Census error and the detection of density dependence

1. Studies aiming to identify the prevalence and nature of density dependence in ecological populations have often used statistical analysis of ecological time-series of population counts. Such time-series are also being used increasingly to parameterize models that may be used in population management. 2. If time-series contain measurement errors, tests that rely on detecting a negative relationship between log population change and population size are biased and prone to spuriously detecting density dependence (Type I error). This is because the measurement error in density for a given year appears in the corresponding change in population density, with equal magnitude but opposite sign. 3. This effect introduces bias that may invalidate comparisons of ecological data with density-independent time-series. Unless census error can be accounted for, time-series may appear to show strongly density-dependent dynamics, even though the density-dependent signal may in reality be weak or absent. 4. We distinguish two forms of census error, both of which have serious consequences for detecting density dependence. 5. First, estimates of population density are based rarely on exact counts, but on samples. Hence there exists sampling error, with the level of error depending on the method employed and the number of replicates on which the population estimate is based. 6. Secondly, the group of organisms measured is often not a truly self-contained population, but part of a wider ecological population, defined in terms of location or behaviour. Consequently, the subpopulation studied may effectively be a sample of the population and spurious density dependence may be detected in the dynamics of a single subpopulation. In this case, density dependence is detected erroneously, even if numbers within the subpopulation are censused without sampling error. 7. In order to illustrate how process variation and measurement error may be distinguished we review data sets (counts of numbers of birds by single observers) for which both census error and long-term variance in population density can be estimated. 8. Tests for density dependence need to obviate the problem that measured population sizes are typically estimates rather than exact counts. It is possible that in some cases it may be possible to test for density dependence in the presence of unknown levels of census error, for example by uncovering nonlinearities in the density response. However, it seems likely that these may lack power compared with analyses that are able to explicitly include census error and we review some recently developed methods.     

A demographic analysis of vole population responses to fragmentation and destruction of habitat

 Fragmentation and destruction of natural habitats is currently considered to be the major threat to wildlife populations. We here perform a comprehensive analysis of the demographic effects of habitat fragmentation and destruction on 14 populations of the root vole. The experiment was divided into two consecutive periods. During the first period, we contrasted populations with the same initial size and structure in continuous and fragmented habitat. During the second period, we fragmented the continuous habitat into the same configuration as the permanently fragmented habitat so that the effect of habitat destruction could be evaluated. We estimated survival and fecundity parameters and combined them into population projection matrices to evaluate their relative impact on population growth. In the first period of the experiment there was no difference in population growth rate between fragmented and continuous populations, although litter size was significantly higher in the continuous populations. In the second period, we found higher population growth rates in populations that had experienced habitat destruction. By applying the transition matrix model to empirical estimates of demographic parameters, we demonstrate that the difference in population growth rate in the second period of the experiment was the result of a nonsignificant difference in adult survival. Movements out of the habitat patches were significantly lower in populations that had experienced habitat destruction. We conclude that predator-caused mortality of animals moving out of the habitat patches was the main determinant of demographic variation in this system. Received: January 31, 2002 / Accepted: March 25, 2003     

When can noninvasive samples provide sufficient information in conservation genetics studies?

Noninvasive sampling, of faeces and hair for example, has enabled many genetic studies of wildlife populations. However, two prevailing problems common to these studies are small sample sizes and high genotyping errors. The first problem stems from the difficulty in collecting noninvasive samples, particularly from populations of rare or elusive species, and the second is caused by the low quantity and quality of DNA extracted from a noninvasive sample. A common question is therefore whether noninvasive sampling provides sufficient information for the analyses commonly conducted in conservation genetics studies. Here, we conducted a simulation study to investigate the effect of small sample sizes and genotyping errors on the precision and accuracy of the most commonly estimated genetic parameters. Our results indicate that small sample sizes cause little bias in measures of expected heterozygosity, pairwise F_ST and population structure, but a large downward bias in estimates of allelic diversity. Allelic dropouts and false alleles had a much smaller effect than missing data, which effectively reduces sample size further. Overall, reasonable estimates of genetic variation and population subdivision are obtainable from noninvasive samples as long as error rates are kept below a frequency of 0.2. Similarly, unbiased estimates of population clustering can be made with genotyping error rates below 0.5 when the populations are highly differentiated. These results provide a useful guide for researchers faced with studying the conservation genetics of small, endangered populations from noninvasive samples.     

