Hierarchical analysis of species distributions and abundance across environmental gradients

Abiotic and biotic processes operate at multiple spatial and temporal scales to shape many ecological processes, including species distributions and demography. Current debate about the relative roles of niche-based and stochastic processes in shaping species distributions and community composition reflects, in part, the challenge of understanding how these processes interact across scales. Traditional statistical models that ignore autocorrelation and spatial hierarchies can result in misidentification of important ecological covariates. Here, we demonstrate the utility of a hierarchical modeling framework for testing hypotheses about the importance of abiotic factors at different spatial scales and local spatial autocorrelation for shaping species distributions and abundances. For the two orchid species studied, understory light availability and soil moisture helped to explain patterns of presence and abundance at a microsite scale (<4 m2), while soil organic content was important at a population scale (<400 m2). The inclusion of spatial autocorrelation is shown to alter the magnitude and certainty of estimated relationships between abundance and abiotic variables, and we suggest that such analysis be used more often to explore the relationships between species life histories and distributions. The hierarchical modeling framework is shown to have great potential for elucidating ecological relationships involving abiotic and biotic processes simultaneously at multiple scales.     

Climate determines upper, but not lower, altitudinal range limits of Pacific Northwest conifers

Does climate determine species' ranges? Rapid rates of anthropogenic warming make this classic ecological question especially relevant. We ask whether climate controls range limits by quantifying relationships between climatic variables (precipitation, temperature) and tree growth across the altitudinal ranges of six Pacific Northwestern conifers on Mt. Rainier, Washington, USA. Results for three species (Abies amabilis, Callitropsis nootkatensis, Tsuga mertensiana) whose upper limits occur at treeline (> 1600 m) imply climatic controls on upper range limits, with low growth in cold and high snowpack years. Annual growth was synchronized among individuals at upper limits for these high-elevation species, further suggesting that stand-level effects such as climate constrain growth more strongly than local processes. By contrast, at lower limits climatic effects on growth were weak for these high-elevation species. Growth-climate relationships for three low-elevation species (Pseudotsuga menziesii, Thuja plicata, Tsuga heterophylla) were not consistent with expectations of climatic controls on upper limits, which are located within closed-canopy forest (< 1200 m). Annual growth of these species was poorly synchronized among individuals. Our results suggest that climate controls altitudinal range limits at treeline, while local drivers (perhaps biotic interactions) influence growth in closed-canopy forests. Climate-change-induced range shifts in closed-canopy forests will therefore be difficult to predict accurately.     

Estimating species-specific survival and movement when species identification is uncertain

Incorporating uncertainty in the investigation of ecological studies has been the topic of an increasing body of research. In particular, mark-recapture methodology has shown that incorporating uncertainty in the probability of detecting individuals in populations enables accurate estimation of population-level processes such as survival, reproduction, and dispersal. Recent advances in mark-recapture methodology have included estimating population-level processes for biologically important groups despite the misassignment of individuals to those groups. Examples include estimating rates of apparent survival despite less than perfect accuracy when identifying individuals to gender or breeding state. Here we introduce a method for estimating apparent survival and dispersal in species that co-occur but that are difficult to distinguish. We use data from co-occurring populations of meadow voles (Microtus pennsylvanicus) and montane voles (M. montanus) in addition to simulated data to show that ignoring species uncertainty can lead to biased estimates of population processes. The incorporation of species uncertainty in mark-recapture studies should aid future research investigating ecological concepts such as interspecific competition, niche differentiation, and spatial population dynamics in sibling species.     

