A noninvasive demographic assessment of sea lions based on stage-specific abundances

A pressing need exists to develop new approaches for obtaining information on demographic rates without causing further threats to imperiled animal populations. In this paper, we illustrate and apply a data-fitting technique based on quadratic programming that uses stage-specific abundance data to estimate demographic rates and asymptotic population growth rates (lambda). We used data from seven breeding colonies of California sea lions (Zalophus californianus) in the Gulf of California, Mexico. Estimates of lambda were similar to those from previous studies relying on a diffusion approximation using trends in total abundance. On average, predicted abundances were within 24% of the observed value for the inverse estimation method and within 29% of the observed value for the diffusion approximation. Our results suggest that three of the seven populations are declining (lambda < 1), but as many as six may be at risk. Elasticity and sensitivity analyses suggest that population management in most sites should focus on the protection of adults, whose survival generally contributes the most to lambda. The quadratic programming approach is a promising noninvasive technique for estimating demographic rates and assessing the viability of populations of imperiled species.     

Spatial statistics for modeling of abundance and distribution of wildlife species in the Masai Mara ecosystem, Kenya

This study illustrates the use of modern statistical procedures for better wildlife management by addressing three key issues: determination of abundance, modeling of animal distributions and variability of diversity in space and time. Prior information in Markov Chain Monte Carlo (MCMC) methods is used to improve estimates of abundance. Measures of autocorrelation are included when modeling distributions of animal counts, and a diversity index to indicate species abundance and richness for large herbivores is developed. Data from the Masai Mara ecosystem in Kenya are used to develop and demonstrate these procedures. The new abundance estimates are up to 35% more accurate than those obtained by existing methods. Significant temporal changes in spatial patterns are found from a space-time analysis of elephant counts over a 20-year period, with strong interactions over 5 km and 6 months space and time separations, respectively. The new diversity index is sensitive to both high abundance and species richness and is also able to capture year to year variation. It indicates an overall marginal decrease in diversity for large herbivores in the Mara ecosystem. The space-time analyses and diversity index can easily be computed thereby providing tools for rapid decision making.     

Abundance of the New Zealand subantarctic southern right whale population estimated from photo-identification and genotype mark-recapture

The abundance of New Zealand subantarctic southern right whales (Eubalaena australis) was estimated for the first time using mark-recapture methods based on photo-identification and microsatellite genotyping (13 loci). Individual identification photographs of 383 whales and microsatellite genotypes of 235 whales were collected during annual austral winter field surveys from 1995 to 1998. Given the 4-year survey period and lack of geographic and demographic closure, we estimated super-population abundance using the POPAN Jolly-Seber model implemented in the software programme MARK. Models with constant survivorship but time-varying capture probability and probability of entry into the population were the most suitable due to the survey design. This provided estimates of abundance in 1998 of 908 non-calf whales (95% C.L. = 755, 1,123) for the photo-identification and 910 non-calf whales (95% C.L. = 641, 1,354) for the microsatellite genotype data sets. The current estimate of 900 whales may represent less than 5% of the pre-whaling abundance in New Zealand waters.     

Simultaneous-count models to estimate abundance from counts of unmarked individuals with imperfect detection

We developed a method to estimate population abundance from simultaneous counts of unmarked individuals over multiple sites. We considered that at each sampling occasion, individuals in a population could be detected at 1 of the survey sites or remain undetected and used either multinomial or binomial simultaneous-count models to estimate abundance, the latter being equivalent to an N-mixture model with one site. We tested model performance with simulations over a range of detection probabilities, population sizes, growth rates, number of years, sampling occasions, and sites. We then applied our method to 3 critically endangered vulture species in Cambodia to demonstrate the real-world applicability of the model and to provide the first abundance estimates for these species in Cambodia. Our new approach works best when existing methods are expected to perform poorly (i.e., few sites and large variation in abundance among sites) and if individuals may move among sites between sampling occasions. The approach performed better when there were >8 sampling occasions and net probability of detection was high (>0.5). We believe our approach will be useful in particular for simultaneous surveys at aggregation sites, such as roosts. The method complements existing approaches for estimating abundance of unmarked individuals and is the first method designed specifically for simultaneous counts.     