Are habitat loss,predation risk and climate related to the drastic decline in a Siberian flying squirrel population? A 15-year study

To devise effective conservation actions, it is important to know which factors are associated with the population parameters of a declining population. Using mark–recapture methods, we estimated the annual population size, growth rate and survival probability of an ear-tagged flying squirrel population over a 15-year period in a 4,500 ha study area in western Finland. The species is considered vulnerable, but detailed knowledge concerning population sizes or trends is lacking. The population parameters and changes therein were regressed against habitat availability, an indicator of predation pressure, and mean winter temperature (an indicator of climate change), to reveal potential reasons for trends in the population. The best-fit models suggested the annual growth rate to be below one, and on average it was 0.93 (±0.06; SE) across the 15-year period. The survival probability was about 0.22 (±0.03) for juveniles and 0.50 (±0.03) for adults. The population size of adult flying squirrels decreased from 65 (±11) individuals in 1995 to 29 (±6) individuals in 2009. The number of flying squirrels was associated with the amount of available habitat, but the decline in population size was more rapid than the loss of habitat area. If the current decreasing trend in habitat availability continues, the population might become extinct by the year 2020. To halt the population decline, it is necessary to refrain from clear-cutting mature spruce stands until new suitable habitats develop from the maturation of younger forests.     

Field surveys of capercaillie (<Emphasis Type="Italic">Tetrao urogallus</Emphasis>) in the Swiss Alps underestimated local abundance of the species as revealed by genetic analyses of non-invasive samples

An increasing number of species are becoming threatened by habitat loss and fragmentation. Therefore, solid estimates of the species’ abundance in the remaining populations are required to develop suitable conservation measures and to monitor their effectiveness. The capercaillie (Tetrao urogallus L.) has experienced a dramatic decline in central Europe and has disappeared from large areas of its former natural range. In Switzerland, the species’ distribution, habitat requirements and demographic status were studied and evaluated in an attempt to support appropriate management decisions to conserve the species. National surveys of the capercaillie in Switzerland have traditionally been obtained from male counts at leks. However, individual attendance to the lek is sex- and age-specific. Thus, male counts at leks may provide a biased estimate of local population sizes. In the present study, we compared two alternative indirect methods to estimate the sizes of local populations at eight study sites situated in the Alps and Prealps of Switzerland. We first assessed the sizes of local populations from the observed density and distribution of direct and indirect evidence of the species’ presence during field surveys. Feather and faeces samples collected during field surveys were genotyped at twelve nuclear microsatellite loci and a sex-specific nuclear gene fragment. Individual genotypes were used as genetic tags to estimate the sizes of the eight local populations using an urn model developed for small populations. The index of local population sizes assessed from field surveys was lower than the number of unique genotypes at each study site, which itself underestimated the abundances of populations in most cases. Based on our results, the genetic tagging method appeared to be less biased than the field survey method. However, an alternative faeces sampling scheme, resulting in 2–3 genotypings per individual, could further improve the accuracy of the size estimates of local populations. Our study confirms that genetic tagging methods are a valuable tool to estimate the sizes of local populations and to monitor the response of rare and elusive species to management actions.     

Long‐term decline despite conservation efforts questions Eurasian Stone‐curlew population viability in intensive farmlands

Agricultural intensification over the past decades has led to a generalized decline in farmland biodiversity. Farmland birds are particularly exposed to rapid changes in habitat and reduced food resources or availability. Understanding how farmland specialists can be preserved and their populations enhanced are major challenges for this century. Based on a long‐term (19‐year) study of a Eurasian Stone‐curlew Burhinus oedicnemus population, we estimated the demographic parameters, including clutch size, egg volume, hatching success, survival rate and apparent population size. Demographic rates found for this French population were, on average, comparable to those found elsewhere in Europe. However, all demographic parameters showed negative trends, including a dramatic decline in the local population (26% decline over 14 years) and a 10% decline in adult survival rate over 11 years. Such a long‐term decline, despite on‐going conservation efforts, calls into question the overall sustainability of arable Stone‐curlew populations. We infer some of the possible causes of this decline, in particular food shortage, and discuss how this pattern could be reversed through conservation measures applicable at very large spatial scales.     