Ecological network determination of sectoral linkages, utility relations and structural characteristics on urban ecological economic system

Analyzing the structure and functioning of the urban system revealed ways to optimize its structure by adjusting the relationships among compartments, thereby demonstrating how ecological network analysis can be used in urban system research. Based on the account of the extended exergy utilization in the sector of urban socio-economic system, which is considered as the composition of extraction (Ex), conversion (Co), agriculture (Ag), industry (In), transportation (Tr), tertiary (Te) and households (Do) sectors, an urban ecological network model is constructed to gain insights into the economic processes oriented to sustainable urban development. Taking Beijing city as the case, the network accounting and related ecological evaluation of a practical urban economy are carried out in this study in the light of flux, efficiency, utility and structure analysis. The results showed that a large quantity of energy and resources have to be consumed to maintain the structure and function of a city. The thermodynamic efficiencies of individual sector in Beijing remain at a low level. The social system in Beijing is a highly competitive network, and there are 8 competitive relations and only two mutualistic ones. The Domestic and Agricultural sector are the major controlling factors of the system. Moreover, the assessment results of Beijing are compared with the other three socio-economic systems, Norway, UK and Italy, and the ecological network function and structure comparisons are correspondingly illuminated and discussed. The conclusions indicate that the exergy-based network analysis can be refined to become an integrative tool for evaluation, policy-making and regulation for urban socio-economic system management concerning structure and efficiency at urban levels.     

Landscape connectivity strengthens local-regional richness relationships in successional plant communities

Local species diversity is maintained over ecological time by a balance between dispersal and species interactions. Local-regional species richness relationships are often used to investigate the relative importance of these two processes and the scales at which they operate. For communities undergoing succession, theory predicts a temporal progression in local-regional species richness relationships: from no relationship to positive linear to saturating. However, observational tests have been mixed, and experiments have been rare. Using a replicated large-scale experiment, we evaluate the impact of two dispersal-governing processes at the regional scale, connectivity and shape of the region (i.e., patches), on the progression of local-regional species richness relationships for plant communities undergoing succession. Regional connectivity accelerates the transition from no relationship to a positive linear relationship, while the shape of the region has no consistent effect nine years post-disturbance. Our results experimentally demonstrate the importance of dispersal in structuring local-regional species richness relationships over time and suggest that conservation corridors among regions can increase local diversity through regional enrichment of plant communities undergoing reassembly.     

Mixed but not admixed: a spatial analysis of genetic variation of an invasive ascidian on natural and artificial substrates

Following the introduction to a new area (pre-border dispersal), post-border processes determine the success in the establishment of non-indigenous species (NIS). However, little is known on how these post-border processes shape the genetic composition of NIS at regional scales. Here, we analyse genetic variation in introduced populations along impacted coastlines to infer demographic and kinship dynamics at the post-border stage. We used as a model system the ascidian species Microcosmus squamiger that has been introduced worldwide. This species can colonize and grow fast on man-made artificial structures, impacting activities such as mariculture. However, it can also establish itself on natural substrates, thus altering natural communities and becoming an ecological problem. We genotyped 302 individuals from eight populations established on natural and artificial substrates in the north-western Mediterranean Sea, using six microsatellite loci. We then compared the resulting genotypes with those found within the native range of the species. We found high levels of genetic diversity and allelic richness in all populations, with an overall deficit of heterozygotes. Autocorrelation analyses showed that there was no within-population genetic structure (at a scale of tens of metres); likewise, no significant differentiation in pairwise comparisons between populations (tens of kilometres apart) and no isolation-by-distance pattern was found. The results suggest that M. squamiger has a natural capacity for high dispersal from one patch of hard substrate to another and no differences whatsoever could be substantiated between natural and artificial substrates. Interestingly, two groups of genetically differentiated individuals were detected that were associated with the two ancestral source areas of the worldwide expansion of the species. Individual assignment tests showed the coexistence of individuals of these two clusters in all populations but with little interbreeding among them as the frequency of admixed individuals was only 15 %. The mechanism responsible for maintaining these genetic pools unmixed is unknown, but it does not appear to compromise post-border colonization of introduced populations.     

