomy with Papio and Theropithecus. The Cercocebus-Mandrillus clade is characterized by skeletal and dental adaptations related to acquisition and processing of hard-object foods that resist decomposition for months on the forest floor. Although species of both mangabey genera can be described as frugivorous seed predators with a strong reliance on hard-object foods, a growing body of evidence indicates that Cercocebus (terrestrial) and Lophocebus (arboreal) mangabeys differ in the hardness of the seeds they consume and the manner in which seeds are processed. The taxa are also distinguished on the basis of dental morphology. Given the purported differences in feeding behaviors of the two mangabey genera, we consider whether there are predictable biomechanical consequences of these behaviors that are reflected in mandibular corpus dimensions. In addition, we present metric data summarizing functional aspects of mangabey mandibular corpus morphology. Mangabey genera are generally not distinguished by differences in relative corpus size, either in postcanine or symphyseal regions. Distinct symphyseal scaling patterns characterize the Papio-Lophocebus clade and the Mandrillus-Cercocebus clade, while the postcanine corpus scales similarly between them. The hypothesis that preferential use of the incisors vs. premolars to initially process these foods results in distinct stress environments is weakly supported, given circumstantial evidence that the relative importance of bending vs. torsion may differ between Cercocebus and Lophocebus.     

Nuclear gene trees and the phylogenetic relationships of the mangabeys (Primates: Papionini)

Phylogenetic relationships of mangabeys within the Old World monkey tribe Papionini are inferred from analyses of nuclear DNA sequences from five unlinked loci. The following conclusions are strongly supported, based on congruence among trees derived for the five separate gene regions: (1) mangabeys are polyphyletic within the Papionini; (2) Cercocebus is the sister taxon to the genus Mandrillus; and (3) Lophocebus belongs to a clade with Papio and Theropithecus, with Papio as its most likely sister taxon. Morphologically based phylogenies positing mangabey monophyly were evaluated by mapping the sequences for each locus on these trees. The data seem to fit these trees poorly in both maximum-parsimony and likelihood analyses. Incongruence among nuclear gene trees occurred in the interrelationships among Lophocebus, Papio, and Theropithecus. Several factors that may account for this incongruence are discussed, including sampling error, random lineage sorting, and introgression.      

Mitochondrial DNA phylogeny of the Old-World monkey tribe Papionini.

The evolution of the Old World monkey tribe Papionini, composed of macaques, baboons, mandrills, drills, and mangabeys, was examined using mitochondrial DNA (mtDNA) sequence data on the cytochrome oxidase subunit II gene. When analyzed cladistically, these data support a baboon clade of savannah (Papio) plus gelada (Theropithecus) baboons, as well as a clade containing drill (Mandrillus) plus mangabey (Cerocebus) genera. This result stands in opposition to most morphological phylogenies, which break up the baboon clade by placing Papio and Mandrillus as sister taxa and Theropithecus as a more distantly related lineage. Analyses of COII gene sequences also suggest that the papionin ancestral stock divided into two lineages, one leading to macaques and the other to the purely African genera. From a molecular evolutionary perspective, the papionin COII gene sequences reveal a pattern of amino acid replacements concentrated in the regions spanning the mitochondrial membrane.     