Demographic models of the northern spotted owl (Strix occidentalis caurina)

Summary Calassical demographic methods applied to life history data on the northern spotted owl yield and estimate of the annual geometric rate of increase for the population of λ=0.96±0.03, which is not significantly different from that for a stable population (λ=1.00). Sensitivity analysis indicates that adult annual survivorship has by far the largest influence on λ, followed by the probability that juveniles survive dispersal, and the adult annual fecundity. Substantial temporal fluctuations in demographic parameters have little effect on the long-run growth rate of the population because of the long adult life expectancy. A model of dispersal and territory occupancy that assumes demographic equilibrium is evaluated using data on the amount of old forest habitat remaining in the Pacific Northwest and the current occupancy of this habitat by northern spotted owls. This model is employed to predict the effect of future habitat loss and fragmentation on the population, implying that extinction will result if the old forest is reduced to less than a proportion 0.21±0.02 of the total area in a large region. The estimated minimum habitat requirement for the population is greater than that allowed in management plants by the USDA Forest Service.     

Are species' responses to global change predicted by past niche evolution?

Predicting how and when adaptive evolution might rescue species from global change, and integrating this process into tools of biodiversity forecasting, has now become an urgent task. Here, we explored whether recent population trends of species can be explained by their past rate of niche evolution, which can be inferred from increasingly available phylogenetic and niche data. We examined the assemblage of 409 European bird species for which estimates of demographic trends between 1970 and 2000 are available, along with a species-level phylogeny and data on climatic, habitat and trophic niches. We found that species'' proneness to demographic decline is associated with slow evolution of the habitat niche in the past, in addition to certain current-day life-history and ecological traits. A similar result was found at a higher taxonomic level, where families prone to decline have had a history of slower evolution of climatic and habitat niches. Our results support the view that niche conservatism can prevent some species from coping with environmental change. Thus, linking patterns of past niche evolution and contemporary species dynamics for large species samples may provide insights into how niche evolution may rescue certain lineages in the face of global change.     

Assessing Landbird Monitoring Programs and Demographic Causes of Population Trends

Abstract: Population trend data from the North American Breeding Bird Survey (BBS) have been used to identify conservation priorities and justify major conservation initiatives. Yet the BBS has been criticized for potential habitat bias and reliance on abundance indices to estimate trends. We compared 1992–2003 BBS trend estimates to trend estimates derived from bird-banding data collected as part of the Monitoring Avian Productivity and Survivorship (MAPS) program for 36 wood warbler species. Similarity in trends between the 2 monitoring programs at the survey-wide and program-wide scales suggested that each program can provide accurate trend information. The MAPS program, however, was designed primarily to complement (rather than duplicate) count-based efforts, such as the BBS, by providing estimates or indices of demographic rates. Demographic data from MAPS can be used to lend insight into proximate (demographic) causes of population trends and inform management. We illustrate this with analyses of 1992–2003 MAPS data for yellow warbler (Dendroica petechia). We used reverse-time capture-recapture models to evaluate importance of new recruits (including immigrating adults and young from the previous year) relative to surviving adults in explaining variation in trend among BBS physiographic strata. We included the number of young per adult captured (an index of productivity) as a covariate in models to assess effects of productivity on trends. Survival was the key demographic driver of recent population trends. Comparison of MAPS productivity indices and adult apparent survival rate estimates to BBS trend estimates largely confirmed this inference. We suggest that increased MAPS coverage, better coordination between MAPS and the BBS, and continued development of analytical methods that link the 2 programs will enhance the value of these monitoring efforts to land managers and conservation planners working at a variety of spatial scales.     

Likelihood based population viability analysis in the presence of observation error

Population viability analysis (PVA) entails calculation of extinction risk, as defined by various extinction metrics, for a study population. These calculations strongly depend on the form of the population growth model and inclusion of demographic and/or environmental stochasticity. Form of the model and its parameters are determined based on observed population time series data. A typical population time series, consisting of estimated population sizes, inevitably has some observation error and likely has missing observations. In this paper, we present a likelihood based PVA in the presence of observation error and missing data. We illustrate the importance of incorporation of observation error in PVA by reanalyzing the population time series of song sparrow Melospiza melodia on Mandarte Island, British Columbia, Canada from 1975–1998. Using Akaike information criterion we show that model with observation error fits the data better than the one without observation error. The extinction risks predicted by with and without observation error models are quite different. Further analysis of possible causes for observation error revealed that some component of the observation error might be due to unreported dispersal. A complete analysis of such data, thus, would require explicit spatial models and data on dispersal along with observation error. Our conclusions are, therefore, two‐fold: 1) observation errors in PVA matter and 2) integrating these errors in PVA is not always enough and can still lead to important biases in parameter estimates if other processes such as dispersal are ignored.     