Species distribution models and ecological theory: A critical assessment and some possible new approaches

Given the importance of knowledge of species distribution for conservation and climate change management, continuous and progressive evaluation of the statistical models predicting species distributions is necessary. Current models are evaluated in terms of ecological theory used, the data model accepted and the statistical methods applied. Focus is restricted to Generalised Linear Models (GLM) and Generalised Additive Models (GAM). Certain currently unused regression methods are reviewed for their possible application to species modelling.A review of recent papers suggests that ecological theory is rarely explicitly considered. Current theory and results support species responses to environmental variables to be unimodal and often skewed though process-based theory is often lacking. Many studies fail to test for unimodal or skewed responses and straight-line relationships are often fitted without justification.Data resolution (size of sampling unit) determines the nature of the environmental niche models that can be fitted. A synthesis of differing ecophysiological ideas and the use of biophysical processes models could improve the selection of predictor variables. A better conceptual framework is needed for selecting variables.Comparison of statistical methods is difficult. Predictive success is insufficient and a test of ecological realism is also needed. Evaluation of methods needs artificial data, as there is no knowledge about the true relationships between variables for field data. However, use of artificial data is limited by lack of comprehensive theory.Three potentially new methods are reviewed. Quantile regression (QR) has potential and a strong theoretical justification in Liebig's law of the minimum. Structural equation modelling (SEM) has an appealing conceptual framework for testing causality but has problems with curvilinear relationships. Geographically weighted regression (GWR) intended to examine spatial non-stationarity of ecological processes requires further evaluation before being used.Synthesis and applications: explicit theory needs to be incorporated into species response models used in conservation. For example, testing for unimodal skewed responses should be a routine procedure. Clear statements of the ecological theory used, the nature of the data model and sufficient details of the statistical method are needed for current models to be evaluated. New statistical methods need to be evaluated for compatibility with ecological theory before use in applied ecology. Some recent work with artificial data suggests the combination of ecological knowledge and statistical skill is more important than the precise statistical method used. The potential exists for a synthesis of current species modelling approaches based on their differing ecological insights not their methodology.     

Breeding systems and seed size in a neotropical flora: testing evolutionary hypotheses

A well-known, but largely untested, prediction in plant reproductive ecology is that dioecious taxa should produce larger, more, higher-quality, or better-defended seeds than cosexual taxa. Using a data set composed of 972 species in 104 families, representing the flora of the Tambopata Wildlife Reserve (Madre de Dios, Peru), we evaluated the first component of this prediction, examining ecological and evolutionary relationships between breeding system and mean seed size with two kinds of tests. First, we conducted cross-species analyses to determine whether species with different breeding systems differed significantly with respect to mean individual seed size. Second, we used a hypothesized phylogeny to identify pairs of the most closely related taxa or clades within the Tambopata community that differed with respect to breeding system. Comparing pair members allowed us to determine whether evolutionary divergence in breeding system (between taxa with unisexual vs. cosexual individuals) was consistently associated with evolutionary change in seed size. In both analyses, we controlled for potentially confounding effects of growth form by examining these relationships within woody and nonwoody taxa. Cross-species analyses revealed that dioecious species produced larger seeds than cosexual species among woody species, shrubs, lianas (each growth form analyzed separately), and all species pooled, but not among trees. Phylogenetically independent contrasts upheld the significant association between breeding system and seed size among woody taxa, lianas, and all taxa pooled, but not among shrubs. We discuss the implications of our findings for evolutionary hypotheses regarding associations between dioecy and seed size.     

Positives and pathologies of natural resource management on private land‐conservation areas

