Early hominid brain evolution: a new look at old endocasts

Early hominid brain morphology is reassessed from endocasts of Australopithecus africanus and three species of Paranthropus, and new endocast reconstructions and cranial capacities are reported for four key specimens from the Paranthropus clade. The brain morphology of Australopithecus africanus appears more human like than that of Paranthropus in terms of overall frontal and temporal lobe shape. These new data do not support the proposal that increased encephalization is a shared feature between Paranthropus and early Homo. Our findings are consistent with the hypothesis that Australopithecus africanus could have been ancestral to Homo, and have implications for assessing the tempo and mode of early hominid neurological and cognitive evolution.     

Comparative context of Plio-Pleistocene hominin brain evolution.

One of the distinguishing features of Homo sapiens is its absolutely and relatively large brain. This feature is also seen in less extreme form in some fossil Homo species. However, are increases in brain size during the Plio-Pleistocene only seen in Homo, and is brain enlargement among Plio-Pleistocene primates confined to hominins? This study examines evidence for changes in brain size for species and lineage samples of three synchronic East African fossil primate groups, the two hominin genera Homo and Paranthropus, and the cercopithecoid genus Theropithecus. Hominin endocranial capacity data were taken from the literature, but it was necessary to develop an indirect method for estimating the endocranial volume of Theropithecus. Bivariate and multivariate regression equations relating measured endocranial volume to three external cranial dimensions were developed from a large (ca. 340) sample of modern African cercopithecoids. These equations were used to estimate the endocranial volumes of 20 Theropithecus specimens from the African Plio-Pleistocene. Spearman's rho and the Hubert nonparametric test were used to search for evidence of temporal trends in both the hominin and Theropithecus data. Endocranial volume apparently increased over time in both Homo and Paranthropus boisei, but there was no evidence for temporal trends in the endocranial volume of Theropithecus. Thus, hypotheses which suggest a mix of environmental, social, dietary, or other factors as catalysts for increasing brain in Plio-Pleistocene primates must accommodate evidence of brain enlargement in both Homo and Paranthropus, and explain why this phenomenon appears to be restricted to hominins.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

The perils of picky eating: dietary breadth is related to extinction risk in insectivorous bats

Several recent papers evaluate the relationship between ecological characteristics and extinction risk in bats. These studies report that extinction risk is negatively related to geographic range size and positively related to habitat specialization. Here, we evaluate the hypothesis that extinction risk is also related to dietary specialization in insectivorous vespertilionid bats using both traditional and phylogenetically-controlled analysis of variance. We collected dietary data and The World Conservation Union (IUCN) rankings for 44 Australian, European, and North American bat species. Our results indicate that species of conservation concern (IUCN ranking near threatened or above) are more likely to have a specialized diet than are species of least concern. Additional analyses show that dietary breadth is not correlated to geographic range size or wing morphology, characteristics previously found to correlate with extinction risk. Therefore, there is likely a direct relationship between dietary specialization and extinction risk; however, the large variation in dietary breadth within species of least concern suggests that diet alone cannot explain extinction risk. Our results may have important implications for the development of predictive models of extinction risk and for the assignment of extinction risk to insectivorous bat species. Similar analyses should be conducted on additional bat families to assess the generality of this relationship between niche breadth and extinction risk.     

Pulsed resource availability changes dietary niche breadth and partitioning between generalist rodent consumers

Identifying the mechanisms that structure niche breadth and overlap between species is important for determining how species interact and assessing their functional role in an ecosystem. Without manipulative experiments, assessing the role of foraging ecology and interspecific competition in structuring diet is challenging. Systems with regular pulses of resources act as a natural experiment to investigate the factors that influence the dietary niches of consumers. We used natural pulses of mast‐fruiting of American beech (Fagus grandifolia) to test whether optimal foraging or competition structure the dietary niche breadth and overlap between two congener rodent species (Peromyscus leucopus and P. maniculatus), both of which are generalist consumers. We reconstructed diets seasonally over a 2‐year period using stable isotope analysis (δ¹³C, δ¹⁵N) of hair and of potential dietary items and measured niche dynamics using standard ellipse area calculated within a Bayesian framework. Changes in niche breadth were generally consistent with predictions of optimal foraging theory, with both species consuming more beechnuts (a high‐quality food resource) and having a narrower niche breadth during masting seasons compared to nonmasting seasons when dietary niches expanded and more fungi (a low‐quality food source) were consumed. In contrast, changes in dietary niche overlap were consistent with competition theory, with higher diet overlap during masting seasons than during nonmasting seasons. Overall, dietary niche dynamics were closely tied to beech masting, underscoring that food availability influences competition. Diet plasticity and niche partitioning between the two Peromyscus species may reflect differences in foraging strategies, thereby reducing competition when food availability is low. Such dietary shifts may have important implications for changes in ecosystem function, including the dispersal of fungal spores.     

