Talocrural joint in African hominoids: implications for Australopithecus afarensis

Talocrural joints of the African apes, modern humans, and A.L.288-1 are compared in order to investigate ankle function in the Hadar hominids. Comparisons between the hominids and African pongids clearly illustrate the anatomical and mechanical changes that occurred in this joint as a consequence of the evolutionary transition to habitual bipedality. Features which are considered include the obliquity of the distal tibial articular surface, the shape of the talar trochlea, and the location and functional implications of the talocrural axis. In every functionally significant feature examined the A.L.288-1 talocrural joint is fully bipedal. Moreover, the Hadar ankle complex also shows the functional constraints which are necessarily imposed by the adaptation to habitual bipedalism.     

Temporal trends and metric variation in the mandibles and dentition of Australopithecus afarensis

The Pliocene hominin samples from Hadar and Laetoli are thought to represent one species, Australopithecus afarensis, that exhibits stasis throughout its temporal range and has high levels of skeletal sexual dimorphism. In this paper, we test the hypothesis of stasis in dental and mandibular dimensions using nonparametric rank correlation methods to detect temporal trends and randomization tests to evaluate their statistical significance. We then use two methods (CV resampling; Fligner-Killeen test) to compare overall levels of variation in the fossil sample to those of extant hominoid species. Together, these analyses allow us to gauge the effects of changes through time on variation in mandibles and teeth of A. afarensis.P(3)mesiodistal length, M(3)size, and canine shape change through time but do not appear unusually variable in the sample as a whole. These temporal trends possibly reflect differences between the Laetoli and Hadar site-samples. For mandibles, a pronounced trend towards greater corpus size occurs late in the temporal sequence and contributes to high levels of variation compared to African apes. These results show that significant directional changes do occur in the A. afarensis mandibles and teeth, and in these elements, at least, the species is not static. Temporal variation is clearly an important component of overall variation in the A. afarensis lineage, even though other factors, such as sexual dimorphism, may also play a part.     

The calcaneus of Australopithecus afarensis and its implications for the evolution of bipedality

Calcanei from African apes, modern humans, and Australopithecus afarensis are compared to investigate the anatomical and mechanical changes that occurred in this bone as a result of the transition to terrestrial bipedality. Features analyzed include the cross-sectional area and volume of the calcaneal tuber, the geometry and orientation of the articular surfaces, and the surface topography of the calcaneal corpus. Calcaneal morphology is unequivocal in its partitioning of quadrupedal pongids and bipedal hominids.     

The taxonomic implications of cranial shape variation in Homo erectus

The taxonomic status of Homo erectus sensu lato has been a source of debate since the early 1980s, when a series of publications suggested that the early African fossils may represent a separate species, H. ergaster. To gain further resolution regarding this debate, 3D geometric morphometric data were used to quantify overall shape variation in the cranial vault within H. erectus using a new metric, the sum of squared pairwise Procrustes distances (SSD). Bootstrapping methods were used to compare the H. erectus SSD to a broad range of human and nonhuman primate samples in order to ascertain whether variation in H. erectus most clearly resembles that seen in one or more species. The reference taxa included relevant phylogenetic, ecological, and temporal analogs including humans, apes, and both extant and extinct papionin monkeys. The mean cranial shapes of different temporogeographic subsets of H. erectus fossils were then tested for significance using exact randomization tests and compared to the distances between regional groups of modern humans and subspecies/species of the ape and papionin monkey taxa. To gauge the influence of sexual dimorphism on levels of variation, comparisons were also made between the mean cranial shapes of single-sex samples for the reference taxa. Results indicate that variation in H. erectus is most comparable to single species of papionin monkeys and the genus Pan, which included two species. However, H. erectus encompasses a limited range of variation given its extensive geographic and temporal range, leading to the conclusion that only one species should be recognized. In addition, there are significant differences between the African/Georgian and Asian H. erectus samples, but not between H. ergaster (Georgia+Africa, excluding OH 9 and Daka) and H. erectus sensu stricto. This finding is in line with expectations for intraspecific variation in a long-lived species with a wide, but probably discontinuous, geographic distribution.     

