Monthly Archives: August 2009

Anthro blog carnival: Four Stone Hearth 74

The 74th Four Stone Hearth anthropology blog carnival is available over at Adam Henne’s Natures/Cultures blog. Catch up on the latest on anthropology blogging. The next Four Stone Hearth will be hosted here on the 9th of September. Send any anthropology submission for the upcoming carnival to Martin Rundkvist or me (be sure to replace [AT] with @ in the email addresses).

Advertisements

Upper Palaeolithic Italy

The early Lower Palaeolithic remains from Ceprano and Monte Poggiolo show that humans arrived in the Italian peninsula sometime prior to 700,000 years ago. However, it is not until the Aurignacian period (~ 38,000-30,000 BP*) of the Upper Palaeolithic that we see signs of widespread occupation. The earliest Aurignacian seems to occur in northern Italy, before expanding southwards (Mussi 1990). The people of the Aurignacian exploited very diverse habitats in both lowland and mountainous areas, as well as on the island of Sicily.
Extrapolating from data by Bocquet-Appel et al. (2005) it is possible that the population of Italy did not exceed 1,000 inhabitants. While this figure is little more than a best guess, it still gives us an idea of the fragile nature of these groups, who were living constantly on the edge of extinction. With such small numbers inbreeding can become a serious problem. However, studies of European Upper Palaeolithic people find them to be surprisingly homogenous as a whole, which would suggest a significant amount of gene flow between far-flung populations (Henke 1989, 1991). In order to understand how mating systems would have operated during the Early Upper Palaeolithic researchers have looked to modern hunter-gatherer groups. Such studies have found hunter-gatherers to have less group closure than agriculturalists. This would have been an important survival strategy for the hunter-gatherer groups of the Upper Palaeolithic, especially when faced with unpredictable environmental conditions and already constrained mating networks. This open network is also reflected in the material culture from this period. Venus figurines, dating chiefly from the Gravettian period, have been found over an area of approximately 2000 km, spanning from modern day Russia in the Northeast to Spain in the Southwest. The Italian Venuses from Savignano, Balzi Rossi and Parabita show the same female form, with an enlarged midsection, pronounced buttocks, large breasts and voluminous thighs, characteristic of the vast majority of these figurines. This suggests that there was an extensive flow of people and ideas during the Early Upper Palaeolithic. This is further supported by the transport of exotic materials, such as flint and shells over distances of several hundred kilometres.

At around 29,000 years ago, the Italian archaeological record falls silent. This hiatus seems to last for around two millennia. Mussi (2000) suggests that Italy may have become a “demographic trap”, due to the combination of low population numbers and small group sizes, leading to the eventual extinction of the Aurignacian people.

Based on their assemblages, is believed that the Gravettian people were a new group, who likely entered the Italian peninsula from south-eastern France. As climatic conditions worsen, more exotic artefacts begin to appear in the archaeological record. A major concentration of sites date to around 25,000 BP, suggesting the arrival of immigrants from northern Europe coinciding with the onset of the Last Glacial Maximum. As new people entered Italy, they had to adapt to strange environments and a more limited variety of animals. We no longer find high altitude sites like in the Aurignacian. This shift to lower altitudes is undoubtedly due to the advancing ice. There is evidence of decreased mobility during the Last Glacial Maximum, which continues until the Mesolithic period (Holt 2003). This may reflect a move to more closed systems, as regionalisation grew due to the ever-increasing population density and greater competition over territories. The burial practices, art, and personal adornments of the Early Upper Palaeolithic, which paralleled those from the rest of the continent disappear at the height of the Last Glacial Maximum, further reflecting a shift to more closed social networks. As temperatures begin to improve towards the end of the Last Glacial Maximum, these cultural artefacts reappear in abundance, once again mirroring the styles and practices seen elsewhere in Europe.

