Niche Partitioning in Sympatric Gorilla and Pan from Cameroon: Implications for Life History Strategies and for Reconstructing the Evolution of Hominin Life History

Factors influencing the hominoid life histories are poorly understood, and little is known about how ecological conditions modulate the pace of their development. Yet our limited understanding of these interactions underpins life history interpretations in extinct hominins. Here we determined the synchronisation of dental mineralization/eruption with brain size in a 20^th century museum collection of sympatric Gorilla gorilla and Pan troglodytes from Central Cameroon. Using δ¹³C and δ¹⁵N of individuals’ hair, we assessed whether and how differences in diet and habitat use may have impacted on ape development. The results show that, overall, gorilla hair δ¹³C and δ¹⁵N values are more variable than those of chimpanzees, and that gorillas are consistently lower in δ¹³C and δ¹⁵N compared to chimpanzees. Within a restricted, isotopically-constrained area, gorilla brain development appears delayed relative to dental mineralization/eruption [or dental development is accelerated relative to brains]: only about 87.8% of adult brain size is attained by the time first permanent molars come into occlusion, whereas it is 92.3% in chimpanzees. Even when M1s are already in full functional occlusion, gorilla brains lag behind those of chimpanzee (91% versus 96.4%), relative to tooth development. Both bootstrap analyses and stable isotope results confirm that these results are unlikely due to sampling error. Rather, δ¹⁵N values imply that gorillas are not fully weaned (physiologically mature) until well after M1 are in full functional occlusion. In chimpanzees the transition from infant to adult feeding appears (a) more gradual and (b) earlier relative to somatic development. Taken together, the findings are consistent with life history theory that predicts delayed development when non-density dependent mortality is low, i.e. in closed habitats, and with the “risk aversion” hypothesis for frugivorous species as a means to avert starvation. Furthermore, the results highlight the complexity and plasticity of hominoid/hominin development.     

Brain Size Growth and Life History in Human Evolution

Increases in endocranial volume (a measure of brain size) play a major role in human evolution. Despite the importance of brain size increase, the developmental bases of human brain size evolution remain poorly characterized. Comparative analyses of endocranial volume size growth illustrate that distinctions between humans and other primates are consequences of differences in rates of brain size growth, with little evidence for differences in growth duration. Evaluation of available juvenile fossils shows that earliest hominins do not differ perceptibly from chimpanzees (Pan). However, rapid and human-like early brain growth apparently characterized Homo erectus at about 1?Ma before present. Neandertals show patterns of brain growth consistent with modern humans during infancy, but reach larger sizes than modern humans as a result of differences in later growth. Growth analyses reveal commonalities in patterns of early brain size growth during the last million years human evolution, despite major increases in adult size. This result implies consistency across hominins in terms of maternal metabolic costs of infancy. Continued size growth past infancy in Neandertals and modern humans, when compared to earlier hominins, may have cognitive implications. Differences between Neandertals and modern humans are implied, but difficult to define with certainty.     

Patterns and Processes in Cultural Evolution

Darwinian models of cultural change have been motivated, in part, by the desire to provide a framework for the unification of the biological and the human sciences. In this paper, drawing upon a distinction between the evolution of enabling mechanisms for the acquisition and dissemination of knowledge (EEM) and the evolution of epistemic theses as cultural products (EET), we propose a model of how culture emerges as a product of biological evolution on the basis of the concept of reaction norms. The goal of this model is to provide a means for conceptualizing how the biological and the cultural realms are connected, when they start to disconnect, and what the key transitions are. We then assess the viability of a Darwinian approach to cultural change. We conclude that the prospects of producing a Darwinian model of cultural change that unifies the human sciences in a way that mirrors the unification of the biological sciences in the light of Darwin’s theory are rather dim.     

The Importance of Population Growth and Regulation in Human Life History Evolution

Explaining the evolution of human life history traits remains an important challenge for evolutionary anthropologists. Progress is hindered by a poor appreciation of how demographic factors affect the action of natural selection. I review life history theory showing that the quantity maximized by selection depends on whether and how population growth is regulated. I show that the common use of R, a strategy’s expected lifetime number of offspring, as a fitness maximand is only appropriate under a strict set of conditions, which are apparently unappreciated by anthropologists. To concretely show how demography-free life history theory can lead to errors, I reanalyze an influential model of human life history evolution, which investigated the coevolution of a long lifespan and late age of maturity. I show that the model’s conclusions do not hold under simple changes to the implicitly assumed mechanism of density dependence, even when stated assumptions remain unchanged. This analysis suggests that progress in human life history theory requires better understanding of the demography of our ancestors.     