In managed natural resource systems, such as fisheries and rangelands, there is a recognized trade‐off between managing for short‐term benefits and managing for longer term resilience. Management actions that stabilize ecological attributes or processes can improve productivity in the supply of ecosystem goods and services in the short term but erode system resilience at longer time scales. For example, fire suppression in rangelands can increase grass biomass initially but ultimately result in an undesirable, shrub‐dominated system. Analyses of this phenomenon have focused largely on how management actions influence slow‐changing biophysical system attributes (such as vegetation composition). Data on the frequency of management actions that reduce natural ecological variation on 66 private land‐conservation areas (PLCAs) in South Africa were used to investigate how management actions are influenced by manager decision‐making approaches, a largely ignored part of the problem. The pathology of natural resource management was evident on some PLCAs: increased focus on revenue‐generation in decision making resulted in an increased frequency of actions to stabilize short‐term variation in large mammal populations, which led to increased revenues from ecotourism or hunting. On many PLCAs, these management actions corresponded with a reduced focus on ecological monitoring and an increase in overstocking of game (i.e., ungulate species) and stocking of extralimitals (i.e., game species outside their historical range). Positives in natural resource management also existed. Some managers monitored slower changing ecological attributes, which resulted in less‐intensive management, fewer extralimital species, and lower stocking rates. Our unique, empirical investigation of monitoring‐management relationships illustrates that management decisions informed by revenue monitoring versus ecological monitoring can have opposing consequences for natural resource productivity and sustainability. Promoting management actions that maintain resilience in natural resource systems therefore requires cognizance of why managers act the way they do and how these actions can gradually shift managers toward unsustainable strategies.     

Social dominance,seasonal movements,and spatial segregation in African elephants: a contribution to conservation behavior

The structure of dominance relationships among individuals in a population is known to influence their fitness, access to resources, risk of predation, and even energy budgets. Recent advances in global positioning system radio telemetry provide data to evaluate the influence of social relationships on population spatial structure and ranging tactics. Using current models of socio-ecology as a framework, we explore the spatial behaviors relating to the maintenance of transitive (i.e., linear) dominance hierarchies between elephant social groups despite the infrequent occurrence of contests over resources and lack of territorial behavior. Data collected from seven families of different rank demonstrate that dominant groups disproportionately use preferred habitats, limit their exposure to predation/conflict with humans by avoiding unprotected areas, and expend less energy than subordinate groups during the dry season. Hence, our data provide strong evidence of rank derived spatial partitioning in this migratory species. These behaviors, however, were not found during the wet season, indicating that spatial segregation of elephants is related to resource availability. Our results indicate the importance of protecting preexisting social mechanisms for mitigating the ecological impacts of high density in this species. This analysis provides an exemplar of how behavioral research in a socio-ecological framework can serve to identify factors salient to the persistence and management of at risk species or populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Adaptive speciation theory: a conceptual review

Speciation—the origin of new species—is the source of the diversity of life. A theory of speciation is essential to link poorly understood macro-evolutionary processes, such as the origin of biodiversity and adaptive radiation, to well understood micro-evolutionary processes, such as allele frequency change due to natural or sexual selection. An important question is whether, and to what extent, the process of speciation is ‘adaptive’, i.e., driven by natural and/or sexual selection. Here, we discuss two main modelling approaches in adaptive speciation theory. Ecological models of speciation focus on the evolution of ecological differentiation through divergent natural selection. These models can explain the stable coexistence of the resulting daughter species in the face of interspecific competition, but they are often vague about the evolution of reproductive isolation. Most sexual selection models of speciation focus on the diversification of mating strategies through divergent sexual selection. These models can explain the evolution of prezygotic reproductive isolation, but they are typically vague on questions like ecological coexistence. By means of an integrated model, incorporating both ecological interactions and sexual selection, we demonstrate that disruptive selection on both ecological and mating strategies is necessary, but not sufficient, for speciation to occur. To achieve speciation, mating must at least partly reflect ecological characteristics. The interaction of natural and sexual selection is also pivotal in a model where sexual selection facilitates ecological speciation even in the absence of diverging female preferences. In view of these results, it is counterproductive to consider ecological and sexual selection models as contrasting and incompatible views on speciation, one being dominant over the other. Instead, an integrative perspective is needed to achieve a thorough and coherent understanding of adaptive speciation.     