A Bayesian approach to line-transect analysis for estimating abundance

Line-transect analysis is a widely used method of estimating plant and animal density and abundance. A Bayesian approach to a basic line-transect analysis is developed for a half-normal detection function. We extend the model of Karunamuni and Quinn [Karunamuni, R.J., Quinn II, T.J., 1995. Bayesian estimation of animal abundance for line-transect sampling. Biometrics 51, 1325–1337] by including a binomial likelihood function for the number of objects detected. The method computes a joint posterior distribution on the effective strip width and the density of objects in the sampled area. Analytical and computational methods for binned and unbinned perpendicular distance data are provided. Existing information about effective strip width and density can be brought into the analysis via prior distributions. The Bayesian approach is compared to a standard line-transect analysis using both real and simulated data. Results of the Bayesian and non-Bayesian analyses are similar when there are no prior data on effective strip width or density, but the Bayesian approach performs better when such data are available from previous or related studies. Practical methods for including prior data on effective strip width and density are suggested. A numerical example shows how the Bayesian approach can provide valid estimates when the sample size is too small for the standard approach to work reliably. The proposed Bayesian approach can form the basis for developing more advanced analyses.     

An integrated approach for predicting fates of reintroductions with demographic data from multiple populations

We devised a novel approach to model reintroduced populations whereby demographic data collected from multiple sites are integrated into a Bayesian hierarchical model. Integrating data from multiple reintroductions allows more precise population-growth projections to be made, especially for populations for which data are sparse, and allows projections that account for random site-to-site variation to be made before new reintroductions are attempted. We used data from reintroductions of the North Island Robin (Petroica longipes), an endemic New Zealand passerine, to 10 sites where non-native mammalian predators are controlled. A comparison of candidate models that we based on deviance information criterion showed that rat-tracking rate (an index of rat density) was a useful predictor of robin fecundity and adult female survival, that landscape connectivity and a binary measure of whether sites were on a peninsula were useful predictors of apparent juvenile survival (probably due to differential dispersal away from reintroduction sites), and that there was unexplained random variation among sites in all demographic rates. We used the two best supported models to estimate the finite rate of increase (λ) for populations at each of the 10 sites, and for a proposed reintroduction site, under different levels of rat control. Only three of the reintroduction sites had λ distributions completely >1 for either model. At two sites, λ was expected to be >1 if rat-tracking rates were <5%. At the other five reintroduction sites, λ was predicted to be close to 1, and it was unclear whether growth was expected. Predictions of λ for the proposed reintroduction site were less precise than for other sites because distributions incorporated the full range of site-to-site random variation in vital rates. Our methods can be applied to any species for which postrelease data on demographic rates are available and potentially can be extended to model multiple species simultaneously.     

Population growth in snow geese: a modeling approach integrating demographic and survey information

There are few analytic tools available to formally integrate information coming from population surveys and demographic studies. The Kalman filter is a procedure that facilitates such integration. Based on a state-space model, we can obtain a likelihood function for the survey data using a Kalman filter, which we may then combine with a likelihood for the demographic data. In this paper, we used this combined approach to analyze the population dynamics of a hunted species, the Greater Snow Goose (Chen caerulescens atlantica), and to examine the extent to which it can improve previous demographic population models. The state equation of the state-space model was a matrix population model with fecundity and regression parameters relating adult survival and harvest rate estimated in a previous capture-recapture study. The observation equation combined the output from this model with estimates from an annual spring photographic survey of the population. The maximum likelihood estimates of the regression parameters from the combined analysis differed little from the values of the original capture-recapture analysis, though their precision improved. The model output was found to be insensitive to a wide range of coefficient of variation (CV) in fecundity parameters. We found a close match between the surveyed and smoothed population size estimates generated by the Kalman filter over an 18-year period, and the estimated CV of the survey (0.078-0.150) was quite compatible with its assumed value (approximately 0.10). When we used the updated parameter values to predict future population size, the model underestimated the surveyed population size by 18% over a three-year period. However, this could be explained by a concurrent change in the survey method. We conclude that the Kalman filter is a promising approach to forecast population change because it incorporates survey information in a formal way compared with ad hoc approaches that either neglect this information or require some parameter or model tuning.     

