Long-term monitoring of tropical bats for anthropogenic impact assessment: Gauging the statistical power to detect population change

Bats are ecologically important mammals in tropical ecosystems; however, their populations face numerous environmental threats related to climate change, habitat loss, fragmentation, hunting, and emerging diseases. Thus, there is a pressing need to develop and implement large-scale networks to monitor trends in bat populations over extended time periods. Using data from a range of Neotropical and Paleotropical bat assemblages, we assessed the ability for long-term monitoring programs to reliably detect temporal trends in species abundance. We explored the magnitude of within-site temporal variation in abundance and evaluated the statistical power of a suite of different sampling designs for several different bat species and ensembles. Despite pronounced temporal variation in abundance of most tropical bat species, power simulations suggest that long-term monitoring programs (?20 years) can detect population trends of 5% per year or more with adequate statistical power (?0.9). However, shorter monitoring programs (?10 years) have insufficient power for trend detection. Overall, our analyses demonstrate that a monitoring program extending over 20 years with four surveys conducted biennially on five plots per monitoring site would have the potential for detecting a 5% annual change in abundance for a suite of bat species from different ensembles. The likelihood of reaching adequate statistical power was sensitive to initial species abundance and the magnitude of count variation, stressing that only the most abundant species in an assemblage and those with generally low variation in abundance should be considered for detailed population monitoring.     

Ruffed grouse population dynamics in the central and southern Appalachians

Ruffed grouse (Bonasa umbellus; hereafter grouse) populations in the central and southern Appalachians are in decline. However, limited information on the dynamics of these populations prevents the development of effective management strategies to reverse these trends. We used radiotelemetry data collected on grouse to parameterize 6 models of population growth to: (1) determine the pattern of growth in these populations, and (2) identify the demographic rates most important to growth. Trend estimates from population models were most similar to trend estimates derived from Breeding Bird Survey and Christmas Bird Count data when models incorporated either a reproductive or survival event. These events randomly increased fecundity or survival, respectively, to their empirical maxima on average once every 5 years. Reproductive events improved estimates on areas dominated by mixed mesophytic forest, while survival events characterized population growth on oak (Quercus spp.)-dominated sites. The finite rate of increase (λ) was most sensitive to brood survival followed by adult and juvenile non-breeding survival on most sites. However, brood survival was low (<0.35 female chicks/hen survived to week 5), and elasticity analyses indicated λ responded more strongly to proportionate change in non-breeding and breeding survival rates of adults and juveniles than any reproductive variable. Life stage analyses corroborated this result. At baseline values, survival of adults and juveniles may be the main determinants of growth in these populations, and reproduction may not be adequate to compensate for these losses. Therefore, population growth above baseline levels may be regularly needed to restock these populations. Researchers have hypothesized that population dynamics may differ between mixed mesopytic and oak-dominated sites due to differences in forage quality and quantity. Thus, a potential mechanism for the increases in λ needed to sustain populations on mixed mesophytic forest sites is the greater fecundity observed during years with high oak or beech (Fagus grandifolia) mast abundance. The availability of this high quality forage allows hens to enter the breeding season in better condition and realize higher fertility. Alternatively, on oak-dominated sites, population growth increases may also be a product of higher non-breeding survival of birds in mast years, when birds do not need to range as far to forage and can limit their exposure to predators.     

Usefulness of volunteer data to measure the large scale decline of “common” toad populations