Intraspecific physiological variation: implications for understanding functional diversity in ectomycorrhizal fungi

 Several thousand fungal species worlwide are thought to form ectomycorrhizas (ECM) with tree hosts and there is currently much interest in determining the functional significance of such diversity in natural and managed ecosystems. While only a few taxa have been investigated in detail, it is clear that ECM fungi display extensive intraspecific variation in a range of physiological and other life-history parameters. Thus, comparative investigations of single (or even a few) isolates of different species are unlikely to provide reliable information on functional capabilities. Extensive screening of taxonomically well-defined isolates is required. This must take into account spatial and temporal variation in gene expression in mycelia growing in axenic culture or in association with a host plant. Accepted: 29 June 1999     

Bootstrap tests of significance and the case for humanlike skeletal-size dimorphism in Australopithecus afarensis

Most estimates of sexual size dimorphism in Australopithecus afarensis indicate that this early hominin was more dimorphic than modern humans. In contrast, a recent study reported that size variation in A. afarensis, as represented by postcranial remains from Hadar and Maka, Ethiopia, is statistically most similar to that of modern humans, indicating a humanlike level of sexual dimorphism. Here, we evaluate the evidence for humanlike dimorphism in A. afarensis. We argue that statistical support for this claim is not as robust as has been asserted for the following reasons: (1) the analysis from which the claim was derived does not distinguish the A. afarensis sample from either the human or chimpanzee samples; (2) for some of the comparisons made, the A. afarensis sample cannot be distinguished from the Gorilla sample using two-tailed tests; and (3) the A. afarensis postcranial sample used in the analysis may contain more male than female specimens, which precludes a straightforward interpretation of the statistical results. Thus, support for humanlike dimorphism is equivocal, and a greater level of dimorphism cannot be ruled out.     

Patterns in taxonomic and functional diversity of lake phytoplankton

1. Patterns in phytoplankton diversity in lakes and their relationships with environmental gradients have been traditionally based on taxonomic analyses and indices, even though measures of functional diversity (FD) might be expected to be more responsive to such gradients. 2. We assessed the influence of water column physical structure, and other components of the overall environment, on lake phytoplankton diversity using two taxonomically based indices [species richness (S) and the Shannon index (H’)] and a FD index, to determine whether these different measures respond in similar ways to habitat structure. The study encompassed 45 lakes in Eastern Canada, within two lake districts [the Eastern Townships Region (ETR) and Laurentians Region (LR)] that vary in geology and landscape and in lake morphometry and chemistry. 3. Across all lakes, S and H’ were higher in lakes having greater vertical temperature heterogeneity and higher susceptibility to wind mixing. In addition, H’ declined with total phosphorus concentration. FD was only related to maximum lake depth, a variable that integrates many other habitat features. 4. Further insight into the factors affecting phytoplankton diversity was obtained by contrasting the two regions. The taxonomically based diversity measures differed little between the regions, while FD was higher in the ETR where more trait variants were present and more evenly distributed amongst species. Whereas factors driving S did not differ between the regions, we found region‐dependent patterns in the relationships of H’ and FD with maximum lake depth: both indices decreased with maximum depth in the region with lakes more exposed to wind (ETR) but increased in the more hilly landscape where lakes are more sheltered from wind mixing (LR). 5. Our study demonstrates that, for phytoplankton communities, a FD index can show simpler and stronger responses to environmental drivers than a taxonomically based index, while shedding further light onto the functional traits that are important in particular lake categories.     

Lucy's length: stature reconstruction in Australopithecus afarensis (A.L.288-1) with implications for other small-bodied hominids

New stature estimates are provided for A.L.288-1 (Australopithecus afarensis) based on (1) the relationship between femur length and stature in separate samples of human pygmies and pygmy chimpanzees and (2) model II regression alternatives to standard least-squares methods. Estimates from the two samples are very similar and converge on a value of approximately 3'6" for "Lucy." These results are compared to prior estimates and extended to other small-bodied hominids such as STS-14 and O.H.62. A new foot-to-stature ratio is also estimated for A.L.288-1, and its potential biomechanical significance for gait is evaluated in comparison to other groups.     

