The carbon isotope ecology and diet of Australopithecus africanus at Sterkfontein,South Africa

The stable carbon isotope ratio of fossil tooth enamel carbonate is determined by the photosynthetic systems of plants at the base of the animal's foodweb. In subtropical Africa, grasses and many sedges have C(4)photosynthesis and transmit their characteristically enriched 13C/(12)C ratios (more positive delta13C values) along the foodchain to consumers. We report here a carbon isotope study of ten specimens of Australopithecus africanus from Member 4, Sterkfontein (ca. 2.5 to 2.0Ma), compared with other fossil mammals from the same deposit. This is the most extensive isotopic study of an early hominin species that has been achieved so far. The results show that this hominin was intensively engaged with the savanna foodweb and that the dietary variation between individuals was more pronounced than for any other early hominin or non-human primate species on record. Suggestions that more than one species have been incuded in this taxon are not supported by the isotopic evidence. We conclude that Australopithecus africanus was highly opportunistic and adaptable in its feeding habits.     

C4 photosynthetic isotope exchange in NAD-ME- and NADP-ME-type grasses

Monitoring photosynthetic isotope exchange is an important tool for predicting the influence of plant communities on the global carbon cycle in response to climate change. C(4) grasses play an important role in the global carbon cycle, but their contribution to the isotopic composition of atmospheric CO(2) is not well understood. Instantaneous measurements of (13)CO(2) (Delta(13)C) and C(18)OO (Delta(18)O) isotope exchange in five NAD-ME and seven NADP-ME C(4) grasses have been conducted to investigate the difference in photosynthetic CO(2) isotopic fractionation in these subgroups. As previously reported, the isotope composition of the leaf material (delta(13)C) was depleted in (13)C in the NAD-ME compared with the NADP-ME grasses. However, Delta(13)C was not different between subtypes at high light, and, although Delta(13)C increased at low light, it did so similarly in both subtypes. This suggests that differences in leaf delta(13)C between the C(4) subtypes are not caused by photosynthetic isotope fractionation and leaf delta(13)C is not a good indicator of bundle sheath leakiness. Additionally, low carbonic anhydrase (CA) in C(4) grasses may influences Delta(13)C and should be considered when estimating the contribution of C(4) grasses to the global isotopic signature of atmospheric CO(2). It was found that measured Delta(18)O values were lower than those predicted from leaf CA activities and Delta(18)O was similar in all species measured. The Delta(18)O in these C(4) grasses is similar to low Delta(18)O previously measured in C(4) dicots which contain 2.5 times the leaf CA activity, suggesting that leaf CA activity is not a predictor of Delta(18)O in C(4) plants.     

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。     

A large male hominin cranium from Sterkfontein, South Africa, and the status ofAustralopithecus africanus

Stw 505 is the most complete hominin cranium discovered in Sterkfontein Member 4 since Broom's excavations. It was found in situ in Member 4 breccia in 1989 and is larger, on the whole, than any other cranium from Sterkfontein that has comparable parts. Displacement due to breakage, as well as plastic deformation, has affected Stw 505 in several areas, especially the face and the vault. Diagnosticmorphology is nevertheless abundant in the specimen. In several areas-the distinct anterior pillar, the straight inferior border of the zygoma, the pattern of cresting on the naso-alveolar clivus, the basal aspect of the temporal bone-Stw 505 closely matches the morphology of specimens of Australopithecus africanus and is distinct from other hominins. Some isolated characters overlap with other groups, mainly early Homo and/or A. robustus. However, only the hypodigm of A. africanus can accommodate the entire suite of morphology.In some cases, Stw 505 introduces more variation into the Sterkfontein sample. For example, prominent superciliary eminences occupy the medial portions of the supraorbital region and flow medially into a strongly protruding glabellar mound. These characteristics are probably attributable to sexual dimorphism. In many respects, Stw 505 highlights similarities between A. africanus and early Homo. Comparison with other species suggests that males of A. africanus do not show derived features of A. robustus that are not also present in females, and that cranial differences between A. afarensis and A. africanus have, if anything, been understated.     

