Removal process estimation of population size for a population with a known sex ratio

A removal model for estimating population size which uses the known population sex ratio is studied. A maximum likelihood estimate and an optimal martingale estimate of the population size are proposed. Their standard errors and large sample properties are obtained. Simulation studies are reported, and the performance of the proposed estimators are compared with the standard maximum likelihood estimator which ignores the sex ratio information. An example on a capture study of deer mice is given.     

Estimating Population Size in a Continuous-time Removal Experiment with a Known Sub-population Size Ratio

We study a continuous-time removal model for estimating the population size for a population in which a sub-population size ratio is known. The maximum likelihood estimate and the optimal martingale estimate of the population size are obtained; these are shown to be equivalent. A comparison between the proposed estimator and the maximum likelihood estimate which ignores the information on the known size ratio is made, using a simulation study. The asymptotic variances of these two estimators are also obtained, and a comparison between them is made. The sensitivity of mis-specification of the known size ratio is examined. We also apply the corresponding discrete-time model to the proposed continuous-time setting, and study the efficiency of the corresponding discrete-time type estimator relative to the proposed estimator.     

A combination line transect and capture-recapture sampling model for multiple observers in aerial surveys

We present a robust sampling methodology to estimate population size using line transect and capture-recapture procedures for aerial surveys. Aerial surveys usually underestimate population density due to animals being missed. A combination of capture-recapture and line transect sampling methods with multiple observers allows violation of the assumption that all animals on the centreline are sighted from the air. We illustrate our method with an example of inanimate objects which shows evidence of failure of the assumption that all objects on the centreline have probability 1 of being detected. A simulation study is implemented to evaluate the performance of three variations of the Lincoln-Petersen estimator: the overall estimator, the stratified estimator, and the general stratified estimator based on the combined likelihood proposed in this paper. The stratified Lincoln-Petersen estimator based on the combined likelihood is found to be generally superior to the other estimators.     

Using the bootstrap with two-phase adaptive stratified samples from multiple populations at multiple locations

Recently the two-phase adaptive stratified sampling design proposed by Francis (1984) has been extended by Manly et al. (2002) for situations where several biological populations are sampled simultaneously, and where this is done at several different geographical locations in order to estimate population totals or means. The method uses the results from a first phase sample to decide how best to allocate a second phase sample to locations and strata, in order to maximise a criterion (based on estimated coefficients of variation) that measures the accuracy of estimation for population totals, for all variables at all locations. One potential problem with this method is bias in the estimators of the population totals and means. In this paper bootstrapping is considered as a means of overcoming these biases. It is shown using model populations of Pacific walrus and shellfish, based on real data, that bootstrapping is a useful tool for removing about half of the bias. This is also confirmed from some simulations using artificial data.     

Estimation of the CDF of a finite population in the presence of acalibration sample

We compare the performance of a number of estimators of the cumulative distribution function (CDF) for the following scenario: imperfect measurements are taken on an initial sample from afinite population and perfect measurements are obtained on a small calibration subset of the initial sample. The estimators we considered include two naive estimators using perfect and imperfect measurements; the ratio, difference and regression estimators for a two-phasesample; a minimum MSE estimator; Stefanski and Bay's SIMEX estimator (1996); and two proposed estimators. The proposed estimators take the form of a weighted average of perfect and imperfect measurements. They are constructed by minimizing variance among the class of weighted averages subject to an unbiasedness constraint. They differ in the manner of estimating the weight parameters. The first one uses direct sample estimates. The second one tunes the unknown parameters to an underlying normal distribution. We compare the root mean square error (RMSE) of the proposed estimator against other potential competitors through computer simulations. Our simulations show that our second estimator has the smallest RMSE among thenine compared and that the reduction in RMSE is substantial when the calibration sample is small and the error is medium or large.     