植物雄蕊合生的多样性、适应意义及分类学意义初探

定义并总结了雄蕊合生现象,对雄蕊合生的类型、适应意义、进化及分类学意义进行了初步的讨论和总结.本文定义"雄蕊合生"为,花内雄蕊与雄蕊之间部分或整体的结合生长,既包括愈合与黏合,也包括结合紧密的贴生和靠生.根据雄蕊合生部位的不同,将雄蕊合牛分为"花丝合生"、"花药合生"和"花丝花药均合生"等三大类.每一类中还存在着合生程度不同的子类型.雄蕊合生的结构还可能进一步与柱头或花柱合生在一起,形成"合蕊柱"一类的复杂结构.雄蕊合生在裸子植物较进化的科如百岁兰科和买麻藤科中就出现了,在种子植物中集中分布在近50个科内,且从系统关系来看,雄蕊合生各个类型可能发生过多次进化上的反复.在被子植物中,花丝合生类型更多地与较早进化的类群联系在一起,且多出现在离瓣花中;花药合生及花丝花药均合生则较晚进化,多与有着明显花冠筒的合瓣花伴随出现.这暗示着,雄蕊的合牛结构有可能与花部其他特征一起影响了植物的繁殖过程,具有一定的适应意义.目前,还没有实验研究针对雄蕊合生各个类型开展实验以揭示其发育机制和适应意义.理论上,花丝合乍能增强雄蕊的强度,有时候还形成了围绕子房与花柱的杯状、管状或环状结构,能对子房和花柱有着保护作用且承受传粉者在花内移动的压力:花丝合生还可将雄蕊固定在一个较为稳定的位置,使得花药接触传粉者身体的部位相对固定,减少了花粉浪费.花药合生能将花内花药都集中到同一个位置,花粉接触传粉者身体的同一部位,从而降低花粉损耗和提高异交授粉的准确性.花丝花药均合生可以同时具有以上两种合生方式的适应意义,而且还极大改变了雄蕊的结构与空间位置,改变了花药与柱头之间的空间位置(雌雄异位),对花内自交可能性和雌雄功能干扰有着潜在的影响.雄蕊合生各个类型的适应意义及其对访花者类型与行为和植物繁殖策略的影响,还需要开展实验进行针对性的研究.由于雄蕊形态与结构较为稳定,雄蕊合生的不同方式以及合生程度可以作为科与种的分类参考.     

Size and shape variation in the proximal femur of Australopithecus africanus

Aside from use as estimates of body mass dimorphism and fore to hind limb joint size comparisons, postcranial elements have not often contributed to assessments of variation in Australopithecus africanus. Meanwhile, cranial, facial, and dental size variation is interpreted to be high or moderately high. Further, the cranial base and face express patterns of structural (shape) variation, which are interpreted by some as evidence for the presence of multiple species. Here, the proximal femur is used to consider postcranial size and shape variation in A. africanus. Original fossils from Makapansgat and Sterkfontein, and samples from Homo, Pan, Gorilla, and Pongo were measured. Size variation was assessed by comparing the A. africanus coefficient of variation to bootstrapped distributions of coefficient of variation samples for each taxon. Shape variation was assessed from isometrically adjusted shape variables. First, the A. africanus standard deviation of log transformed shape variables was compared to bootstrapped distributions of logged standard deviations in each taxon. Second, shape variable based Euclidean distances between fossil pairs were compared to pairwise Euclidean distance distributions in each reference taxon. The degree of size variation in the A. africanus proximal femur is consistent with that of a single species, and is most comparable to Homo and Pan, lower than A. afarensis, and lower than some estimates of cranial and dental variation. Some, but not all, shape variables show more variation in A. africanus than in extant taxa. The degree of shape difference between some fossils exceeds the majority of pairwise differences in the reference taxa. Proximal femoral shape, but not size, variation is consistent with high estimates of A. africanus cranial variation.     

Fruit size and shape: Allometry at different taxonomic levels in bird-dispersed plants

Summary The likelihood that a plant's seeds will be dispersed by fruit-eating birds may depend upon the size and shape of its fruits. Assuming that elongate fruits can be swallowed more easily than spherical fruits of equal volume and that plant fitness is enhanced by seed dispersal by many individuals and species of birds, natural selection should favour increasing fruit elongation with increasing fruit size in bird-dispersed plants. According to this view, this allometric pattern would be adaptive. Alternatively, fruit shape in bird-dispersed plants may be constrained by development or phylogeny. To determine whether there was any evidence to support the adaptive allometry hypothesis, we examined allometric relationships between length and diameter in fruits and seeds in a group of neotropical bird-dispersed plant species. Using the major axis technique, we regressed ln(diameter) on ln(length) for fruits and seeds at various taxonomic levels: (1) within individual trees ofOcotea tenera (Lauraceae) (2) among 19 trees within a population ofO. tenera, as well as among pooled fruits from multiple trees within 20 other species in the Lauraceae, (3) among 25 sympatric species within a plant family (Lauraceae) and (4) among 167 species representing 63 angiosperm families within a plant community in Monteverde, Costa Rica. At most taxonomic levels, a tendency for fruit length to increase more rapidly than fruit diameter among fruits (negative allometry) occurred more frequently than expected by chance. Estimated slopes of the regressions of fruit length on fruit diameter were < 1 within 15 of the 19 individualO. tenera trees, among tree means withinO. tenera, among pooled fruits within 16 of the 20 other species in the Lauraceae, among species means within the Lauraceae and among means of all bird-dispersed species in the lower montane forests of Monteverde. Seed allometry showed similar patterns, although for both fruits and seeds the broad confidence intervals of the slopes estimated by major axis regression overlapped 1 in many cases. Among the 63 Monteverde family means, fruit length and diameter scaled isometrically. Based on measurements of ontogenetic changes in fruit shape in a single species,O. viridifolia, we found no evidence that negative allometry in fruit shape within the Lauraceae was an inevitable consequence of developmental constraints. Instead, increasing elongation of fruits and seeds in certain plant taxa is consistent with adaptation to gape-limited avian seed dispersers. Contrary results from vertebrate-dispersed species from Malawi and Spain may reflect differences between the New and Old World in plant taxa, seed dispersers or evolutionary history.     

