Cranial morphology and adaptations in Eocene Adapidae. I. Sexual dimorphism in Adapis magnus and Adapis parisiensis

Adapis is one of the best known lemuriform fossil primates. Quantitative analysis of all well-preserved crania of Adapis magnus (n = 8) and Adapis parisiensis (n = 12) together with maxillary and mandibular dentitions preserving canines corroborates Stehlin's hypothesis that Adapis was sexually dimorphic. Males are from 13% to 16% larger than females in cranial length, corresponding to a weight dimorphism estimated at 44% to 56%, and have relatively broader skulls with more prominent sagittal and nuchal crests. Canine dimorphism ranges from 13% to 19%, which is equal to or only slightly greater than that expected as a result of body size dimorphism (i.e., relative canine dimorphism is slight or nonexistent). By comparison with living primates, the observed body size dimorphism in Adapis implies a polygynous breeding system. Cebus apella is a diurnal arboreal living primate with moderate body size dimorphism and slight relative canine dimorphism and one can speculate that Adapis lived in polygynous multimale troops of moderate size like those of C. appella. Adapis extends the geological history of sexual dimorphism and polygyny in primates back to the Eocene. Extant lemuriform primates are generally not dimorphic or polygynous and they clearly do not adequately represent the range of social adaptations present in Eocene primates. The evolutionary lineage from Adapis magnus to Adapis parisiensis exhibits reduction in body size and in relative canine size, and phyletic dwarfing in Adapis is possibly an adaptive response to increasing climatic seasonality and environmental instability in the late Eocene and early Oligocene.     

Using extant patterns of dental variation to identify species in the primate fossil record: a case study of middle Eocene <Emphasis Type="Italic">Omomys</Emphasis> from the Bridger Basin,southwestern Wyoming

Patterns of extant primate dental variation provide important data for interpreting taxonomic boundaries in fossil forms. Here I use dental data from several well-known living primates (as well as data from selected Eocene forms) to evaluate dental variation in Middle Eocene Omomys, the first North American fossil primate identified by paleontologists. Measurements were collected from a sample of 148 omomyid dental specimens recovered from Bridger B localities in the Bridger Basin, Wyoming. Most of these specimens have not previously been described. Nonmetric traits were also scored for this sample. Lower molar coefficients of variation range from 4.01 for M₂ length (n = 80) to 6.73 for M₃ talonid width (n = 57). All of the nonmetric traits scored exhibit less than 100% presence in the overall sample, including traits previously described as representative of Omomys (e.g., P₄ metaconids present in 91%, n = 55; M² pericones present in 80%, n = 15). Dental traits also vary in a set of spatially restricted localities from the same fossil horizon and in a separate, single fossil locality (DMNH 868, P₄ metaconids present in 67%, n = 6). An increasing frequency in several premolar traits across time in these more restricted samples suggests an anagenetic change in Bridger B Omomys. However, this degree of morphological variability is consistent with that seen in extant primate species from single locations. Metric variation in this sample is comparable to that seen in other Eocene primates, such as new data presented here for the omomyid Arapahovius gazini from the Washakie Basin, southern Wyoming. Omomys metric variation is also comparable to that found in several samples of well-known extant primates from single localities (e.g., ring-tailed lemurs and gray–brown mouse lemurs). These metric data also correspond to the patterns of variability described in previously published studies of Omomys carteri. In sum, a single species interpretation (O. carteri) for this new Bridger B Omomys sample from southern Wyoming is affirmed, and this study illustrates the usefulness of dental data from extant primates for evaluating primate fossil samples.     