Density-Dependent Growth and Life History Evolution of Polycyclic Leaf Pathogens: A Continuous Time Growth Model

A density-dependent growth model of a polycyclic leaf pathogen is analysed. Viability and fecundity of the pathogen are regulated by the current population density. Spores are produced continuously from a sporulating infection until death of the infection, and consequently, all age classes of infections are present at a certain point in time. The leaf area of the host varies with time. Evolution of life history strategies are studied by letting different pathogen genotypes compete with each other. Evolution of life history strategies and evolution of impact of disease are discussed in relation to the ecology of the host-pathogen system. The model is exemplified by Erysiphe graminis f. sp. hordei growing on Hordeum vulgare.     

Life History Evolution in Amphicarpic Plants

Abstract Plants with dimorphic flowers or seeds provide excellent material for the study of life history evolution because the dimorphism often involves measurable differences in morphology, size, number, or genetic relatedness. For amphicarpic plants, the proportion of aerial: subterranean morphs produced is highly variable (from 0 to > 100) and related to both environmental and genetic factors. Plants from aerial seeds produce lower ratios of aerial: subterranean morphs than those from subterranean seeds. Despite substantial variation, subterranean seeds are generally heavier than aerial seeds (but fewer) and produce vigorous seedlings with high survivorship and high fitness. Adaptive advantages of subterranean seeds include retention of offspring in favorable parental microhabitats, protection of seeds from herbivory, predation, or fire, and avoidance of desiccating conditions on the soil surface; potential disadvantages include lack of gene exchange, high energy costs, limited dispersal, and sibling competition. For the few species studied, aerial reproduction is more plastic than subterranean reproduction and more likely to be affected by environmental conditions. Quantitative genetic analyses of a population of the annual grass Amphicarpum purshii have revealed lower heritabilites for subterranean relative to aerial reproductive traits. Subterranean seed number and mass show genetic correlations with shoot mass while aerial seed number and mass do not; seed set percentages of both seed types as well as percentage allocation to both reproductive morphs show negative genetic correlations with shoot mass. In this Amphicarpum population, directional selection on shoot mass may indirectly select for increased subterranean (but not aerial) seed output.     

Lemur Biorhythms and Life History Evolution

Skeletal histology supports the hypothesis that primate life histories are regulated by a neuroendocrine rhythm, the Havers-Halberg Oscillation (HHO). Interestingly, subfossil lemurs are outliers in HHO scaling relationships that have been discovered for haplorhine primates and other mammals. We present new data to determine whether these species represent the general lemur or strepsirrhine condition and to inform models about neuroendocrine-mediated life history evolution. We gathered the largest sample to date of HHO data from histological sections of primate teeth (including the subfossil lemurs) to assess the relationship of these chronobiological measures with life history-related variables including body mass, brain size, age at first female reproduction, and activity level. For anthropoids, these variables show strong correlations with HHO conforming to predictions, though body mass and endocranial volume are strongly correlated with HHO periodicity in this group. However, lemurs (possibly excepting Daubentonia) do not follow this pattern and show markedly less variability in HHO periodicity and lower correlation coefficients and slopes. Moreover, body mass is uncorrelated, and brain size and activity levels are more strongly correlated with HHO periodicity in these animals. We argue that lemurs evolved this pattern due to selection for risk-averse life histories driven by the unpredictability of the environment in Madagascar. These results reinforce the idea that HHO influences life history evolution differently in response to specific ecological selection regimes.     

Hominin Obstetrics and the Evolution of Constraints

Birth is significantly more complicated and dangerous in modern humans than in other great apes. This disparity is often hypothesized to be the result of evolutionary constraints on obstetric dimensions related to bipedalism and/or thermoregulation in later hominins. Previous attempts to test such hypotheses have used biomechanical methods and results have been mixed. But evolutionary constraints, restrictions or limitations on the course or outcome of evolution, are the result of an interaction between selective pressures and genetic constraints—the latter revealed in patterns of integration. Integration between traits can result in directional or stabilizing selection on one trait leading to correlated responses in other traits, which can bias and constrain evolutionary trajectories. Therefore, trait evolution may be constrained for reasons separate from those that can be estimated using biomechanical models, and to study evolutionary constraints it is necessary to understand the role genetic constraints play in morphological change. The results presented here show that genetic constraints can significantly reduce the evolutionary potential of the birth canal to evolve in humans, apes, and likely earlier hominins, but also point to an overall reduction in the level of constraints during hominin evolution. These findings suggest that divergent selection pressures for obstetric requirements and other pelvic functions in hominins reduced levels of genetic constraint on birth canal evolution, likely lowering the amount of time needed for evolutionary change, and permitting morphological evolution along a trajectory that might have previously been difficult or impossible to traverse.     