Sicily comes in for special attention. The island has long been of interest to anthropologists, who saw the it as a stepping stone between North Africa and Europe. Ferembach (1986) postulated that Sicily may have served as an entry point for the North African Aterians (or their ancestors) around 50,000 years ago, who later became the “Cro-Magnon race”. However, this idea finds little archaeological or skeletal support. Evidence of occupation prior to the Last Glacial Maximum in Sicily is patchy at best. At the site of Fontana Nuova, Aurignacian tools and isolated fragments of human bone have been found and estimated to date to around 30,000 years ago, making it the earliest record of occupation for the island (Chilardi et al 1996). However, it is the only evidence we have for the colonisation of the island during the Aurignacian and it is not until the Epigravettian (~20,000-10,000 BP) that we have further evidence of humans in Sicily. This suggests that the earliest settlers probably went extinct due their small numbers, limited resources and restricted gene flow with the mainland. It also reflects the pattern seen in the rest of Italy, since there are no Aurignacian sites known after 30,000 BP.

At the site of Grotta di San Teodoro the skeletal remains of seven individuals were found, making it the single largest Upper Palaeolithic sample in Sicily. An unpublished direct AMS 14C date situates the skeletal remains at around 14,800 BP (D’Amore et al. 2009). A recent craniometric study of the San Teodoro skeletons shows that they have higher affinities with the Late Upper Palaeolithic and Mesolithic populations, rather than Early Upper Palaeolithic ones (ibid.). These results suggest one of two possible scenarios (a) an early arrival with gene flow, thus explaining the homogeneity with the mainland Italian groups or (b) the late arrival of the direct descendants of the San Teodoro population on the island of Sicily. At the time of the Last Glacial Maximum there was probably a land bridge between Sicily and the Italian mainland, with sea levels being some 120 metres lower. The appearance of exotic faunal evidence further suggest this land connection. While Sicily was occupied as early as the Aurignacian, it may not have been until the Late Epigravettian that the island had a stable population able to overcome the threat of extinction. Like the rest of Italy, Sicily gives us a unique insight the challenges faced by Europe’s latest inhabitants.

*BP is used to indicated uncalibrated radiometric years before present.

References
Bocquet-Appel et al. Estimates of Upper Palaeolithic meta-population size in Europe from archaeological data. Journal of Archaeological Science (2005) vol. 32 (11) pp. 1656-1668.

Chilardi S, Frayer DW, Gioia P, Macchiarelli R, Mussi M (1996) Fontana Nuova di Ragusa (Sicily, Italy): southernmost Aurignacian site in Europe 70: 553-563.

D’Amore G, Marco SD, Tartarelli G, Bigazzi R, Sineo L (2009) Late Pleistocene human evolution in Sicily: comparative morphometric analysis of Grotta di San Teodoro craniofacial remains 56: 537-550.

Ferembach. Les Hommes du Paléolithique Supérieur. Autour du Bassin Méditerraneen. L’Anthropologie (1986) vol. 90 (3) pp. 579-587.

Henke W (1991) Biological Distances in Late Pleistocene and Early Holocene Human Populations in Europe. In: Variability and Evolution. Poznan, Poland: Miskiewicz University Press. pp. 39-64.

Henke (1989) Jungpaläolithiker und Mesolithiker: Beiträge zur Anthropologie. Habilitationsschrift, FB Biologie, Mainz, 1701 S.

Holt BM (2003) Mobility in Upper Paleolithic and Mesolithic Europe: Evidence from the lower limb. Am. J. Phys. Anthropol.

Mussi (1990) Continuity and change in Italy at the Last Glacial Maximum.. In: Soffer, Gamble, editors. The world at 18000: high latitudes. London: . pp. 126-147.

Mussi M (2000) Heading south: the gravettian colonisation of Italy. In: Roebroeks, Mussi, Svoboda, Fennema, editors. Hunters of the Golden Age: The Mid Upper Palaeolithic of Eurasia 30,000–20,000 BP. Leiden: Leiden University Press. pp. 355-374.

var addthis_pub=”4a1c2d6a388b86af”;
Bookmark and Share

Measuring cranial variation using geography as a proxy for neutral genetic distances

Certain anatomical features of the human skeleton are known to vary with geography and climate. To what extent each variable contributes to our physical makeup is less clear. The problem is that populations with similar climate are geographically close to one another. Even if we find shared traits among populations from similar climates it may be just as a result of geographic proximity (and thus clinal gene flow), rather than shared common ancestry.

