The Importance of Analysis Method for Breeding Bird Survey Population Trend Estimates

Population trends from the Breeding Bird Survey are widely used to focus conservation efforts on species thought to be in decline and to test preliminary hypotheses regarding the causes of these declines. A number of statistical methods have been used to estimate population trends, but there is no consensus as to which is the most reliable. We quantified differences in trend estimates or different analysis methods applied to the same subset of Breeding Bird Survey data. We estimated trends for 115 species in British Columbia using three analysis methods: U.S. National Biological Service route regression, Canadian Wildlife Service route regression, and nonparametric rank-trends analysis. Overall, the number of species estimated to be declining was similar among the three methods, but the number of statistically significant declines was not similar (15, 8, and 29 respectively). In addition, many differences existed among methods in the trend estimates assigned to individual species. Comparing the two route regression methods, Canadian Wildlife Service estimates had a greater absolute magnitude on average than those of the U.S. National Biological Service method. U.S. National Biological Service estimates were on average more positive than the Canadian Wildlife Service estimates when the respective agency's data selection criteria were applied separately. These results imply that our ability to detect population declines and to prioritize species of conservation concern depend strongly upon the analysis method used. This highlights the need for further research to determine how best to accurately estimate trends from the data. We suggest a method for evaluating the performance of the analysis methods by using simulated Breeding Bird Survey data.     

Reliability of Indicators of Decline in Abundance

Abstract: Although there are many indicators of endangerment (i.e., whether populations or species meet criteria that justify conservation action), their reliability has rarely been tested. Such indicators may fail to identify that a population or species meets criteria for conservation action (false negative) or may incorrectly show that such criteria have been met (false positive). To quantify the rate of both types of error for 20 commonly used indicators of declining abundance (threat indicators), we used receiver operating characteristic curves derived from historical (1938–2007) data for 18 sockeye salmon (Oncorhynchus nerka) populations in the Fraser River, British Columbia, Canada. We retrospectively determined each population's yearly status (reflected by change in abundance over time) on the basis of each indicator. We then compared that population's status in a given year with the status in subsequent years (determined by the magnitude of decline in abundance across those years). For each sockeye population, we calculated how often each indicator of past status matched subsequent status. No single threat indicator provided error‐free estimates of status, but indicators that reflected the extent (i.e., magnitude) of past decline in abundance (through comparison of current abundance with some historical baseline abundance) tended to better reflect status in subsequent years than the rate of decline over the previous 3 generations (a widely used indicator). We recommend that when possible, the reliability of various threat indicators be evaluated with empirical analyses before such indicators are used to determine the need for conservation action. These indicators should include estimates from the entire data set to take into account a historical baseline.     

Bayesian Decision Analysis for Evaluating Management Options to Promote Recovery of a Depleted Salmon Population

Abstract:  The endangered population of sockeye salmon (Oncorhynchus nerka) in Cultus Lake, British Columbia, Canada, migrates through commercial fishing areas along with other, much more abundant sockeye salmon populations, but it is not feasible to selectively harvest only the latter, abundant populations. This situation creates controversial trade-offs between recovery actions and economic revenue. We conducted a Bayesian decision analysis to evaluate options for recovery of Cultus Lake sockeye salmon. We used a stochastic population model that included 2 sources of uncertainty that are often omitted from such analyses: structural uncertainty in the magnitude of a potential Allee effect and implementation uncertainty (the deviation between targets and actual outcomes of management actions). Numerous state-dependent, time-independent management actions meet recovery objectives. These actions prescribe limitations on commercial harvest rates as a function of abundance of Cultus Lake sockeye salmon. We also quantified how much reduction in economic value of commercial harvests of the more abundant sockeye salmon populations would be expected for a given increase in the probability of recovery of the Cultus population. Such results illustrate how Bayesian decision analysis can rank options for dealing with conservation risks and can help inform trade-off discussions among decision makers and among groups that have competing objectives.     