Logging Concessions Can Extend the Conservation Estate for Central African Tropical Forests

Abstract:  The management of tropical forest in timber concessions has been proposed as a solution to prevent further biodiversity loss. The effectiveness of this strategy will likely depend on species-specific, population-level responses to logging. We conducted a survey (749 line transects over 3450 km) in logging concessions (1.2 million ha) in the northern Republic of Congo to examine the impact of logging on large mammal populations, including endangered species such as the elephant ( Loxodonta africana ), gorilla ( Gorilla gorilla ), chimpanzee ( Pan troglodytes ), and bongo ( Tragelaphus eurycerus ). When we estimated species abundance without consideration of transect characteristics, species abundances in logged and unlogged forests were not different for most species. When we modeled the data with a hurdle model approach, however, analyzing species presence and conditional abundance separately with generalized additive models and then combining them to calculate the mean species abundance, species abundance varied strongly depending on transect characteristics. The mean species abundance was often related to the distance to unlogged forest, which suggests that intact forest serves as source habitat for several species. The mean species abundance responded nonlinearly to logging history, changing over 30 years as the forest recovered from logging. Finally the distance away from roads, natural forest clearings, and villages also determined the abundance of mammals. Our results suggest that logged forest can extend the conservation estate for many of Central Africa's most threatened species if managed appropriately. In addition to limiting hunting, logging concessions must be large, contain patches of unlogged forest, and include forest with different logging histories.     

Compositional analysis of overdispersed counts using generalized estimating equations

Multivariate abundance data are commonly collected in ecology, and used to explore questions of “community composition”—how relative abundance of different taxa changes with environmental conditions. In this paper, we propose a log-linear marginal modeling approach for analyzing such compositional count data, via generalized estimating equations. This method exploits the multiplicative nature of log-linear models for counts, by reparameterizing models that describe marginal effects on mean abundance. This allows partitioning into “main effects” and compositional effects, which is appealing for interpretation. We apply the proposed approach to reanalyze compositional counts of benthic invertebrates from Delaware Bay, and data of invertebrate communities inhabiting Acacia plants in eastern Australia. In both cases we resort to a resampling approach to make inferences about regression parameters, because the number of clusters was not large compared to cluster size.     

Random versus stratified location of transects or points in distance sampling: theoretical results and practical considerations

A composite approach mixing design-based and model-based inference is considered for analyzing line-transect or point-transect data. In this setting, the properties of the animal abundance estimator stem from the sampling scheme adopted to locate transects or points on the study region, as well as from the modeled detection probabilities. Moreover, the abundance estimation can be viewed as a “generalized” version of Monte Carlo integration. This approach permits to prove the superiority of the stratified placement of transects or points (based on a regular tessellation of the study region) over the uniform random placement. Even if the result was already established for the fixed-area sampling, i.e., when a perfect detection takes place, it was lacking in distance sampling. Comparisons with other widely-applied schemes pursuing an even placement of transects or points are also considered.     

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.     

Chilean Blue Whales as a Case Study to Illustrate Methods to Estimate Abundance and Evaluate Conservation Status of Rare Species

Abstract: Often abundance of rare species cannot be estimated with conventional design‐based methods, so we illustrate with a population of blue whales (Balaenoptera musculus) a spatial model‐based method to estimate abundance. We analyzed data from line‐transect surveys of blue whales off the coast of Chile, where the population was hunted to low levels. Field protocols allowed deviation from planned track lines to collect identification photographs and tissue samples for genetic analyses, which resulted in an ad hoc sampling design with increased effort in areas of higher densities. Thus, we used spatial modeling methods to estimate abundance. Spatial models are increasingly being used to analyze data from surveys of marine, aquatic, and terrestrial species, but estimation of uncertainty from such models is often problematic. We developed a new, broadly applicable variance estimator that showed there were likely 303 whales (95% CI 176–625) in the study area. The survey did not span the whales' entire range, so this is a minimum estimate. We estimated current minimum abundance relative to pre‐exploitation abundance (i.e., status) with a population dynamics model that incorporated our minimum abundance estimate, likely population growth rates from a meta‐analysis of rates of increase in large baleen whales, and two alternative assumptions about historic catches. From this model, we estimated that the population was at a minimum of 9.5% (95% CI 4.9–18.0%) of pre‐exploitation levels in 1998 under one catch assumption and 7.2% (CI 3.7–13.7%) of pre‐exploitation levels under the other. Thus, although Chilean blue whales are probably still at a small fraction of pre‐exploitation abundance, even these minimum abundance estimates demonstrate that their status is better than that of Antarctic blue whales, which are still <1% of pre‐exploitation population size. We anticipate our methods will be broadly applicable in aquatic and terrestrial surveys for rarely encountered species, especially when the surveys are intended to maximize encounter rates and estimate abundance.     