Ontogeny and homoplasy in the papionin monkey face

SUMMARY Recent molecular research has provided a consistent estimate of phylogeny for the living papionin monkeys (Cercocebus, Lophocebus, Macaca, Mandrillus, Papio, and Theropithecus). This phylogeny differs from morphological phylogenies regarding the relationships of the mangabeys (Cercocebus and Lophocebus) and baboons (Mandrillus, Papio, and Theropithecus). Under the likely assumption that the molecular estimate is correct, the incongruence between the molecular and morphological data sets indicates that the latter include numerous homoplasies. Knowledge of how these homoplasies emerge through development is important for understanding the morphological evolution of the living papionins, and also for reconstructing the phylogenetic relationships and adaptations of their fossil relatives. Accordingly, we have used geometric morphometric techniques and the molecular phylogeny to investigate the ontogeny of a key area of morphological homoplasy in papionins, the face. Two analyses were carried out. The first compared allometric vectors of Cercocebus, Lophocebus, Macaca, Mandrillus, and Papio to determine which of the facial resemblances among the genera are homoplasic and which are plesiomorphic. The second analysis focused on early post‐natal facial form in order to establish whether the facial homoplasies exhibited by the adult papionins are to some degree present early in the post‐natal period or whether they develop only later in ontogeny. The results of our analyses go some way to resolving the debate over which papionin genera display homoplasic facial similarities. They strongly suggest that the homoplasic facial similarities are exhibited by Mandrillus and Papio and not by Cercocebus and Lophocebus, which share the putative primitive state with Macaca. Our results also indicate that Mandrillus and Papio achieve their homoplasic similarities in facial form not through simple extension of the ancestral allometric trajectory but through a combination of an extension of allometry into larger size ranges and a change in direction of allometry away from the ancestral trajectory. Thus, the face of Mandrillus is not simply a hypermorphic version of the face of its sister taxon, Cercocebus, and the face of Papio is not merely a scaled‐up version of the face of its sister taxon, Lophocebus. Lastly, our results show that facial homoplasy is not restricted to adult papionins; it is also manifest in infant and juvenile papionins. This suggests that the homoplasic facial similarities between Mandrillus and Papio are unlikely to be a result of sexual selection.     

Phylogenetic analysis of the African papionin basicranium using 3‐D geometric morphometrics: The need for improved methods to account for allometric effects

The basicranium has been argued to contain a strong phylogenetic signal in previous analyses of primate cranial morphology. Therefore, further study of basicranial morphology may offer new insights into controversial phylogenetic relationships within primate groups. In this study, I apply 3‐D geometric morphometric techniques in a phylogenetic analysis of the African papionin basicranium. The effects of allometry strongly influence African papionin basicranial morphology and, unless these size effects are controlled or eliminated, phylogenetic analyses suggest traditional phylogenetic groupings of small taxa (mangabeys) and large taxa (geladas, mandrills, drills, and baboons). When the effects of allometry are eliminated by excluding size‐correlated principal components (PCs) or by regression analysis with retention of residuals, phylogenetic analyses of African papionin basicranial morphology are incongruent with recent molecular and morphological studies. By contrast, a cladistic analysis of basicranial characters using the narrow allometric coding method suggests the same phylogenetic relationships as recent molecular and morphological studies. These results suggest that important phylogenetic information is contained within the size‐correlated data, and this information is being discarded during the attempt to eliminate the effects of body size. Future 3‐D morphometric studies of phylogeny should focus on the development of new methodologies to adjust for allometric effects, as current techniques appear to be ill‐equipped to deal with the case of a size‐disparate, lower‐level taxonomic group. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Craniomandibular morphology supporting the diphyletic origin of mangabeys and a new genus of the Cercocebus/Mandrillus clade, Procercocebus

Previous studies have noted skeletal and dental differences supporting the diphyletic origin of the mangabeys. Documented postcranial and dental characters are congruent with molecular data and thus support a close relationship between Cercocebus and Mandrillus (mandrills and drills) on the one hand and Lophocebus, Papio (baboons), and Theropithecus (geladas) on the other. Most of these characters, however, are postcranial and difficult to assess in the papionin fossil record because associated material is rare. In order to assess the African papionin fossil record and determine the evolutionary history of this group, cranial characters are critical. Here, a set of craniomandibular morphologies are documented that support the diphyletic origin of the mangabeys and more broadly support the molecular African papionin clades (i.e., Cercocebus/Mandrillus vs. Lophocebus/Papio/Theropithecus). These characters are then used to identify a series of fossil crania from Taung as representative of a new member of the Cercocebus/Mandrillus clade, Procercocebus antiquus. Procercocebus antiquus is closest in morphology to the extant taxon Cercocebus torquatus, and a probable ancestor-descendant relationship between Procercocebus and Cercocebus is suggested. Paleoecological reconstructions also suggest that a predator-prey relationship between African crowned eagles and the Procercocebus-Cercocebus lineage has existed for approximately the last two million years. Implications for Cercocebus biogeography and evolution are discussed.     

