Experimental and environmental factors affect spurious detection of ecological thresholds

Threshold detection methods are increasingly popular for assessing nonlinear responses to environmental change, but their statistical performance remains poorly understood. We simulated linear change in stream benthic macroinvertebrate communities and evaluated the performance of commonly used threshold detection methods based on model fitting (piecewise quantile regression [PQR]), data partitioning (nonparametric change point analysis [NCPA]), and a hybrid approach (significant zero crossings [SiZer]). We demonstrated that false detection of ecological thresholds (type I errors) and inferences on threshold locations are influenced by sample size, rate of linear change, and frequency of observations across the environmental gradient (i.e., sample-environment distribution, SED). However, the relative importance of these factors varied among statistical methods and between inference types. False detection rates were influenced primarily by user-selected parameters for PQR (tau) and SiZer (bandwidth) and secondarily by sample size (for PQR) and SED (for SiZer). In contrast, the location of reported thresholds was influenced primarily by SED. Bootstrapped confidence intervals for NCPA threshold locations revealed strong correspondence to SED. We conclude that the choice of statistical methods for threshold detection should be matched to experimental and environmental constraints to minimize false detection rates and avoid spurious inferences regarding threshold location.     

Exploring alternative states in ecological systems with a qualitative analysis of community feedback

Demonstrating and predicting the existence of alternative states in natural communities remains a challenge for ecologists and is essential for resource managers. Positive feedback is often presented as central in maintaining alternative ecosystem states, but no formal approach relates this part of theory to real world applications. Through qualitative modelling of community response to long-term perturbations, we define generic mechanistic links between positive feedback and the occurrence of alternative states. Positive feedback diminishes a system's overall resistance to change, and can create and maintain correlations in the relative abundance of variables that coincide with alternative states.Through specific models of the dynamics of Tasmanian rocky-reef communities, which are affected by climate and fishing and persist within alternative states, we demonstrate the ability of our theoretical framework to predict alternative states in ecosystems and inform management intervention. A qualitative knowledge of community structure permits a thorough analysis of system feedback and an assessment of the potential for an ecosystem to exhibit alternative states. We illustrate the usefulness of the approach to inform management priorities, and to focus monitoring and field research on the key drivers of ecosystem dynamics.     

The apportionment of quadratic entropy: a useful alternative for partitioning diversity in ecological data

Many methods that study the diversity within hierarchically structured populations have been developed in genetics. Among them, the analysis of molecular variance (AMOVA) (Excoffier et al., 1992) has the advantage of including evolutionary distances between individuals. AMOVA is a special case of a far more general statistical scheme produced by Rao (1982a; 1986) and called the apportionment of quadratic entropy (APQE). It links diversity and dissimilarity and allows the decomposition of diversity according to a given hierarchy. We apply this framework to ecological data showing that APQE may be very useful for studying diversity at various spatial scales. Moreover, the quadratic entropy has a critical advantage over usual diversity indices because it takes into account differences between species. Finally, the differences that can be incorporated in APQE may be either taxonomic or functional (biological traits), which may be of critical interest for ecologists.     

The value of small size: loss of forest patches and ecological thresholds in southern Madagascar.

Many services generated by forest ecosystems provide essential support for human well-being. However, the vulnerability of these services to environmental change such as forest fragmentation are still poorly understood. We present spatial modeling of the generation of ecosystem services in a human-dominated landscape where forest habitat patches, protected by local taboos, are located in a matrix of cultivated land in southern Madagascar. Two ecosystem services dependent on the forest habitats were addressed: (1) crop pollination services by wild and semidomesticated bees (Apoidea), essential for local crop production of, for example, beans, and (2) seed dispersal services based on the presence of ring-tailed lemurs (Lemur catta). We studied the vulnerability of these ecosystem services to a plausible scenario of successive destruction of the smallest habitat patches. Our results indicate that, in spite of the fragmented nature of the landscape, the fraction of the landscape presently covered by both crop pollination and seed dispersal services is surprisingly high. It seems that the taboo system, though indirectly and unintentionally, contributes to upholding the generation of these services by protecting the forest patches. Both services are, however, predicted to be very vulnerable to the successive removal of small patches. For crop pollination, the rate of decrease in cover was significant even when only the smallest habitat patches were removed. The capacity for seed dispersal across the landscape displayed several thresholds with habitat patch removal. Our results suggest that, in order to maintain capacity for seed dispersal across the landscape and crop pollination cover in southern Androy, the geographical location of the remaining forest patches is more crucial than their size. We argue that in heavily fragmented production landscapes, small forest patches should increasingly be viewed as essential for maintaining ecosystem services, such as agricultural production, and also should be considered in the ongoing process of tripling the area of protected habitats in Madagascar.     