A structural equation model to integrate changes in functional strategies during old-field succession

From a functional perspective, changes in abundance, and ultimately species replacement, during succession are a consequence of integrated suites of traits conferring different relative ecological advantages as the environment changes over time. Here we use structural equations to model the interspecific relationships between these integrated functional traits using 34 herbaceous species from a Mediterranean old-field succession and thus quantify the notion of a plant strategy. We measured plant traits related to plant vegetative and reproductive size, leaf functioning, reproductive phenology, seed mass, and production on 15 individuals per species monitored during one growing season. The resulting structural equation model successfully accounts for the pattern of trait covariation during the first 45 years post-abandonment using just two forcing variables: time since site abandonment and seed mass; no association between time since field abandonment and seed mass was observed over these herbaceous stages of secondary succession. All other predicted traits values are determined by these two variables and the cause-effect linkage between them. Adding pre-reproductive vegetative mass as a third forcing variable noticeably increased the predictive power of the model. Increasing the time after abandonment favors species with increasing life span and pre-reproductive biomass and decreasing specific leaf area. Allometric coefficients relating vegetative and reproductive components of plant size were in accordance with allometry theory. The model confirmed the trade-off between seed mass and seed number. Maximum plant height and seed mass were major determinants of reproductive phenology. Our results show that beyond verbal conceptualization, plant ecological strategies can be quantified and modeled.     

Competitive regimes and female bonding in two species of squirrel monkeys (Saimiri oerstedi and S. sciureus)

Summary Ecological and behavioral data from long-term field studies of known individuals in two closely related squirrel monkey species (Saimiri oerstedi and S. sciureus) were used to examine hypotheses about the source of variation in female bonding among group-living primates. Social relationships in species which live in cohesive groups are thought to depend on the nature of competition for resources. S. oerstedi and S. sciureus both live in large groups and are subject to intense predation. Direct feeding competition both between and within groups is extremely low in S. oerstedi; in this species female relationships are undifferentiated, no female dominance hierarchy is evident and females disperse from their natal group. S. sciureus also experiences very low levels of between-group competition, but within-group direct competition for resources is frequent; this species demonstrates differentiated female relationships, a female dominance hierarchy, and female philopatry. The correlated ecological and social variables found in these two congeners further minimize the minor effects of phylogenetic differences and emphasize the importance of food distribution in determining social characteristics. Offprint requests to: S. Boinski     

Use of a sensitive indicator species in the assessment of biological effects of sewage disposal in fjords near Bergen,Norway

Coordinated environmental, ecological and biochemical studies have been applied to assess the impact of sewage disposal in a fjordic system near Bergen, Norway. The ecological and biochemical effects were studied in 1983 at four sampling locations situated along a spatial gradient of effects of the sewage on conditions in the sediments. Two of the locations, near Dolviken, were found to be considerably affected by the sewage. Relatively few species of macrobenthic invertebrate fauna were present at these locations, and analysis of the distribution of individuals among species indicated distortion of the benthic community structure. On the basis of its distribution along spatial gradients of organic enrichment and various criteria relating to its suitability for biochemical analysis, the polychaete Glycera alba (Müller) was selected as the most suitable pollution-sensitive indicator species for use in the biochemical studies. In individuals from the two affected locations near Dolviken, maximal activities of the regulatory glycolytic enzyme, phosphofructokinase, and the pyruvate oxidoreductase, alanopine dehydrogenase, were very low. Activities of several other enzymes associated with carbohydrate catabolism were also lower in these groups than in the reference group collected from Raunefjorden. The ecological and biochemical measures both corresponded closely with the changes in environmental conditions along the gradient of sewage effects. The results are discussed with reference to earlier coordinated ecological and biochemical investigations carried out in Scotland and Norway and to experimental studies of the effects of pollutants and hypoxia on energy-yielding metabolism of polychaetes. It is suggested that the enzymatic changes in G. alba may be a sensitive component of an integrated metabolic response, which may involve a decrease in glycolytic energy production for the fuelling of muscular activity. Further development of this coordinated ecological and biochemical approach is discussed, with emphasis on its potential utility in the assessment of biological effects of the disposal of organically rich waste materials in coastal waters.     

