Problems of body-weight estimation in fossil primates

Body-weight estimates of fossil primates are commonly used to infer many important aspects of primate paleobiology, including diet, ecology, and relative encephalization. It is important to examine carefully the methodologies and problems associated with such estimates and the degree to which one can have confidence in them. New regression equations for predicting body weight in fossil primates are given which provide body-weight estimates for most nonhominid primate species in the fossil record. The consequences of using different subgroups (evolutionary “grades”) of primate species to estimate fossil-primate body weights are explored and the implications of these results for interpreting the primate fossil record are discussed. All species (fossil and extant) were separated into the following “grades”: prosimian grade, monkey grade, ape grade, anthropoid grade, and all-primates grade. Regression equations relating lower molar size to body weight for each of these grades were then calculated. In addition, a female-anthropoid grade regression was also calculated for predicting body weight infernales of extinct, sexually dimorphic anthropoid species. These equations were then used to generate the fossil-primate body weights. In many instances, the predicted fossil-primate body weights differ substantially from previous estimates.     

New estimates of early hominid body size

Body weights for 12 early hominid specimens are estimated based on an analysis of four variables shown to have high correlation with body size in living Old World primates. Average size estimates of around 36 kg are suggested for gracile early hominids and around 59 kg for robust early hominids. Size variation is considerably more pronounced in the robust group than in the gracile group, suggesting substantially greater sexual dimorphism in the former.     

Estimation of body weights from craniometric variables in baboons

Body weights of adult baboons (genera Papio, Mandrillus, and Theropithecus) were gathered from notes of collectors and museum records. However, these data were insufficient to establish mean body weights for all baboon groups. Thus, log cube roots of mean body weights were regressed as functions of the logs of several cranial and dental variables. The resulting least squares regression coefficients were used to estimate weights for 503 adult baboons from cranial measurements. The ability of the various regression functions to assess baboon body weight was determined by comparing reported and estimated mean and individual body weights. The best estimator of baboon body weights was the function derived from the factor scores of a principal components analysis of seven craniometric variables regressed on body weight. However, each of these craniometric variables singly was nearly as precise an estimator of body weight as the multivariate combination of all seven. Other measurements such as dental dimensions and foramen magnum area estimated weight less accurately. Body weight estimates derived from the regression analyses coupled with museum and literature records allowed an assessment of size relationships among all baboon groups.     

Allometric scaling in the dentition of primates and prediction of body weight from tooth size in fossils

Tooth size varies exponentially with body weight in primates. Logarithmic transformation of tooth crown area and body weight yields a linear model of slope 0.67 as an isometric (geometric) baseline for study of dental allometry. This model is compared with that predicted by metabolic scaling (slope = 0.75). Tarsius and other insectivores have larger teeth for their body size than generalized primates do, and they are not included in this analysis. Among generalized primates, tooth size is highly correlated with body size. Correlations of upper and lower cheek teeth with body size range from 0.90–0.97, depending on tooth position. Central cheek teeth (P and M) have allometric coefficients ranging from 0.57–0.65, falling well below geometric scaling. Anterior and posterior cheek teeth scale at or above metabolic scaling. Considered individually or as a group, upper cheek teeth scale allometrically with lower coefficients than corresponding lower cheek teeth; the reverse is true for incisors. The sum of crown areas for all upper cheek teeth scales significantly below geometric scaling, while the sum of crown areas for all lower cheek teeth approximates geometric scaling. Tooth size can be used to predict the body weight of generalized fossil primates. This is illustrated for Aegyptopithecus and other Eocene, Oligocene, and Miocene primates. Regressions based on tooth size in generalized primates yield reasonable estimates of body weight, but much remains to be learned about tooth size and body size scaling in more restricted systematic groups and dietary guilds.     

ORIGINAL ARTICLE: Eyes wide shut: can hypometabolism really explain the primate colonization of Madagascar?