Measuring a species decline is pivotal to evaluate their conservation status, but an accurate assessment of demographic trends requires observations collected across broad spatial and temporal scales. Volunteers can help to collect information over large scales, but their data may be affected by heterogeneity for sampling efforts and protocols, which may influence detection probability. Ignoring this issue may conduct to misleading conclusions. Here we show that data collected by different volunteer groups can be integrated with measures of sampling efforts, to obtain information on large scale demographic trends. We collected data on 33 common toad (Bufo bufo) populations across Italy for the period 1993–2010. We used two approaches (meta-analysis; analysis of average change in population size) to evaluate the overall demographic trend. We incorporated measures of volunteer sampling efforts into analyses, to take into account changes in detection probability. Toad abundance significantly declined in the last decade. From 2000 to 2010, 70% of populations showed a strong decline, and only 10% increased. Trends were heterogeneous among populations, but taking into account sampling effort reduced heterogeneity by 40%. We detected a 76% cumulative average decline of toad populations, despite an increasing mean sampling effort. The widespread toad decline rises concern for its future, also because the causes remain unclear. Volunteer data can be extremely useful to identify large scale population trends, if information on sampling effort are recorded and used to adjust counts.     

Incorporating parametric uncertainty into population viability analysis models

Uncertainty in parameter estimates from sampling variation or expert judgment can introduce substantial uncertainty into ecological predictions based on those estimates. However, in standard population viability analyses, one of the most widely used tools for managing plant, fish and wildlife populations, parametric uncertainty is often ignored in or discarded from model projections. We present a method for explicitly incorporating this source of uncertainty into population models to fully account for risk in management and decision contexts. Our method involves a two-step simulation process where parametric uncertainty is incorporated into the replication loop of the model and temporal variance is incorporated into the loop for time steps in the model. Using the piping plover, a federally threatened shorebird in the USA and Canada, as an example, we compare abundance projections and extinction probabilities from simulations that exclude and include parametric uncertainty. Although final abundance was very low for all sets of simulations, estimated extinction risk was much greater for the simulation that incorporated parametric uncertainty in the replication loop. Decisions about species conservation (e.g., listing, delisting, and jeopardy) might differ greatly depending on the treatment of parametric uncertainty in population models.     

Numbers of wintering waterbirds in Great Britain and the Isle of Man (1994/1995-1998/1999): II. Coastal waders (Charadrii)

Great Britain's wintering coastal wader populations have been estimated for the period 1994/1995-1998/1999 from data provided from two sources: the Wetland Bird Survey (WeBS) and the Non-estuarine Coastal Waterfowl Survey (UK-NEWS). New methodology for estimating the number of wintering waders is applied. It imputes (fills in) the value of missing counts before estimating the population size of each species as being the mean of the largest annual count made between November and March over the relevant 5-year period. This methodology has led to an 11% larger estimate of the number of waders present on Great Britain's coasts than the traditional approach based on averaging just January counts over a 5-year period, and it suggests that Great Britain's coastline supports ca. 2.1 million waders. Updated values are presented from which site evaluations based on 1% of the national population can be derived. Great Britain is of considerable international importance for waders. It holds >25 and >50% of the flyway populations of nine and four species of wader, respectively, but for the first time since the start of monitoring in the early 1970s, the historical increase in the number of predominantly coastal waders wintering in Great Britain is coming to an end. Seven of the 14 species that have shown population changes of >5% since the last set of 1987/1988-1991/1992 population estimates have declined in numbers. The possible causes of the fluctuations in wader populations, such as climate change and changing nutrient inputs to coastal waters are discussed. There is an urgent need to identify the causative factors leading to these declines, and to use the new population estimates to identify new sites that should be afforded legal protection, an action that should help Great Britain maintain its internationally important wader populations. The decrease in the updated population estimate of Eurasian oystercatcher, for example, has made it possible to determine that 19 rather than 17 sites in Great Britain are worthy of statutory protection on the basis of holding 1% or more of its national population.     

Combining data from state and national monitoring surveys to assess large-scale impacts of agricultural policy

An increasing number of state and national databases are available to assess agricultural and environmental trends in natural resource populations. We use a case study approach to consider methodologies for combining state and national data to assess the impact of agricultural policy on state wildlife populations. The scientific question is to assess the impact of the Conservation Reserve Program on pheasant populations in Iowa, using land cover/use data from the National Resources Inventory and count data from an annual state pheasant population survey. Our approach involves identifying a common spatial polygon for linking summaries from each of two datasets, and then estimating parameters that describe temporal trends in land cover and in pheasant populations over a common time period within each polygon. Estimated pheasant population parameters are regressed on land cover summaries to investigate the impact of the Conservation Reserve Program on pheasant populations in regions of the state. Results reveal that the population response to the Conservation Reserve Program varies by region in relation to the physiography and agricultural use of the region, in ways that were not anticipated by policy developers. Statistical considerations for developing appropriate models for combining data are discussed.     