Movement ecology: size-specific behavioral response of an invasive snail to food availability

Immigration, emigration, migration, and redistribution describe processes that involve movement of individuals. These movements are an essential part of contemporary ecological models, and understanding how movement is affected by biotic and abiotic factors is important for effectively modeling ecological processes that depend on movement. We asked how phenotypic heterogeneity (body size) and environmental heterogeneity (food resource level) affect the movement behavior of an aquatic snail (Tarebia granifera), and whether including these phenotypic and environmental effects improves advection-diffusion models of movement. We postulated various elaborations of the basic advection diffusion model as a priori working hypotheses. To test our hypotheses we measured individual snail movements in experimental streams at high- and low-food resource treatments. Using these experimental movement data, we examined the dependency of model selection on resource level and body size using Akaike's Information Criterion (AIC). At low resources, large individuals moved faster than small individuals, producing a platykurtic movement distribution; including size dependency in the model improved model performance. In stark contrast, at high resources, individuals moved upstream together as a wave, and body size differences largely disappeared. The model selection exercise indicated that population heterogeneity is best described by the advection component of movement for this species, because the top-ranked model included size dependency in advection, but not diffusion. Also, all probable models included resource dependency. Thus population and environmental heterogeneities both influence individual movement behaviors and the population-level distribution kernels, and their interaction may drive variation in movement behaviors in terms of both advection rates and diffusion rates. A behaviorally informed modeling framework will integrate the sentient response of individuals in terms of movement and enhance our ability to accurately model ecological processes that depend on animal movement.     

巨枝人工林灌木层优势种的种间联结性分析

种间联结的研究对于人工林群落的生物多样性保育和可持续经营与管理具有重要的意义,目前对巨桉人工林灌木层物种之间的作用关系知之甚少。根据野外60个样方的调查数据,运用x^2检验、Jaccard指数和Spearman秩相关系数研究了巨桉（Eucalyptusgrandis）人工林灌木层18个优势种的种间关联和相关关系。结果表明,灌木层优势种具有总体上的正联结性,但在153个种对中,具有显著正联结性的种对数约占总种对数的10％,绝大数种对表现为弱联结性或无联结性,种间关系较为松散,表明巨桉林下灌木种群具有相对独立分布的特性。研究种间关系时,种间关联结合种间相关得出的结果较好。根据分析结果将18个优势种划分为4个生态种组,生态种组内的种其资源利用方式和生态要求相似,而组间表现出明显的差异。生态种组的划分能为巨桉人工林林下灌木层的管理提供理论依据。     

A critical review of the precautionary approach of the IUCN impact classification for non-native taxa

The International Union for the Conservation of Nature (IUCN) proposes the use of the Environmental Impact Classification for Alien Taxa to standardize the classification of introduced species (IS) based on their environmental impact. The IUCN invoked the precautionary principle (PP) via 2 rules: the impact assigned to a taxon must be the maximum recorded impact across different impact assessments, and when the main driver of environmental damage is unclear, it must be assumed to be caused by the IS. The validity of PP is conditioned on the degree of emergency that warrants urgent decisions and on the scientific evidence demonstrating the advantages of applying a preventive measure. The application of an impact classification system does not arise in the context of an emergency that requires management; it occurs before the decision-making phase. Thus, PP should not be used in early steps of the risk analysis process. The IUCN also did not provide enough scientific basis to justify the use of PP. Instead, the PP rules appear to be rooted primarily in the ethical value system underlying conservation science. Conservationists assign intrinsic value to native species by virtue of their roles and relationships within ecological and evolutionary systems and processes; thus, individuals introduced in new environments not only cease to have value because they are no longer part of that natural diversity and lack those links with the rest of the ecosystem, but they become a threat to what conservationists value most. The consequence of this belief is that all introduced taxa will have an impact at some level, suggesting that values justify the PP rules.     