Aim We investigate the hypothesis that it was the capacity for torpor or hibernation that enabled the lemuriforms (and possibly other mammal groups, e.g. tenrecs and rodents) to invade Madagascar by means of over‐water dispersal. Location Madagascar, East Africa and the Mozambique Channel. Methods We consider the body weights and life‐style features of living primate taxa that employ heterothermy. Using this information as a standard for comparison, we summarize the available information on the Palaeogene strepsirrhine radiation (i.e. members of the infraorder Adapiformes, the extinct sister‐taxon to the lemuriforms, as well as putative stem lemuriforms), particularly with respect to possible trends in body weight among early, middle and late Eocene adapiforms. We discuss Eocene climatic conditions in the northern hemisphere and Africa, and assess the likelihood of adaptations for heterothermy in adapiforms. Finally, we estimate the body weights of the common ancestors to the living Lemuriformes and Lemuroidea using the method of phylogenetically independent contrasts. Results The mean body weights estimated for the early, middle and late Eocene strepsirrhine faunas remain at approximately 2 kg, outside of the range of living primates using heterothermy. The adapiforms’ appearance coincided with the ‘initial Eocene thermal maximum’, an unusually hot period of global warming, and their demise corresponded with a major cooling event coincident with the appearance of ice sheets on Antarctica. They show no evidence of having evolved adaptations that allowed them to withstand climatic deterioration. The body weights of the ancestral lemuriform and ancestral lemuroid are of a similar order to the mean body weight estimated for the adapiforms, i.e. approximately 1.8 and 2.1 kg, respectively. Main conclusions The available evidence argues against the widespread use of heterothermy by adapiforms. The adapiform–lemuriform divergence may have been the result of the lemuriforms adapting to the drier and more seasonal environments that characterized the African Eocene, and this suite of adaptations may have included the use of heterothermy, but there is as yet no substantial evidence to confirm the presence of either group on the continent prior to c. 40 Ma. The estimated body weights of the common lemuriform and lemuroid ancestors are well outside the size range of living mammals that employ heterothermy. We conclude that the hypothesis that it was the ability to use heterothermy that enabled the strepsirrhines, and not the haplorhines, to invade Madagascar, is unlikely. Alternative explanations for this anomaly should be sought.     

L’orang-outan fossile de Hoà Binh (Viêt-nam) : poids corporel et hypothèse locomotrice

The orang-utan of Hoà Binh (Vietnam): body weight and locomotor hypothesis. The discovery of the first complete skeleton of an adult orang-utan (Pongo pygmaeus), probably a female, in the Hoà Binh province, Vietnam, brings original data about the evolution of these primates. We have predicted the body weight of this orang-utan by using two parameters : dental (crown area of cheek teeth) and skeletal (lengths of femur, radius and tibia). The dental dimensions give plausible predicted estimates of body weights that range between 70.9–112.4 kg for M¹ (with a mean of 89.3 kg) and 67.9–98.1 kg for M₁ (with a mean of 81.6 kg). From the observations of Sugardjito and van Hoof [21] on modern orang-utans, heavy body weights have some influence both on the employment of certain types of locomotion and on travelling heights. To cite this article : A.-M. Bacon, V.T. Long, C. R. Palevol 2 (2003).     

Limb bone proportions and body mass of the cave bear (Ursus spelaeus)

The European cave bear evolved during the Middle Pleistocene and adapted to mountain environments. Earlier workers have described the cave bear as a robust bear. In this study the cave bears limb bone morphology is compared to the limb bone morphology of extant bears. Body mass estimates for the cave bear are made both based on different limb bone characters and based on dental and cranial characters. The shafts are wider in the cave bear limb bones than in the extant bear limb bones, and consequently the shaft widths give higher weight estimates to the cave bear than the other dimensions. The widened shafts are suggested to be a special adaptation (of presently unknown significance) rather than an indicator of an increased body mass.     

Allometric patterns of cranial bone thickness in fossil hominids

The interspecific allometry of five measures of total cranial bone thickness is examined in 10 extant catarrhine genera and two fossil hominid samples representing A. africanus and Asian H. erectus. Analysis of the modern sample shows that most interspecific variation in vault thickness can be accounted for by variation in body size. Correlation values are moderate to high (r = 0.75–0.98), and all variables exhibit positive allometry. The bone thickness:body mass relationship of modern humans broadly conforms with that of other primates. However, in the distribution of relative thickness throughout the skull, H. sapiens is distinguished by relative thickening of the parietal and extreme relative thinning of the temporal squama. The bone thickness:body mass relationship in the two early hominid species is examined using published mean body weight estimates generated from post-cranial predictor variables. A. africanus exhibits great similarity to modern humans in its relation to the catarrhine regression data and in the distribution of relative thickness throughout the skull. H. erectus also shows a modern human-like pattern in the distribution of its relative thickness; however, its bone thickness:body mass relationship is dissimilar to that displayed by all other taxa, including the other hominid species. On the basis of these results, it is suggested that the published body weight estimate assigned to H. erectus greatly underestimates actual mean body size for Asian members of this species. © 1996 Wiley-Liss, Inc.     