Mark-recapture estimators for dual frame population size of prominent nesting structures: the effect of uncertain detection probability

The combined mark-recapture and line transect sampling methodology proposed by Alpizar-Jara and Pollock [Journal of Environmental and Ecological Statistics, 3(4), 311–327, 1996; In Marine Mammal Survey and Assessment Methods Symposium. G.W. Garner, S.C. Amstrup, J.L. Laake, B.F.J. Manly, L.L. McDonald, and D.C. Robertson (Eds.), A.A. Balkema, Rotterdam, Netherlands, pp. 99–114, 1999] is used to illustrate the estimation of population size for populations with prominent nesting structures (i.e., bald eagle nests). In the context of a bald eagle population, the number of nests in a list frame corresponds to a pre-marked sample of nests, and an area frame corresponds to a set of transect strips that could be regularly monitored. Unlike previous methods based on dual frame methodology using the screening estimator [Haines and Pollock (Journal of Environmental and Ecological Statistics, 5, 245–256, 1998a; Survey Methodology, 24(1), 79–88, 1998b)], we no longer need to assume that the area frame is complete (i.e., all the nests in the sampled sites do not need to be seen). One may use line transect sampling to estimate the probability of detection in a sampled area. Combining information from list and area frames provides more efficient estimators than those obtained by using data from only one frame. We derive an estimator for detection probability and generalize the screening estimator. A simulation study is carried out to compare the performance of the Chapman modification of the Lincoln–Petersen estimator to the screening estimator. Simulation results show that although the Chapman estimator is generally less precise than the screening estimator, the latter can be severely biased in presence of uncertain detection. The screening estimator outperforms the Chapman estimator in terms of mean squared error when detection probability is near 1 wheareas the Chapman estimator outperforms the screening estimator when detection probability is lower than a certain threshold value depending on particular scenarios.     

Offspring sex ratio response to host size in the parasitoid wasp Spalangia endius

The host size model, an adaptive model for maternal manipulation of offspring sex ratio, was examined for the parasitoid wasp Spalangia endius. In a Florida strain, as the model predicts, daughters emerged from larger hosts than sons, but only when mothers received both small and large hosts simultaneously. The pattern appeared to result from the mother's ovipositional choice and not from differential mortality of the sexes during development. If sex ratio manipulation is adaptive in the Florida strain, it appears to be through a benefit to daughters of developing on large hosts rather than through a benefit to sons of developing on small hosts. Both female and male parasitoids were larger when they developed on larger hosts. For females, developing on a larger host (1) increased offspring production, except for the largest hosts, (2) increased longevity, (3) lengthened development, and (4) had no effect on wing loading. For males, development on a larger host had no effect on any measure of male fitness – mating success, longevity, development duration, or wing loading. In contrast, a strain from India showed no difference in the size of hosts from which daughters versus sons emerged, although both female and male parasitoids were larger when they developed on larger hosts. These results together with previous studies of Spalangia reveal no consistent connection between host-size-dependent sex ratio and host-size-dependent parasitoid size among strains of S. endius or among species of Spalangia. Received: 28 October 1998 / Received in revised form: 20 May 1999 / Accepted: 30 May 1999     

Estimating an homogeneous series of a population abundance indicator despite changes in data collection procedure: A hierarchical Bayesian modelling approach

Abundance indicators are required both to assess and to manage wild populations. As new techniques are developed and teams in charge of gathering the data change, data collection procedures (DCPs) can evolve in space and time. How to estimate an homogeneous series of abundance indicator despite changes in DCP? To tackle this question a hierarchical Bayesian modelling (HBM) approach is proposed. It integrates multiple DCPs in order to derive a single abundance indicator that can be compared over space and time irrespective of the DCP used. Compared to single DCP models, it takes further advantage for abundance estimation of the joint treatment of a larger set of spatio-temporal units. After presenting the general formulation of our HBM approach, it is applied to the juvenile Atlantic salmon (Salmo salar L.) population of the River Nivelle (France). Posterior model checking, using χ² discrepancy measure, do not reveal any inadequacy between the model and the data. Despite a change in the DCP used (successive removals to catch-per-unit of effort), a unique abundance indicator for the 425 spatio-temporal units (site × year) sampled over twenty-four years (1985-2008) is estimated. The HBM approach allows the assessment of precision of the abundance estimates and shows variation between DCPs: a reduction in precision is observed during the most recent years (2005-2008) when only the catch-per-unit of effort DCP was used. The merits and generality of our HBM approach are discussed. We contend it extends previous single DCP models or inter-calibration of two DCPs, and it could be applied to a wide range of specific situations (taxon and DCPs).     