Molecular systematics of the old world monkey tribe papionini: analysis of the total available genetic sequences

The phylogenetic relationships among the genera of the tribe Papionini are inferred using a taxonomic congruence approach in which gene trees derived for eight unlinked genetic sequence datasets are compared. Population genetics theory predicts that species relationships will be revealed with greater probability when the topology of gene trees from many unlinked loci are found to be congruent. The theory underlying this approach is described. Monophyly of the mangabeys is not supported by any of the gene trees; instead, they are polyphyletic with Cercocebus found to be the sister taxon to Mandrillus in five gene trees (with no conflicting trees), and Lophocebus found to be closely related to Papio and/or Theropithecus in all trees. Theropithecus and Papio are not strongly supported as sister taxa (present in one or two trees only);Lophocebus and Papio are supported as sister taxa in the majority of trees. A close relationship between Mandrillus and Papio is not supported in any of the trees.The relationships among Papio, Lophocebus, and Theropithecus cannot be resolved by congruence, probably due to the short time interval estimated between their divergences. The mtDNA COII sequences are used to estimate divergence dates within the papionins. The internode between the divergences of these species is estimated to be between 290 ka and 370 ka. Lastly, the evolution of morphological features such as long faces, suborbital facial fossae, and terrestrial skeletal adaptations is discussed.     

Morphometric variation in the papionin muzzle and the biochronology of the South African Plio‐Pleistocene karst cave deposits

Papionin monkeys are widespread, relatively common members of Plio‐Pleistocene faunal assemblages across Africa. For these reasons, papionin taxa have been used as biochronological indicators by which to infer the ages of the South African karst cave deposits. A recent morphometric study of South African fossil papionin muzzle shape concluded that its variation attests to a substantial and greater time depth for these sites than is generally estimated. This inference is significant, because accurate dating of the South African cave sites is critical to our knowledge of hominin evolution and mammalian biogeographic history. We here report the results of a comparative analysis of extant papionin monkeys by which variability of the South African fossil papionins may be assessed. The muzzles of 106 specimens representing six extant papionin genera were digitized and interlandmark distances were calculated. Results demonstrate that the overall amount of morphological variation present within the fossil assemblage fits comfortably within the range exhibited by the extant sample. We also performed a statistical experiment to assess the limitations imposed by small sample sizes, such as typically encountered in the fossil record. Results suggest that 15 specimens are sufficient to accurately represent the population mean for a given phenotype, but small sample sizes are insufficient to permit the accurate estimation of the population standard deviation, variance, and range. The suggestion that the muzzle morphology of fossil papionins attests to a considerable and previously unrecognized temporal depth of the South African karst cave sites is unwarranted. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Chondrocranial development in larval Rana sylvatica (Anura: Ranidae): morphometric analysis of cranial allometry and ontogenetic shape change

This study provides baseline quantitative data on the morphological development of the chondrocranium in a larval anuran. Both linear and geometric morphometric methods are used to quantitatively analyze size-related shape change in a complete developmental series of larvae of the wood frog, Rana sylvatica. The null hypothesis of isometry was rejected in all geometric morphometric and most linear morphometric analyses. Reduced major axis regressions of 11 linear chondrocranial measurements on size indicate a mixture of allometric and isometric scaling. Measurements in the otic and oral regions tend to scale with negative allometry and those associated with the palatoquadrate and muscular process scale with isometry or positive allometry. Geometric morphometric analyses, based on a set of 11 chondrocranial landmarks, include linear regression of relative warp scores and multivariate regression of partial warp scores and uniform components on log centroid size. Body size explains about one-quarter to one-third of the total shape variation found in the sample. Areas of regional shape transformation (e.g., palatoquadrate, otic region, trabecular horns) are identified by thin-plate spline deformation grids and are concordant with linear morphometric results. Thus, the anuran chondrocranium is not a static structure during premetamorphic stages and allometric patterns generally follow scaling predictions for tetrapod cranial development. Potential implications regarding larval functional morphology, cranial development, and chondrocranial evolution in anurans are discussed.     

