Linking noninvasive genetic sampling and traditional monitoring to aid management of a trans-border carnivore population

Noninvasive genetic sampling has been embraced by wildlife managers and ecologists, especially those charged with monitoring rare and elusive species over large areas. Challenges arise when desired population measures are not directly attainable from genetic data and when monitoring targets trans-border populations. Norwegian management authorities count individual brown bears (Ursus arctos) using noninvasive genetic sampling but express management goals in the annual number of bear reproductions (females that produce cubs), a measure that is not directly available from genetic data. We combine noninvasive genetic sampling data with information obtained from a long-term intensive monitoring study in neighboring Sweden to estimate the number of annual reproductions by females detected within Norway. Most female brown bears in Norway occur near the border with neighboring countries (Sweden, Finland, and Russia) and their potential reproduction can therefore only partially be credited to Norway. Our model includes a simulation-based method that corrects census data to account for this. We estimated that 4.3 and 5.7 reproductions can be credited to females detected with noninvasive genetic sampling in Norway in 2008 and 2009, respectively. These numbers fall substantially short of the national target (15 annual reproductions). Ignoring the potential for home ranges to extend beyond Norway's borders leads to an increase in the estimate of the number of reproductions by -30%. Our study shows that combining noninvasive genetic sampling with information obtained from traditional intensive/invasive monitoring can help answer contemporary management questions in the currency desired by managers and policy makers. Furthermore, combining methodologies and thereby accounting for space use increases the accuracy of the information on which decisions are based. It is important that the information derived from multiple approaches is applicable to the same focal population and that predictions are cross-validated. When monitoring and management are constrained to administrative units, census data should be adjusted by discounting portions of individual space utilization that extend beyond the focal jurisdiction. Our simulation-based approach for making such an adjustment may be useful in other situations where management authorities target portions of trans-border populations.     

Potential effects of environmental contamination on Yuma Myotis demography and population growth.

Unplanned natural and anthropogenic disasters provide unique opportunities for investigating the influence of perturbations on population vital rates and species recovery times. We investigated the potential effects of a major pesticide spill by comparing annual survival rates using mark-recapture techniques on a riparian bat species, Yuma Myotis (Myotis yumanensis). Demography and population dynamics for most bat species remain poorly understood despite advances in mark-recapture estimation and modeling techniques. We compared survival and population growth rates of two roost populations exposed to a large chemical (metam sodium) spill in the upper Sacramento River in Northern California with two roost populations outside the contaminated area from 1992 to 1996. Hypotheses about long-term effects of the spill on female juvenile and adult survival were tested using an information-theoretic approach (AIC). Working hypotheses included effects of age, chemical spill, and time trend on survival. Female adult survival was higher than female juvenile survival across all sites, suggesting stage-specific mortality risks. Model-averaged estimates of female juvenile survival in the contaminated area (0.50-0.74) were lower than in control roosts (0.60-0.78) for each year in the study, suggesting that the spill may have reduced juvenile survival for several years. Female adult survival (0.73-0.89) did not appear to be strongly affected by the spill during the years of the study. There was an increase in survival for both stage-classes across all populations during the study period, which may have been caused by the end of an extended drought in California in the winter of 1993. The spill-affected population was in decline for the first year of the study as indicated by an estimated growth rate (lambda) < 1, but population growth rates increased during the four-year period.     

Albatross populations in peril: a population trajectory for black-browed albatrosses at south Georgia.

Simulation modeling was used to reconstruct Black-browed Albatross (Diomedea melanophris) population trends. Close approximations to observed data were accomplished by annually varying survival rates, reproductive success, and probabilities of returning to breed given success in previous years. The temporal shift in annual values coincided with the start of longline fishing at South Georgia and potential changes in krill abundance. We used 23 years of demographic data from long-term studies of a breeding colony of this species at Bird Island, South Georgia, to validate our model. When we used annual parameter estimates for survival, reproductive success, and probabilities of returning to breed given success in previous years, our model trajectory closely followed the observed changes in breeding population size over time. Population growth rate was below replacement (lambda < 1) in most years and was most sensitive to changes in adult survival. This supports the recent IUCN uplisting of this species from "Vulnerable" to "Endangered." Comparison of pre-1988 and post-1988 demography (before and after the inception of a longline fishery in the breeding area) reveals a decrease in lambda from 0.963 to 0.910. A life table response experiment (LTRE) showed that this decline in lambda was caused mostly by declines in survival of adults. If 1988-1998 demographic rates are maintained, the model predicts a 98% chance of a population of fewer than 25 pairs within 78 years. For this population to recover to a status under which it could be "delisted," a 10% increase in survival of all age classes would be needed.     