乳山湾外海夏季大型底栖动物分布与环境因子的典范对应分析

应用典范对应分析(Canonical correspondence analysis,CCA)研究了乳山湾外海21个样点中的32种大型底栖动物与包括化学耗氧量、溶解氧等在内的11个沉积物和底层水体环境因子间的相关关系.作出种类分布与环境因子关系的二维排序图,排序图直观地反映了主要大型底栖动物种类分布与环境因子间的关系.排序图中环境因子与前两个排序轴的相关系数大小表明,总氮、溶解氧、盐度、pH、水深和总磷是影响乳山湾外海大型底栖动物群落分布的主导因素.排序图还揭示了大型底栖动物物种对生态环境的需求以及它们在不同资源维上的生态分化现象,不同生境中大型底栖动物分布的种类不同.根据对环境因子的生态需求不同,32种大型底栖动物可分为6组:组Ⅰ,8种,分布在高总氮生境;组Ⅱ,4种,分布在高溶解氧、高盐度、低总氮生境;组Ⅲ,10种,分布在水浅、高pH生境;组Ⅳ,4种,分布在水深、低pH生境;组Ⅴ,仅1种,分布在高总磷生境;组Ⅵ,5种,分布在水深、低总磷生境.图2表5参29     

Climatic and Biotic Stochasticity: Disparate Causes of Convergent Demographies in Rare, Sympatric Plants

Abstract:  Species with known demographies may be used as proxies, or approximate models, to predict vital rates and ecological properties of target species that either have not been studied or are species for which data may be difficult to obtain. These extrapolations assume that model and target species with similar properties respond in the same ways to the same ecological factors, that they have similar population dynamics, and that the similarity of vital rates reflects analogous responses to the same factors. I used two rare, sympatric annual plants (sand gilia [ Gilia tenuiflora arenaria ] and Monterey spineflower [ Chorizanthe pungens pungens ]) to test these assumptions experimentally. The vital rates of these species are similar and strongly correlated with rainfall, and I added water and/or prevented herbivore access to experimental plots. Their survival and reproduction were driven by different, largely stochastic factors and processes: sand gilia by herbivory and Monterey spineflower by rainfall. Because the causal agents and processes generating similar demographic patterns were species specific, these results demonstrate, both theoretically and empirically, that it is critical to identify the ecological processes generating observed effects and that experimental manipulations are usually needed to determine causal mechanisms. Without such evidence to identify mechanisms, extrapolations among species may lead to counterproductive management and conservation practices.     

The evolution of female social relationships in nonhuman primates

Considerable interspecific variation in female social relationships occurs in gregarious primates, particularly with regard to agonism and cooperation between females and to the quality of female relationships with males. This variation exists alongside variation in female philopatry and dispersal. Socioecological theories have tried to explain variation in female-female social relationships from an evolutionary perspective focused on ecological factors, notably predation and food distribution. According to the current “ecological model”, predation risk forces females of most diurnal primate species to live in groups; the strength of the contest component of competition for resources within and between groups then largely determines social relationships between females. Social relationships among gregarious females are here characterized as Dispersal-Egalitarian, Resident-Nepotistic, Resident-Nepotistic-Tolerant, or Resident-Egalitarian. This ecological model has successfully explained differences in the occurrence of formal submission signals, decided dominance relationships, coalitions and female philopatry. Group size and female rank generally affect female reproduction success as the model predicts, and studies of closely related species in different ecological circumstances underscore the importance of the model. Some cases, however, can only be explained when we extend the model to incorporate the effects of infanticide risk and habitat saturation. We review evidence in support of the ecological model and test the power of alternative models that invoke between-group competition, forced female philopatry, demographic female recruitment, male interventions into female aggression, and male harassment. Not one of these models can replace the ecological model, which already encompasses the between-group competition. Currently the best model, which explains several phenomena that the ecological model does not, is a “socioecological model” based on the combined importance of ecological factors, habitat saturation and infanticide avoidance. We note some points of similarity and divergence with other mammalian taxa; these remain to be explored in detail. Received: 30 September 1996 / Accepted after revision: 20 July 1997