The hunters and the hunted revisited

The dietary niches of extinct animals, including hominids and predators, may be constrained using stable carbon isotope ratios in fossil tooth enamel.(13)C/(12)C ratios of many of the primates abundant in the faunal assemblages of Members 1 and 2 at Swartkrans, including cercopithecoids and Australopithecus (Paranthropus) robustus, and a range of other possible prey species, have been reported previously. Resulting suggestions of a mixed, or omnivorous, diet for A. robustus raise questions about niche overlap with coeval, larger brained Homo. Here we present(13)C/(12)C data from Homo and several large predators including Panthera pardus, Dinofelis sp., Megantereon cultridens and Chasmoporthetes nitidula in Member 1, and P. pardus and P. leo in Member 2, in order to compare the two hominid species and to determine likely predators of the various primates and other macrovertebrates. Results for three Homo cf. ergaster individuals are indistinguishable from those of A. robustus, showing that proportions of C(3)- and C(4)-based foods in their diets did not differ. P. pardus, Megantereon and Crocuta are shown to be likely predators of the hominids and Papio baboons in Member 1, while the Dinofelis individual concentrated on prey which consumed C(4)grasses. The hunting hyaenid C. nitidula preyed on either mixed feeders or on a range of animals across the spectrum of C(3)and C(4)variation. The data from Members 1 and 2 confirm a shift in leopard diets towards animals that consumed C(4)grasses.     

Muscle delta13C change in Nile tilapia (Oreochromis niloticus) fingerlings fed on C3-or C4-cycle plants grain-based diets

The effects of grain-based diets from C3 or C4-cycle plants on muscle delta(13)C change process in Nile tilapia (Oreochromis niloticus) fingerlings were investigated. Two groups of sex reversal males Nile tilapia fingerlings were fed with isoproteic (32.0% DP) and isocaloric (3200 kcal DE/kg) diets, differing from each other by their delta(13)C. Muscle samples were collected and the carbon isotopic composition was measured. For C4 diet, the formula for the muscle delta(13)C change related to the intake time of a new diet was delta(13)C=-14.88-9.21e(-0.0209t) and the half-life (T) of the muscle carbon was 33.2 days. For C3 diet, the formula was delta(13)C=-25.43+8.59e(-0.0533t) with T=13 days. The C3 diet was considered more appropriate based on its palatability and consequent larger food intake than the C4 diet, resulting in an increased muscle delta(13)C change rate. However, for future studies, would be necessary to mix both the C3 and C4 feedstuffs to formulate diets nutritionally appropriated, with contrasting stable isotopes signatures. Tissue delta(13)C change rate is therefore indicated as a promising tool to better understand the biotic and abiotic factors that influence nutrients utilization from the diet and animal growth.     

Hard-object feeding in sooty mangabeys (Cercocebus atys) and interpretation of early hominin feeding ecology

Morphology of the dentofacial complex of early hominins has figured prominently in the inference of their dietary adaptations. Recent theoretical analysis of craniofacial morphology of Australopithecus africanus proposes that skull form in this taxon represents adaptation to feeding on large, hard objects. A modern analog for this specific dietary specialization is provided by the West African sooty mangabey, Cercocebus atys. This species habitually feeds on the large, exceptionally hard nuts of Sacoglottis gabonensis, stereotypically crushing the seed casings using their premolars and molars. This type of behavior has been inferred for A. africanus based on mathematical stress analysis and aspects of dental wear and morphology. While postcanine megadontia, premolar enlargement and thick molar enamel characterize both A. africanus and C. atys, these features are not universally associated with durophagy among living anthropoids. Occlusal microwear analysis reveals complex microwear textures in C. atys unlike those observed in A. africanus, but more closely resembling textures observed in Paranthropus robustus. Since sooty mangabeys process hard objects in a manner similar to that proposed for A. africanus, yet do so without the craniofacial buttressing characteristic of this hominin, it follows that derived features of the australopith skull are sufficient but not necessary for the consumption of large, hard objects. The adaptive significance of australopith craniofacial morphology may instead be related to the toughness, rather than the hardness, of ingested foods.     