Molecular phylogeny of Old World monkeys (Cercopithecidae) as inferred from gamma-globin DNA sequences

DNA sequence data of the nuclear-encoded gamma1-gamma2-globin duplication region were used to examine the phylogenetic relationships of 16 cercopithecid (Old World monkey) species representing 12 extant genera. Morphology- and molecular-based hypotheses of Old World monkey branching patterns are generally congruent, except for generic relationships within the subtribe Papionina. The cercopithecids divide into colobines (leaf-eating monkeys) and cercopithecines (cheek-pouched monkeys). The colobines examined by the DNA data divide into an Asian clade (Nasalis, proboscis monkeys; Trachypithecus, langurs) and an African clade (Colobus, colobus monkeys). The cercopithecines divide into tribes Cercopithecini (Erythrocebus, patas monkey; Chlorocebus, green monkeys; Cercopithecus, guenons) and Papionini. Papionins divide into subtribes Macacina (Macaca, macaques) and Papionina (Papio, hamadryas baboons; Mandrillus, drills and mandrills; Theropithecus, gelada baboons; Lophocebus, arboreal mangabeys; Cercocebus, terrestrial mangabeys). In a morphologically based classification, Mandrillus is a subgenus of Papio, whereas Lophocebus is a subgenus of Cercocebus. In contrast, the molecular evidence treats Mandrillus as a subgenus of Cercocebus, and treats both Theropithecus and Lophocebus as subgenera of Papio. Local molecular clock divergence time estimates were used as a yardstick in a "rank equals age" system to propose a reduction in taxonomic rank for most clades within Cercopithecidae.     

Cercopithecine Y-chromosome data provide a test of competing morphological evolutionary hypotheses

We report here the results of the first molecular evolutionary analysis to include members of all 10 extant genera of cercopithecine monkeys. A total of 44 individuals were surveyed for approximately 2.2 kb of the testis-specific protein, Y-chromosome (TSPY). The TSPY sequences were subjected to parsimony analyses in PAUP 4.0, followed by tree comparison tests designed to assess existing morphological hypotheses of cercopithecine evolution. The results of these tests show that the present Y-chromosome dataset unambiguously supports: (1) monophyly of Macaca, (2) polyphyly of the mangabeys (Cercocebus and Lophocebus), (3) paraphyly of Cercopithecus, and (4) inclusion of Allenopithecus and Miopithecus in the tribe Cercopithecini. A number of unexpected Y-chromosome relationships are also discussed, including a pattern suggesting resurrection of the genus Chlorocebus for the guenons currently identified as Erythrocebus patas, Cercopithecus aethiops, and Cercopithecus lhoesti. Relative rate tests reveal significant difference in the TSPY substitution rate across numerous lineages in the tribe Cercopithecini. Because the rate differences follow no obvious phylogenetic pattern, "local" molecular clocks were not employed and divergence dates were not estimated for this tribe. In contrast, similar analysis of the Papionini reveals rate heterogeneity between a single pair of taxonomic groups: Macaca vs. the "African papionins." Divergence dates were therefore calculated for the tribe by calibrating TSPY clocks specific to each of these two clades.     

