A 3D geometric morphometric analysis of the bovid distal humerus, with special reference to Rusingoryx atopocranion (Pleistocene, Eastern Africa).

The family Bovidae [Mammalia: Artiodactyla] is speciose and has extant representatives on every continent, forming key components of mammal communities. For these reasons, bovids are ideal candidates for studies of ecomorphology. In particular, the morphology of the bovid humerus has been identified as highly related to functional variables such as body mass and habitat. This study investigates the functional morphology of the bovid distal humerus in isolation due to its increased likelihood of preservation in the fossil record, and the resulting opportunity for a better understanding of the ecomorphology of extinct bovids. A landmark scheme of 30 landmarks was used to capture the 3D distal humerus morphology in 111 extant bovid specimens. We find that the distal humerus has identifiable morphologies associated with body mass, habitat preference and tribe affiliation and that some characteristics are shared between high body mass bovids and those living on hard, flat terrain which is likely due to the high stress on the bone in both cases. We directly apply our findings regarding extant bovids to the extinct alcelaphine bovid, Rusingoryx atopocranion from the mid to late Pleistocene (>33-45 ka) Lake Victoria region of Kenya. This species is known for some peculiar morphologies including a domed cranium with hollow nasal crests, and having small hooves for a bovid of its size. Another interesting aspect of Rusingoryx's skeletal morphology which has not been addressed is an unusual protrusion on the lateral epicondyle of the distal humerus. Despite considerable individual variation in the Rusingoryx specimens, we find evidence to support its historical assignment to the tribe Alcelaphini, and that it likely preferred open grassland habitats, which is consistent with independent reconstructions of the palaeoenvironment. We also provide the most accurate body mass estimate for Rusingoryx to date, based on distal humerus centroid size. Overall, we are able to conclude that the distal humerus in extant bovids is highly informative regarding body mass, habitat preference and tribe, and that this can be applied directly to a fossil taxon with promising results.

Ecomorphology of the cervid intermediate phalanx and its implications for palaeoenvironmental reconstruction.

This paper reports on newly developed ecomorphological models for the cervid intermediate phalanx. Using a geometric morphometric approach, we quantitatively assess the overall gracility of the bone, the depth and concavity of the proximal articulation and the roundness and symmetry of the distal articulation in the intermediate phalanx, to establish relationships between morphology, locomotor behavior and environment. The morphology of the phalanx was found to vary along a gradient from gracile phalanges with shallow proximal articulations in forms adapted to yielding substrate, to robust phalanges with deeper proximal articulations in taxa adapted to firm substrate. Phylogeny and allometry are accounted for using regressions and phylogenetic comparative methods. Although the results indicate phylogeny explains part of the morphological variation, overall the shape of the intermediate phalanx appears mainly driven by differences in function. Consequently, this element promises to be a useful palaeoenvironmental proxy that can be applied on fossil assemblages with cervid remains.

Ecomorphology of the cervid calcaneus as a proxy for paleoenvironmental reconstruction.

This study presents new ecomorphological models for the cervid calcaneus that can be used to make predictions about the nature of ancient environments. Using geometric morphometrics to quantitatively assess the length of the articular surface supporting the malleolus, the length and orientation of the tuber calcanei, and the position of the articular facets, we aimed to establish correlations between morphological traits, locomotor behavior, and environmental parameters in extant cervids. The morphology of the calcaneus was found to primarily vary with locomotor strategy and habitat, along a continuum from habitats with an open vegetation structure to habitats with a closed vegetation structure. Confounding factors, including sexual dimorphism, allometry, and phylogeny were accounted for using Principal Component Analysis, regressions and phylogenetic comparative methods. The results of our analyses suggested that these factors did not substantially obscure habitat predictions. As such, the calcaneus provides a valuable proxy for paleoenvironmental reconstruction that is broadly applicable to Quaternary fossil assemblages with a sufficiently large sample of cervids.

Developmental instability in wild Nigerian olive baboons (Papio anubis).