Quantifying population declines based on presence‐only records for red‐list assessments

Accurate trend estimates are necessary for understanding which species are declining and which are most in need of conservation action. Imperfect species detection may result in unreliable trend estimates because this may lead to the overestimation of declines. Because many management decisions are based on population trend estimates, such biases could have severe consequences for conservation policy. We used an occupancy‐modeling framework to estimate detectability and calculate nationwide population trends for 14 Swiss amphibian species both accounting for and ignoring imperfect detection. Through the application of International Union for Conservation of Nature Red List criteria to the different trend estimates, we assessed whether ignoring imperfect detection could affect conservation policy. Imperfect detection occurred for all species and detection varied substantially among species, which led to the overestimation of population declines when detectability was ignored. Consequently, accounting for imperfect detection lowered the red‐list risk category for 5 of the 14 species assessed. We demonstrate that failing to consider species detectability can have serious consequences for species management and that occupancy modeling provides a flexible framework to account for observation bias and improve assessments of conservation status. A problem inherent to most historical records is that they contain presence‐only data from which only relative declines can be estimated. A move toward the routine recording of nonobservation and absence data is essential if conservation practitioners are to move beyond this toward accurate population trend estimation.     

Variability in Population Abundance and the Classification of Extinction Risk

Abstract: Classifying species according to their risk of extinction is a common practice and underpins much conservation activity. The reliability of such classifications rests on the accuracy of threat categorizations, but very little is known about the magnitude and types of errors that might be expected. The process of risk classification involves combining information from many sources, and understanding the quality of each source is critical to evaluating the overall status of the species. One common criterion used to classify extinction risk is a decline in abundance. Because abundance is a direct measure of conservation status, counts of individuals are generally the preferred method of evaluating whether populations are declining. Using the thresholds from criterion A of the International Union for Conservation of Nature (IUCN) Red List (critically endangered, decline in abundance of >80% over 10 years or 3 generations; endangered, decline in abundance of 50–80%; vulnerable, decline in abundance of 30–50%; least concern or near threatened, decline in abundance of 0–30%), we assessed 3 methods used to detect declines solely from estimates of abundance: use of just 2 estimates of abundance; use of linear regression on a time series of abundance; and use of state‐space models on a time series of abundance. We generated simulation data from empirical estimates of the typical variability in abundance and assessed the 3 methods for classification errors. The estimates of the proportion of falsely detected declines for linear regression and the state‐space models were low (maximum 3–14%), but 33–75% of small declines (30–50% over 15 years) were not detected. Ignoring uncertainty in estimates of abundance (with just 2 estimates of abundance) allowed more power to detect small declines (95%), but there was a high percentage (50%) of false detections. For all 3 methods, the proportion of declines estimated to be >80% was higher than the true proportion. Use of abundance data to detect species at risk of extinction may either fail to detect initial declines in abundance or have a high error rate.     

Lifting a veil on diversity: a Bayesian approach to fitting relative-abundance models.

Bayesian methods incorporate prior knowledge into a statistical analysis. This prior knowledge is usually restricted to assumptions regarding the form of probability distributions of the parameters of interest, leaving their values to be determined mainly through the data. Here we show how a Bayesian approach can be applied to the problem of drawing inference regarding species abundance distributions and comparing diversity indices between sites. The classic log series and the lognormal models of relative- abundance distribution are apparently quite different in form. The first is a sampling distribution while the other is a model of abundance of the underlying population. Bayesian methods help unite these two models in a common framework. Markov chain Monte Carlo simulation can be used to fit both distributions as small hierarchical models with shared common assumptions. Sampling error can be assumed to follow a Poisson distribution. Species not found in a sample, but suspected to be present in the region or community of interest, can be given zero abundance. This not only simplifies the process of model fitting, but also provides a convenient way of calculating confidence intervals for diversity indices. The method is especially useful when a comparison of species diversity between sites with different sample sizes is the key motivation behind the research. We illustrate the potential of the approach using data on fruit-feeding butterflies in southern Mexico. We conclude that, once all assumptions have been made transparent, a single data set may provide support for the belief that diversity is negatively affected by anthropogenic forest disturbance. Bayesian methods help to apply theory regarding the distribution of abundance in ecological communities to applied conservation.     