Understanding the ecology of mammalian carnivorans and herbivores from Valdegoba cave (Burgos, northern Spain) through stable isotope analysis

Stable carbon and oxygen isotope analyses of mammalian carnivoran and herbivore species from the late Pleistocene Valdegoba cave site in northern Spain imply competition and partitioning in resource use. In general, the data support the previously recognized ecology for the analyzed species. δ¹³C values show that the ecosystem around the cave was dominated by C₃ plants. The observed δ¹⁸O values are similar to what is found in modern environments. The analyzed bovids, Bos primigenius, Capra pyrenaica, and Rupicapra rupicapra, showed the most positive δ¹³C values. Bos primigenius had the most positive mean carbon isotope value and is suggested to feed on grasses in open environments. Values for Capra pyrenaica primarily indicate grass feeding, while Rupicapra rupicapra had the widest diet for the analyzed species, likely including grass and browse. Cervus elaphus, Equus ferus, Equus hydruntinus, and Stephanorhinus hemitoechus displayed more negative δ¹³C values indicating the use of similar resources. The smallest species analyzed, Castor fiber, displayed the most negative δ¹³C and δ¹⁸O values, implying a preference for eating C₃ plants and being semi-aquatic. The canids, Canis lupus and Vulpes vulpes, displayed the most positive δ¹³C and δ¹⁸O values, and overlap many of the sampled ungulate species. Positive δ¹⁸O values in canids implies that this group obtains much of its water from its prey, uses a different water source, or has physiological differences from the other carnivorans that influence oxygen isotope values. Lynx pardinus had values similar to the canids. Crocuta crocuta had δ¹³C values more negative than expected for a generalist predator. These values are likely due to concentration of diet on taxa from more forested environments. The most negative δ¹³C values are observed in the bears, Ursus arctos and Ursus spelaeus. These values are the likely result of hibernation and the inclusion of significant vegetation in bear diets.     

Compound-specific deltaD-delta13C analyses of n-alkanes extracted from terrestrial and aquatic plants

Stable hydrogen and carbon isotopic compositions of individual n-alkanes were determined for various terrestrial plants (33 samples including 27 species) and aquatic plants (six species) in natural environments from Japan and Thailand. In C3 plants, n-alkanes extracted from angiosperms have a deltaD value of -152+/-26 per thousand (relative to Standard Mean Ocean Water [SMOW]) and delta13C value of -36.1+/-2.7 per thousand (relative to Peedde Belemnite [PDB]), and those from gymnosperms have a deltaD value of -149+/-16 per thousand and delta13C value of -31.6+/-1.7 per thousand. Angiosperms have n-alkanes depleted in 13C relative to gymnosperms. n-Alkanes from C4 plants have a deltaD value of -171+/-12 per thousand and delta13C value of -20.5+/-2.1 per thousand, being a little depleted in D and much enriched in 13C compared to C3 plants. n-Alkanes of CAM plants are a little depleted in D and vary widely in delta13C relative to those of C3 and C4 plants. In aquatic plants, n-alkanes from freshwater plants have a deltaD value of -187+/-16 per thousand and delta13C value of -25.3+/-1.9 per thousand, and those from seaweeds have a deltaD value of -155+/-34 per thousand and delta13C value of -22.8+/-1.0 per thousand. All n-alkanes from various plant classes are more depleted in D and 13C relative to environmental water and bulk tissue, respectively. In addition, the hydrogen and carbon isotopic fractionations during n-alkane synthesis are distinctive for these various plant classes. While C3 plants have smaller isotopic fractionations in both D and 13C, seaweed has larger isotopic fractionations.     