Model of coral population response to accelerated bleaching and mass mortality in a changing climate

We model coral community response to bleaching and mass mortality events which are predicted to increase in frequency with climate change. The model was parameterized for the Arabian/Persian Gulf, but is generally applicable. We assume three species groups (Acropora, faviids, and Porites) in two life-stages each where the juveniles are in competition but the adults can enter a size-refuge in which they cannot be competitively displaced. An aggressive group (Acropora species) dominates at equilibrium, which is not reached due to mass mortality events that primarily disadvantage this group (compensatory mortality, >90% versus 25% in faviids and Porites) roughly every 15 years. Population parameters (N individuals, carrying capacity) were calculated from satellite imagery and in situ transects, vital rates (fecundity, mortality, and survival) were derived from the model, field observations, and literature. It is shown that populations and unaltered community structure can persist despite repeated 90% mortality, given sufficiently high fecundity of the remaining population or import from connected populations. The frequency of disturbance determines the dominant group—in low frequency Acropora, in high frequency Porites. This is congruent with field observations. The model of an isolated population was more sensitive to parameter changes than that of connected populations. Highest sensitivity was to mortality rate and recruitment rate. Community composition was sensitive to spacing of disturbances and level of catastrophic mortality. Decreased mortality led to Acropora dominance, increased mortality led to Acropora extinction. In nature, closely spaced disturbances have severely disadvantaged Acropora populations over the last decade. Unless a longer (>10 years) disturbance-free interval can be maintained, a permanent shift away from Acropora dominance will be observed. A mortality rate of 99% in Acropora, as observed in 1996, is not sustainable if repetitive and neither is a disturbance frequency <15 years—each leading to population collapse. This shows that the severity and/or the spacing of the 1996–1998–2002 disturbances were unusual in frequency and duration.     

Testing the effectiveness of capture mark recapture population estimation techniques using a computer simulation with known population size

Estimation of small mammal population sizes is important for monitoring ecosystem condition and for conservation. Here, we test the accuracy of standard methods of population size estimation using Capture-Mark-Recapture (CMR) on a simulated population of agents. The use of a computer simulation allows complete control of population sizes and behaviors, thereby avoiding assumptions that may be violated in real populations. We find that the recommended protocol for CMR sampling, using uniformly distributed traps, consistently overestimates population sizes by as much as 100% when studies are conducted over only two trapping periods. More than 20 trapping periods are required before this method, or that of placing traps randomly, gives an accurate estimation of population size (i.e., within a 95% confidence limit of the actual value). Non-random sampling, by placing traps on runways used by small mammals, produces the most accurate, and least variable, estimates of population. However, we show that around 10 trapping periods are still required to produce an accurate population estimate using this method. Given that most real populations do not comply with the ‘ideal’ assumptions made by CMR, we suggest that population estimates based on CMR may be fundamentally flawed, and recommend that protocols for CMR population estimation methods may need revising.     

Changes in the cell size of the diatom Cylindrotheca closterium in a hyperhaline pond

Cell length of the diatom Cylindrotheca closterium (Ehr.) Reimann & Lewin was investigated on a fortnightly basis from January to April 2000 in a hyperhaline pond (Sardinia, Mediterranean Sea) and relationships, with changes in temperature, salinity, and nitrite concentrations were assessed using single and multiple regression analyses. Results of our study indicate that C. closterium cell size was inversely related to temperature changes, but that salinity and nitrite concentrations may have a major idiosyncratic effect on cell size variability.     

No reduction in aggression after loss of a broodmate: a test of the brood size hypothesis

In some vertebrate species, parents create a large brood or litter then, in the event of unfavourable ecological conditions, apparently allow the number of offspring to be adaptively reduced through siblicide. But how is sibling aggression regulated so that deaths occur only in unfavourable conditions? One proposed mechanism is brood size-dependent aggression. Two experiments tested for this mechanism by reducing three-chick broods of blue-footed boobies either during or after the period of dominance hierarchy establishment. In neither experiment did aggression of the two eldest and highest ranking chicks decline after removal of the youngest broodmate, in comparison with controls. These results suggest that dominant booby chicks do not become less aggressive to each other after disappearance of their youngest broodmate and that this species does not show brood size dependent aggression. Elder blue-footed booby chicks increase their attacks on broodmates when they receive less food, and this mechanism may be sufficient to tailor brood size to food availability.     