Assessing conservation status with extensive but low-resolution data: Application of frequentist and Bayesian models to endangered Athabasca River rainbow trout

Use of extensive but low-resolution abundance data is common in the assessment of species at-risk status based on quantitative decline criteria under International Union for Conservation of Nature (IUCN) and national endangered species legislation. Such data can be problematic for 3 reasons. First, statistical power to reject the null hypothesis of no change is often low because of small sample size and high sampling uncertainty leading to a high frequency of type II errors. Second, range-wide assessments composed of multiple site-specific observations do not effectively weight site-specific trends into global trends. Third, uncertainty in site-specific temporal trends and relative abundance are not propagated at the appropriate spatial scale. A common result is the propensity to underestimate the magnitude of declines and therefore fail to identify the appropriate at-risk status for a species. We used 3 statistical approaches, from simple to more complex, to estimate temporal decline rates for a designatable unit (DU) of rainbow trout in the Athabasca River watershed in western Canada. This DU is considered a native species for purposes of listing because of its genetic composition characterized as >0.95 indigenous origin in the face of continuing introgressive hybridization with introduced populations in the watershed. Analysis of abundance trends from 57 time series with a fixed-effects model identified 33 sites with negative trends, but only 2 were statistically significant. By contrast, a hierarchical linear mixed model weighted by site-specific abundance provided a DU-wide decline estimate of 16.4% per year and a 3-generation decline of 93.2%. A hierarchical Bayesian mixed model yielded a similar 3-generation decline trend of 91.3% and the posterior distribution showed that the estimate had a >99% probability of exceeding thresholds for an endangered listing. We conclude that the Bayesian approach was the most useful because it provided a probabilistic statement of threshold exceedance in support of an at-risk status recommendation.     

The Importance of Being Spatial (and Reserved): Assessing Northern Spotted Owl Habitat Relationships with Hierarchical Bayesian Models

Abstract:  Regional conservation planning increasingly draws on habitat suitability models to support decisions regarding land allocation and management. Nevertheless, statistical techniques commonly used for developing such models may give misleading results because they fail to account for 3 factors common in data sets of species distribution: spatial autocorrelation, the large number of sites where the species is absent (zero inflation), and uneven survey effort. We used spatial autoregressive models fit with Bayesian Markov Chain Monte Carlo techniques to assess the relationship between older coniferous forest and the abundance of Northern Spotted Owl nest and activity sites throughout the species' range. The spatial random-effect term incorporated in the autoregressive models successfully accounted for zero inflation and reduced the effect of survey bias on estimates of species–habitat associations. Our results support the hypothesis that the relationship between owl distribution and older forest varies with latitude. A quadratic relationship between owl abundance and older forest was evident in the southern portion of the range, and a pseudothreshold relationship was evident in the northern portion of the range. Our results suggest that proposed changes to the network of owl habitat reserves would reduce the proportion of the population protected by up to one-third, and that proposed guidelines for forest management within reserves underestimate the proportion of older forest associated with maximum owl abundance and inappropriately generalize threshold relationships among subregions. Bayesian spatial models can greatly enhance the utility of habitat analysis for conservation planning because they add the statistical flexibility necessary for analyzing regional survey data while retaining the interpretability of simpler models.     

Response of Bird Populations to Farmland Set-Aside Programs

Abstract:  The role of agricultural landscapes in biodiversity conservation has been largely ignored despite their potential role in conserving declining species. Within agricultural landscapes, set-aside programs may offer the most promising conservation opportunities due to the large area involved in these programs. I explored the relationship between set-aside effect size—on the basis of data from field studies of birds in cropland and set-aside fields—and population changes following establishment of these fields. Species whose abundance was most strongly influenced by the establishment of set-aside lands also tended to show positive changes in population trends following broad-scale implementation of the set-aside program. This relationship was strongest for grassland obligate birds, a group of birds experiencing broad-scale population declines throughout North America. There is now increasing evidence that set-aside lands within the United States are providing population-level benefits to grassland birds. Nevertheless, there are also increasing concerns about the stability of these set-aside lands in the face of increased demand for crops and rising commodity prices. If set-aside area within the United States is allowed to decline, additional declines of birds and perhaps other taxa within agricultural landscapes seem likely.     