Stable isotopic variations in modern herbivore tooth enamel, plants and water on the Tibetan Plateau: Implications for paleoclimate and paleoelevation reconstructions

Stable isotope analyses of fossil teeth and other authigenic minerals have been used to reconstruct the paleoenvironment and paleoelevation of the Tibetan Plateau. The accuracy of such reconstructions is limited by the lack of a comprehensive modern comparative database from the region. We analyzed the carbon and oxygen isotopic compositions (δ¹³C and δ¹⁸O values) of tooth enamel from modern herbivores, the δ¹³C values of grasses and the δ¹⁸O values of water samples collected from various elevations within the Tibetan Plateau to examine their relationships with modern environment/elevation. The δ¹³C values of enamel samples from horses, yaks and goats display a narrow range of variation, with a mean of − 10.7 ± 1.4‰ (n = 301), indicating that these modern herbivores were feeding predominantly on C3 plants, consistent with the current dominance of C3 vegetation in the region. Some of the samples have δ¹³C values between − 7.3 and − 10‰. Although these higher δ¹³C values could suggest consumption of some C4 plants by the animals, the lack of significant seasonal δ¹³C variations within individual teeth indicates that these higher enamel δ¹³C values are due to consumption of C3 plants experiencing water stress and/or some CAM plants rather than C4 plants. Our data show that the conservative “cut-off” δ¹³C value for a pure C3 diet within the Tibetan Plateau should be − 8‰ for modern herbivores and − 7‰ (or even − 6.5‰) for fossils if the region was as arid in the past as today. In contrast to the small intra-tooth δ¹³C variations within individual teeth, serial enamel samples display large intra-tooth δ¹⁸O variations, reflecting seasonal variations in the δ¹⁸O of meteoric water. The mean δ¹⁸O values of tooth enamel from yaks and horses show a strong correlation with water δ¹⁸O values, confirming that the δ¹⁸O of tooth enamel from obligate drinker generally tracks the δ¹⁸O of meteoric water. Unfortunately, elevation alone cannot explain most of the variance in the δ¹⁸O of precipitation and tooth enamel, suggesting that quantitative reconstruction of the paleoelevation of the Tibetan Plateau using re-constructed δ¹⁸O values of paleo-meteoric water from fossil enamel or other oxygen-bearing minerals is not warranted. For a given environment, horses have the lowest enamel–δ¹⁸O values while goats display the highest enamel–δ¹⁸O values among the species studied. The large inter-species δ¹⁸O variations are likely due to differences in physiology and diet/drinking behavior of the animals. This underscores the importance of species-specific studies when interpreting δ¹⁸O data of fossil mammalian teeth in a stratigraphic sequence as a record of paleoclimate changes.     

Stable isotopes in fossil hominin tooth enamel suggest a fundamental dietary shift in the Pliocene

Accumulating isotopic evidence from fossil hominin tooth enamel has provided unexpected insights into early hominin dietary ecology. Among the South African australopiths, these data demonstrate significant contributions to the diet of carbon originally fixed by C₄ photosynthesis, consisting of C₄ tropical/savannah grasses and certain sedges, and/or animals eating C₄ foods. Moreover, high-resolution analysis of tooth enamel reveals strong intra-tooth variability in many cases, suggesting seasonal-scale dietary shifts. This pattern is quite unlike that seen in any great apes, even ‘savannah’ chimpanzees. The overall proportions of C₄ input persisted for well over a million years, even while environments shifted from relatively closed (ca 3 Ma) to open conditions after ca 1.8 Ma. Data from East Africa suggest a more extreme scenario, where results for Paranthropus boisei indicate a diet dominated (approx. 80%) by C₄ plants, in spite of indications from their powerful ‘nutcracker’ morphology for diets of hard objects. We argue that such evidence for engagement with C₄ food resources may mark a fundamental transition in the evolution of hominin lineages, and that the pattern had antecedents prior to the emergence of Australopithecus africanus. Since new isotopic evidence from Aramis suggests that it was not present in Ardipithecus ramidus at 4.4 Ma, we suggest that the origins lie in the period between 3 and 4 Myr ago.     

Dietary and environmental reconstruction with stable isotope analyses of herbivore tooth enamel from the Miocene locality of Fort Ternan, Kenya

Tooth enamel of nine Middle Miocene mammalian herbivores from Fort Ternan, Kenya, was analyzed for δ¹³C and δ¹⁸O. The δ¹⁸O values of the tooth enamel compared with pedogenic and diagenetic carbonate confirm the use of stable isotope analysis of fossil tooth enamel as a paleoenvironmental indicator. Furthermore, the δ¹⁸O of tooth enamel indicates differences in water sources between some of the mammals. The δ¹³C values of tooth enamel ranged from −8·6–−13·0‰ which is compatible with a pure C₃diet, though the possibility of a small C₄fraction in the diet of a few of the specimens sampled is not precluded. The carbon isotopic data do not support environmental reconstructions of a Serengeti-typed wooded grassland with a significant proportion of C₄grasses. This study does not preclude the presence of C₃grasses at Fort Ternan; it is possible that C₃grasses could have had a wider geographic range if atmospheric CO₂levels were higher than the present values.     