Evolution of the Vertebrate Central Nervous System: Patterns and Processes

AS brains do not fossilize, most proposed phylogenetic sequencesfor central nervous system characters must be based on the patternsof variation of those characters in living organisms. Similarly,hypotheses regarding how brains change through time, and theevolutionary processes that produce these changes, are ultimatelybased on the character patterns recognized. It is critical inthese analyses to distinguish between homologous and homoplasouscharacters if errors in the reconstruction and interpretationof phylogenies are to be minimized. Definitions of homologyand homoplasy are reviewed, as are the concepts that bear ontheir application. Cladistic definitions are adopted, and criteriafor distinguishing homologous from homoplasous characters arediscussed. Analysis of a number of CNS characters that are usuallyassumed to be homologous reveals that homoplasous charactersappear among them. As in other organ systems, homoplasous charactersare actually common. A number of previous hypotheses regardingCNS evolution are reviewed in the context of new data on neuralconnections and their cladistic analysis. Some of these hypothesesmay be falsified by a cladistic treatment of CNS characters,whereas sufficient data do not exist to evaluate others.     

植物生活史对策的进化

植物生活史对策的进化班勇（中国林业科学院林业研究所,北京１０００９１）ＥｖｏｌｕｔｉｏｎｏｆＬｉｆｅＨｉｓｔｏｒｙＳｔｒａｔｅｇｙｉｎＰｌａｎｔｓ￥ＢａｎＹｏｎｇ（ＲｅｓｅａｒｃｈＩｎｓｔｉｔｕｔｅｏｆＦｏｒｅｓｔｒｙ,ＣｈｉｎｅｓｅＡｃａｄｅｍｙｏ...     

The Origin of Life: A Problem of History,Chemistry, and Evolution

The origin of life is a field full of controversies, not only because of our vague understanding concerning the relevant issues, but also, perhaps more often, owing to our dim conceptual framework throughout the whole field. To improve this situation, an in‐depth conceptual dissection is presented here. It is elucidated that, at its core, the origin of life has three aspects. The facts involved in the process are taken as the historical aspect, which is destined to be uncertain and often irrelevant to debate regarding details. The rules involved include two distinct aspects: chemical mechanisms operated in the whole process, while evolutionary mechanisms joined in only after the emergence of the first Darwinian entities – and then accounted for the subsequent buildup of complexity (this cannot be explained solely by natural selection). Basically, we can ask about the possibility of any assumed event in the origin of life: ‘Is it evolutionarily plausible, chemically feasible, and historically likely?’ Clues from any of the three aspects may be quite valuable in directing our explorations on the other two. This conceptual dissection provides a clearer context for the field, which may even be more useful than any sort of specific research.     

Oviposition Site Choice and Life History Evolution

SYNOPSIS. Studies of life history evolution, as well as muchof life history theory, have typically focused on "hard" componentsof life histories; phenotypic characteristics that can be readilyobserved, quantified, and ultimately, connected rather directlyto fitness. Typical of these are propagule size, propagule number,and age and size at maturity. What is largely missing from thestudy of life history evolution is consideration of the roleof behavior, principally female oviposition site choice, inthe evolution of life histories. For oviparous organisms, naturalselection cannot produce locally optimized "hard" componentsof life history phenotypes without a consistent environmentalcontext (whether invariant orvariable); in a variable environment,that consistent environmental context can be most effectivelyprovided by interactive oviposition site choice. I present amodel of selection on oviposition site choice in the contextof the evolution of "hard" components of life history phenotypes,along with some experimental data illustrating oviposition sitechoice in response to predators. The model and data are thenrelated to the overall question of the role of oviposition sitechoice in life history evolution. The conclusion is that ovipositionsite choice must be under equally strong selection with eggsize, egg number and the other hard components of life historiesin order to generate and optimize locally adapted or ecologicallyspecialized life history phenotypes, and must therefore, playa significant role in the evolution of life histories.     

Loss of Olfactory Receptor Function in Hominin Evolution

The mammalian sense of smell is governed by the largest gene family, which encodes the olfactory receptors (ORs). The gain and loss of OR genes is typically correlated with adaptations to various ecological niches. Modern humans have 853 OR genes but 55% of these have lost their function. Here we show evidence of additional OR loss of function in the Neanderthal and Denisovan hominin genomes using comparative genomic methodologies. Ten Neanderthal and 8 Denisovan ORs show evidence of loss of function that differ from the reference modern human OR genome. Some of these losses are also present in a subset of modern humans, while some are unique to each lineage. Morphological changes in the cranium of Neanderthals suggest different sensory arrangements to that of modern humans. We identify differences in functional olfactory receptor genes among modern humans, Neanderthals and Denisovans, suggesting varied loss of function across all three taxa and we highlight the utility of using genomic information to elucidate the sensory niches of extinct species.     