New primate hind limb elements from the middle Eocene of China

The continued washing, sorting, and identification of middle Eocene (45 Mya) primates from the Shanghuang fissure-fillings (Jiangsu Province, China) have produced additional hind limb elements. All are isolated elements. The strepsirhine hind limb elements include a first metatarsal and a talus, which are appropriate in size and morphology to pertain to Adapoides troglodytes. Adapoides is interpreted as a quadrupedal-climbing (nonleaping) primate with similarities to living lorises and the fossil primate Adapis. The haplorhine hind limb elements are estimated to span a range of adult body sizes from tiny (17 g) to small (200 g). Included among the new sample of haplorhine hind limb specimens is the smallest primate talus reported thus far. These new postcranial specimens expand our understanding of early haplorhine hind limb anatomy and demonstrate additional similarities between Shanghuang eosimiids and other anthropoids.     

Taxonomic variation in the patterns of craniofacial dimorphism in primates

Understanding sexual dimorphism in living primates is important for interpreting the biological and taxonomic significance of variation in the primate fossil record. In the past two decades, there has been an increasing emphasis on the fact that sexual dimorphism varies in both magnitude and pattern among species. Several studies have suggested that distinct patterns of dimorphism may assist in species recognition and perhaps phylogenetic analysis. This study evaluates patterns of craniofacial dimorphism in samples of 82 anthropoid primates. Dimensions of the viscerocranium tend to be more dimorphic than those of the neurocranium and orbits. Principal components analysis of phylogenetically controlled data demonstrates a basic pattern of dimorphism in overall skull proportions, and a distinction between length and breadth measurements. For any given species there can be substantial variation in the magnitude of dimorphism among dimensions, and different species can show substantially different patterns of dimorphism within and between regions of the skull and jaws. Patterns of dimorphism are clearly associated with phylogeny. Pattern similarity is not dependent on the overall magnitude of craniofacial dimorphism, or body mass dimorphism. Among all anthropoids, there are few combinations of characters that consistently show greater or lesser degrees of dimorphism. Such "stability" of patterns increases within genera. Patterns of dimorphism are likely to be useful for interpreting the taxonomic significance of variation in the fossil record. However, phylogenetic propinquity alone is not reason to use an extant species as a model for variation in an extinct species. Rather, care must be taken to identify stable patterns of dimorphism within a group of closely related extant species.     

Sexual differences in the anterior dentition in African primates

A study was made of ranges of variation of the anterior dentition of various African nonhuman primates. Comparisons of the dentitions were made between different species and sex differences within each species were determined. Among the nonhuman primate groups studied were: Gorilla gorilla gorilla, Pan troglodytes, Pan paniscus, Cercopithecus nictitans, Cercocebus albigena and Colobus badius. In monkeys the canine teeth of the males are considerably larger than those of the females. There are also considerable differences in size in the rest of the anterior dentition. In apes, and specifically only gorillas, distinct sex differences are only found in the maxillary canines. In the chimpanzees, sex differences in the dentition are much smaller and there is considerable overlap in the ranges of variation. There are no fundamental differences in the size of the rest of the anterior dentition in the apes. The present study shows that differences due to sex in the anterior dentition, excluding the canine, are not as great as has been considered. If we consider the fossil record of man, whose morphological complex includes a much reduced canine, the probability will be that sex differences in the rest of the dentition will be negligible. Given the fragmentary nature of the fossil record, it is, therefore, highly unlikely that the determination of the sex of any fossil hominid specimen can be accurately made based solely on the evidence of its dentition.     

Metric variation and sexual dimorphism in the dentition of Ouranopithecus macedoniensis

The fossil sample attributed to the late Miocene hominoid taxon Ouranopithecus macedoniensis is characterized by a high degree of dental metric variation. As a result, some researchers support a multiple-species taxonomy for this sample. Other researchers do not think that the sample variation is too great to be accommodated within one species. This study examines variation and sexual dimorphism in mandibular canine and postcanine dental metrics of an Ouranopithecus sample. Bootstrapping (resampling with replacement) of extant hominoid dental metric data is performed to test the hypothesis that the coefficients of variation (CV) and the indices of sexual dimorphism (ISD) of the fossil sample are not significantly different from those of modern great apes. Variation and sexual dimorphism in Ouranopithecus M(1) dimensions were statistically different from those of all extant ape samples; however, most of the dental metrics of Ouranopithecus were neither more variable nor more sexually dimorphic than those of Gorilla and Pongo. Similarly high levels of mandibular molar variation are known to characterize other fossil hominoid species. The Ouranopithecus specimens are morphologically homogeneous and it is probable that all but one specimen included in this study are from a single population. It is unlikely that the sample includes specimens of two sympatric large-bodied hominoid species. For these reasons, a single-species hypothesis is not rejected for the Ouranopithecus macedoniensis material. Correlations between mandibular first molar tooth size dimorphism and body size dimorphism indicate that O. macedoniensis and other extinct hominoids were more sexually size dimorphic than any living great apes, which suggests that social behaviors and life history profiles of these species may have been different from those of living species.     