Size variation in the postcranium ofAustralopithecus afarensis and extant species of hominoidea

The postcranial sample ofA. afarensis can be divided into two size groups. Among the best preserved elements which are represented by both morphs are the distal femur, proximal ulna, and capitate. The difference between the large and small fossil femora is similar to the difference between average male and femaleG. gorilla andP. pygmaeus. The distal femora ofH. sapiens are less sexually dimorphic while those ofP. paniscus, P. troglodytes, andH. lar are not significantly dimorphic at all. Large and small capitates and proximal ulnae ofA. afarensis differ slightly more than the highly dimorphic species of extant Hominoidea. In my sample of Amerindians, the capitate and proximal ulna are also strongly dimorphic. The two species ofPan have insignificant sexual dimorphism in these traits. There results imply that strong sexual dimorphism in body size is the primitive condition for the large bodied hominoids.     

Comparison of inverse-dynamics musculo-skeletal models of AL 288-1 Australopithecus afarensis and KNM-WT 15000 Homo ergaster to modern humans, with implications for the evolution of bipedalism

Size and proportions of the postcranial skeleton differ markedly between Australopithecus afarensis and Homo ergaster, and between the latter and modern Homo sapiens. This study uses computer simulations of gait in models derived from the best-known skeletons of these species (AL 288-1, Australopithecus afarensis, 3.18 million year ago) and KNM-WT 15000 (Homo ergaster, 1.5-1.8 million year ago) compared to models of adult human males and females, to estimate the required muscle power during bipedal walking, and to compare this with those in modern humans. Skeletal measurements were carried out on a cast of KNM-WT 15000, but for AL 288-1 were taken from the literature. Muscle attachments were applied to the models based on their position relative to the bone in modern humans. Joint motions and moments from experiments on human walking were input into the models to calculate muscle stress and power. The models were tested in erect walking and 'bent-hip bent-knee' gait. Calculated muscle forces were verified against EMG activity phases from experimental data, with reference to reasonable activation/force delays. Calculated muscle powers are reasonably comparable to experimentally derived metabolic values from the literature, given likely values for muscle efficiency. The results show that: 1) if evaluated by the power expenditure per unit of mass (W/kg) in walking, AL 288-1 and KNM-WT 15000 would need similar power to modern humans; however, 2) with distance-specific parameters as the criteria, AL 288-1 would require to expend relatively more muscle power (W/kg.m(-1)) in comparison to modern humans. The results imply that in the evolution of bipedalism, body proportions, for example those of KNM-WT 15000, may have evolved to obtain an effective application of muscle power to bipedal walking over a long distance, or at high speed.     

Intermittency in processing explains the diversity and shape of functional grazing responses

Central to theoretical studies of trophic interactions is the formulation of the consumer response to varying food availability. Response functions, however, are only rarely derived in mechanistic ways. As a consequence, the uncertainty in the functional representation of feeding remains large, as, e.g., evident from the ongoing debate on the usage of Ivlev, or Holling type I, II, and III functions in aquatic ecosystem models. Here, I refer to the work of Sj?berg in Ecol Model 10:215-225 (1980) who proposed to apply elements of the queuing theory developed in operational research to plankton-plankton interactions. Within this frame, food item processing is subdivided into two major stages which may operate with variable synchronicity. Asynchronous phasing of the two stages enhances the probability of long total processing times. This phenomenon is here termed feeding intermittency. Intermittency is assumed to determine the functional form of grazing kinetics, for which a novel grazing function containing a "shape" parameter is derived. Using this function, I evaluate the hypotheses that intermittency is influenced by (1) patchiness in the prey field (e.g., related to turbulence), and (2) the ratio of actual prey size to optimal prey size. Evidence for the first hypothesis arises from explaining reported variations in clearance rates of Acartia tonsa under different turbulence regimes. Further model applications to ingestion data for rotifers, copepods, and ciliates support the view that an increasing food size enhances intermittency and, this way, affects functional grazing responses. In the application to ciliate grazing, a possible prey density effect appears, possibly due to an intermittent activation of a feeding sub-stage. Queueing theory offers mechanistic explanations for transitions between Holling I-, II-, and Ivlev-type grazing. In doing so for variable prey size ratios, it may also refine size-based ecosystem models which are increasingly emerging in plankton ecology.