Toward accurate and precise estimates of lion density

Reliable estimates of animal density are fundamental to understanding ecological processes and population dynamics. Furthermore, their accuracy is vital to conservation because wildlife authorities rely on estimates to make decisions. However, it is notoriously difficult to accurately estimate density for wide‐ranging carnivores that occur at low densities. In recent years, significant progress has been made in density estimation of Asian carnivores, but the methods have not been widely adapted to African carnivores, such as lions (Panthera leo). Although abundance indices for lions may produce poor inferences, they continue to be used to estimate density and inform management and policy. We used sighting data from a 3‐month survey and adapted a Bayesian spatially explicit capture‐recapture (SECR) model to estimate spatial lion density in the Maasai Mara National Reserve and surrounding conservancies in Kenya. Our unstructured spatial capture‐recapture sampling design incorporated search effort to explicitly estimate detection probability and density on a fine spatial scale, making our approach robust in the context of varying detection probabilities. Overall posterior mean lion density was estimated to be 17.08 (posterior SD 1.310) lions >1 year old/100 km², and the sex ratio was estimated at 2.2 females to 1 male. Our modeling framework and narrow posterior SD demonstrate that SECR methods can produce statistically rigorous and precise estimates of population parameters, and we argue that they should be favored over less reliable abundance indices. Furthermore, our approach is flexible enough to incorporate different data types, which enables robust population estimates over relatively short survey periods in a variety of systems. Trend analyses are essential to guide conservation decisions but are frequently based on surveys of differing reliability. We therefore call for a unified framework to assess lion numbers in key populations to improve management and policy decisions.     

Site-selection bias and apparent population declines in long-term studies

Detecting population declines is a critical task for conservation biology. Logistical difficulties and the spatiotemporal variability of populations make estimation of population declines difficult. For statistical reasons, estimates of population decline may be biased when study sites are chosen based on abundance of the focal species. In this situation, apparent population declines are likely to be detected even if there is no decline. This site-selection bias is mentioned in the literature but is not well known. We used simulations and real population data to examine the effects of site-selection biases on inferences about population trends. We used a left-censoring method to detect population-size patterns consistent with site-selection bias. The site-selection bias is an important consideration for conservation biologists, and we offer suggestions for minimizing or mitigating it in study design and analysis. Article impact statement: Estimates of population declines are biased if studies begin in large populations, and time-series data show a signature of such an effect.     

Risk-Based Viable Population Monitoring

Abstract:  We describe risk-based viable population monitoring, in which the monitoring indicator is a yearly prediction of the probability that, within a given timeframe, the population abundance will decline below a prespecified level. Common abundance-based monitoring strategies usually have low power to detect declines in threatened and endangered species and are largely reactive to declines. Comparisons of the population's estimated risk of decline over time will help determine status in a more defensible manner than current monitoring methods. Monitoring risk is a more proactive approach; critical changes in the population's status are more likely to be demonstrated before a devastating decline than with abundance-based monitoring methods. In this framework, recovery is defined not as a single evaluation of long-term viability but as maintaining low risk of decline for the next several generations. Effects of errors in risk prediction techniques are mitigated through shorter prediction intervals, setting threshold abundances near current abundance, and explicitly incorporating uncertainty in risk estimates. Viable population monitoring also intrinsically adjusts monitoring effort relative to the population's true status and exhibits considerable robustness to model misspecification. We present simulations showing that risk predictions made with a simple exponential growth model can be effective monitoring indicators for population dynamics ranging from random walk to density dependence with stable, decreasing, or increasing equilibrium. In analyses of time-series data for five species, risk-based monitoring warned of future declines and demonstrated secure status more effectively than statistical tests for trend.     

Using a Bayesian network to clarify areas requiring research in a host–pathogen system

Bayesian network analyses can be used to interactively change the strength of effect of variables in a model to explore complex relationships in new ways. In doing so, they allow one to identify influential nodes that are not well studied empirically so that future research can be prioritized. We identified relationships in host and pathogen biology to examine disease‐driven declines of amphibians associated with amphibian chytrid fungus (Batrachochytrium dendrobatidis). We constructed a Bayesian network consisting of behavioral, genetic, physiological, and environmental variables that influence disease and used them to predict host population trends. We varied the impacts of specific variables in the model to reveal factors with the most influence on host population trend. The behavior of the nodes (the way in which the variables probabilistically responded to changes in states of the parents, which are the nodes or variables that directly influenced them in the graphical model) was consistent with published results. The frog population had a 49% probability of decline when all states were set at their original values, and this probability increased when body temperatures were cold, the immune system was not suppressing infection, and the ambient environment was conducive to growth of B. dendrobatidis. These findings suggest the construction of our model reflected the complex relationships characteristic of host–pathogen interactions. Changes to climatic variables alone did not strongly influence the probability of population decline, which suggests that climate interacts with other factors such as the capacity of the frog immune system to suppress disease. Changes to the adaptive immune system and disease reservoirs had a large effect on the population trend, but there was little empirical information available for model construction. Our model inputs can be used as a base to examine other systems, and our results show that such analyses are useful tools for reviewing existing literature, identifying links poorly supported by evidence, and understanding complexities in emerging infectious‐disease systems.     