Cost of reproduction in the Queensland fruit fly: Y-model versus lethal protein hypothesis

The trade-off between lifespan and reproduction is commonly explained by differential allocation of limited resources. Recent research has shown that the ratio of protein to carbohydrate (P : C) of a fly''s diet mediates the lifespan–reproduction trade-off, with higher P : C diets increasing egg production but decreasing lifespan. To test whether this P : C effect is because of changing allocation strategies (Y-model hypothesis) or detrimental effects of protein ingestion on lifespan (lethal protein hypothesis), we measured lifespan and egg production in Queensland fruit flies varying in reproductive status (mated, virgin and sterilized females, virgin males) that were fed one of 18 diets varying in protein and carbohydrate amounts. The Y-model predicts that for sterilized females and for males, which require little protein for reproduction, there will be no effect of P : C ratio on lifespan; the lethal protein hypothesis predicts that the effect of P : C ratio should be similar in all groups. In support of the lethal protein hypothesis, and counter to the Y-model, the P : C ratio of the ingested diets had similar effects for all groups. We conclude that the trade-off between lifespan and reproduction is mediated by the detrimental side-effects of protein ingestion on lifespan.     

Shallow-water habitats as sources of fallback foods for hominins

Underground storage organs (USOs) have been proposed as critical fallback foods for early hominins in savanna, but there has been little discussion as to which habitats would have been important sources of USOs. USOs consumed by hominins could have included both underwater and underground storage organs, i.e., from both aquatic and terrestrial habitats. Shallow aquatic habitats tend to offer high plant growth rates, high USO densities, and relatively continuous USO availability throughout the year. Baboons in the Okavango delta use aquatic USOs as a fallback food, and aquatic or semiaquatic USOs support high-density human populations in various parts of the world. As expected given fossilization requisites, the African early- to mid-Pleistocene shows an association of Homo and Paranthropus fossils with shallow-water and flooded habitats where high densities of plant-bearing USOs are likely to have occurred. Given that early hominins in the tropics lived in relatively dry habitats, while others occupied temperate latitudes, ripe, fleshy fruits of the type preferred by African apes would not normally have been available year round. We therefore suggest that water-associated USOs were likely to have been key fallback foods, and that dry-season access to aquatic habitats would have been an important predictor of hominin home range quality. This study differs from traditional savanna chimpanzee models of hominin origins by proposing that access to aquatic habitats was a necessary condition for adaptation to savanna habitats. It also raises the possibility that harvesting efficiency in shallow water promoted adaptations for habitual bipedality in early hominins. Am J Phys Anthropol 140:630–642, 2009. © 2009 Wiley-Liss, Inc.     

Muscle delta13C change in Nile tilapia (Oreochromis niloticus): effects of growth and carbon turnover

The contribution of growth and turnover to the muscle delta(13)C change process was investigated using mathematical models which associate delta(13)C change to time of intake of a new diet or increase in body mass. Two groups of Nile tilapia (Oreochromis niloticus) were fed on diets based on C3 (delta(13)C=-25.64+/-0.06 per thousand) or C4 (delta(13)C=-16.01+/-0.06 per thousand) photosynthetic cycle plants to standardize the muscle delta(13)C. After establishing the carbon isotopic equilibrium, fish (mean mass 24.12+/-6.79 g) then received the other treatment diet until a new carbon isotopic equilibrium could be established, characterizing T1 (C3-C4) and T2 (C4-C3) treatments. No significant differences were observed in fish productive performance. Good fits were obtained for the models that associated the delta(13)C change to time, resulting in carbon half-life values of 23.33 days for T1 and 25.96 days for T2. Based on values found for the muscle delta(13)C change rate from growth (0.0263 day(-1) and 0.0254 day(-1)) and turnover (0.0034 day(-1) and 0.0013 day(-1)), our results indicate that most of the delta(13)C change could be attributed to growth. The application of model that associated the delta(13)C change to body mass increase seems to produce results with no apparent biological explanation. The delta(13)C change rate could directly reflect the daily ration and growth rate, and consequently the isotopic change rates of carbon and other tissue elements can be properly used to assess different factors that may interfere in nutrient utilization and growth.     