The use of cranial variables for the estimation of body mass in fossil hominins

Estimating body mass/size/weight remains a crucial precursor to the evaluation of relative brain size and to achieving an understanding of brain evolution in fossil species. Despite the obvious close association between the metrics of postcranial elements and body mass a number of factors combine to reduce their utility. This study examines the feasibility of cranial variables for predicting body mass. The use of traditional regression procedures, independent contrasts analysis, and variance partitioning all support the hypothesis that cranial variables are correlated with body mass even when taking phylogeny into account, with r values typically ranging between 0.52 and 0.98. Body mass estimates derived for fossil hominins using cranial variables are similar to those obtained from previous studies using either cranial or postcranial elements. In particular, upper facial breadth and orbital height display strong predictive capability. Average body masses derived from Least Squares Regression (LSR) equations were used to calculate estimates of body mass for three hominin species. This resulted in estimates of between 30 kg and 47 kg for Australopithecus africanus, 48 kg and 52 kg for Paranthropus robustus, and 75 kg for Homo neanderthalensis. It is proposed that regression equations derived for the order primates are used to estimate body mass for archaic hominins, while hominoid based equations are most suited for Homo.     

Genetic variance and covariance patterns for body weight and energy balance characters in an advanced intercross population of mice

We estimated heritabilities and genetic correlations for a suite of 15 characters in five functional groups in an advanced intercross population of over 2000 mice derived from a cross of inbred lines selected for high and low heat loss. Heritabilities averaged 0.56 for three body weights, 0.23 for two energy balance characters, 0.48 for three bone characters, 0.35 for four measures of adiposity, and 0.27 for three organ weights, all of which were generally consistent in magnitude with estimates derived in previous studies. Genetic correlations varied from -0.65 to +0.98, and were higher within these functional groups than between groups. These correlations generally conformed to a priori expectations, being positive in sign for energy expenditure and consumption (+0.24) and negative in sign for energy expenditure and adiposity (-0.17). The genetic correlations of adiposity with body weight at 3, 6, and 12 weeks of age (-0.29, -0.22, -0.26) all were negative in sign but not statistically significant. The independence of body weight and adiposity suggests that this advanced intercross population is ideal for a comprehensive discovery of genes controlling regulation of mammalian adiposity that are distinct from those for body weight.     

Diets of fossil primates from the Fayum Depression of Egypt: a quantitative analysis of molar shearing

Over the last 90 years, Eocene and Oligocene aged sediments in the Fayum Depression of Egypt have yielded at least 17 genera of fossil primates. However, of this diverse sample the diets of only four early Oligocene anthropoid genera have been previously studied using quantitative methods. Here we present dietary assessments for 11 additional Fayum primate genera based on the analysis of body mass and molar shearing crest development. These studies reveal that all late Eocene Fayum anthropoids were probably frugivorous despite marked subfamilial differences in dental morphology. By contrast, late Eocene Fayum prosimians demonstrated remarkable dietary diversity, including specialized insectivory (Anchomomys), generalized frugivory (Plesiopithecus), frugivory+insectivory (Wadilemur), and strict folivory (Aframonius). This evidence that sympatric prosimians and early anthropoids jointly occupied frugivorous niches during the late Eocene reinforces the hypothesis that changes in diet did not form the primary ecological impetus for the origin of the Anthropoidea. Early Oligocene Fayum localities differ from late Eocene Fayum localities in lacking large-bodied frugivorous and folivorous prosimians, and may document the first appearance of primate communities with trophic structures like those of extant primate communities in continental Africa. A similar change in primate community structure during the Eocene-Oligocene transition is not evident in the Asian fossil record. Putative large anthropoids from the Eocene of Asia, such as Amphipithecus mogaungensis, Pondaungia cotteri, and Siamopithecus eocaenus, share with early Oligocene Fayum anthropoids derived features of molar anatomy related to an emphasis on crushing and grinding during mastication. However, these dental specializations are not seen in late Eocene Fayum anthropoids that are broadly ancestral to the later-occurring anthropoids of the Fayum's upper sequence. This lack of resemblance to undisputed Eocene African anthropoids suggests that the "progressive" anthropoid-like dental features of some large-bodied Eocene Asian primates may be the result of dietary convergence rather than close phyletic affinity with the Anthropoidea.     