Mandibular shape variation in mainland and insular hylobatids

Although hylobatids are the most speciose of the living apes, their morphological interspecies and intraspecies variation remains poorly understood. Here, we assess mandibular shape variation in two species of Hylobates, white-handed (Hylobates lar) and black-handed (Hylobates agilis) gibbons. Using 71 three-dimensional landmarks to quantify mandibular shape, interspecies and intraspecies variation and geographic patterns of mandibular shape are examined in a mixed sex sample of adult H. lar and H. agilis through generalized Procrustes analysis, Procrustes analysis of variance, and principal components analysis. We find that relative to H. agilis, H. lar exhibits a higher amount of variation in mandibular shape. Both species demonstrate similar allometric patterns in mandibular shape. We also highlight a geographic pattern in mandibular shape variation. Compared to mainland hylobatids, insular hylobatids have relatively lower, more posteriorly oriented, and anteroposteriorly wider mandibular condyles, with an increased distance between the condyles and the coronoid processes. This geographic pattern could reflect differences in functional demands on the mandible during mastication and/or could be driven by factors often associated with evolutionary pressures of island populations relative to mainland populations. The findings of this study highlight how little is known about Hylobates morphological variation and how important this is for using Hylobates to help interpret the primate fossil record. Understanding interspecific and intraspecific variation in extant primates is vital to interpreting variation in the primate fossil record.     

Tail length variation inMacaca fascicularis andM. mulatta

Based on collectors' measurements of 1,040 specimens, variation of relative tail length [RTL = (tail length/head and body length) × 100] has been studied inMacaca fascicularis andM. mulatta, two closely related species that replace one another in tropical Asia and subtropical Asia, respectively. RTL usually is greater than 90 inM. fascicularis and usually is less than 60 inM. mulatta; intermediate values occur in only 3.5% of specimens studied. Within each species, RTL is approximately equal in females and males. From infancy to adulthood, RTL tends to decline in both species. InM. fascicularis, RTL generally decreases with increasing latitude; inM. mulatta, RTL is approximately constant latitudinally. Where the geographic ranges of these two species meet at ca. 15°N in the Indochinese Peninsula, a few specimens have been collected in which RTL and dorsal pelage color are intermediate between those inM. fascicularis andM. mulatta. The observed pattern of variation suggests that the ranges ofM. fascicularis andM. mulatta formerly were separated by a zoogeographic barrier — perhaps during a Pleistocene glacial interval. After disappearance of the postulated barrier, the ranges of these two species apparently became contiguous and limited hybridization has occurred.     

Evaluation of the fossil fish‐specific diversity in a chadian continental assemblage: Exploration of morphological continuous variation in Synodontis (Ostariophysi,Siluriformes)