Experimental test of parasitism hypothesis for population cycles of a forest lepidopteran

Population cycles of herbivores are thought to be driven by trophic interaction mechanisms, either between food plant and herbivore or between the herbivorous prey and its natural enemies. Observational data have indicated that hymenopteran parasitoids cause delayed density-dependent mortality in cyclic autumnal moth (Epirrita autumnata) populations. We experimentally tested the parasitism hypothesis of moth population cycles by establishing a four-year parasitoid-exclusion experiment, with parasitoid-proof exclosures, parasitoid-permeable exclosures, and control plots. The exclusion of parasitoids led to high autumnal moth abundances, while the declining abundance in both the parasitoid-permeable exclosures and the control plots paralleled the naturally declining density in the study area and could be explained by high rates of parasitism. Our results provide firm experimental support for the hypothesis that hymenopteran parasitoids have a causal relationship with the delayed density-dependent component required in the generation of autumnal moth population cycles.     

Evaluation of Local Ecological Knowledge as a Method for Collecting Extensive Data on Animal Abundance

Abstract:  The use of local ecological knowledge (LEK) has been advocated for biodiversity monitoring and management. To date, however, it has been underused in studying wild populations of animals and, particularly, in obtaining quantitative abundance estimates. We evaluated LEK as a tool for collecting extensive data on local animal abundance and population trends. We interviewed shepherds in southeastern Spain, asking them to estimate the local abundance of the terrestrial tortoise Testudo graeca . We quantified reliability of abundance estimates derived from interviews by comparing them with those obtained from standard field-sampling protocols (distance sampling). We also explored the complementarity of these 2 approaches. LEK provided high-quality and low-cost information about both distribution and abundance of T. graeca . Interviews with shepherds yielded abundance estimates in a much wider range than linear transects, which only detected the species in the upper two-thirds of its abundance range. Abundance estimates from both methodologies showed a close relationship. Analysis of confidence intervals indicated local knowledge could be used to estimate mean local abundances and to detect mean population trends. A cost analysis determined that the information derived from LEK was 100 times cheaper than that obtained through linear-transect surveys. Our results should further the use of LEK as a standard tool for sampling the quantitative abundance of a great variety of taxa, particularly when population densities are low and traditional sampling methods are expensive or difficult to implement.     

Choosing a Management Strategy: Two Structured Decision-Making Methods for Evaluating the Predictions of Stochastic Simulation Models

We describe two structured decision-making methods—one using a hierarchy of goals and a second using ranking on the sum of weighted criteria—that may be useful for many practical conservation problems, particularly when advisory groups evaluate the output of simulation models. We illustrate both methods by applying them to the problem of choosing a management strategy to address the "mobbing" problem in endangered Hawaiian monk seals. Both methods require estimates of the probabilities of various outcomes, such as a population size of more than 400 seals after 20 years under a specific management regime. We used a simulation model of a small monk seal population to generate these probabilities. Both methods provide an explicit, well-documented, and reproducible decision process that helps justify the decision. Furthermore, they are easy for those untrained in decision analysis to understand and use, they focus discussion on management objectives, they facilitate an examination of trade-offs in the light of multiple and sometimes conflicting objectives, they are suitable for use in workshops, and, at least in our example, they lead to management recommendations that are not highly sensitive to minor changes in probability estimates or other factors.     

Population level effects of reduced fecundity in the fish species perch (Perca fluviatilis) and the implications for environmental monitoring

A 40% reduction in relative gonad size in perch (Perca fluviatilis) has been observed over that past two decades at the Swedish national reference site Kvädöfjärden. This biomarker response could be interpreted as a reduction in fecundity and increased risk of local extinction. However, abundance estimates from the same area has not provided any evidence of a reduction in population size. In the present study, a matrix population model was developed to investigate if a reduction in fecundity can be expected to have long term effects on population viability for perch and to evaluate the probability to detect such effects through abundance estimates. The model was parameterized from 17 years of population data from Kvädöfjärden as well as from other studies on perch. The model included density dependence and environmental stochasticity. The results indicated that a reduction in fecundity that is in level with the observed reduction in relative gonad size in Kvädöfjärden will cause a substantial risk for local extinction. The risk that the population will fall below 20% of the carrying capacity within 50 years is 44% when the fecundity is reduced by 40%. However, due to variability in abundance measurements it will take some time before a reduction in gonad size leads to statistically significant effects on the population. If the fecundity is reduced by 40% successively over a 10-year period, the probability to detect this through abundance estimates within 10 years is less than 50%. The results of the present study clearly show that relevant biomarkers have an important role in environmental monitoring as early warning signals, preferably in combination with measurements at higher levels of biological organization.     