Kangaroo tooth enamel oxygen and carbon isotope variation on a latitudinal transect in southern Australia: implications for palaeoenvironmental reconstruction

Tooth enamel apatite carbonate carbon and oxygen isotope ratios of modern kangaroos (Macropus spp.) collected on a 900-km latitudinal transect spanning a C₃–C₄ transition zone were analysed to create a reference set for palaeoenvironmental reconstruction in southern Australia. The carbon isotope composition of enamel carbonate reflects the proportional intake of C₃ and C₄ vegetation, and its oxygen isotope composition reflects that of ingested water. Tooth enamel forms incrementally, recording dietary and environmental changes during mineralisation. Analyses show only weak correlations between climate records and latitudinal changes in δ¹³C and δ¹⁸O. No species achieved the δ¹³C values (~?1.0 ‰) expected for 100 % C₄ grazing diets; kangaroos at low latitudes that are classified as feeding primarily on C₄ grasses (grazers) have δ¹³C of up to ?3.5 ‰. In these areas, δ¹³C below ?12 ‰ suggests a 100 % C₃ grass and/or leafy plant (browse) diet while animals from higher latitude have lower δ¹³C. Animals from semi-arid areas have δ¹⁸O of 34–40 ‰, while grazers from temperate areas have lower values (~28–30 ‰). Three patterns with implications for palaeoenvironmental reconstruction emerge: (1) all species in semi-arid areas regularly browse to supplement limited grass resources; (2) all species within an environmental zone have similar carbon and oxygen isotope compositions, meaning data from different kangaroo species can be pooled for palaeoenvironmental investigations; (3) relatively small regional environmental differences can be distinguished when δ¹³C and δ¹⁸O data are used together. These data demonstrate that diet–isotope and climate–isotope relationships should be evaluated in modern ecosystems before application to the regional fossil record.     

Assessing fluctuating odontometric asymmetry among fossil hominin taxa through alternative measures of central tendency: effect of outliers and directional components on reported results.

Preliminary inquires into the distribution and expression of fluctuating odontometric asymmetry (FOA), among selected fossil hominins, have revealed results that may be serviceable within studies that assess, among others, palaeobiological, evolutionary processes and events. Though several intricate statistical applications have aided in the advancement of FOA to the hominin fossil record, little is known regarding the influence of outliers and directional components on reported results. Moreover, most methods employed to test homogeneity among FOA datasets are sensitive to the assumption that underlying samples reflect Gaussian distributions. Because this assumption is often violated, alternative formulations of Levene's test statistic, which have been shown to be robust under non-normality, have been suggested. Unfortunately, previous FOA studies have failed to address their potential. Given this, we considered two areas that may influence interpretations of FOA among fossil hominin studies. Firstly, we assessed distributions of signed data (d(u)) among samples of Australopithecus africanus, Paranthropus robustus and Homo habilis for outliers and directional asymmetry to evaluate their influence on reported heterogeneity. Secondly, in an attempt to decrease the probability of falsely rejecting H(0) due to non-normality, we considered alternative estimates of central tendency for comparisons of FOA. Our study confirms the need for intrinsic scrutiny of data, as the removal of one extreme value within the buccolingual H. habilis sample produced statistically significant outcomes at the sample level, while directional asymmetry was exposed within an expanded buccolingual P. robustus sample. However, though servicing alternative measures of central tendency remains informative, except for the buccolingual P. robustus sample before the correction of directional asymmetry, replacement of the mean was not required herein. Consistent with previous investigations, significant differences between buccolingual values in apposing arcades were unique among A. africanus and P. robustus, with the latter expressing greater FOA overall. Finally, our results strengthen the assertion that the individuals sampled among H. habilis may indicate an episode of developmental compromise where external and/or internal noises are lessened through internal homeostasis.     