Dissecting the drivers of population cycles: Interactions between parasites and mountain hare demography

There is a growing awareness that cyclic population dynamics in vertebrate species are driven by a complex set of interactions rather than a single causal factor. While theory suggests that direct host-parasite interactions may destabilise population dynamics, the interaction between host and parasite may also influence population dynamics through indirect effects that result in delayed responses to either density or to life-history traits. Using empirical data on mountain hares (Lepus timidus) infected with a nematode parasite (Trichostrongylus retortaeformis), we developed an individual-based model (IBM) that incorporated direct effects and delayed life-history effects (DLHEs) of a macroparasite, alternative transmission mechanisms and seasonality in host population dynamics. The full model describes mean characteristics of observed mountain hare time series and parasite abundance, but by systematically removing model structure we dissect out dynamic influences of DLHEs. The DLHEs were weakly destabilising, increasing the propensity for cyclic dynamics and suggesting DLHEs could be important processes in host-parasite systems. Further, by modifying model structure we identify a strong influence of parasite transmission mechanism on host population stability, and discuss the implications for parasite aggregation mechanisms, host movement and natural geographical variation in host population dynamics. The effect of T. retortaeformis on mountain hares likely forms part of a complex set of interactions that lead to population cycles.     

Making a stand: five centuries of population growth in colonizing populations of Pinus ponderosa

The processes underlying the development of new populations are important for understanding how species colonize new territory and form viable long-term populations. Life-history-mediated processes such as Allee effects and dispersal capability may interact with climate variability and site-specific factors to govern population success and failure over extended time frames. We studied four disjunct populations of ponderosa pine in the Bighorn Basin of north-central Wyoming to examine population growth spanning more than five centuries. The study populations are separated from continuous ponderosa pine forest by distances ranging from 15 to >100 km. Strong evidence indicates that the initial colonizing individuals are still present, yielding a nearly complete record of population history. All trees in each population were aged using dendroecological techniques. The populations were all founded between 1530 and 1655 cal yr CE. All show logistic growth patterns, with initial exponential growth followed by a slowing during the mid to late 20th century. Initial population growth was slower than expectations from a logistic regression model at all four populations, but increased during the mid-18th century. Initial lags in population growth may have been due to strong Allee effects. A combination of overcoming Allee effects and a transition to favorable climate conditions may have facilitated a mid-18th century pulse in population growth rate.     

On costs and decisions in population management

Many populations have a negative impact on their habitat, or upon other species in the environment, if their numbers become too large. For this reason they are often managed using some form of control. The objective is to keep numbers at a sustainable level, while ensuring survival of the population. Here we present models that allow population management programs to be assessed. Two common control regimes will be considered: reduction and suppression. Under the suppression regime the population is maintained close to a particular threshold through near continuous control, while under the reduction regime, control begins once the population reaches a certain threshold and continues until it falls below a lower pre-defined level. We discuss how to best choose the control parameters, and we provide tools that allow population managers to select reduction levels and control rates. Additional tools will be provided to assess the effect of different control regimes, in terms of population persistence and cost. In particular we consider the effects of each regime on the probability of extinction and the expected time to extinction, and compare the control methods in terms of the expected total cost of each regime over the life of the population. The usefulness of our results will be illustrated with reference to the control of a koala population inhabiting Kangaroo Island, Australia.     

Dynamics and management of infectious disease in colonizing populations

The introduction of chronic, infectious diseases by colonizing populations (invasive or reintroduced) is a serious hazard in conservation biology, threatening the original host and other spillover species. Most research on spatial invasion of diseases has pertained to established host populations, either at steady state or fluctuating through time. Within a colonizing population, however, the spread of disease may be influenced by the expansion process of the population itself. Here we explore the simultaneous expansion of a colonizing population and a chronic, nonlethal disease introduced with it, describing basic patterns in homogeneous and structured landscapes and discussing implications for disease management. We describe expected outcomes of such introductions for three qualitatively distinct cases, depending on the relative velocities at which the population and epidemic expand. (1) If transmissibility is low the disease cannot be sustained, although it may first expand its range somewhat around the point of introduction. (2) If transmissibility is moderate but the wave-front velocity for the population, vp, is higher than that for the disease, vd, the disease wave front lags behind that of the population. (3) A highly transmissible disease, with vd > vp, will invade sufficiently rapidly to track the spread of the host. To test these elementary theoretical predictions, we simulated disease outbreaks in a spatially structured host population occupying a real landscape. We used a spatially explicit, individual-based model of Persian fallow deer (Dama mesopotamica) reintroduced in northern Israel, considering a hypothetical introduction of bovine tuberculosis. Basic patterns of disease expansion in this realistic setting were similar to our conceptual predictions for homogeneous landscapes. Landscape heterogeneity, however, induced the establishment of population activity centers and disease foci within them, leading to jagged wave fronts and causing local variation in the relative velocities at which the population and epidemic expanded. Based on predictions from simple theory and simulations of managed outbreaks, we suggest that the relative velocities at which the population and epidemic expand have important implications for the impact of different management strategies. Recognizing which of our three general cases best describes a particular outbreak will aid in planning an efficient strategy to contain the disease.     