Integrated data analysis in the presence of emigration and mark loss

Integrated data analyses are becoming increasingly common in studies of wild animal populations where two or more separate sources of data contain information about common parameters. These types of analyses provide robust parameter estimates which fully reflect all available information, as well as estimates of parameters that would be unidentifiable in a separate analysis. In this article we present an integrated Bayesian analysis of four long-term datasets (counts, two mark-recapture-recovery time series, and productivity) relating to a colony of common guillemots (Uria aalge) on the Isle of May, southeast Scotland. A complication when considering the dynamics of populations of this kind is the unobservable emigration of immature animals. In the analysis of mark-recapture-recovery data, the rate of emigration is frequently confounded with that of mark loss and it is only possible to estimate the product of these parameters. By combining all available data for the Isle of May guillemots in an integrated population model, we are able to estimate these parameters separately and thus obtain improved estimates of prerecruitment emigration.     

A sensitivity analysis to guide research and management for Hector's dolphin

Hector's dolphin (Cephalorhynchus hectori) is an uncommon endemic of New Zealand which is suspected to be in decline due to entanglement mortality. However, uncertainty in available data has led to a dispute between the New Zealand Ministry of Fisheries and the New Zealand Department of Conservation over the status of this species. We use a density-dependent deterministic model to predict the future abundance and geographic distribution of Hector's dolphin under different scenarios of fisheries management. We then examine the sensitivity of this model to a number of parameters for which few or no data are available. We find that two populations of Hector's dolphins are predicted to decline in the future even when the most optimistic parameter estimates are used. The status of the third population is dependent upon the estimate of maximum annual population growth rate. Because of the dependence of final abundance estimates on the estimates of entanglement mortality rates and maximum population growth rate, research efforts should be concentrated on estimating these parameters.     

Brown hyaenas on roads: Estimating carnivore occupancy and abundance using spatially auto-correlated sign survey replicates

Carnivore survey protocols that properly address spatial sampling and detectability issues are seldom feasible at a landscape-scale. This limits knowledge of large-scale patterns in distribution, abundance and their underlying determinants, hindering conservation of globally threatened carnivore populations. Occupancy analysis of data from logistically efficient sign surveys along consecutive road segments (spatially auto-correlated replicates) offers a potential solution. We adapted and applied this newly-developed method over 62,979 km² of human-modified land in South Africa. Our aims were to (1) generate unbiased estimates of brown hyaena occupancy and abundance (2) investigate two suspected determinants of occupancy using a combination of biological and socio-economic sampling techniques, and (3) use simulations to evaluate the effort required for abundance and occupancy estimates with acceptable bias, precision and power. Brown hyaena occupancy was estimated at 0.748 (±SE 0.1), and estimated overall density in agricultural land (0.15/100 km², ±SE 0.08) was an order of magnitude lower than in protected areas. Positive attitudes to carnivores and presence of wildlife farms exerted strong positive effects on occupancy, so changes in these factors may well exert monotonic impacts on local metapopulation status. Producing reliable occupancy and abundance estimates would require ?6 replicates and ?12 replicates per site respectively. Detecting 50% and 30% declines in brown hyaena occupancy with adequate power would require five annual surveys at ?65 sites and ?125 sites respectively. Our results suggest that protocols based on spatially auto-correlated sign survey replicates could be used to monitor carnivore populations at large, and possibly even country-wide spatial scales.     