Phylogeny, Life History Evolution and Biogeography of the Rhinanthoid Orobanchaceae

Rhinanthoid Orobanchaceae form a monophyletic lineage that include the hemiparasitic genera Euphrasia, Melampyrum, Tozzia, Bartsia, Nothobartsia, Odontites (s.l.), Rhinanthus, Rhynchocorys, Parentucellia, Hedbergia and holoparasitic Lathraea. In this study, we aimed to reconstruct the phylogeny, evolution of life-history traits (life cycle and seed size) and explain the extant biogeographical patterns in this group. For phylogenetic reconstruction, we used molecular data obtained by sequencing the nuclear ITS region and the chloroplast trnT-trnL intergenic spacer and matK?+?trnK regions. The genus Melampyrum was found to occupy the sister position to the rest of the group. The other genera were assembled in the sister Rhinanthus-Rhynchocorys-Lathraea and Bartsia-Euphrasia-Odontites subclades. The reconstruction of life-cycle evolution yielded ambiguous results suggesting nonetheless a substantially higher likelihood of perenniality compared to annuality in most ancestor lineages. Seed size varied across two orders of magnitude (average weight per seed: 0.02–7.22 mg) and tended to decrease in the Bartsia-Euphrasia-Odontites subclade compared to the rest of the group. Seed-size evolution was correlated with life-history evolution in the group if the generally small-seeded Bartsia-Euphrasia-Odontites subclade is excluded. We formulated hypotheses relating the extant biogeographical affinities of individual genera to the geological history of the Euro-Caucasian diversity center of the group. Notable dispersal events in Euphrasia and Bartsia were hypothesized to be allowed or at least facilitated by a specific combination of the life-history traits.     

A Spring Forward for Hominin Evolution in East Africa

Groundwater is essential to modern human survival during drought periods. There is also growing geological evidence of springs associated with stone tools and hominin fossils in the East African Rift System (EARS) during a critical period for hominin evolution (from 1.8 Ma). However it is not known how vulnerable these springs may have been to climate variability and whether groundwater availability may have played a part in human evolution. Recent interdisciplinary research at Olduvai Gorge, Tanzania, has documented climate fluctuations attributable to astronomic forcing and the presence of paleosprings directly associated with archaeological sites. Using palaeogeological reconstruction and groundwater modelling of the Olduvai Gorge paleo-catchment, we show how spring discharge was likely linked to East African climate variability of annual to Milankovitch cycle timescales. Under decadal to centennial timescales, spring flow would have been relatively invariant providing good water resource resilience through long droughts. For multi-millennial periods, modelled spring flows lag groundwater recharge by 100 s to 1000 years. The lag creates long buffer periods allowing hominins to adapt to new habitats as potable surface water from rivers or lakes became increasingly scarce. Localised groundwater systems are likely to have been widespread within the EARS providing refugia and intense competition during dry periods, thus being an important factor in natural selection and evolution, as well as a vital resource during hominin dispersal within and out of Africa.     

The Ontogeny of Encephalization: Tradeoffs Between Brain Growth,Somatic Growth,and Life History in Hominoids and Platyrrhines

This study examines variation in brain growth relative somatic growth in four hominoids and three platyrrhines to determine whether there is a trade-off during ontogeny. I predicted that somatic growth would be reduced during periods of extensive brain growth, and species with larger degrees of encephalization would reach a smaller body size at brain growth completion because more energy is directed towards the brain. I measured cranial capacity and skeletal size in over 500 skeletal specimens from wild populations. I calculated nonlinear growth curves and velocity curves to determine brain/body growth allometry during ontogeny. In addition, I calculated linear regressions to describe the brain/body allometry during the postnatal period prior to brain size reaching an asymptote. The results showed that somatic growth is not substantially reduced in species with extensive brain growth, and body size at brain growth completion was larger in species with greater degrees of encephalization. Furthermore, large body size at brain growth completion was not correlated with interbirth interval, but was significantly correlated with prolonged juvenile periods and late age at maturity when data were corrected for phylogeny. These results indicate that neither reduction in body growth nor reproductive rate are compensatory mechanisms for the energetic costs of brain growth. Other avenues for meeting energetic costs must be in effect. In addition, the results show that somatic growth in encephalized species is particularly slow during the juvenile period after brain growth at or near completion, suggesting that these growth patterns are explained by reasons other than energetic costs.     

四川55种鱼生活史型的研究

四川５５种鱼生活史型的研究刁晓明,罗一兵李波（西南农业大学水产系,重庆６３０７１６）（重庆大学计算机系,６３００００）ＬｉｆｅＨｉｓｔｏｒｙＰａｔｔｅｒｎｓｏｆ５５ＦｉｓｈＳｐｅｃｉｅｓｉｎＳｉｃｈｕａｎ￥ＤｉａｏＸｉａｏｍｉｎｇ;ＬｕｏＹｉｂｉｎｇ...