As I mentioned in my previous post, anthropologists often compared cranial data to matched microsatellite datasets. However, it is rarely possible to get an exact match between the cranial and microsatellite populations. The anthropologist will instead use populations that are genetically similar and which may or may not be representative of the target population. Another option is to substitute microsatellite data with geographic distances, since studies have found a strong correlation between genetic distance and geographic distance (Manica et al. 2005; Ramachandran et al. 2005; Romero et al. 2008). This allows us to get around the need to match phenotypic data with genetic datasets.

A recent paper by Betti et al. used geographic distance as a proxy for neutral genetic distance. They set out to test the extent to which cranial differences can be explained by geographic proximity, by comparing pairwise phenotypic distances among populations and pairwise geographic distances using isolation by distance (IBD) models, as well as comparing pairwise cranial distances with climatic variables after correcting for IBD. Geographic distances were calculated as the shortest distance over land between populations while avoiding areas greater than 2000 metres above sea level. Intercontinental land bridges were also factored into their model.

Their study found geographic distance (and by extension genetic distance) to be a strong predictor of cranial variation. Minimum and maximum temperatures were also a significant predictor of cranial differentiation but not as strong as geographic distance. It also appears that much of this climate-related variation is influenced by the populations from exceptionally cold climates. A previous study by Roseman also found that populations living in extremely cold climates showed greater selection. Betti et al. suggest that this may be due to culture acting as an environmental buffer, with the buffer breaking down at extremely cold climates, after which cranial plasticity takes over.

Since climate and geographic distance covary, not considering isolation by distance leads to an overestimation of the effect of climate on cranial differences between populations. Not surprisingly facial traits showed the strongest correlation with climate. In summary, this study suggests that cranial measurements are predominately influenced by neutral evolutionary processes, especially in populations that do not live in extremely cold climates.

References:
Betti et al. 2009. The relative role of drift and selection in shaping the human skull. Am. J. Phys. Anthropol. in press.

Manica A, Prugnolle F, Balloux F. 2005. Geography is a better determinant of human genetic differentiation than ethnicity. Hum Genet 118:366–371.

Ramachandran S, Deshpande O, Roseman CC, Rosenberg NA, Feldman MW, Cavalli-Sforza LL. 2005. Support from the relationship of genetic and geographic distance in human populations for a serial founder effect originating in Africa. Proc Natl Acad Sci USA 102:15942–15947.

Romero IG, Manica A, Goudet J, Handley LL, Balloux F. 2008. How accurate is the current picture of human genetic variation? Heredity 102:120–126.

var addthis_pub=”4a1c2d6a388b86af”;
Bookmark and Share
Above photo by jacorbett70 under creative commons license.

Examining cranial robusticity

Palaeoanthropologist Darren Curnoe (2009) gives the following biological definition of the term ‘robust’:

…a descriptive anatomical term referring to individuals, complexes, organs, structures or traits which are heavily built, rugged, well defined or corpulent.

Bones tend to more robust where muscles, tendons or ligaments insert into the periosteum. When these insertion sites are subjected to stress, blood flow increases. This in turn stimulates the production of osteoblasts, which lay down extra bone. With respect to the skull the term robust is generally used to refer to so-called superstructures, such as the supraorbital ridges, occipital crests or zygomaxillary tuberosities. Anthropologists often classify robusticity based on the relative expression of a particular trait, or indeed its absence. Given that robusticity is related to physical stress, traits tend to be more pronounced in males and in certain populations (e.g. Aboriginal Australians and Fuegians).

The retention of robust features in certain populations, particularly Aboriginal Australians, has been used to support the multiregional hypothesis of human origins (e.g. Wolpoff et al. 2001; Frayer et al., 1993). On the other hand, proponents of a replacement model see robust traits (e.g. in Australian Aboriginal populations) as retained plesiomorphies and argue that these traits cannot be used to show continuity (Lieberman 2000). In response, many multiregionalists have revised their position to suggest that the reduction of the browridge in later Neandertals, such as St Césaire and Vindija, represents a synapomorphy between Neandertals and modern humans, likely due to interbreeding. The underlying assumption here is that these robust traits have a strong genetic component. Furthermore, there is a notable decrease in cranial robusticity from the early Upper Palaeolithic to late Upper Palaeolithic. It has been suggested that this may reflect changes in diet. Transition from hunter-gather to agricultural lifestyle is associated with a reduction in cranial robusticity, although correlation does not necessarily prove causation. However, not all hunter-gather groups are universally more robust than argriculturalists, which might suggest some other factors at play.