Resolving conflicts between butterfly host resource abundance and genet population size estimates for a vegetatively spreading, threatened grassland legume

Population size estimates are an integral part of rare plant conservation, but common abundance measurements such as cover and ramet number may not accurately index genet population size for vegetatively spreading species. Population monitoring of Kincaid’s lupine (threatened species) populations occurs through genet-anonymous leaf cover and raceme counts despite extensive, non-adventitious rhizome growth. While the current monitoring scheme provides important resource abundance measurements for the endangered Fender’s blue butterfly, whose larvae feed on Kincaid’s lupine leaves, the methods are not appropriate for estimating lupine genet number. Major axis regression revealed well supported statistical relationships between cover, raceme, and plantlet (a measurement of modular plant growth) density within six study patches (n = 3 populations) of Kincaid’s lupine (R² > 0.90) and when all patch data were combined (R² > 0.91). Genet population size estimates from genotype only data with ACE (an estimator used to infer species richness) were similar to estimates derived from a combination of plantlet density and genet to plantlet ratios (genotype derived) in small, more thoroughly genotyped lupine patches. However, genet number estimates from ACE were 3–5-fold greater in less intensely genotyped patches. Genet-anonymous plant abundance measurements, such as cover, can be used to estimate genet number in populations of vegetatively spreading plants provided they are calibrated with a unit of modular plant growth. Calibration of vegetative measurements, consistency of between population relationships, and closer scrutiny of highly supported statistical models may be necessary to develop more pertinent monitoring methods for rare, vegetatively spreading plants.     

Large mammal population declines in Africa’s protected areas

Protected areas (PAs) are the cornerstone of global conservation efforts but their performance in maintaining populations of their key species remains poorly documented. Here, we address this gap using a new database of 583 population abundance time series for 69 species of large mammals in 78 African PAs. Population abundance time series were aggregated to form a multi-species index of overall change in population abundance. The index reveals on average a 59% decline in population abundance between 1970 and 2005. Indices for different parts of Africa demonstrate large regional differences, with southern African PAs typically maintaining their populations and western African PAs suffering the most severe declines. These results indicate that African PAs have generally failed to mitigate human-induced threats to African large mammal populations, but they also show some successes. Further development of our index could help to measure future progress towards post-2010 targets for reducing biodiversity loss.     

Influence of forest fragmentation on an endangered large-bodied lemur in northwestern Madagascar

Madagascar’s diverse and mostly endemic fauna and flora suffer from recent landscape changes that are primarily caused by high levels of human activities. The loss and fragmentation of forest habitats are well known consequences of human activities. In this study, we investigate the effects of forest fragmentation on presence, abundance and genetic diversity in a larger-bodied lemur species, Lepilemur edwardsi, in northwestern Madagascar. In addition, we characterized the genetic differentiation among populations and demographic changes. We found L. edwardsi at only 13 (76.5%) of 17 visited sites, 11 of which were situated in the Ankarafantsika National Park (ANP). We captured between two and 17 individuals per site. We sequenced the mtDNA d-loop of all samples and genotyped 14 microsatellite loci in two exemplary populations for demographic analyses. A negative influence on forest fragmentation could be detected, since the fragments had a lower genetic diversity than sites in the ANP. Genetic differentiation between populations ranged from low to high but was almost always significant. A typical pattern of isolation-by-distance could not be detected and the data could rather be interpreted as results of random genetic drift. The data furthermore revealed signals of a demographic collapse of about two orders of magnitude in the two exemplary sites. This decline probably started during the last few hundred years of intensified human disturbances and population growth. Given the results of this study, urgent conservation actions are needed and should concentrate on an effective protection of the few remaining populations in order to ensure the long-term survival of L. edwardsi.     

Demography of Asian elephants (Elephas maximus) at Uda Walawe National Park, Sri Lanka based on identified individuals

We provide estimates of population size and other demographic variables for individually-identified Asian elephants (Elephas maximus) in Uda Walawe National Park (UWNP), Sri Lanka based on systematic year-round observations. Two hundred and eighty-six adult females and 241 adult males were identified, of which four adults (2% of males) had tusks. Sightings-based demographic models showed seasonal immigration and emigration from the study area. The total population, including non-adults, was between 804 and 1160 individuals. Density ranged from 102 to 116 adult females per 100 km² and remains at this level throughout the year. This large, un-fragmented population of Asian elephants should be of high conservation priority. We find that estimates of survivorship and migration rates should be based on long sampling intervals when possible, but estimates of density and population size can still be made when observations are constrained to shorter intervals, if spatial data are available. We offer suggestions to guide census design for other elephant populations or cryptic species. We urge that other locations be systematically surveyed as well using photographic identification.     