Local adaptation and sexual selection: a reciprocal transfer experiment with the grasshopper Chorthippus biguttulus

Immigration into locally adapted populations has been suggested, among other potential causes, to maintain genetic variance in fitness necessary for good-genes models. Using a reciprocal transplant experiment we examined whether females prefer native to transferred males in the grasshopper Chorthippus biguttulus. On average, native and transferred males did not differ in their attractiveness, measured as female response rate to playbacks of male acoustic courtship signals. In line with this result, we found no significant effect of transfer on body size, condition, fluctuating asymmetry or song traits. However, the reciprocal transplant experiment showed that environmental conditions did influence body condition and maximum loudness of the calling song, but that the genetic origin of male grasshoppers had no significant effect on any of the analysed traits.Communicated by L. Simmons     

Modeling the brown bear population in Slovenia: A tool in the conservation management of a threatened species

In this paper, we address three aspects of the brown bear population in Slovenia: its size (and its evolution over time), its spatial expansion out of the core area, and its potential habitat based on natural habitat suitability. Data collected through measurement/observation of the bear population and from the literature are used. A model is developed for each aspect. The results are estimates of population size, a picture of the spatial expansion of the population and maps of its optimal and maximal potential habitat (based on natural suitability). Overall, the brown bear population has been increasing since the establishment of a core protective area and has been expanding outside this area. The habitat suitability maps show that there is room for further expansion. Based on habitat suitability and bear population density, as well as human activity and current damage reports, we recommend that the Alps should be temporarily kept free of the bears, until the necessary mitigation measures regarding human–bear conflicts are carried out. On the other hand it is of crucial importance to adapt human activities and improve bear management in the optimal habitat, with which the goals of successful conservation of the species might be achieved.     

Solitary behavior in a high-altitude population of the social sweat bee Halictus rubicundus (Hymenoptera: Halictidae)

In the subalpine region of the Rocky Mountains of Colorado, United States, Halictus rubicundus has a solitary life cycle, but it is social in other parts of its known range. The brood is protandrous, with a nearly equal investment in the sexes. Productivity averages 6.5 offspring per foundress female, similar to the second brood of social nests in New York, but less than the combined productivity of both New York broods. Leucophora sp. (Diptera: Anthomyiidae) is the principal cause of brood mortality in Colorado. Foundress females in about half the nests survive until brood emerge as adults. Retention of these foundresses decreases offspring mortality by 68%. Comparable abilities to express solitary behavior with a single brood may characterize other eusocial halictine lineages that have successfully invaded high altitudes in the Rocky Mountains. The apparent inability to do this may help explain the absence of other eusocial halictine bees and polistine wasps at high altitudes, despite their success at lower elevations in the same mountains. Presence or absence of this ability may help explain latitudinal distributions of these lineages in North America. Holarctic distributions of lineages with eusocial behavior can be explained by migration as solitary populations from Eurasia to North America across Pleistocene Bering land bridges, with re-expression of double-brooded, eusocial behavior when the species then extended their ranges southward in North America. Received: 4 November 1994/Accepted after revision: 23 October 1995     

Revealing life‐history traits by contrasting genetic estimations with predictions of effective population size