A recent in press paper by Baab et al. sets out to examine the possible mechanisms behind robust cranial characters. The null hypothesis in their study is that neutral evolutionary processes (e.g. genetic drift) were responsible for the pattern of cranial robusticity in modern humans – the rejection of which would suggest selection acting on these traits. To test the null hypothesis of neutral evolution of cranial robusticity Mahalonobis D2 distances for robust characters were compared to Ddm distances derived from microsatellite data. Microsatellites are useful in reconstructing evolutionary relationships due to their unusually high mutation rates, which result in largely selectively neutral polymorphisms.

Of the variables examined, only cranial shape was significantly correlated with robusticity, while cranial size, climate and neutral genetic distances were not. This is at odds with an earlier study by Mirazón Lahr and Wright (1996) (1996) who found the strongest correlation between cranial robusticity and cranial size. This finding may be due to use of geometric morphometrics by Baab and colleagues, which is better at separating size and shape compared to the linear morphometrics used by Mirazón Lahr and Wright (1996).

Cranial robusticity was not correlated with neutral genetic distances, suggesting that neutral evolutionary processes (e.g. genetic drift) were not responsible for the pattern of cranial robusticity in the populations studied. As noted by the authors, this finding could also be explained by a non-perfect match of populations among some of the cranial and molecular samples. In studies such as this one, it is often difficult to find an exact match between the populations from which we derive our cranial and molecular data. In such cases, we are left with the choice of eliminating samples or using another genetically similar population. The authors choose the latter but neither option is ideal and both have their own disadvantages. Unfortunately, the reason for including Upper Palaeolithic and Neolithic samples in this study is never fully explained and the assumption that modern genetic populations are appropriate proxies for such populations is never justified. Setting this aside, the findings of this study caution the use of robust traits in constructing phylogenetic relationships in modern humans.

The strongest correlations were found between cranial robusticity and either cranial or masticatory shape. This lends support to the hypothesis that robusticity is in some part functionally determined. The study also found crania with more prognathic faces, longer skulls, expanded glabellar and occipital regions to be more robust. Mirazón Lahr and Wright (1996) noted a similar tendency of longer skulls to have superstructures, while further emphasising their tendency to be associated with narrow skulls and a large palatal region.

While most of the robust variables in this study were areas of muscle insertions, the supraorbital region has a distinct aetiology. While many have interpreted the supraorbital region as an area of stress reinforcement in the skull (the so-called beam model) which is strongly influenced by mastication (Endo 1966, 1970; Russell 1985), there is a strong evidence to suggest that this is not its primary purpose. Supraorbital development begins early in life, suggesting that the supraorbital ridge may be part of the overall craniofacial complex and is likely under genetic control. While the beam model is intuitive, it is unsupported by empirical data. Hylander and colleagues (Hylander et al. 1991a, 1991b, 1992; Hylander and Ravosa 1992) conducted in vivo strain gauge experiments in different primates to assess the amount of strain magnitudes generated during mastication. They found these levels to be low to induce bone deposition in all the species they studied, even when chewing hard food. Moreover, anthropoids do not show a correlation between the browridge and the moment arms of the masticatory muscles, as the beam model would predict (Ravosa 1991). These researchers adopt the model proposed by Moss and Young (1960), which views supraorbital development as the result of placement of the brain and eyes. They postulated that the reduction of the brow ridge in modern humans was related to the expansion of the frontal lobe in our species. In hominins with orbits positioned well in front of the frontal lobes, as in chimpanzees or the erectines, the space between the orbits and the brain case is bridged by a brow ridge. If the supraorbital region is under genetic control, as the research of Hylander and Ravosa suggests, it would be of interest to examine this region in isolation to assess if it correlates with neutral evolutionary processes, particularly in light of a recent paper by Von Cramen-Taubedal which found the shape of the frontal bone to be consistent with neutral genetic expectation.

References

Baab KL, SE Freidline, SL Wang, T Hanson. 2009. Relationship of cranial robusticity to cranial form, geography and climate in Homo sapiens (in press). Am. J. Phys. Anthropol.

Curnoe D. 2009. Possible causes and significance of cranial robusticity among Pleistocene-Early Holocene Australians. Journal of Archaeological Science (2009) vol. 36 (4): 980-990.