Density and population size estimates for North Cascade grizzly bears using DNA hair-sampling techniques

We used non-invasive DNA hair-sampling and catch per unit effort (CPUE: grizzly bears detected per 1000 trap nights) to estimate relative density and population size for a threatened grizzly bear population in the North Cascade Ecosystem of Washington and British Columbia. We used linear, logistic, and linear through the origin regression analyses to estimate the relationship between catch per unit effort and grizzly bear density for seven other grizzly populations. One grizzly bear was detected during 5304 trap nights (CPUE=0.19) over 3 years in the North Cascades. This CPUE was much lower than in the other seven populations, including two threatened grizzly populations in the Cabinet-Yaak and Selkirk Mountain Ecosystems. The logistic model (curvilinear relationship) best fit the data (R²=0.927), and yielded density and population size estimates of 0.15 bears/100 km² (90% CI=0.03-0.71) and six bears (90% CI=1-27), respectively. Natural recovery seems unlikely for the North Cascade grizzly bear population because the population has a high likelihood of extinction due to demographic and environmental stochastic effects associated with extremely small population numbers. We recommend population augmentation. DNA hair-sampling and catch per unit effort models can be a useful method to evaluate relative densities and numbers of animals in small, threatened grizzly bear populations when sample sizes are too small to yield traditional mark-recapture analysis.     

Potential of a national monitoring program for forests to assess change in high-latitude ecosystems

Broad-scale monitoring in Alaska has become of increasing interest due to uncertainty about the potential impacts of changing climate on high-latitude ecosystems. The Forest Inventory and Analysis (FIA) program is a national monitoring program for all public and private forestlands in the US, but the program is not currently implemented in the boreal region of Alaska. We provide an overview of the strengths and weaknesses of the FIA system for monitoring the potential impact of climate change on Alaska’s species, communities, and ecosystems. The primary strength of the system is a scientifically rigorous design-based statistical estimation method that produces estimates of forest attributes with known sampling error and quantifiable measurement error. The weaknesses of the system include low power for small area estimates, lack of spatial context and contiguity, and difficulty in inferring causality of factors when changes in monitored attributes are detected.Climate change is expected to impact many components of boreal ecosystems, but for most indicators the direction and magnitude of change are difficult to predict because of complex interactions among system components. Status and trend information provided by FIA monitoring that could be helpful to conservation decisions includes abundance and rarity of vascular plants, invasive species, biomass and carbon content of vegetation, shifting vegetation species distribution, disturbance frequency, type, and impact, and wildlife habitat characteristics. Because of unique factors such as the low level of infrastructure, modifications to the FIA monitoring system used in the conterminous US have been proposed for Alaska. Remote sensing data would play a greater role in meeting monitoring objectives, and sampling intensity of field plots would be reduced. Coordination with other national, regional, and local monitoring efforts provides potential for increased understanding of change in boreal ecosystems at multiple scales.     

A method for assessing changes in the abundance of butterflies

A method based on transect count has been developed to assess changes in abundance of butterflies from year to year. The method involves weekly walks around atransect route making counts of butterflies seen within defined limits. The transects are divided into sections related to habitat or management units. Walks are made only when weather conditions satisfy specified minimum requirements. The method has been tested for three years at Monks Wood and for two years at a number of other sites.The basis for annual comparisons is an index of abundance which is produced for each brood of each species, except when separation of broods is not possible. This index is correlated with abundance, although the precise nature of the relationship will vary from species to species. Evidence on this presented for two species.The method makes it possible to monitor the abundance of butterflies at selected sites, using recorders, such as nature reserve wardens, who can fit in one or two hours recording each week when the weather is suitable. Such a scheme, based on the methods described in this paper, began in 1976. In addition to the monitoring of fluctuations of abundance, the method provides considerable information on the phenology and ecology of butterflies. The division of the transects into sections makes some assessment of the effects of habitat change, due to management or other factors, possible.     