Postnatal Cranial Development in Papionin Primates: An Alternative Model for Hominin Evolutionary Development

The evolution of hominin growth and life history has long been a subject of intensive research, but it is only recently that paleoanthropologists have considered the ontogenetic basis of human morphological evolution. To date, most human EvoDevo studies have focused on developmental patterns in extant African apes and humans. However, the Old World monkey tribe Papionini, a diverse clade whose members resemble hominins in their ecology and population structure, has been proposed as an alternative model for human craniofacial evolution. This paper reviews prior studies of papionin development and socioecology and presents new analyses of juvenile shape variation and ontogeny to address fundamental questions concerning primate cranial development, including: (1) When are cranial shape differences between species established? (2) How do epigenetic influences modulate early-arising pattern differences? (3) How much do postnatal developmental trajectories vary? (4) What is the impact of developmental variation on adult cranial shape? and, (5) What role do environmental factors play in establishing adult cranial form? Results of this inquiry suggest that species differences in cranial morphology arise during prenatal or earliest postnatal development. This is true even for late-arising features that develop under the influence of epigenetic factors such as mechanical loading. Papionins largely retain a shared, ancestral pattern of ontogenetic shape change, but large size and sexual dimorphism are associated with divergent developmental trajectories, suggesting differences in cranial integration. Developmental simulation studies indicate that postnatal ontogenetic variation has a limited influence on adult cranial morphology, leaving early morphogenesis as the primary determinant of cranial shape. The ability of social factors to influence craniofacial development in Mandrillus suggests a possible role for phentotypic plasticity in the diversification of primate cranial form. The implications of these findings for taxonomic attribution of juvenile fossils, the developmental basis of early hominin characters, and hominin cranial diversity are discussed.     

Skeletal and dental morphology of African papionins: unmasking a cryptic clade

One of the more perplexing problems in primate systematics concerns the phyletic relationships of the large African monkeys--Mandrillus (including drills), Papio, Lophocebus and Cercocebus. For over twenty years, there has been molecular evidence that mangabeys are an unnatural group and that the terrestrial forms--Cercocebus--are the sister taxon of Mandrillus, while the arboreal forms--Lophocebus--are more closely allied with Papio. Nevertheless, most systematists have been reluctant to accept this scheme due to the lack of morphological evidence. In this paper, we undertake a detailed analysis of the scapula, humerus, radius, ulna, pelvis, femur and dentition of papionin primates. We identify a host of features shared by Cercocebus and Mandrillus to the exclusion of Lophocebus and Papio. The polarity of characters is established by examining an outgroup comprised of several species of Macaca. The features shared by Cercocebus and Mandrillus are functionally related to specific feeding and locomotor behaviors that include aggressive manual foraging, the processing of hard-object foods and the climbing of vertical trunks. We hypothesize that the ability to subsist on hard seeds and nuts gleaned from the forest floor is a key adaptation for the Cercocebus-Mandrillus clade.     

Does allometry account for shape variability in <Emphasis Type="Italic">Ephedrus persicae</Emphasis> Froggatt (Hymenoptera: Braconidae: Aphidiinae) parasitic wasps?

We analysed linear measurements on various parts of the body and the configuration of 11 landmarks on the wing in a large sample of Ephedrus persicae that had emerged from 13 aphid host species, to assess whether static allometry (a measure of the scaling relationship between traits in a population of individuals at the same ontogenetic stage) accounts for variation in body shape. The analysed specimens came from several localities in Europe, Asia Minor, Japan and South America, and cover a large portion of the distribution area of E. persicae. We found that allometry accounts for variation in body shape among different biotypes within the E. persicae group. The allometric slopes for head size (HD), petiolus width (PETW), mesoscutum width (MSC), and ovipositor sheath length (OVPL) diverged significantly among biotypes, indicating biotype-specific allometries. The analysis of allometric variation in wing shape showed that the pattern and direction of allometric changes also differed among individuals that had emerged from different hosts. Our results (observed divergences in the directions of allometric slopes of particular morphometric traits and wing shape) suggest that allometric relations within E. persicae are not conserved, so that allometry itself changes, evolving differently in aphid parasitoids that emerge from different hosts.