Simulation model of universal law of school size distribution applied to southern bluefin tuna (Thunnus maccoyii) in the Great Australian Bight

A universal law of animal group size distribution correlates well to observed fish school size distribution from fisheries catch data. I applied the law to fisheries independent aerial survey data of southern bluefin tuna (Thunnus maccoyii) collected over a 10-year period in the Great Australian Bight. The law does not correlate to the observed school size distribution. A computer model originally demonstrated the formation of the universal law from simple rules. I redesigned this model as an individual-based simulation model calibrated from acoustic tag observations and state a mathematical formula for a resultant new family of transient group size distributions. The new formula correlates accurately to the simulation and to the aerial survey data. I use the mathematical model to estimate area of aggregation and total abundance. This approach is new as it does not seek stationary states of group size distribution and because it demonstrates a quantitative relationship between individual behaviour and group size distribution. This work elevates the pattern of group size distribution from a curiosity to a useful tool, and introduces a new family of transient distributions that may have a general application to other grouping phenomena.     

Modeling Abundance Index Data from Anuran Calling Surveys

Abstract:  Evaluation of anuran populations is commonly based on calling surveys that report categorical abundance index data. I present a statistical model for abundance index data that are observations representing ordered abundance classes (e.g., none, some, many). The proposed model provides a formal treatment of detection probability, factors that affect detection, and variation in abundance. The model can be viewed as a generalization of that proposed by MacKenzie et al. (2002) for estimating site-occupancy rates in that it allows for more than two abundance classes. Because the abundance distribution is characterized by multiple abundance classes, it may be more sensitive to subtle changes in the underlying abundance that may go undetected with simple occupancy estimates under which sites are characterized merely as occupied or not. The method is most immediately applicable to surveys of anurans in which index data related to the intensity of calling activity are collected. I applied the proposed method to calling index data from the green frog (  Rana clamitans ) collected as part of the North American Amphibian Monitoring Program. The best model indicated considerable variation in detectability over time and in response to temperature. The resulting adjusted (for detectability) abundance-state distribution demonstrates the negative bias in abundance state obtained from simplistic summaries of calling index data that disregard these sources of variation in detectability.     

A framework for adapting survey design through time for wildlife population assessment

Sampling strategies for monitoring the status and trends in wildlife populations are often determined before the first survey is undertaken. However, there may be little information about the distribution of the population and so the sample design may be inefficient. Through time, as data are collected, more information about the distribution of animals in the survey region is obtained but it can be difficult to incorporate this information in the survey design. This paper introduces a framework for monitoring motile wildlife populations within which the design of future surveys can be adapted using data from past surveys whilst ensuring consistency in design-based estimates of status and trends through time. In each survey, part of the sample is selected from the previous survey sample using simple random sampling. The rest is selected with inclusion probability proportional to predicted abundance. Abundance is predicted using a model constructed from previous survey data and covariates for the whole survey region. Unbiased design-based estimators of status and trends and their variances are derived from two-phase sampling theory. Simulations over the short and long-term indicate that in general more precise estimates of status and trends are obtained using this mixed strategy than a strategy in which all of the sample is retained or all selected with probability proportional to predicted abundance. Furthermore the mixed strategy is robust to poor predictions of abundance. Estimates of status are more precise than those obtained from a rotating panel design.     

Estimating species occurrence, abundance, and detection probability using zero-inflated distributions

Researchers have developed methods to account for imperfect detection of species with either occupancy (presence absence) or count data using replicated sampling. We show how these approaches can be combined to simultaneously estimate occurrence, abundance, and detection probability by specifying a zero-inflated distribution for abundance. This approach may be particularly appropriate when patterns of occurrence and abundance arise from distinct processes operating at differing spatial or temporal scales. We apply the model to two data sets: (1) previously published data for a species of duck, Anas platyrhynchos, and (2) data for a stream fish species, Etheostoma scotti. We show that in these cases, an incomplete-detection zero-inflated modeling approach yields a superior fit to the data than other models. We propose that zero-inflated abundance models accounting for incomplete detection be considered when replicate count data are available.     