In the fossil record, the quantification of continuous morphological variation has become a central issue when dealing with species identification and speciation. In this context, fossil taxa with living representatives hold great promise, because of the potential to characterise patterns of intraspecific morphological variation in extant species prior to any interpretation in the fossil record. The vast majority of catfish families fulfil this prerequisite, as most of them are represented by extant genera. However, although they constitute a major fish group in terms of distribution, and ecological and taxonomic diversity, the quantitative study of their past morphological variation has been neglected, as fossil specimens are generally identified based on the scarcest remains, that is, complete neurocrania that bear discrete characters. Consequently, a part of freshwater catfish history is unprospected and unknown. In this study, we explored the morphological continuous variation of the humeral plate shape in Synodontis catfishes using Elliptic Fourier Analysis (EFA), and compared extant members and fossil counterparts. We analysed 153 extant specimens of 11 Synodontis species present in the Chad basin, in addition to 23 fossil specimens from the Chadian fossiliferous area of Toros Menalla which is dated around 7 Ma. This highly speciose genus, which is one of the most diversified in Africa, exhibits a rich fossil record with several hundred remains mostly identified as Synodontis sp. The analysis of the outline of the humeral plate reveals that some living morphological types were already represented in the Chad Basin 7 My ago, and allows for the discovery of extinct species. Beside illuminating the complex Neogene evolutionary history of Synodontis, these results underline the interest in the ability of isolated remains to reconstruct a past dynamic history and to validate the relevance of EFA as a tool to explore specific diversity through time. J. Morphol. 277:1486–1496, 2016. © 2016 Wiley Periodicals, Inc.     

Morphological variation in adult chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d'Ivoire

Twenty five adult chimpanzee skeletons (Pan troglodytes verus) of known age and sex (15 females, 10 males) from a long‐term study site in Taï National Park, Cote d'Ivoire present new data on variation. These skeletons provide a rare opportunity to measure the cranium and postcranium from the same individuals. We compare measurements and indices of the Taï sample with those of relatively complete Pan troglodytes schweinfurthii skeletons from Gombe National Park, Tanzania. Measurements of Pan paniscus are included as an outside comparison. The Taï and Gombe samples are analyzed by sex; combined sex samples are compared between the two groups, and the two sexes to each other. Taï females and males do not differ in most long bone lengths or in pelvic dimensions, but do differ significantly in cranial capacity, facial measurements, clavicle length, scapular breadth, and femur length. Gombe females and males differ significantly in some facial measurements and in scapular breadth. In combined sex samples, Taï individuals have lower cranial capacity, longer palate and mandible, and greater dimensions in the trunk and limb lengths. Taï females account for most of the variation; males differ from each other only in greater length of humerus and femur. The Taï skeletons provide new data for assessing individual variation and sexual dimorphism within and between populations and species. The combination of cranial and postcranial data provides a clearer picture of chimpanzee intraspecific and interspecific variation than can be gained from either data set alone. Am J Phys Anthropol, 2008. © 2007 Wiley‐Liss, Inc.     

Metric variation and species recognition in the fossil record

One of the first and most important tasks of the paleontologist is classifying specimens into species. Species recognition commonly involves sorting specimens on the basis of qualitative and quantitative similarities and differences. Often, however, variation in simple metric characters like tooth size or jaw length plays an important role in debates about whether a sample comprises a single species or more than one morphologically similar species. For example, Simpson, Roe, and Lewontin 1 suggested that a fossil sample showing a coefficient of variation greater than 10.0 was likely to comprise more than one species. Well‐known controversies over species recognition in which metric sample variation has been important have simmered for years, focusing on hominids, hominoids, and other extinct primates. Some of these have been resolved; others have not. For example, Pilbeam and Zwell 2 convincingly demonstrated multiple species among South African hominids by showing that metric tooth size variation was too great to be reasonably interpreted as sexual dimorphism. But metric variation continues to play a role in debates about whether Australopithecus afarensis 3 , 4 and Homo habilis 5 - 9 each comprise a single species or two or more separate species. Similarly, there has been steady debate about the number of species present in African Proconsul. Some favor an interpretation of a single extremely dimorphic species, 10 - 12 while others favor an interpretation of two or more species. 13     