BACKGROUND: Developmental instability in archaeological samples can be detected through analysis of skeletal and dental remains. During life, disruptions to biological internal homeostasis that occur during growth and development redirect bodily resources to returning to homeostasis and away from normal processes such as symmetrical development. Because dental enamel does not remodel in life, any deviations from normal development are left behind. Even subtle disturbances to developmental trajectory may be detected in asymmetrical development of traits, specifically a random variation in sides termed fluctuating asymmetry. Human dental fluctuating asymmetry studies are common, but here we investigate the permanent dentition of a non-human primate Papio anubis, for potential fluctuating asymmetry relative to sex, weaning, and reproductive maturity. The sample stems from an outlier population that lives in the wettest and most humid habitat of any studied baboon group. METHODS: The skulls of adult baboons were collected after their natural death in Gashaka Gumti National Park, Nigeria. The permanent dentition of antimeric teeth (paired) were measured for maximum length and breadth using standard methods. The metrics were analyzed to assess the presence of fluctuating asymmetry in adult permanent mandibular and maxillary dentition. Measurement error and other forms of asymmetry (antisymmetry, directional asymmetry) were considered and dental measures expressing true fluctuating asymmetry were used to address three research questions. RESULTS: Males exhibit greater fluctuating asymmetry than females, suggesting that males experience greater overall instability during the developmental period. While weaning is not more stressful than other life history stages for males and females (using the first molar fluctuating asymmetry index as a proxy compared to other teeth), it is more stressful for females than males. The onset of reproduction is also not more stressful than other life history stages for males and females (using the third molar fluctuating asymmetry index as a proxy compared to other teeth), but it is more stressful for males than females. We explore possible explanations for these findings in the discussion.

Ontogenetic and morphological variation in primate long bones reflects signals of size and behavior.

OBJECTIVES: Many primates change their locomotor behavior as they mature from infancy to adulthood. Here we investigate how long bone cross-sectional geometry in Pan, Gorilla, Pongo, Hylobatidae, and Macaca varies in shape and form over ontogeny, including whether specific diaphyseal cross sections exhibit signals of periosteal adaptation or canalization. MATERIALS AND METHODS: Diaphyseal cross sections were analyzed in an ontogenetic series across infant, juvenile, and adult subgroups. Three-dimensional laser-scanned long bone models were sectioned at midshaft (50% of biomechanical length) and distally (20%) along the humerus and femur. Traditional axis ratios acted as indices of cross-sectional circularity, while geometric morphometric techniques were used to study cross-sectional allometry and ontogenetic trajectory. RESULTS: The humeral midshaft is a strong indicator of posture and locomotor profile in the sample across development, while the mid-femur appears more reflective of shifts in size. By comparison, the distal diaphyses of both limb elements are more ontogenetically constrained, where periosteal shape is largely static across development relative to size, irrespective of a given taxon's behavior or ecology. DISCUSSION: Primate limb shape is not only highly variable between taxa over development, but at discrete humeral and femoral diaphyseal locations. Overall, periosteal shape of the humeral and femoral midshaft cross sections closely reflects ontogenetic transitions in behavior and size, respectively, while distal shape in both bones appears more genetically constrained across intraspecific development, regardless of posture or size. These findings support prior research on tradeoffs between function and safety along the limbs.

Fire in the Plio-Pleistocene: the functions of hominin fire use, and the mechanistic, developmental and evolutionary consequences.

Fire is a powerful natural force that can change landscapes extremely quickly. Hominins have harnessed this resource for their own purposes, with mechanistic and developmental physiological consequences. In addition, the use of fire has niche constructive effects, altering selective environments for genetic and cultural evolution. We review the record for hominin fire use in the Plio-Pleistocene, before considering the various functions for its use, and the resultant mechanistic and developmental consequences. We also adopt the niche construction framework to consider how the use of fire can modify selective environments, and thus have evolutionary consequences at genetic and cultural levels. The light that fire produces may influence photoperiodicity and alter hormonally-controlled bodily rhythms. Fire used for cooking could have extended the range of foods hominins were able to consume, and reduced digestion costs. This may have contributed to the expansion of the hominin brain and facial anatomy, influenced by a higher quality cooked diet. Fire may also have allowed dispersal into northern areas with much cooler climates than the hominin African origin, posing novel problems that affected diet and social behaviour.

Ecological continuity between Lower and Upper Bed II, Olduvai Gorge, Tanzania.