Conservation value of roadside verges for stenotopic heathland Carabidae: corridors or refugia?

It has been shown that, in The Netherlands, sandy roadside verges may aid the survival of stenotopic Carabidae associated with open habitats such as heathland and sandy grassland. It is suggested that roadsides serve as corridors linking heathland fragments, thus maintaining a metapopulation structure, which helps avoid regional extinction. However, a counter-argument is that verges encourage dispersal into unsuitable habitats, and function as sinks in which individuals die without reproducing. The stenotopic species recorded from roadside verges but not in adjacent open habitats all had high powers of dispersal. The least successful dispersers were found only in open habitats and in verges closely connected to them. From this, the conclusion that verges function as corridors can be drawn. We reinterpret these data to suggest that stenotopic species were breeding on the roadside verges, and that they acted as refugia rather than corridors. We also present evidence from a roadside verge in the Breckland region of East Anglia, England. We show that many stenotopic species breed on the verge, which thus functions as a refugium. For Carabidae dependent on early successional habitats, survival may be more likely on disturbed roadside verges than on nature reserves in the absence of regular management disturbance.     

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species (Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.     

Taxonomic attribution of the Olduvai hominid 7 manual remains and the functional interpretation of hand morphology in robust australopithecines

In this paper, we test the currently accepted taxonomic hypothesis that the hand of the Homo habilis holotype Olduvai hominid 7 (OH7) from Olduvai Gorge can be unambiguously assigned to Homo. Morphometric and morphological comparison with humans and australopithecines (Australopithecus and Paranthropus) indicate that the OH7 hand most likely belongs to P. boisei. The morphological adaptations of Paranthropus are thus further evaluated in the light of the alternative taxonomic hypothesis for OH7. Functional analyses suggest that morphological features related to human-like precision grasping, previously considered diagnostic of toolmaking by some, may be alternatively attributed to specialized manual feeding techniques in robust australopithecines.     

Dental microwear and stable isotopes inform the paleoecology of extinct hominins

Determining the diet of an extinct species is paramount in any attempt to reconstruct its paleoecology. Because the distribution and mechanical properties of food items may impact postcranial, cranial, mandibular, and dental morphologies related to their procurement, ingestion, and mastication, these anatomical attributes have been studied intensively. However, while mechanical environments influence skeletal and dental features, it is not clear to what extent they dictate particular morphologies. Although biomechanical explanations have been widely applied to extinct hominins in attempts to retrodict dietary proclivities, morphology may say as much about what they were capable of eating, and perhaps more about phylogenetic history, than about the nature of the diet. Anatomical attributes may establish boundary limits, but direct evidence left by the foods that were actually (rather than hypothetically) consumed is required to reconstruct diet. Dental microwear and the stable light isotope chemistry of tooth enamel provide such evidence, and are especially powerful when used in tandem. We review the foundations for microwear and biogeochemistry in diet reconstruction, and discuss this evidence for six early hominin species (Ardipithecus ramidus, Australopithecus anamensis, Au. afarensis, Au. africanus, Paranthropus robustus, and P. boisei). The dietary signals derived from microwear and isotope chemistry are sometimes at odds with inferences from biomechanical approaches, a potentially disquieting conundrum that is particularly evident for several species.     

Early hominin speciation at the Plio/Pleistocene transition

Over the last half-decade or so, there has been an explosion in the recognition of hominin genera and species. We now have the late Miocene genera Orrorin and Sahelanthropus, the mid Pliocene genus Kenyanthropus, three new Pliocene species of Australopithecus (A. anamensis, A. garhi and A. bahrelghazali) and a sub species of Ardipithecus (Ar. r. kadabba) to contend with. Excepting also the more traditional species allocated to Paranthropus, Australopithecus and early Homo we are approaching around 15 species over 5 million years (excluding hominin evolution over the last one million years). Can such a large number of hominin species be justified? An examination of extant hominid (Gorilla gorilla, Pan troglodytes, and Pan paniscus) anatomical variability indicates that the range of fossil hominin variability supports the recognition of this large number of fossil species. It is also shown that not all hominins are directly related to the emergence of early Homo and as such have become extinct. Indeed the traditional australopithecine species 'A'. anamensis, 'A'. afarensis and 'A'. garhi are considered here to belong to a distinct genus Praeanthropus. They are also argued not be hominins, but rather an as yet undefined hominid group from which the more derived hominins evolved. The first hominin is represented by A. africanus or a hominin very much like it. The Paranthropus clade is defined by a derived heterochronic condition of peramorphosis, associated with sequential progenesis (contraction of successive growth stages) in brain and dental development, but a mixture of peramorphic and paedomorphic features in its craniofacial anatomy. Conversely, Kenyanthropus and Homo both share a pattern of peramorphosis, associated with sequential hypermorphosis (prolongation of successive growth stages) in brain development, and paedomorphosis processes in cranial, facial and dental development. This suggests, that these two clades share an important synapomorphy not recognised in the parsimony analyses, suggesting that they may form a sister group relationship to the exclusion of Paranthropus. This highlights the need to re-interpret phylogenetic results in terms of function and development. The rapid speciation and extinction as argued here is in keeping with other fossil groups in Africa at the Plio/Pleistocene transition. This emphasises that we must approach the pre-australopithecines and hominins as part of the endemic African fauna, and not in isolation to the evolutionary and climatic processes that were operating all around them.     