Baculum length and copulatory behavior in primates

The length of the baculum (os penis) was measured in 74 adult males representing 46 primate species. These data, and a review of previously published measurements, indicate that variation in baculum length among primates is related to taxonomic and behavioral differences. Thus, many New World monkeys have shorter bacula, relative to body weight, than Old World monkeys. The baculum is shorter in colobine monkeys than in cercopithecines. Among the great apes, reduction of the baculum is more pronounced in Pan and Gorilla than in Pongo. Very long bacula are found in some nocturnal prosimians (eg, Lorisidae) and also in Macaca arctoides. A comparison of baculum length relative to body weight was made in 34 species for which detailed information on copulatory behavi or was available. The presence of an elongated baculum was shown to correlate with copulatory patterns involving prolonged intromission and/or the maintenance of intromission during the postejaculatory interval (eg, Galago crassicaudatus, Loris tardigradus, M, arctoides). The evolutionary significance of these observations is discussed and it is suggeted that similar copulatory patterns may occur in species with elongated bacula (eg, Daubentonia, Perodicticus) for which behavioral data are lacking at present. The same hypothesis also applies to an extinct adapid primate which possessed a very large baculum.     

A solution to the human paradox: fundamental ontogenies and heterochronies

Solving the human paradox means explaining how a genetic difference of a mere 1% can be consistent with 5 million years of anatomical transformation from great apes to present-dayHomo sapiens. The solution proposed here is that of the internal history of ontogenetic change. A concept of “fundamental ontogeny” is developed and deduced from comparison between living and fossil primates. The fossil human lineage can be summarized into five fundamental ontogenies corresponding to successive skull plans (bauplans) resulting from five major phases of craniofacial contraction: prosimians (adapiforms), monkey apes (propliopithecidae), great apes (dryopithecidae), australopithecines andHomo. The morphological areas defined by these skull plans include more-or-less numerous species. This concept leads to renewed debate about (i) the relationship between speciation and bauplans, and (ii) the mechanisms involved in the successive steps of cranio-facial contraction and the correlated morphological changes. It is suggested that, from great apes to modern man, numerous heterochronies (hypermorphosis, hypomorphosis and post-displacements) have occurred during ontogeny, allowing the acquisition of permanent bipedalism inAustralopithecus andHomo, the increased cranial capacity of primitive forms ofHomo, and the disappearance of simian characters associated with renewed increase in cranial capacity inH. sapiens.     

Sexual Dimorphism in the Pelvis of <Emphasis Type="Italic">Microcebus</Emphasis>

Pelvic sexual dimorphism occurs in many anthropoid species and is often attributed to obstetric selection on female pelvic morphology. Few studies of pelvic dimorphism have included strepsirrhine taxa, which typically have relatively smaller infants than those of anthropoids. Because smaller female primates give birth to relatively larger infants, it is possible that the pelves of Microcebus, the smallest extant primate genus, will show some evidence of selection on obstetric adequacy. A comparison of adult female and neonatal body masses indicates that individual neonatal Microcebus are relatively large compared to adult female body mass, even though members of the taxon frequently produce twins. I examined variation in the bony pelvis within a sample of Microcebus. I measured specimens from a single locality, which probably represent 1 population. I measured 8 pelvic and 3 femoral variables to investigate skeletal size and pelvic size and shape dimorphism. Females significantly exceed males in absolute values of sacral width, pelvic height, pubic length, and distances from the pubic symphysis to the ischial tuberosity and points on the sacrum. Measurements of the femur are not significantly greater in females, suggesting that the pelvic differences are not due to skeletal size dimorphism. Significant pelvic shape or ratio differences, calculated via the geometric mean of 5 variables as the denominator, included greater relative pubic length and sacral width in females. Hence selection for obstetric adequacy may occur in the extremely small-bodied Microcebus.     

The role of character displacement in the molarization of hominin mandibular premolars

Closely related species are likely to experience resource competition in areas where their ranges overlap. Fossil evidence suggests that hominins in East Africa c. 2–1.5 million years ago may have lived synchronically and sympatrically, and that competition may have contributed to the different tooth sizes observed in Homo and Paranthropus. To assess the likelihood that these taxa overlapped, we applied a character displacement model to the postcanine tooth size of fossil hominins and validated this model in populations of living primates. Mandibular fourth premolar (P₄) crown size was measured from fossil taxa and from living primate species where dietary overlap is established. Dimensions of the P₄ crown were fitted to a character matrix and described as the response variables of a generalized linear model that took taxon and location as input variables. The model recovered significant divergence in samples of closely related, living primates. When applied to fossil hominins the same model detected strong indications of character displacement between early Homo and Paranthropus (P = 0.002) on the basis of their P₄ crown size. Our study is an example of how ecologically informed morphologies measured in appropriate extant referents can provide a comparative context for assessing community and ecological evolution in the fossil record.     