The sediments of Bed II at Olduvai Gorge, Tanzania, have received less scientific scrutiny than Bed I strata, likely due to the lower density of fossil hominins in the younger layers. Nevertheless, Bed II provides important contextual information about changes over time in early hominin environments. Mammal fossils from Bed II are separated into older and younger faunas by a regional disconformity under which lies an aeolian tuff, the Lemuta Member, dated to ∼1.74 Ma (millions of years ago). Differences between the faunal communities above and below the Lemuta Member have been previously noted and interpreted as evidence for distinct environmental change. Here, we investigate the palaeoecological conditions that characterised the transition between older and younger Bed II sites via analysis of their mammalian community ecological niche exploitation profiles, comparing these to known modern habitats. We present a comprehensive species list for both Bed II faunas. An initial correspondence analysis points to palaeoecological differences in pre- and post-Lemuta Member assemblages, although neither is representative of a forest habitat. When taphonomic differences in body mass profiles are taken into account, however, ecological differences appear far less important. Based on a resampling procedure to generate modern locality data with body mass profiles similar to the Bed II faunas, discriminant correspondence analyses liken both fossil assemblages to modern woodland habitats, with Lower Bed II having some affinity to floodplain and marshy woodlands, likely related to the presence of the Olduvai palaeolake in this stratigraphic interval. A comparison of the Bed II faunas to each other suggests that their differences could reasonably be sampled from within a single modern habitat. Although additional evidence points to an increase in aridity subsequent to the Lemuta Member, the importance of woodland habitats throughout Bed II is clear.

Bovid postcranial ecomorphological survey of the Laetoli paleoenvironment.

Here we report on a bovid postcranial ecomorphological survey of the fossil assemblages from the Plio-Pleistocene site of Laetoli, Tanzania. A global sample of extant bovids (n=205), cervids (n=14), and tragulids (n=5) from seven known habitat types constitutes the comparative data set. All long bones, carpals, tarsals, and phalanges were measured. Discriminant function analyses (DFA) were conducted in order to evaluate the ability of each element to accurately predict habitat affiliation. The baseline of chance accuracy for DFAs (i.e., the percentage of correct predictions that can be expected when habitat assignments are randomized) served as the cut-off point between good and bad habitat predictors. A total of 22 elements yielded percentages of correct classification over the baseline of accuracy, and these were extended to the Laetoli fossil assemblages. Summaries of the number of specimens predicted to belong to each habitat type were used to reconstruct the paleoenvironment. The results indicate that, at the time of the deposition of the Laetolil Beds, the area had heavy woodland-bushland cover with some lighter tree and bush cover and grass available. These results lend strong support to recent suggestions that the area was on the more wooded end of the habitat spectrum, contra initial conclusions that it represented a mosaic of more open habitats. The results also indicate that, during the deposition of the Ndolanya Beds, the environment had become more open and the grassland component of the environment had increased significantly. Light woodland-bushland and an abundance of grass cover dominated the landscape, although tracts of land with denser vegetation likely existed. This conclusion agrees with earlier suggestions that the area was a semiarid bushland.

The palaeoecology of the Upper Ndolanya Beds at Laetoli, Tanzania.

The palaeoecology of the fauna from the Ndolanya Beds, Laetoli, Tanzania, has been analysed to reconstruct the environment of this 2.6 Ma site. Community profiles have been constructed in relation to three variables that carry ecological meaning: body weight, locomotor adaptations and feeding preferences. Comparing the similarities and differences in the structure of the fossil faunas with those of modern environments allows us to draw inferences about palaeoenvironmental conditions, and this is based on a comparative dataset of 15 modern environments (44 localities) covering a wide range of climatic and ecological conditions across Asia, Africa and Central America. In addition, 16 fossil sites in East and South Africa have been analysed in the same way, and both sets of comparative data have been used as the basis for comparison with an ecological diversity analysis of the fauna from the Upper Ndolanya Beds. The Ndolanya fauna is characterized by a predominance of medium to large sized terrestrial and herbivorous species. There is evidence of taphonomic bias that eliminated many of the smaller species. A comparison of multivariate analyses of 23 selected modern localities conducted both with and without the small species included, indicates that the loss of these species does not adversely influence the results of an ecological diversity analysis. The evidence suggests that at the time of the deposition of the Ndolanya Beds the Laetoli region was a semi-arid bushland. This is considerably drier and more open than is suggested for the earlier Laetolil Beds.