Decomposition and soil biota after reclamation of coal mine spoils in an arid region

Summary We compared soil biota and buried wheat straw decomposition on sites subject to topsoiling plus straw mulch reclamation procedures 1–4 years prior to our study and on an unmined site. Rates of straw decomposition were highest on the unmined site. Decomposition rates were higher on the 1- and 2-year-old sites than on the 3- and 4-year-old reclaimed spoil. Microarthropod population densities and number of taxa were greater from decomposing straw on the unmined site and lowest from straw on the 1- and 2-year-old areas. Soil bacteria, fungi, and protozoan populations on buried straw on the oldest reclaimed sites were generally equal to those on the unmined area. Nematode populations on buried straw in the unmined site and 1- and 2-year-old reclaimed sites were similar. Populations of denitrifying bacteria were larger on recently reclaimed sites than on 3- and 4-year-old sites. Decomposition and nitrogen mineralization varied as a function of the diversity and abundance of soil microarthropods. Soil microfauna can serve as an index of soil development in a disturbed arid soil.     

Long-term trends in lobster populations in a partially protected vs. no-take Marine Park

Increasing the level of protection afforded to the marine environment requires assessment of the efficacy of existing marine protected areas (MPAs) in protecting exploited species. Long-term data from before and after the establishment of MPAs provide a rare but valuable opportunity to assess these effects. In this study we present long-term data (1977-2005) from before and after park establishment, on the abundance of spiny lobster Jasus edwardsii from fixed sites in a no-take marine park and a recreationally fished marine park, to assess the efficacy of no-take vs. partial protection. Lobster densities were comparable between both marine parks prior to park establishment, but the response of lobster populations differed markedly following protection. On average, legal-sized lobster were eleven times more abundant and biomass 25 times higher in the no-take marine park following park establishment, while in the partially protected marine park there has been no significant change in lobster numbers. Furthermore, no difference was found in densities of legal-sized lobster between the partially protected marine park and nearby fully-fished sites (<1 per 500 m²). Long-term data from fully fished and partially protected sites suggest long-term declines in lobster populations and reflect regional patterns in catch per unit effort estimates for the fishery. The long-term patterns presented provide an unequivocal example of the recovery of lobster populations in no-take MPAs, but clearly demonstrate that allowing recreational fishing in MPAs has little benefit to populations of exploited species such as J. edwardsii.     

Large scale predator control improves the productivity of a rare New Zealand riverine duck

Declines in avian populations are often attributed to the presence of introduced predators but conservation managers frequently lack good information about the effectiveness of potential predator control regimes for protecting threatened species. Whio, Hymenolaimus malacorhynchos, are a threatened New Zealand waterfowl that has been declining in both distribution and abundance. We conducted a six-year study using a paired-catchment experiment in New Zealand Nothofagus forest as part of an adaptive management programme to assess whether whio populations responded positively to stoat (Mustela erminea) control. Video monitoring identified stoats as the primary nest predator. Year-round low-intensity stoat control (10 traps per linear km) significantly reduced the stoat abundance index in trapped sites compared with untrapped sites. As a result, whio nesting success and productivity, the number of fledglings produced per pair, increased significantly in the trapped compared to the untrapped area. However, survival rates and the number of pairs did not change significantly between the treatments. These findings indicate stoats are the primary agent of decline for whio in this Nothofagus forest system, and show that low-intensity stoat control is sufficient to improve the productivity of whio populations. Overall, the study demonstrates the value of an adaptive management approach whereby management techniques can be evaluated to ensure that the primary agent of decline is clearly identified and that predator densities are kept sufficiently low.