Genetic population structure as indirect measure of dispersal ability in a Lake Tanganyika cichlid

Diversification and speciation processes are influenced by intrinsic (ecological specialization, dispersal) and extrinsic (habitat structure and instability) factors, but the effect of ecological characteristics on dispersal is difficult to assess. This study uses mitochondrial control region sequences to investigate the population structure and demographic history of the endemic Lake Tanganyika cichlid Neolamprologus caudopunctatus with a preference for the rock-sand interface along two stretches of continuous, rocky shoreline, and across a sandy bay representing a potential dispersal barrier. Populations along uninterrupted habitat were not differentiated; whereas, the sandy bay separated two reciprocally monophyletic clades. The split between the two clades between 170,000 and 260,000 years BP coincides with a period of rising water level following a major lowstand, and indicates that clades remained isolated throughout subsequent lake level fluctuations. Low long-term effective population sizes were inferred from modest genetic diversity estimates, and may be due to recent population expansions starting from small population sizes 45,000–60,000 years BP. Comparisons with available data from specialized rock-dwelling species of the␣same area suggest that habitat structure and lake level fluctuations determine phylogeographic patterns on large scales, while fine-scale population structure and demography are modulated by species-specific ecologies.     

Historical demography of a wild lemur population (Propithecus verreauxi) in southwest Madagascar

The human colonization of Madagascar is associated with the extinction of numerous lemur species. However, the degree to which humans have negatively influenced the historical population dynamics of extant lemur species is not well understood. This study employs genetic and demographic analyses to estimate demographic parameters relating to the historical population dynamics of a wild lemur population, Verreaux’s sifaka (Propithecus verreauxi). The genetic analyses are used to determine whether this population experienced a historically recent (i.e., within the last 2000 years) population bottleneck, as well as to estimate the historical population growth rate and the timing of any changes in population size in the past. In addition, a retrospective demographic analysis is used to determine sources of variation and covariation in the sifaka life cycle and how variation in life-cycle transitions contributes to variation in population growth rate. The genetic analyses indicate that the sifaka population did not experience a recent population bottleneck; however, the historical population growth rate was negative, indicating that the ancestral population size was much larger than the current size. The timing of the ancestral population decline has a point estimate of 2300 years ago, but with large credible intervals: 3611–1736 years ago. This point estimate corresponds with the first evidence for human arrival to Madagascar. Climatic variation has also likely influenced past (and current) population dynamics due to stochastic annual rainfall patterns and climatic desiccation, the latter of which began in southwestern Madagascar around 4000 years ago. Variation in the survival of 2-year-old animals as well as large adult females makes the largest contribution to variation in population growth rate. In the absence of more explicit models pertaining to historical population dynamics, it is difficult to attribute the negative population growth rate of this species solely to a single factor (e.g., hunting, habitat destruction).     

Modeling observation error and its effects in a random walk/extinction model

This paper examines the consequences of observation errors for the "random walk with drift", a model that incorporates density independence and is frequently used in population viability analysis. Exact expressions are given for biases in estimates of the mean, variance and growth parameters under very general models for the observation errors. For other quantities, such as the finite rate of increase, and probabilities about population size in the future we provide and evaluate approximate expressions. These expressions explain the biases induced by observation error without relying exclusively on simulations, and also suggest ways to correct for observation error. A secondary contribution is a careful discussion of observation error models, presented in terms of either log-abundance or abundance. This discussion recognizes that the bias and variance in observation errors may change over time, the result of changing sampling effort or dependence on the underlying population being sampled.     