The structure of Marine Ecosystems: An alternative

In The Structure of Marine Ecosystems (1974), Steele developed a simulation model which suggested that the stability of planktonic communities was dependent on a threshold response allowing phytoplankton a refuge in concentration below which herbivorous zooplankton did not feed. Observations from the North Sea were presented to support this hypothesis. In order to more realistically represent the simulated herbivores, Calanus spp., several of Steele's assumptions have been modified. The results of the revised model indicate that feeding thresholds are not essential for the long-term realism and stability of the simulated system and suggest an alternative, ecosystem control through predation.Contribution no. 861 from the Department of Oceanography, University of Washington. This study was supported by NSF Grant DES 74-22640 to B.W. Frost.     

Paleobiology, community ecology, and scales of ecological pattern

The fossil record provides a wealth of data on the role of regional processes and historical events in shaping biological communities over a variety of time scales. The Quaternary record with its evidence of repeated climatic change shows that both terrestrial and marine species shifted independently rather than as cohesive assemblages over scales of thousands of years. Larger scale patterns also show a strong individualistic component to taxon dynamics; assemblage stability, when it occurs, is difficult to separate from shared responses to low rates of environmental change. Nevertheless, the fossil record does suggest that some biotic interactions influence large-scale ecological and evolutionary patterns, albeit in more diffuse and protracted fashions than those generally studied by community ecologists. These include: (1) the resistance by incumbents to the establishment of new or invading taxa, with episodes of explosive diversification often appearing contingent on the removal of incumbents at extinction events; (2) steady states of within-habitat and global diversity at longer time scales (10(7)-l0(8) yr), despite enormous turnover of taxa; and (3) morphological and biogeographic responses to increased intensities of predation and substratum disturbance over similarly long time scales. The behavior of species and communities over the array of temporal and spatial scales in the fossil record takes on additional significance for framing conservation strategies, and for understanding recovery of species, lineages, and communities from environmental changes.     

Sensitivity of ecological models to their climate drivers: statistical ensembles for forcing.

Global and regional numerical models for terrestrial ecosystem dynamics require fine spatial resolution and temporally complete historical climate fields as input variables. However, because climate observations are unevenly spaced and have incomplete records, such fields need to be estimated. In addition, uncertainty in these fields associated with their estimation are rarely assessed. Ecological models are usually driven with a geostatistical model's mean estimate (kriging) of these fields without accounting for this uncertainty, much less evaluating such errors in terms of their propagation in ecological simulations. We introduce a Bayesian statistical framework to model climate observations to create spatially uniform and temporally complete fields, taking into account correlation in time and space, spatial heterogeneity, lack of normality, and uncertainty about all these factors. A key benefit of the Bayesian model is that it generates uncertainty measures for the generated fields. To demonstrate this method, we reconstruct historical monthly precipitation fields (a driver for ecological models) on a fine resolution grid for a climatically heterogeneous region in the western United States. The main goal of this work is to evaluate the sensitivity of ecological models to the uncertainty associated with prediction of their climate drivers. To assess their numerical sensitivity to predicted input variables, we generate a set of ecological model simulations run using an ensemble of different versions of the reconstructed fields. We construct such an ensemble by sampling from the posterior predictive distribution of the climate field. We demonstrate that the estimated prediction error of the climate field can be very high. We evaluate the importance of such errors in ecological model experiments using an ensemble of historical precipitation time series in simulations of grassland biogeochemical dynamics with an ecological numerical model, Century. We show how uncertainty in predicted precipitation fields is propagated into ecological model results and that this propagation had different modes. Depending on output variable, the response of model dynamics to uncertainty in inputs ranged from uncertainty in outputs that matched that of inputs to those that were muted or that were biased, as well as uncertainty that was persistent in time after input errors dropped.     