Endo B. 1966. Experimental studies on the mechanical significance of the form of the human facial skeleton. J Faculty Sci Univ Tokyo (Section V, Anthropol) 3:1–106.

Endo B. 1970. Analysis of stress around the orbit due to masseter and temporalis muscles respectively. J Anthropol Soc Nippon 78:251–266.

Frayer DW, MH Wolpoff, AG Thorne, FH Smith, GG Pope. Theories of modern human origins: the paleontological test. American Anthropologist (1993) vol. 95 (1): 14-50.

Hylander WL, Picq PG, Johnson KR. 1991a. Masticatory–stress hypotheses and the supraorbital region of primates. Am J Phys Anthropol 86:1–36.

Hylander WL, Picq PG, Johnson KR. 1991b. Function of the supraorbital region of primates. Arch Oral Biol 36:273– 281.

Hylander WL, Ravosa MJ. 1992. An analysis of the supraorbital region of primates: a morphometric and experimental approach. In: Smith P, Tchernov E,
editors. Structure, function and evolution of teeth. Tel Aviv: Freund Publishing. p 223–255.

Lieberman, DE. (2000) Ontogeny, homology, and phylogeny in the Hominid craniofacial skeleton: the problem of the browridge. In P. O’Higgins and M. Cohn (eds.) Development, Growth and Evolution: implications for the study of hominid skeletal evolution. London: Academic Press, pp. 85-122.

Moss ML, RW Young. 1960. A functional approach to craniology. Am. J. Phys. Anthropol. 18:281-292.

Mirazón Lahr M, RVS Wright. 1996. The question of robusticity and the relationship between cranial size and shape in Homo sapiens. Journal of Human Evolution.

Ravosa MJ. 1991. Interspecific perspective on mechanical and nonmechanical models of primate circumorbital morphology. Am J Phys Anthropol. 86(3):369-96.

Russell MD. 1985. The Supraorbital Torus:” A Most Remarkable Peculiarity”. Current Anthropology. vol. 26 (3) pp. 337

Wolpoff MH, J Hawks, DW Frayer, K Hunley. 2001. Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory. Science. vol. 291 (5502):293-297.

Bookmark and Share
Above photo modified from original by Thomas Hawk under creative commons license.

Why the Aquatic Ape Hypothesis doesn’t hold water

Among this week’s new videos from TED, was a talk given by Elaine Morgan – the chief promoter of the Aquatic Ape Hypothesis (AAH). The AAH was first formulated by Alister Hardy and is the idea that human evolution went through an aquatic stage, which in turn explains many of the features of the human physiology. For anybody with a poor understanding of evolutionary biology the AAH arguments can seem quite compelling. Instead of repeating the numerous reasons why the AAH fails (Jim Moore has an entire website dedicated to this), I wish to address some of the specific arguments made in this video.

Morgan starts off my stating that “… there’s one aspect of this story which they [evolutionists] have thrown no light on and they seem anxious to skirt around and step over it and talk about something else. So I’m going to talk about it. It’s the question of why are we so different from the chimpanzees?”. Either Morgan has not been reading the hundreds of research papers that have addressed these important questions or she is trying to hoodwink her audience. Palaeoanthropologists and primatologists have long recognised the value of studying human and chimp differences in order to understand our shared evolutionary history. In fact, it is impossible to talk about functional anatomy and phylogenetic history in humans without reference to our closest extinct and living hominin relatives.

She continues: “Yet when you look at the phenotypes. There’s a chimp, there’s a man. They’re astoundingly different. No resemblance at all.” I am hearing this correctly? No resemblance at all? Even a five year old can see the striking similarities between chimpanzees and humans. To suggest that there is “no resemblance at all” is laughable. It was the similarities of humans to other non-human primates that led Charles Darwin to argue for common ancestry between humans and the great apes. Even without the fossil record and the unambiguous molecular evidence, the morphological similarities alone would be enough to suggest a shared common ancestry of chimps and humans.