Contrasting natural experiments confirm competition between House Finches and House Sparrows

After House Finches were introduced from the western to the eastern United States and rapidly increased in numbers, House Sparrows declined, leading to suggestions that the decline was caused by interspecific competition. However, other potential causes were not excluded. The rapid decline in House Finches following the emergence of a new disease (mycoplasmal conjunctivitis) caused by a novel strain of Mycoplasma gallisepticum (MG) in 1994 has provided a natural experiment and an opportunity to revisit the hypothesis that interspecific competition from House Finches drives population changes in House Sparrows. If true, the recent decline in House Finches should lead to an increase in House Sparrows. In this paper we test the hypothesis that House Sparrow and House Finch numbers in the northeastern United States vary inversely by examining data from three independent volunteer programs that monitor bird species' abundance and distribution (Christmas Bird Count, Project FeederWatch, and Breeding Bird Survey). In the first analysis we found that House Sparrow and House Finch numbers varied inversely during a time interval when House Finches were increasing and a time interval when House Finches were decreasing. In the second analysis, we found that the rates of geometric change in House Sparrow abundance (ln[HOSP(t+1)/HOSP(t)]) were negatively correlated with initial House Finch (HOFI(t)) and sparrow (HOSP(t)) abundances at individual sites, irrespective of the time period. Given that finch range expansion and subsequent declines in abundance are the result of two very different phenomena, it would be very unlikely for apparent competition or spurious correlations to cause the observed concomitant changes in House Sparrow abundance. We conclude that interspecific competition exists between these two species.     

Monitoring,imperfect detection,and risk optimization of a Tasmanian devil insurance population

Most species are imperfectly detected during biological surveys, which creates uncertainty around their abundance or presence at a given location. Decision makers managing threatened or pest species are regularly faced with this uncertainty. Wildlife diseases can drive species to extinction; thus, managing species with disease is an important part of conservation. Devil facial tumor disease (DFTD) is one such disease that led to the listing of the Tasmanian devil (Sarcophilus harrisii) as endangered. Managers aim to maintain devils in the wild by establishing disease‐free insurance populations at isolated sites. Often a resident DFTD‐affected population must first be removed. In a successful collaboration between decision scientists and wildlife managers, we used an accessible population model to inform monitoring decisions and facilitate the establishment of an insurance population of devils on Forestier Peninsula. We used a Bayesian catch‐effort model to estimate population size of a diseased population from removal and camera trap data. We also analyzed the costs and benefits of declaring the area disease‐free prior to reintroduction and establishment of a healthy insurance population. After the monitoring session in May–June 2015, the probability that all devils had been successfully removed was close to 1, even when we accounted for a possible introduction of a devil to the site. Given this high probability and the baseline cost of declaring population absence prematurely, we found it was not cost‐effective to carry out any additional monitoring before introducing the insurance population. Considering these results within the broader context of Tasmanian devil management, managers ultimately decided to implement an additional monitoring session before the introduction. This was a conservative decision that accounted for uncertainty in model estimates and for the broader nonmonetary costs of mistakenly declaring the area disease‐free.     

Judicious Use of Multiple Hypothesis Tests

Abstract:  When analyzing a table of statistical results, one must first decide whether adjustment of significance levels is appropriate. If the main goal is hypothesis generation or initial screening for potential conservation problems, then it may be appropriate to use the standard comparisonwise significance level to avoid Type II errors (not detecting real differences or trends). If the main goal is rigorous testing of a hypothesis, however, then an adjustment for multiple tests is needed. To control the familywise Type I error rate (the probability of rejecting at least one true null hypothesis), sequential modifications of the standard Bonferroni method, such as Holm's method, will provide more statistical power than the standard Bonferroni method. Additional power may be achieved through procedures that control the false discovery rate (FDR) (the expected proportion of false positives among tests found to be significant). Holm's sequential Bonferroni method and two FDR-controlling procedures were applied to the results of multiple-regression analyses of the relationship between habitat variables and the abundance of 25 species of forest birds in Japan, and the FDR-controlling procedures provided considerably greater statistical power.