An ecological model for heavy metal contamination of crops and ground water

A model for heavy metal contamination of crops and ground water was set up and tested. It was found possible to use the model to predict the heavy metal concentration in crops.The model indicates that Pb is limiting the use of sludge as a soil conditioner, and that in most cases, contamination of crops seems to be a greater danger than contamination of ground water. However, when soil with a high humus or clay content is used at a pH close to 7.00, municipal sludge can be used in considerable quantities without danger.     

An individual-oriented model for ecological risk assessment of wading birds

The contribution of certain contaminants to reproductive failure in many avian species has been an ongoing concern. Appropriate quantitative techniques have focused either on the individual organisms by providing explicit bioaccumulation dynamics or on whole ecosystems by looking at the fate of the contaminant but fail to make the necessary link via population dynamics of interacting individuals. We used the individual-oriented approach in an effort to quantify effects of chronic contaminant exposure on individual birds. This was made possible by the use of an object-oriented model, where individual birds are interacting objects, and their actions are implemented by passing to them appropriate messages. Using this modeling approach a breeding colony of Great Blue Herons (Ardea herodias) is simulated as an assemblage of interacting individuals whose daily actions (foraging, growth, feeding of the young) are simultaneously followed over short time intervals for a nesting season. Spatial distribution of the contaminants in prey resources is used on a cell by cell basis and their effects on certain behavior characteristics of adult birds (e.g. foraging efficiency, effects on flying efficiency, parental care) are taken into account. Results showed that sublethal effects could have a considerable effect on colony success. Appropriate selection of endpoints for risk assessment yields a variety of scenarios for colony success.     

Ecological network analyses and their use for establishing reference domain in functional assessment of an estuary

We developed a framework to use ecological network analysis for functional assessment of large aquatic ecosystems in the context of ecosystem-based management. We established a reference domain for the Neuse River Estuary, North Carolina, USA, from changes over time. Four reference network models of the trophic structure of the estuary during early and late summers of 1997 and 1998 were constructed and analyzed. The estuary has experienced various symptoms of eutrophication during the past 20 years, including summer-time hypoxia and fish kills. The networks were used to quantify indices of nominal trophic dynamics and their variation. The ratio of biomass of nekton to that of macrobenthos, derived from network construction, was used to index severity of eutrophication and to promote accessibility of ecological network analysis to environmental management. The ratio increased from early to late summer, and network metrics demonstrated a variety of responses in association with that change. Some variables from network analysis, especially related to consumers, reflected some but not all of this change. Others reflected the most severe increase in the ratio in late summer 1997 when hypoxia was most extensive. We evaluated uncertainty and the modulating effects of hierarchy by comparing variation of input biomasses with integrative response variables. Relative variation in input variables was generally greater than that of the integrative response variables as predicted by hierarchy theory. Ecological network analysis has previously served as support for ecosystem-based management of large aquatic systems with some success. However, its use can be enhanced by making it more accessible to environmental managers and policy makers. Ways to do this include promoting simple metrics from network construction and explicitly associating network analysis to concepts familiar to the management community, such as functional assessment and reference.     

Social and ecological influences on activity budgets of vervet monkeys,and their implications for group living

Summary Activity patterns were documented over a 20-month period in six groups of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya. Group size varied both among groups and within groups through time. The home ranges of two groups were located in Acacia xanthophloea habitat whereas those of four adjacent groups were located in A. tortilis habitat. Repeated measures analysis of variance, three-way analysis of variance, and meta-analysis of group size effects were carried out on feeding, moving, resting, scanning, allo-grooming, and other behavior. Time spent feeding varied seasonally, peaking during periods when A. tortilis seeds were eaten. However, time spent feeding did not vary within or among groups, or between habitats, males and females, or dominants and subordinates. Males spend more time scanning and less time allo-grooming than females, and high-ranking individuals of both sexes spent more time scanning than low-ranking individuals. Among females, subordinates spent more time moving, whereas among males, dominants spent more time moving. Groups living in A. xanthophloea habitat spent more time moving and allo-grooming than groups in A. tortilis habitat but this may have been confounded by the fact that these two groups were on average larger than the others. Significant differences in activity budgets among groups demonstrate that activity patterns observed in one group are not always readily generalized to other groups even when they come from the same population. Group size analyses examined potential costs and benefits of group living. Increased intragroup competition, measured by time spent moving and feeding, was only weakly evident in larger groups. Individuals in larger groups allo-groomed more than individuals in smaller groups. Self-grooming also increased in larger groups. Individuals in larger groups spent less time scanning than individuals in smaller groups, but there was more scanning per group in larger groups than in smaller groups. Correspondence to: L.A. Isbell     