Throughout the talk she constantly refers to humans as naked, as if to suggest we are hairless. One need only look at a shirtless Alec Baldwin, Robin Williams or Andy Garcia to know this is not the case. While it is true that humans are less hairy than the rest of our primate kin there are far more compelling hypotheses to explain our lack of hair (thermoregulation, defence against parasites or sexual selection). She claims that hairlessness is an aquatic trait when in fact most aquatic/semi-acquatic mammals are in fact hairy. Otters, polar bears, seals, and walruses are but some examples that spring to mind.

Regarding the failed savannah hypothesis of bipedalism Morgan has this to say: “What do scientists do when a paradigm fails… carry on as though nothing had ever happened… If they haven’t got a paradigm they can’t ask the questions… The only other option open to them is to stop asking the questions. So that is what they have done now. That is why you don’t hear them talking about it.”

When paradigms fail science marches on. When was the last time you heard a scientist defending the merits of Lamarckism, psychoanalysis and phrenology? Morgan is correct that the savannah hypothesis doesn’t weigh up against the evidence but she mistakenly claims that the anthropologists haven’t let go of this idea. The savannah hypothesis was formulated in a time when there was a dearth of palaeoecological data for the most important African archaeological sites. As more data came in, anthropologists changed their models correspondingly. No serious anthropologist still adheres to the savannah hypothesis. Morgan chooses to ignore this fact, instead preferring the easier route of attacking a strawman. In the references section below, you will find just a spattering of the work anthropologists have been doing on palaeoecological reconstructions of the environments occupied by our forebearers. These papers address the very questions Morgan asserts that scientists have stopped asking. Does she not read the anthropological literature or does she just choose to ignore it? She implies that because the savannah hypothesis is false it somehow offers support to the AAH. In fact, the consensus opinion suggests that neither savannah nor aquatic environments were very important in the early stages of human evolution, but rather our hominin ancestors exploited wood and forest habitats. A number of anthropologists have proposed an arboreal origin of hominin bipedality (Crompton 2008; Pickerford 2006; Senut 2003, 2006). In fact, the best known human ancestor Lucy shows clear arboreal adaptations.

Morgans proceeds by stating that “there is only one circumstance in which they always, all of them [non human primates], walk on two legs and that is when they are wading through water.” Contrary to Morgan’s claim, the data have shown apes to be bipedal more often on land than in the water.

She follows this up by saying that the fat in humans is similar to that seen in aquatic mammals. Humans have a similar number of fat cells compared with other primates. The increased subcutaneous fat seen in humans is most likely a result of diet rather than an evolutionary adaptation. Non-human primate obesity is well documented, particularly in primates kept in captivity (Videan 2007; Altmann et al 1993; Kemnitz et al 1989; Schwartz et al 1993). Moreover, the distribution of fat in humans runs contrary to need aquatic mammals have for streamlining.

Ten minutes into the talk she states that “the only creatures that have got conscious control of their breath are the diving animals and the birds”. Humans are not the only non-aquatic mammal which can hold its breath. Various monkeys, for instance, can and do hold their breath, as well as dogs.

Finally, she asserts that “we are streamlined.” Humans are anything but streamlined. Our motion in the water is generally quite wasteful. Ask any swimming coach. Fish have a fusiform shape (tapered at both ends), which is ideal for moving through the water with the least amount of resistance. Let’s put this into perspective. The sailfish records speeds of up to 116 km/hr (72 mph), while Michael Phelps can average a measly about 6.5 km/hr (4 mph) on a good day! Our body shape is a consequence of our adaptation to bipedalism, the requirements of childbirth in women, sexual dimorphism and sexual selection.

While I generally enjoy listening to the speakers at TED, I think this is an idea NOT worth spreading.

References

Altmann J, Alberts SC, Altmann SA, Roy SB (2002) Dramatic change in local climate patterns in the Amboseli Basin, Kenya. Afr J Ecol 40, 248–251.

Altmann J, Schoeller D, Altmann SA, Muruthi P, Sapolsky RM (1993) Body size and fatness of free-living baboons reflect food availability and activity levels. Am J Primatol 30: 149–61.

Andrews P (1996) Palaeoecology and hominoid palaeoenviron-ments. Biol Rev 71, 257–300.

Andrews P, Humphrey L (1999) African Miocene environments and the transition to early hominines. In African Biogeography, Climate Change and Early Hominid Evolution (eds Bromage TG, Schrenk F), pp. 282–300. New York: Oxford University Press.