Brief communication: evidence bearing on the status of Homo habilis at Olduvai Gorge

Students of the early hominin career have debated the status of Homo habilis since its discovery in 1960. Today discussion centers on which specimens should be included in the species and what constitutes the holotype. Recent reviews of early Homo suggest that the Olduvai Hominid 8 foot may sample Paranthropus while the OH 7 skull bones, mandible, and hand sample H. habilis. Moreover, some suggest that while H. habilis in Middle Bed I at Olduvai is craniodentally Homo-like, the postcranial skeleton of H. habilis is more like that of Australopithecus. Evidence presented here indicates not only that OH 7 and OH 8 represent H. habilis but also that they come from a single individual. The association of OH 35 with OH 7 and OH 8 is less certain. Morphological, pathological, and taphonomic evidence favors the inclusion of OH 35 in the holotype. However, stratigraphic evidence suggests that OH 35 and OH 8 are not coterminous. With or without OH 35, the holotype of H. habilis ranks as one of the most complete early hominin skeletons and the most complete and functionally informative specimen of early Homo.     

Co-existence and niche segregation of three small bovid species in southern Mozambique

Niche segregation among three small antelopes – red duiker, common duiker and suni – was investigated in a coastal savanna woodland/forest mosaic. It was expected that these similar‐sized concentrate selectors would show differentiation in diet choice to decrease competition. Diet composition did not vary significantly among the different vegetation types. For all three antelope species, the number of dietary items was large, with a minimum of 70 different food items per species. Dietary specialization was low, with only 10% of the food items being exclusively used by each of the species. The ranks of food items were positively correlated among species in the wet season, but not in the dry season. Diet breadth significantly decreased in the dry season. The use of exclusive species was significantly larger in the dry season with lowest values recorded for the common duiker. Diet overlap in the wet season was considerable, but significantly decreased in the dry season, the time of food scarcity. The dry season data showed evidence for niche segregation, although this was not based on displacement. As niche segregation in the dry season was coupled to a random apportionment of diet items among antelope species, it cannot be interpreted as the result of competitive displacement.     

陕西公王岭蓝田直立人内耳迷路的复原及形态特点

1964年在陕西公王岭发现的蓝田人头骨的形态比周口店直立人和印度尼西亚爪哇直立人原始,其厚重的骨壁及较小的脑量,落入了早期人属成员的变异范围。最新测年结果将蓝田人的生存年代从原先普遍接受的距今115万年提早到大约163万年前,接近能人和南方古猿生存年代变异范围的下限,蓝田人是迄今为止我国发现的有确定年代数据的最早的古人类化石。本文采用高分辨率CT技术对蓝田人的颞骨岩部进行了扫描,对骨性内耳迷路进行了3D虚拟复原,通过与和县直立人、欧洲古老型智人、早期人属成员、南方古猿非洲种、粗壮傍人和现代人内耳迷路的21项测量项目的对比和分析,结果显示蓝田人内耳迷路的测量数据与南方古猿非洲种最接近,其次为现代人和欧洲古老型智人,而与早期人属成员和粗壮傍人相差较大。主成分分析结果显示,蓝田人内耳迷路与早期人属成员、欧洲古老型智人、南方古猿非洲种及现代人都有重叠区域,距离最近的是南方古猿非洲种Sts 5,其次为和县直立人和南方古猿非洲种Sts 19,而与粗壮傍人距离较远。本文研究提供了中更新世中国古人类内耳迷路的形态数据,为进一步探讨蓝田人体质特征演化上的意义提供了参考资料。     

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.     