Dental development in Megaladapis edwardsi (Primates, Lemuriformes): implications for understanding life history variation in subfossil lemurs

Teeth grow incrementally and preserve within them a record of that incremental growth in the form of microscopic growth lines. Studying dental development in extinct and extant primates, and its relationship to adult brain and body size as well as other life history and ecological parameters (e.g., diet, somatic growth rates, gestation length, age at weaning), holds the potential to yield unparalleled insights into the life history profiles of fossil primates. Here, we address the absolute pace of dental development in Megaladapis edwardsi, a giant extinct lemur of Madagascar. By examining the microstructure of the first and developing second molars in a juvenile individual, we establish a chronology of molar crown development for this specimen (M1 CFT = 1.04 years; M2 CFT = 1.42 years) and determine its age at death (1.39 years). Microstructural data on prenatal M1 crown formation time allow us to calculate a minimum gestation length of 0.54 years for this species. Postnatal crown and root formation data allow us to estimate its age at M1 emergence (approximately 0.9 years) and to establish a minimum age for M2 emergence (>1.39 years). Finally, using reconstructions or estimates (drawn elsewhere) of adult body mass, brain size, and diet in Megaladapis, as well as the eruption sequence of its permanent teeth, we explore the efficacy of these variables in predicting the absolute pace of dental development in this fossil species. We test competing explanations of variation in crown formation timing across the order Primates. Brain size is the best single predictor of crown formation time in primates, but other variables help to explain the variation.     

Early hominid body weight and encephalization

The body weight of the Plio-Pleistocene hominids of Africa is estimated by predicting equations derived from the Terry Collection of human skeletons with known body weights. About 50% of the variance in body weight can be accounted for by vertebral and femoral size. Predicted early hominid weights range from 27.6 kg (61 lb) to 54.3 kg (119 lb). The average weight for Australopithecus is 43.2 kg (95 lb) and for Homo sp. indet. from East Rudolf, Kenya, is 52.8 kg (116 lb). These estimates are consistent even if pongid proportions are assumed. Indices of encephalization show that the brain to body weight ratio in Australopithecus is above the great ape averages but well below Homo sapiens. The Homo sp. indet. represented by the KNM-ER 1470, O.H. 7 and O.H. 13 crania have encephalization indices above Australopithecus despite the greater body weight of the former.     

Protoanthropoidea (Primates, Simiiformes): A New Primate Higher Taxon and a Solution to the Rooneyia Problem

As a derivative of the hypothesis that anthropoids evolved from omomyid-like primates, the enigmatic North American fossil Rooneyia viejaensis, from the latest Eocene of Texas, is placed in a new higher taxon, Protoanthropoidea, which is proposed as the sister-group of Anthropoidea. Rooneyia and anthropoids share synapomorphically a pattern of character states relating to the unique orbital morphology of higher primates, including; highly convergent and frontated orbits roofed above by an extended frontal bone; funnel-shaped orbital fossae; orbital apices that are recessed beneath the forebrain; a deep, large lateral process of the frontal bone (upper portion of the postorbital bar) that may presage closure of the orbit by an enlarged ascending process of the zygomatic. If the sister-group of anthropoids occupied North America as part of a Laurasian geographic distribution during the Paleogene, as some primate genera did, ancestral anthropoids may likewise have occurred across Laurasia, prestaging them to enter Africa and Central/South America in two independent episodes of dispersal—without having the ancestral platyrrhines crossing the daunting Atlantic Ocean.     

Alternative spatial sampling in studies of plant demography: consequences for estimates of population growth rate

Ecologists commonly use matrix models to study the population dynamics of plants. Most studies of plant demography use plot-based methods to collect data, in part, because mapped individuals are easier to relocate in subsequent surveys and survey methods can be standardized among sites. However, there is tremendous variation among studies, both in terms of plot arrangement and the total area sampled. In addition, there has been little discussion of how alternative sampling arrangements influence estimates of population growth rates (λ) calculated with matrix models. We surveyed the literature to determine what sampling designs are most used in studies of plant demography using matrix models. We then used simulations of three common sampling techniques—using a single randomly placed plot, multiple randomly placed plots, and systematically distributed plots—to evaluate how these alternative strategies influenced the precision of estimates of λ. These simulations were based on long-term demographic data collected on 13 populations of the Amazonian understory herb Heliconia acuminate (Heliconiaceae). We found that the method used to collect data did not affect the bias or precision of estimates in our system—a surprising result, since the advantage in efficiency that is gained from systematic sampling is a well-known result from sampling theory. Because the statistical advantage of systematic sampling is most evident when there is spatial structure in demographic vital rates, we attribute this result to the lack of spatially structured vital rates in our focal populations. Given the likelihood of spatial autocorrelation in most ecological systems, we advocate sampling with a systematic grid of plots in each study site, as well as that researchers ensure that enough area is sampled—both within and across sites—to encompass the range of spatial variation in plant survival, growth, and reproduction.     