Genetic and phenotypic variation for flight ability in the cactophilicDrosophila species,D. aldrichi andD. buzzatii

The flight ability ofDrosophila aldrichi (Patterson & Crow) andD. buzzatii (Patterson & Wheeler) using tethered flights, was measured with respect to age-related changes, genetic variation and adult body size variation induced by rearing at different larval densities.Drosophila buzzatii flew for much longer thanD. aldrichi, especially females, but age-related changes in flight duration were significant only forD. aldrichi. Effects of body size on flight ability were significant inD. buzzatii, but not inD. aldrichi. InD. buzzatii, there was a significant genotype-environment interaction (larval density × line) for flight duration, with short and average flight duration isofemale lines showing longer flights, but a long flight duration line shorter flights as body size decreased (i.e., as larval density increased). Heritability estimates for flight duration were similar in the two species, but flight duration showed no significant genetic correlations with developmental time, body size or wing dimensions (except for one wing dimension inD. buzzatii). Although not significantly different between the species, heritabilities for life-history traits (adult size and developmental time) showed contrasting patterns — with higher heritability for body size (body weight and thorax length) inD. buzzatii, and higher for developmental time inD. aldrichi. In agreement with limited previous field evidence,D. buzzatii is better adapted for colonization than isD. aldrichi.     

Multivariate discriminant analysis of dental data bearing on early hominid affinities

The dimensions of hominoid dentitions are compared by multiple discriminant analysis. By this technique the fossil taxa are compared with living pongids and modern man in a multivariate framework. This enables the classification of the fossils to be made consistent with that of the living forms. H. africanus and H. erectus generally form the most compact grouping within the hominids, thus suporting the argument that these two species can indeed be lumped into a single genus. The degree of separation between H. africanus and Paranthropus is found to be at least as great as that between the genera of modern apes. Gigantopithecus sorts with the hominids rather than with the pongids and seems to be most closely related to Paranthropus.     

Diversity of lobate phytoliths in grass leaves from the Sahel region, West Tropical Africa: Tribe Paniceae

Phytoliths are microscopic amorphous silicon dioxide (SiO2.H2O) particles occurring in leaves, internodes, glumes and inflorescence within all members of the grass family Poaceae. Phytoliths of grasses are of particular interest, as they possess morphological features which have encouraged many investigators to identify these plants from which fossil phytoliths might have originated. The present study is a step towards preparing a systematic inventory of grass phytoliths in western tropical Africa. Morphology and dimensions of phytoliths from 66 species belonging to the tribe Paniceae have been studied. Four shape categories of lobate phytoliths have been determined in leaf blade spodograms: bilobate, nodular bilobate, polylobate, quadra-lobate. Bilobate shaped phytoliths are frequently represented in all genera of Paniceae. 13 groups of lobate phytoliths have been distinguished based on significant morphological criteria like shape of outer margins, shape of the shank and number of lobes. A size category system of lobate phytolith dimensions (length, width; length and width of shanks) has been developed by the analysis of average, minimum and maximum values of these dimensions. Application of the size category system results on classifying the major groups into 25 subgroups. The study proves that size and shape can be used to assign some of the lobate phytoliths to their respective genera. Some rarely produced lobate shapes like nodular bilobate and polylobate types could be used together on assemblage basis as markers for definite genera in the tribe Paniceae, e.g. Brachiaria, Panicum, Pennisetum and Setaria. Also, bilobate phytoliths with concave margins have been recorded in five species. Bilobate phytoliths with flattened and convex margins and quadra-lobate shapes are produced by almost all species which therefore resulted in an inconsistent and indefinite relationship with the taxa that produce them. The study shows a correlation between width dimensions of bilobate shapes and their shanks. Greater width dimensions usually connected to thick shanks while short ones are attached to thin shanks. A spectrum on percentages of species producing each type of lobate phytolith has been designed. It is recommended that such spectrum should be carried out for all tribes of Poaceae on phyto-geographical basis which might eliminate the effect of redundancy and multiplicity on the classification of grass phytoliths.     