Incorporating harvest rates into the sex-age-kill model for white-tailed deer

Although monitoring population trends is an essential component of game species management, wildlife managers rarely have complete counts of abundance. Often, they rely on population models to monitor population trends. As imperfect representations of real-world populations, models must be rigorously evaluated to be applied appropriately. Previous research has evaluated population models for white-tailed deer (Odocoileus virginianus); however, the precision and reliability of these models when tested against empirical measures of variability and bias largely is untested. We were able to statistically evaluate the Pennsylvania sex-age-kill (PASAK) population model using realistic error measured using data from 1,131 radiocollared white-tailed deer in Pennsylvania from 2002 to 2008. We used these data and harvest data (number killed, age-sex structure, etc.) to estimate precision of abundance estimates, identify the most efficient harvest data collection with respect to precision of parameter estimates, and evaluate PASAK model robustness to violation of assumptions. Median coefficient of variation (CV) estimates by Wildlife Management Unit, 13.2% in the most recent year, were slightly above benchmarks recommended for managing game species populations. Doubling reporting rates by hunters or doubling the number of deer checked by personnel in the field reduced median CVs to recommended levels. The PASAK model was robust to errors in estimates for adult male harvest rates but was sensitive to errors in subadult male harvest rates, especially in populations with lower harvest rates. In particular, an error in subadult (1.5-yr-old) male harvest rates resulted in the opposite error in subadult male, adult female, and juvenile population estimates. Also, evidence of a greater harvest probability for subadult female deer when compared with adult (≥2.5-yr-old) female deer resulted in a 9.5% underestimate of the population using the PASAK model. Because obtaining appropriate sample sizes, by management unit, to estimate harvest rate parameters each year may be too expensive, assumptions of constant annual harvest rates may be necessary. However, if changes in harvest regulations or hunter behavior influence subadult male harvest rates, the PASAK model could provide an unreliable index to population changes. © 2012 The Wildlife Society.     

Below-threshold mortality: implications for studies in evolution,ecology and demography

Evolutionary biologists, ecologists and experimental gerontologists have increasingly used estimates of age-specific mortality as a critical component in studies of a range of important biological processes. However, the analysis of age-specific mortality rates is plagued by specific statistical challenges caused by sampling error. Here we discuss the nature of this ‘demographic sampling error’, and the way in which it can bias our estimates of (1) rates of ageing, (2) age at onset of senescence, (3) costs of reproduction and (4) demographic tests of evolutionary models of ageing. We conducted simulations which suggest that using standard statistical techniques, we would need sample sizes on the order of tens of thousands in most experiments to effectively remove any bias due to sampling error. We argue that biologists should use much larger sample sizes than have previously been used. However, we also present simple maximum likelihood models that effectively remove biases due to demographic sampling error even at relatively small sample sizes.     

Comparing survey and multiple recruitment–mortality models to assess growth rates and population projections

Estimation of population trends and demographic parameters is important to our understanding of fundamental ecology and species management, yet these data are often difficult to obtain without the use of data from population surveys or marking animals. The northeastern Minnesota moose (Alces alces Linnaeus, 1758) population declined 58% during 2006–2017, yet aerial surveys indicated stability during 2012–2017. In response to the decline, the Minnesota Department of Natural Resources (MNDNR) initiated studies of adult and calf survival to better understand cause‐specific mortality, calf recruitment, and factors influencing the population trajectory. We estimated population growth rate (λ) using adult survival and calf recruitment data from demographic studies and the recruitment–mortality (R‐M) Equation and compared these estimates to those calculated using data from aerial surveys. We then projected population dynamics 50 years using each resulting λ and used a stochastic model to project population dynamics 30 years using data from the MNDNR's studies. Calculations of λ derived from 2012 to 2017 survey data, and the R‐M Equation indicated growth (1.02 ± 0.16 [SE] and 1.01 ± 0.04, respectively). However, the stochastic model indicated a decline in the population over 30 years (λ = 0.91 ± 0.004; 2014–2044). The R‐M Equation has utility for estimating λ, and the supporting information from demographic collaring studies also helps to better address management questions. Furthermore, estimates of λ calculated using collaring data were more certain and reflective of current conditions. Long‐term monitoring using collars would better inform population performance predictions and demographic responses to environmental variability.     

Can effective population size estimates be used to monitor population trends of woodland bats? A case study of Myotis bechsteinii

Molecular approaches to calculate effective population size estimates (Ne) are increasingly used as an alternative to long‐term demographic monitoring of wildlife populations. However, the complex ecology of most long‐lived species and the consequent uncertainties in model assumptions means that effective population size estimates are often imprecise. Although methods exist to incorporate age structure into Ne estimations for long‐lived species with overlapping generations, they are rarely used owing to the lack of relevant information for most wild populations. Here, we performed a case study on an elusive woodland bat, Myotis bechsteinii, to compare the use of the parentage assignment Ne estimator (EPA) with the more commonly used linkage disequilibrium (LD) Ne estimator in detecting long‐term population trends, and assessed the impacts of deploying different overall sample sizes. We used genotypic data from a previously published study, and simulated 48 contrasting demographic scenarios over 150 years using the life history characteristics of this species The LD method strongly outperformed the EPA method. As expected, smaller sample sizes resulted in a reduced ability to detect population trends. Nevertheless, even the smallest sample size tested (n = 30) could detect important changes (60%–80% decline) with the LD method. These results demonstrate that genetic approaches can be an effective way to monitor long‐lived species, such as bats, provided that they are undertaken over multiple decades.     