Estimating the functional form for the density dependence from life history data

Two contrasting approaches to the analysis of population dynamics are currently popular: demographic approaches where the associations between demographic rates and statistics summarizing the population dynamics are identified; and time series approaches where the associations between population dynamics, population density, and environmental covariates are investigated. In this paper, we develop an approach to combine these methods and apply it to detailed data from Soay sheep (Ovis aries). We examine how density dependence and climate contribute to fluctuations in population size via age- and sex-specific demographic rates, and how fluctuations in demographic structure influence population dynamics. Density dependence contributes most, followed by climatic variation, age structure fluctuations and interactions between density and climate. We then simplify the density-dependent, stochastic, age-structured demographic model and derive a new phenomenological time series which captures the dynamics better than previously selected functions. The simple method we develop has potential to provide substantial insight into the relative contributions of population and individual-level processes to the dynamics of populations in stochastic environments.     

Modelling skewed data with many zeros: A simple approach combining ordinary and logistic regression

We discuss a method for analyzing data that are positively skewed and contain a substantial proportion of zeros. Such data commonly arise in ecological applications, when the focus is on the abundance of a species. The form of the distribution is then due to the patchy nature of the environment and/or the inherent heterogeneity of the species. The method can be used whenever we wish to model the data as a response variable in terms of one or more explanatory variables. The analysis consists of three stages. The first involves creating two sets of data from the original: one shows whether or not the species is present; the other indicates the logarithm of the abundance when it is present. These are referred to as the presence data and the log-abundance data, respectively. The second stage involves modelling the presence data using logistic regression, and separately modelling the log-abundance data using ordinary regression. Finally, the third stage involves combining the two models in order to estimate the expected abundance for a specific set of values of the explanatory variables. A common approach to analyzing this sort of data is to use a ln (y+c) transformation, where c is some constant (usually one). The method we use here avoids the need for an arbitrary choice of the value of c, and allows the modelling to be carried out in a natural and straightforward manner, using well-known regression techniques. The approach we put forward is not original, having been used in both conservation biology and fisheries. Our objectives in this paper are to (a) promote the application of this approach in a wide range of settings and (b) suggest that parametric bootstrapping be used to provide confidence limits for the estimate of expected abundance.     

Daphne: a new multimetric benthic index for the quality assessment of marine coastal environment in the Northern Adriatic Sea

A new benthic index, named Daphne, is proposed for the Northern Adriatic coastal area, near the Po river delta. It is based on six characteristics of the community that do not require in-depth taxonomic expertise: number of mollusc species, % of bivalves, % of polychaetes, abundance of the opportunistic species Corbula gibba, % of amphipods and number of 'typical mollusc species' that are individuated by multivariate analysis. The application of the index in selected stations along a gradient of decreasing disturbance shows that it is simple to use in regular monitoring campaigns and that the results are consistent with environmental quality data in the special conditions of this area subjected to considerable river runoff. The index can be used in addition and as a complement to more widespread indices (such as M-AMBI); a comparison of the two indices performance is discussed.     

Bridging information domains to improve ecological understanding of biological invasions in a marine ecosystem

We devised a practical method for integrating information on 2 marine invasive species using 3 different approaches: standardized ecological monitoring, online-reporting databases, and surveys of anglers and crabbers. Focusing on 2 recently introduced species with different characteristics, the Asian shore crab (Hemigrapsus sanguineus) and Chinese mitten crab (Eriocheir sinensis), in the Hudson-Raritan watershed of New York and New Jersey, we used sensitivity analyses to explore the relative contribution of each information source to knowledge of species abundance and distribution. All 3 information sources contributed something unique to understanding abundance and distribution of the introduced crabs. Online and survey data on Asian shore crabs significantly affected predictions of abundance, whereas monitoring data did not. When survey data were omitted, abundance estimates were unchanged over time, but when they were included, the model predicted an increased abundance in 2012. All 3 data sets for the Asian shore crab significantly affected estimates of species coverage; surveys had the biggest influence, increasing range size by 4097.25 km². For the catadromous Chinese mitten crab, ecological monitoring data collected in freshwater shortly after the original sighting significantly shaped model estimates for abundance and documented the establishment phase of the mitten crab in an area outside the spatial scope of the surveyed resource users. However, the survey data significantly enlarged mitten crab range-size estimates by 6498.01 km². By demonstrating that data integration produced an image of the invasion process that would not have emerged had we used any 1 method individually, model results provide evidence for the advantages of an interdisciplinary approach.