Major stages in the evolution of primate neurocrania

An important constraint on the evolution of primate skeletons is the isometric relationship which exists between skeletal weight and body weight. The evolution of primate skulls during the Tertiary and Quaternary periods indicates that redeployment of bone mass took place largely within the skull (i.e. between proximate ossifying centres) and that the major vector was from the splanchnocranium towards the neurocranium. This vector of bone mass redeployment accords well with the general treand within primates of increased encephalisation over time. There are however, several interesting examples of vectors which were oriented in the opposite sense, in particular in the robust australopithecine lineage, in which the emphasis on bone mass deployment was towards the splanchnocranium and away from the neurocranium. A fuller understanding of skeletal isometry, and a wider application in comparative anatomy may throw much light on the evolution of skeletal systems, and it may resolve somelong-standing debates. Among these may be identification of the selection pressures which have led to dental and alveolar reduction inHomo sapiens sapiens (bone mass redeployment into the neurocranium) and perhaps an explanatation for some types of osteoporosis in old people whose body weight decreases may result in isometric skeletal mass decreases (for every 100 gms muscle tissue loss, there will be about 7 gms of bone tissue loss).     

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.     

On the definition of variables in studies of primate dental allometry

Cranial and dental measurements are taken on 253 adult female primates from 32 species. Regression equations are calculated to determine allometric relationships between anterior tooth size, posterior tooth size, and body size. When cranial length or skull length is used as the measure of general size, the results of the equations differ from when body weight is the reference dimension. Similarly, using different definitions of posterior tooth size (such as mandibular second molar length and maxillary postcanine area) alters results substantially. The same occurs with different definitions of anterior tooth size. It has been common in studies of primate dental allometry to generalize from the specific variables measured to broad functional interpretations. However, highly correlated variables cannot be substituted one for another in allometric analyses without important changes in the results of the equation. Interpretation of allometric data is more highly restricted to the precise variables measured in a particular study than has been generally recognized.     

Sexual dimorphism and direct and maternal genetic effects on body weight in mice

Summary Genetic and phenotypic parameters for three-, six- and eight-week body weight and for weight gain between three and six weeks of age were estimated from data collected over 14 generations in a randombred control population. Genetic parameters were also estimated for sexual dimorphism in body weight and gain. Heritability estimates were substantial for body weight at all ages and for body weight gain. Additive maternal variances were also large. Estimates of the covariance between direct and maternal genetic effects were negative and substantial for three- and six-week weights and gain. Also the covariance between maternal effects on weaning weight and direct genetic effects on six- and eight-week weights were negative. These results indicate a consistent antagonism between maternal and direct genetic effects in this population.The analysis of sexual dimorphism yielded estimates of 0.87±.09 and 0.71±.14 for the correlation between additive direct effects on males and females for six-week weight and body weight gain respectively. Corresponding heritability estimates were 0.07±.09 and 0.11±.09. Heritability estimates for sexual dimorphism in three- and eight-week weights were negative.Journal Paper No. 3687 of the North Carolina State University Agricultural Experiment Station. This investigation was supported in part by NIH Grant No. GM11546.     

An economic assessment of the contribution of biological control to the management of invasive alien plants and to the protection of ecosystem services in South Africa

This study is a first attempt at a holistic economic evaluation of South African endeavours to manage invasive alien plants using biological control. Our focus was on the delivery of ecosystem services from habitats that are invaded by groups of weeds, rather than by each individual weed species. We established the net present value of the weed biological control efforts, and derived benefit:cost ratios by comparing this value (a cost) to the estimated value of ecosystem services protected by weed biological control. We identified four major functional groupings of invading alien plants, and assessed their impact on water resources, grazing and biodiversity. We estimated the area that remained free of invasions due to all historic control efforts in South Africa, and the proportion that remained free of invasion as a result of biological control (which was initiated in 1913). The estimated value of potential ecosystem services amounted to 152 billion South African rands (ZAR—presently, about US$ 19.7 billion) annually. Although an estimated ZAR 6.5 billion was lost every year due to invading alien plants, this would have amounted to an estimated additional ZAR 41.7 billion had no control been carried out, and 5–75% of this protection was due to biological control. The benefit:cost ratios ranged from 50:1 for invasive sub-tropical shrubs to 3,726:1 for invasive Australian trees. Benefit:cost ratios remained positive and our conclusion, that biological control has brought about a considerable level of protection of ecosystem services, remains robust even when our estimates of the economic impacts of key variables (i.e. sensitivity analyses of indeterminate variables) were substantially reduced.