东北森林土壤动物同功能种团及其生态分布

对位于小兴安岭的东北林业大学凉水自然保护区和位于张广才岭的东北林业大学帽儿山实验林场的土壤动物进行了研究。结果表明,该地区土壤动物可划分为腐食性、植食性,捕食性动物三个同功能种团,土壤动物各同功能种团在组成、个体数量、生物量等方面均具有相对稳定性,群落土壤动物同功能种团的数量分布状况能在很大程度上反映群落环境的相似与差异程度;群落中各土壤动物同功能种团的个体数量间无明显的相关性,但腐食性动物与捕食性动物两个同功能种团的生物量之间具有的相关性。图1表2参13     

Geographic assignment of seabirds to their origin: combining morphologic, genetic, and biogeochemical analyses.

Longline fisheries, oil spills, and offshore wind farms are some of the major threats increasing seabird mortality at sea, but the impact of these threats on specific populations has been difficult to determine so far. We tested the use of molecular markers, morphometric measures, and stable isotope (delta15N and delta13C) and trace element concentrations in the first primary feather (grown at the end of the breeding period) to assign the geographic origin of Calonectris shearwaters. Overall, we sampled birds from three taxa: 13 Mediterranean Cory's Shearwater (Calonectris diomedea diomedea) breeding sites, 10 Atlantic Cory's Shearwater (Calonectris diomedea borealis) breeding sites, and one Cape Verde Shearwater (C. edwardsii) breeding site. Assignment rates were investigated at three spatial scales: breeding colony, breeding archipelago, and taxa levels. Genetic analyses based on the mitochondrial control region (198 birds from 21 breeding colonies) correctly assigned 100% of birds to the three main taxa but failed in detecting geographic structuring at lower scales. Discriminant analyses based on trace elements composition achieved the best rate of correct assignment to colony (77.5%). Body measurements or stable isotopes mainly succeeded in assigning individuals among taxa (87.9% and 89.9%, respectively) but failed at the colony level (27.1% and 38.0%, respectively). Combining all three approaches (morphometrics, isotopes, and trace elements on 186 birds from 15 breeding colonies) substantially improved correct classifications (86.0%, 90.7%, and 100% among colonies, archipelagos, and taxa, respectively). Validations using two independent data sets and jackknife cross-validation confirmed the robustness of the combined approach in the colony assignment (62.5%, 58.8%, and 69.8% for each validation test, respectively). A preliminary application of the discriminant model based on stable isotope delta15N and delta13C values and trace elements (219 birds from 17 breeding sites) showed that 41 Cory's Shearwaters caught by western Mediterranean long-liners came mainly from breeding colonies in Menorca (48.8%), Ibiza (14.6%), and Crete (31.7%). Our findings show that combining analyses of trace elements and stable isotopes on feathers can achieve high rates of correct geographic assignment of birds in the marine environment, opening new prospects for the study of seabird mortality at sea.     

Implications of ecological data constraints for integrated policy and livelihoods modelling: An example from East Kalimantan, Indonesia

Policy and human livelihoods modelling increasingly demands integrated research which requires ecological expertise. However, contributions from ecologists are often based on sparse data. Rather than discounting such data, in this paper we demonstrate how ecological modelling can effectively contribute to the development of policy recommendations in spite of data constraints. In a petrol subsidy analysis in East Kalimantan, Indonesia, we accounted for ecological data uncertainty by (a) assuming large parameter value ranges and (b) conducting a robustness test for policy recommendations. In addition to data scarcity, counter-intuitive results emerged emphasising the need for model validation. These counter-intuitive results indicated that decreasing petrol prices led to increased poverty. This informed a policy recommendation to prevent the reduction of petrol prices below IDR 5500 per litre. Using two key livelihood resources (fish and honey), we found that while a traditional sensitivity analysis suggested highly robust results, a robustness test indicated that policy recommendations would change if the incorrectness of parameter values approached 50%. The results show that ecological modelling can contribute effectively in spite of sparse data to guide policy, as well as identifying future research priorities.     