Hominid environments at Hadar from paleosol studies in a framework of Ethiopian climate change

The amount and seasonal distribution of paleo-rainfall is a major concern of paleoanthropology because they determine the nature of the vegetation and the structure of the ecosystem, particularly in eastern Africa. The δ¹⁸O and δ¹³C of paleosol carbonates are quantitative proxies of these critical features of the paleoenvironment. The Afar region of Ethiopia lies between the African and Indian summer monsoons, and is prone to profound climate change. In the western Afar, the dominant paleoenvironment of the Hadar Formation during the late Pliocene was a major meandering river's distal low, flat floodplain, on which muds accreted that were continuously transformed into vegetated soils with Bk horizons rich in CaCO₃. The mean δ¹³C of paleosols throughout the Hadar Formation translates to an average vegetative cover across the extensive floodplain of about 30% of the C₄ grasses and 70% of unspecified C₃ plants. The character of the paleosols, such as the one at Locality 333, and their δ¹⁸O_Carbonate argue for a highly seasonal rainfall of about twice today's amount, implying that the C₃ plants were mostly sizeable trees and that the biome for Australopithecus afarensis was a grassy woodland. The amount of grasses abruptly increased in the lower Busidima Formation with its early Homo and artifacts to a more open grassy woodland of ca. 50% grasses. However, this transition in δ¹³C is not mirrored in the δ¹⁸O, which persists at a quite negative average value of −6.4‰ over the entire >2-Myr duration of both formations. This value for the carbonate means that the paleosoil water was a quite negative −4.1‰, a significant 5‰ more negative than our estimate of modern rain at Hadar. We put the negative δ¹⁸O of paleo-Hadar's rainfall into an isotopic framework of the dynamic history of climate change in sub-Saharan northern Africa. There have been two end-member climate regimes: (1) an earlier persistently pluvial Pliocene regime, with its strong summer monsoon, as registered in the Hadar Formation; and (2) the modern cyclical, mostly arid regime that began ca. 1 Myr ago, which has been punctuated by about ten cyclically predictable brief millennia-long pluvial episodes. The best known pluvial of the latter regime is the latest one, the African Humid Period (AHP), just 9.0-6.5 kyr ago, whose δ¹⁸O_Rainfall matches that for paleo-Hadar. The known climatological factors that brought on the AHP are probably the same ones that were persistently present for the Afar of the Pliocene. This dynamic rainfall history undoubtedly has influenced hominid occupation of the keystone Afar area at the gateway out of, and into, Africa.     

Photosynthetic Gas Exchange and Discrimination against 13CO2 and C18O16O in Tobacco Plants Modified by an Antisense Construct to Have Low Chloroplastic Carbonic Anhydrase

The physiological role of chloroplastic carbonic anhydrase (CA) was examined by antisense suppression of chloroplastic CA (on average 8% of wild type) in Nicotiana tabacum. Photosynthetic gas-exchange characteristics of low-CA and wild-type plants were measured concurrently with short-term, on-line stable isotope discrimination at varying vapor pressure deficit (VPD) and light intensity. Low-CA and wild-type plants were indistinguishable in the responses of assimilation, transpiration, stomatal conductance, and intercellular CO2 concentration to changing VPD or light intensity. At saturating light intensity, low-CA plants had lower discrimination against 13CO2 than wild-type plants by 1.2 to 1.8[per mille (thousand) sign]. Consequently, tissue of the low-CA plants was higher in 13C than the control plants. It was calculated that low-CA plants had chloroplast CO2 concentrations 13 to 22 [mu]mol mol-1 lower than wild-type plants. Discrimination against C18O16O in low-CA plants was 20% of that of the wild type, confirming a role of chloroplastic CA in the mechanism of discrimination against C18O16O ([delta]C18O16O). As VPD increased, stomatal closure caused a reduction in chloroplastic C02 concentration, and since VPD and chloroplastic CO2 concentration act in opposing directions on [delta]C18O16O, no effect of VPD was seen on [delta]C18O16O.