Effective population size, a central concept in conservation biology, is now routinely estimated from genetic surveys and can also be theoretically predicted from demographic, life‐history, and mating‐system data. By evaluating the consistency of theoretical predictions with empirically estimated effective size, insights can be gained regarding life‐history characteristics and the relative impact of different life‐history traits on genetic drift. These insights can be used to design and inform management strategies aimed at increasing effective population size. We demonstrated this approach by addressing the conservation of a reintroduced population of Asiatic wild ass (Equus hemionus). We estimated the variance effective size (N_ev) from genetic data () and formulated predictions for the impacts on N_ev of demography, polygyny, female variance in lifetime reproductive success (RS), and heritability of female RS. By contrasting the genetic estimation with theoretical predictions, we found that polygyny was the strongest factor affecting genetic drift because only when accounting for polygyny were predictions consistent with the genetically measured N_ev. The comparison of effective‐size estimation and predictions indicated that 10.6% of the males mated per generation when heritability of female RS was unaccounted for (polygyny responsible for 81% decrease in N_ev) and 19.5% mated when female RS was accounted for (polygyny responsible for 67% decrease in N_ev). Heritability of female RS also affected N_ev; (heritability responsible for 41% decrease in N_ev). The low effective size is of concern, and we suggest that management actions focus on factors identified as strongly affecting , namely, increasing the availability of artificial water sources to increase number of dominant males contributing to the gene pool. This approach, evaluating life‐history hypotheses in light of their impact on effective population size, and contrasting predictions with genetic measurements, is a general, applicable strategy that can be used to inform conservation practice.     

Estimating animal densities and home range in regions with irregular boundaries and holes: A lattice-based alternative to the kernel density estimator

Density estimates based on point processes are often restrained to regions with irregular boundaries or holes. We propose a density estimator, the lattice-based density estimator, which produces reasonable density estimates under these circumstances. The estimation process starts with overlaying the region with nodes, linking these together in a lattice and then computing the density of random walks of length k on the lattice. We use an approximation to the unbiased crossvalidation criterion to find the optimal walk length k. The technique is illustrated using walleye (Sander vitreus) radiotelemetry relocations in Lake Monroe, Indiana. We also use simulation to compare the technique to the traditional kernel density estimate in the situation where there are no significant boundary effects.     

Structure and resilience of the social network in an insect colony as a function of colony size

Social interactions are critical to the organization of worker activities in insect colonies and their consequent ecological success. The structure of this interaction network is therefore crucial to our understanding of colony organization and functioning. In this paper, I study the properties of the interaction network in the colonies of the social wasp Ropalidia marginata. I find that the network is characterized by a uniform connectivity among individuals with increasing heterogeneity as colonies become larger. Important network parameters are found to be correlated with colony size and I investigate how this is reflected in the organization of work in colonies of different sizes. Finally, I test the resilience of these interaction networks by experimental removal of individuals from the colony and discuss the structural properties of the network that are related to resilience in a social network. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau, and R. James).     

Inter- and intra-sexual patterns of fluctuating asymmetry in the red-billed streamertail: should symmetry always increase with ornament size?

Recent work on fluctuating asymmetry has suggested that ornaments should have higher levels of fluctuating asymmetry than (1) non-ornaments and (2) homologous structures in the non-ornamented sex. In addition, as both ornament size and symmetry should increase with individual quality there should be a tendency for ornament symmetry to increase with ornament size. In non-ornaments, a U-shaped relationship between symmetry and size is expected, with the individuals at the extremes being more asymmetrical than individuals around the optimum. We tested these predictions in the red-billed streamertail (Trochilus polytmus), a sexually dimorphic endemic Jamaican hummingbird. The lengths of four bilaterally symmetrical traits (first and second outermost tail feathers, tarsi and wings) in 43 adult males and 42 females were measured. The second outermost tail feathers of adult males (which are elongated into streamers) were absolutely but not relatively more asymmetrical than non-ornaments (including the homologous feathers in females). When character size was controlled for, wings were shown to be relatively more symmetrical than other traits. Symmetry did not increase with increasing trait size in any of the morphological traits measured. There was a U-shaped relationship between asymmetry and trait size for four traits (adult male streamers, adult male wings and female outer tail feathers). These results do not support any of the predictions made by fluctuating asymmetry hypotheses and suggest that stabilising selection may act on ornaments as well as non-ornaments. These predictions have been supported in swallows and peafowl but not in sunbirds; this may be due to differences in female perception of tail ornaments. Perhaps male tails do not convey information about quality in some species, or there may be inter-specific differences in the relative costs of tail ornaments and the benefit of marginal increases in tail length and symmetry.