Types of larval development in marine bottom invertebrates,their distribution and ecological significance: a re-evaluation

The distribution of various types of larval development among marine bottom invertebrates has been discussed on the basis of ecological evidence by Thorson (1936, 1946, 1950, 1952) and Mileikovsky (1961b, 1965). The information at hand is reviewed anew in this paper and is re-evaluated in the light of modern pertinent literature. The interrelationships between certain larval types and their distribution are not as rigid and direct as originally assumed. This can be proved even by the copy book example of the distribution of the various forms of development among species of the coastal gastropod genus Littorina. Especially among species with wide distributional areas, local populations may exhibit greater diversity in larval types than has previously been thought. Different types of larval development have now become known to exist in different populations of opisthobranch gastropods and lamellibranchs, i.e., in invertebrate groups in which such variability had been ruled out by Thorson. Variability in the type of larval development within given species — as a function of geographical, seasonal and other environmental parameters —is also more common in other marine bottom invertebrates than formerly considered. Marine bottom invertebrates are characterized not only by the 3 main different types of larval development proposed by Thorson (pelagic, direct, viviparous), but also by a fourth type: demersal (free non-pelagic) development. This fourth type occurs at all water depths and in all geographic zones of the oceans. The most important of the 4 types is pelagic (planktotrophic) development. Thorson's rule (decrease in numbers of species possessing pelagic development from the Equator towards the Poles, and from shallow-shelf waters to greater oceanic depths) is well substantiated by new data. However, one correction is necessary: pelagic development is not completely absent in the abyssal zone, as was proposed by Thorson (1950, and later), but is represented in it by at least several species belonging to various groups of invertebrates, and is also fairly common in the bathyal zone. A detailed analysis of the distributional pattern of the different types of development of marine bottom invertebrates must further take into consideration asexual reproduction with all its different modifications. Asexual reproduction in benthonic invertebrates is ecologically significant because of its common occurrence in nature; in numerous species it is also important as a biological supplement to sexual reproduction. The vast majority of species inhabiting the shallow-shelf zone and, partly, the higher levels of the slope zone of ocean areas located roughly between the polar circles, reveals development by means of planktotrophic larval stages. In the highest latitudes and on the slopes to abyssal depths—characterized by low water temperatures, scarcity of food, increasing hydrostatic pressure and other environmental peculiarities—other types of larval development prevail and, progressively, replace pelagic development with increasing latitude or depth. The distributional patterns of the various types of development among marine bottom invertebrates form one of the most important factors determining the basic distributional dynamics of the whole benthos in all oceans, both in the geological past and at the present time.Dedicated to the memory of Professor G. Thorson —founder of modern reproductive and larval ecology of marine bottom invertebrates.     

Modelling the responses of wildlife to human disturbance: An evaluation of alternative management scenarios for black-crowned night-herons

The impact of anthropogenic disturbance on wildlife is increasing becoming a source of concern as the popularity of outdoor recreation rises. There is now more pressure on site managers to simultaneously ensure the continued persistence of wildlife and provide recreational opportunities. Using ‘Simulation of Disturbance Activities’, a model designed to investigate the impact of recreational disturbance on wildlife, we demonstrate how a simulation modelling approach can effectively inform such management decisions. As an example, we explored the implications of various design and management options for a proposed recreational area containing a historic breeding bird colony. By manipulating the proximity, orientation and intensity of recreation, we were able to evaluate the impact of recreational activities on the behaviour of black-crowned night-heron nestlings (Nycticorax nycticorax). Using a classification and regression tree (CART) procedure to analyse simulation output, we explored the dynamics of multiple strategies in concert. Our analysis revealed that there are inherent advantages in implementing multiple strategies as opposed to any single strategy. Nestlings were not disturbed by recreation when bird-watching facility placement (proximity and orientation) and type were considered in combination. In comparison, proximity alone only led to a <10% reduction in disturbance. Thus we demonstrate how simulation models based on customised empirical data can bridge the gap between field studies and active management, enabling users to test novel management scenarios that are otherwise logistically difficult. Furthermore, such models potentially have broad application in understanding human-wildlife interactions (e.g. exploring the implications of roads on wildlife, probability of bird strikes around airports, etc.). They therefore represent a valuable decision-making tool in the ecological design of urban infrastructures.