Variation in the Nutritional Value of Primate Foods: Among Trees, Time Periods, and Areas

The study of nutritional ecology has proven to be useful for understanding many aspects of primate behavior and ecology and is a valuable tool in primate conservation. However, to date this approach has had limited application since chemical analyses of food items is very time-consuming and collections of perishable food material are often made in remote field locations. Such logistic difficulties have led to plant material being collected in a variety of fashions, and it is not known how variation in collection method might influence our understanding of the chemical basis of dietary selection. A standardization of collection methods is greatly needed to allow for direct comparison among studies. To develop an appropriate standardized method and to evaluate past research, it is necessary to understand along what dimensions plant chemistry varies. We evaluated variation in nutritional value—protein, fiber, digestibility, alkaloids, saponins, cyanogenic glycocides, and minerals—of leaf material from species eaten by red colobus (Piliocolobus tephrosceles) and black-and-white colobus (Colobus guereza) of Kibale National Park, Uganda. We consider variation at 3-levels: among trees, time periods, and areas. While there was considerable variation among species with respect to protein, digestibility, and saponins, there was also variation among individuals of the same species; in fact, individuals may vary by as much as 20%. The average coefficients of variation (CV) among individuals of the same species are 13.4 for protein, 12 for digestibility, and 43 for saponins, while the average CV among species are 35, 31.3, and 82.4, respectively. No species showed a variable response with respect to testing for the presence or absence of cyanogenic glycocides, while 2 of 11 species tested for alkaloids showed a variable response. Over 2 years there was evidence of variation among time periods in the chemical composition of the same food items. The protein-to-fiber ratio of mature leaves of the same species collected from 4 sites separated by 12 km within Kibale was also variable and in some cases the variation among sites was greater than the differences among species. For example, while Funtumia latifolia had little variation in protein-to-fiber ratio at 3 sites (0.44 at all sites), the remaining site was 28% greater. Because temporal variation is less than variation among individuals, it is likely more important to sample from multiple trees at a single point in time than to sample across time. However, the most accurate assessment of nutrient intake is obtained by collecting plant material from the specific trees selected for consumption.     

Differences in Growth Generate the Diverse Palate Shapes of New World Leaf-Nosed Bats (Order Chiroptera,Family Phyllostomidae)

New World leaf-nosed bats (Family Phyllostomidae) display incredible craniofacial diversity that is associated with their broad range of dietary preferences. The short and broad palates of highly frugivorous bats are functionally linked to high bite forces, and the long and narrow palates of nectarivorous bats to flower feeding. Although the functional correlates and evolutionary history of shape variation in phyllostomid palates are beginning to be understood, the specific developmental processes that govern palate diversification remain unknown. To begin to resolve this issue, this study quantified palate morphology in seven phyllostomid species from a range of developmental stages and in adults. This sample includes species with short and broad, long and narrow, and intermediate palate shapes, and thereby covers the range of palate shapes displayed by phyllostomids. Results indicate that while initial palate shape (i.e., width vs. length) varies among species, the pattern of this variation does not match that observed in adults. In contrast, the relative growth of palate width and length in developing phyllostomids and the ratio of these axes in adults are significantly correlated. These and other results suggest that evolutionary alterations in patterns of palate growth have governed the diversification of palate shapes in adult phyllostomids. This implies that the diverse palate shapes of phyllostomids are the result of relatively subtle evolutionary changes in later rather than earlier development events.