Morphological variation in some lower palaeozoic dalmanellid brachiopods

Three predictions arising from the niche-variation model concerning morphological variation are tested. These are: increasing diversity accompanied by reduced species variability, variation directly proportional to species abundance and eurytopic species more variable than stenotopic forms. Using Ordovician and Silurian dalmanellid brachiopods, measurements were taken at different intervals through ontogeny of 5 brachial characters on 11 species ofBancroftina, Dalmanella andOnniella (all Ordovician andIsorthis (Silurian). Morphological variability decreases through ontogeny but is not related to species abundance as predicted. Indeed, evidence suggests the opposite. Less abundant species are generally more variable, this is again the opposite of the predictions. There is no apparent relationship between morphological variability, eurytopy and stenotopy. These results considered with other data suggest that other Lower Palaeozoic animals, particularly suspension feeders, have similar niche requirements and these may be fairly broad.     

Dietary niche breadth in a local community of passerine birds: an analysis using phylogenetic contrasts

The analysis of a local community of forest passerines (13 species) using phylogenetic contrasts shows a correlation between body size of bird species and mean prey size, minimum prey size, maximum prey size and the size range of dietary items. This suggests that larger birds drop small prey taxa from their prey list, because of the difficulty of capturing very small prey, for energetic reasons or because of microhabitat usage. We find some support for the third hypothesis. Dietary niche breadth calculated across prey taxa is not related to body size. Dietary niche breadth, however, is correlated with size-corrected measurements of the bill and locomotor apparatus. Long and slender bills increase the dietary niche breadth. Thus subtle differences constrain foraging and the techniques of extracting certain prey taxa form crevices. Dietary niche breadth and foraging diversity are positively correlated with population density: at least locally dietary generalists occur at higher breeding densities than specialists.     

Enamel microstructure of the hominid KB 5223 from Kromdraai, South Africa

The Plio-Pleistocene site of Kromdraai, South Africa, is well known for the recovery of the holotype of Paranthropus robustus, one of nine individual hominids recovered from this site to date. Among the Kromdraai sample, the specimen KB 5223 comprises several isolated deciduous and permanent lower teeth assigned to Paranthropus, the only recognized genus at this site. However, a more recent analysis of this specimen suggested that it should be classified as Homo. The lower right first permanent molar of KB 5223 had been previously sectioned along the tips of the mesial cusps, exposing its enamel microstructure. Previous studies had indicated differences between Homo and Paranthropus at the microstructural level. A portable confocal scanning microscope was used to describe details of the enamel microstructure of the M1 and I1 of this specimen. Angles formed between the striae of Retzius and the enamel dentine junction (EDJ), daily secretion rates in cuspal enamel of the protoconid and metaconid and crown formation time of the RM1 are provided. The number of perikymata on the right I1 was counted. Results indicate that some features recorded in the KB 5223 molar differ from those of Paranthropus. However, the number of perikymata on the I1 is lower than values so far reported for early Homo but similar to Paranthropus. Crown formation time of KB 5223 M1 was markedly lower than mean values of M1 in H. sapiens, but similar to other early hominids. Daily secretion rates in the cuspal enamel of KB 5223 M1 were higher than in modern humans.     

Analysis and significance of linear enamel hypoplasia in Plio-Pleistocene hominins

This study of linear enamel hypoplasia (LEH) in Plio-Pleistocene hominins builds on a previous study (Guatelli-Steinberg [2003] Am. J. Phys. Anthropol. 120:309-322) that focused on LEH in early South African hominins. The present study is more comprehensive, encompassing dental specimens of hominins from East Africa as well, including early Homo. As a developmental defect of enamel, LEH is used in anthropological contexts to reveal information about physiological stress. However, intrinsic aspects of enamel development and morphology can affect the expression of LEH, complicating efforts to understand the significance of these defects. In this study, the analysis of LEH is conducted with respect to enamel development and morphology. It is predicted that Paranthropus should have fewer defects on its canine teeth than Australopithecus and Homo, owing to its abbreviated period of enamel formation. This prediction is supported: Paranthropus has statistically significantly fewer defects per canine than Australopithecus and Homo. The previous study demonstrated that despite the wider spacing of perikymata on the teeth of South African Paranthropus, defects on the canine teeth of this genus were not wider than those of Australopithecus. A multiple linear regression analysis in that study, as well as a separate analysis in the present study, indicate that the number of perikymata within defects is a better predictor of defect width than perikymata spacing. In this study, it was additionally found that the average number of perikymata within Australopithecus defects is statistically significantly greater than it is in Paranthropus, thus explaining why Paranthropus defects are not wider than those of Australopithecus. The biological significance of this difference in the number of perikymata within the defects of Australopithecus and Paranthropus is considered in light of several factors, including: 1) the possibility that other intrinsic attributes of enamel morphology may be involved (specifically the faster extension rates of Paranthropus that result in shallower defects), 2) generic differences in the canalization of enamel development, and 3) generic differences in the duration of disruptions to enamel growth.