Andrews P (2007) The biogeography of hominid evolution. J Biogeogr 34, 381–382.

Andrews P, Kelley J (2007) Middle Miocene dispersals of apes. Folia Primatol 78, 328–343.

Andrews P, Bamford M (2008) Past and present vegetation ecology of Laetoli, Tanzania. J Hum Evol 54, 78–98.

Codron D, Luyt J, Lee-Thorp JA, Sponheimer M, De Ruiter D, Codron J (2005) Utilization Of Savanna-Based Resources By Plio-Pleistocene Baboons. S Afr J Sci 101, 245–248.

Crompton RH, EE Vereecke, SKS Thorpe (2008) Locomotion and posture from the common hominoid ancestor to fully modern hominins, with special reference to the last common panin/hominin ancestor. J Anat 212, 501–543.

Demenocal PB (2004) African Climate Change And Faunal Evolution During The Pliocene-Pleistocene. Earth Planet Sci Lett 220, 3–24.

Denton G (1999) Cenozoic climate change. In African Biogeography, Climate Change and Early Hominid Evolution (eds Bromage TG, Schrenk F), pp. 94–114. New York: Oxford University Press.

Dowsett HJ, Barron JA, Poore RZ, et al. (1999) Middle Pliocene paleoenvironmental reconstruction: PRISM2. US Geol Surv, Reston, Va, Open File Rep 99–535.

Elton S (2000) Ecomorphology and evolutionary biology of African Cercopithecoids: providing an ecological context for hominin evolution. PhD dissertation, University of Cambridge.

Jacobs BF (2004) Palaeobotanical studies from tropical Africa: relevance to the evolution of forest, woodland and savannah biomes. Phil Trans R Soc Lond B359, 1573–1583.

Kemnitz JW, Goy RW, Flitsch TJ, Lohmiller JJ, Robinson JA (1989) Obesity in male and female rhesus monkeys: fat distribution, glucoregulation, and serum androgen levels. J Clin Endocrinol Metab 69:287–93.

Kingston J, Harrison T (2007) Isotopic dietary reconstructions of Pliocene herbivores at Laetoli: implications for hominin paleo- ecology. Palaeogeog Palaeoclimatol Palaeoecol 243, 272–306.

Kovarovic KM, Andrews P, Aiello L (2002). The palaeoecology of the Upper Ndolanya Beds, Laetoli, Tanzania. J Hum Evol 43, 395–418.

Pickford M (2006) Paleoenvironments, Paleoecology, Adaptations and the Origins of Bipedalism in Hominidae. In Human Origins and Environmental Backgrounds (eds Ishida H, Tuttle RH, Pick- ford M, Ogihara M, Nakatsukasa M), pp. 175–198. Heidelberg: Springer.

Schwartz SM, Kemnitz JW, Howard CF Jr (1993) Obesity in free-ranging rhesus macaques. Int J Obes 17:1–9.

Senut B (2003) Palaeontological approach to the evolution of hominid bipedalism: the evidence revisited. Cour Forsch-Inst Senkenberg 243, 125–134.

Senut B (2006) Arboreal Origins of Bipedalism. In Human Origins and Environmental Backgrounds (eds Ishida H, Tuttle RH, Pickford M, Ogihara N, Nakatsukasa M), pp. 199–208. Heidelberg: Springer.

Sikes N (1999) Plio-Pleistocene floral context and habitat prefer- ences of sympatric hominid species in East Africa. In African Bio- geography, Climate Change and Early Hominid Evolution (eds Bromage TG, Schrenk F), pp. 301–315. New York: Oxford Univer- sity Press.

Videan EN, J Fritz, J Murphy (2007) Development of guidelines for assessing obesity in captive chimpanzees (Pan troglodytes). Zoo Biology 26: 93–104.

Vincens A, Garcin Y, Buchet G. (2007) Influence of rainfall seasonality on African lowland vegetation during the Late Quaternary: pollen evidence from Lake Masoko, Tanzania. J Biogeogr 34, 1274–1288.

WoldeGabriel G, Haile-Selassie Y, Renne P, et al. (2001) Geology and palaeontology of the Late Miocene Middle Awash valley, Afar rift, Ethiopia. Nature 412, 175–178.

Bookmark and Share