Evaluation of target strength–fish length equation choices for estimating estuarine fish biomass

In the Gulf of Mexico (GOM), fish biomass estimates are necessary for the evaluation of habitat use and function following the mandate for ecosystem-based fisheries management in the recently reauthorized Sustainable Fisheries Act of 2007. Acoustic surveys have emerged as a potential tool to estimate fish biomass in shallow-water estuaries, however, the transformation of acoustic data into an index of fish biomass is not straightforward. In this article, we examine the consequences of equation selection for target strength (TS) to fish length relationships on potential error generation in hydroacoustic fish biomass estimates. We applied structural equation models (SEMs) to evaluate how our choice of an acoustic TS–fish length equation affected our biomass estimates, and how error occurred and propagated during this process. To demonstrate the magnitude of the error when applied to field data, we used SEMs on normally distributed simulated data to better understand the sources of error involved with converting acoustic data to fish biomass. As such, we describe where, and to what magnitude, error propagates when estimating fish biomass. Estimates of fish lengths were affected by measurement errors of TS, and from inexact relationships between fish length and TS. Differences in parameter estimates resulted in significant differences in fish biomass estimates and led to the conclusion that in the absence of known TS–fish length relationships, Love’s (J Acoust Soc Am 46:746–752, 1969) lateral-aspect equation may be an acceptable substitute for an ecosystem-specific TS–fish length relationship. Based upon SEMs applied to simulated data, perhaps the most important, yet most variable, component is the mean volume backscattering strength, which significantly inflated biomass errors in approximately 10% of the cases. Handling editor: M. Power     

Changes in bird populations on sample lowland English farms in relation to loss of hedgerows and other non-crop habitats

Farmland bird population trends were examined on a sample of lowland English farms to assess the relative importance of habitat loss and habitat degradation. Data were extracted from 11 farms surveyed by territory mapping between 1966 and 1986 as part of the British Trust for Ornithology's Common Birds Census. The population size of 38 bird species was quantified for each farm in each year. The extents of five non-crop habitats were measured at 4-yearly intervals on each farm. The farms were selected because some had undergone extensive removal of non-crop habitats while others had undergone little or none. Although declines were commonest on farms where the severest habitat loss had taken place, we found no evidence that habitat loss was the main factor causing population declines: all 11 farms had significant numbers of declining species, even where habitat loss was minimal. Furthermore, general linear modelling found no significant effects of habitat loss on population trends and principal-components analysis found limited effects of habitat extent on community composition. These results suggest that habitat loss is of secondary importance in causing farmland bird population declines. We suggest that other processes, such as habitat degradation, may have caused a baseline population decline in at least 10 farmland bird species and that declines may have been exacerbated by localised habitat loss. Received: 4 February 1998 / Accepted: 1 April 1998     

Impact of Habitat Fragmentation on the Demography of Lion-tailed Macaque (<Emphasis Type="Italic">Macaca silenus</Emphasis>) Populations in the Rainforests of Anamalai Hills,Western Ghats,India

Habitat fragmentation is considered the most serious threat to primate conservation in the tropics, and understanding it effects on lion-tailed macaque is very important because most of the populations live in fragmented habitats. We examined demographic parameters of 9 lion-tailed macaque groups in 8 rain forest fragments with reference to fragment area, tree density, canopy cover, tree height, and total basal area of food trees. Group size ranged from 7 to 90 individuals but was not related to habitat variables. Birth and growth rates of groups did not differ significantly between small (n = 4) and large (n = 4) fragments. Tree density, canopy cover, and total basal of food trees all show strong positive correlations with fragment area. Growth rate correlates with tree density, but there are no other significant relationships between birth or growth rate and habitat variables. The percentage of immature individuals in the group is significantly positively associated with the total basal area of food trees, but not with any other habitat variable. Comparison of our data from this study with data available for the same population in 1996 indicates a slight decline in birth rate but an increase in total number of individuals, from 154 to 242. Of the 5 small fragment groups, 3 have increased in size since 1996 while the sizes of the other 2 groups have remained the same. Based on this study, we advocate that to manage the fragile lion-tailed macaque groups the following steps need to be taken: 1) create dispersal corridors between the fragments using fruit trees to facilitate male dispersal, 2) construct canopy bridges across the prevailing roads, 3) protect the fragments from further degradation, and 4) periodically monitor these populations for long-term conservation.