Body size,demography and foraging in a socially plastic sweat bee: a common garden experiment

Phenotypic plasticity may evolve when conditions vary temporally or spatially on a small enough scale. Plasticity is thought to play a central role in the early stages of evolutionary transitions, including major transitions such as those between non-sociality and sociality. The sweat bee Halictus rubicundus is of special interest in this respect, because it is socially plastic in the British Isles: Nests are social or non-social depending on the environment. However, sociality comprises a complex suite of inter-related traits. To further investigate social plasticity in H. rubicundus, we measured traits that are potentially integral to social phenotype at a northern site, where nests are non-social, and a southern site where nests can be social. We found that foundresses at non-social sites were smaller, produced offspring of a size more similar to themselves, initiated nesting later, and took longer to produce their first female offspring. They began provisioning earlier in the day, finished earlier, and collected more pollen loads. Common garden experiments suggested that these differences represent mainly plasticity, as expected for traits involved in the overall plastic social phenotype, with only limited evidence for fixed genetic differences in foraging. Conditions during overwintering did not have major effects on a foundress' subsequent behaviour.     

The optimal harvest and management in the models of animal populations

We extended zero-cost optimization model for population of domestic animals. Also the model of hiring of labor in cattle-breeding farm is constructed and investigated.     

Determinants of reproductive potential and population size in open populations of Patella vulgata

The importance of external and internal population processes in determining variation in reproductive output and variation in population size were quantified with model simulations for open populations of the sequentially hermaphroditic limpet Patella vulgata using field data from the Isle of Man and South West Ireland. Cross-correlation analyses of model outputs and elasticity analyses show that population dynamics are dominated by the effects of large females, and that recruitment adds little to reproductive output. However, populations experiencing low but highly variable recruitment appear male limited and recruitment pulses carrying young males into the population are correlated to reproductive output with a 2–5-year lag. We conclude that pulses in recruitment can be a major structuring force in these limpet populations, but site-specific post-recruitment processes will determine the relative importance of recruitment to population dynamics and the lag between recruitment and reproductive output.     

Cooperation in breeding by nonreproductive wrens: kinship,reciprocity, and demography

Summary Stripe-backed wrens (Campylorhynchus nuchalis) often live as adults in large groups on permanent, communally defended territories. Nonbreeding adults cooperate in rearing the young of a single breeding pair; this aid substantially increases the reproductive success of the breeders. In a 6-year study in Venezuela of a completely colorbanded population of 25–30 groups, most adults participated in breeding only as helpers and priority to breeding status was strictly age-determined. Detailed behavioral observations at breeding nests with nestlings showed that, in a sample of 100, helpers nearly always contributed as much to the care of young as breeders. Further, aid-giving does not vary systematically with relatedness of ycung to helpers or with probability of future reciprocation by young. Young being raised are most often at least half siblings of helpers, but seldom return aid to adults that helped raise them. Even adopted helpers collaborate fully. Patterns of demography and dispersal show slow turnover of breeders, delayed reproduction, and a viscous population structure.Application of Hamilton's condition for selection for aid-giving reveals that most individuals in this population can maximize inclusive fitness in the first 2 years by helping instead of breeding. Variation in helping effort and in age of first breeding is related to variation in natal group size and competition resulting from variable demographic neighborhoods in different years or in different parts of the population. Because reciprocation in the form of specific alliance formation among nonreproductives is uncommon, nonspecific reciprocity between cohorts and kin selection account well for the observed pattern of age-dependence in first breeding. Nondiscriminating helping in this population is associated with stable monogamous pair boncs, stable territory boundaries and group membership, strict seniortiy for breeding position, high viscosity and consistent effectiveness of aid. Under these circumstances, very simple behavioral rules amounting to nearly automatic helping seem sufficient to confer critical inclusive fitness gain on helpers.     

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.     

Connectivity or demography: Defining sources and sinks in coral reef fish metapopulations

The identity of an individual patch as a source or a sink within a metapopulation is a function of its ability to produce individuals and to disperse them to other patches. In marine systems patch identity is very often defined by dispersal ability alone—upstream patches are sources—while issues of variable habitat quality (which affects local production) are ignored. This can have important ramifications for the science of marine reserve siting. This study develops a spatially explicit source–sink metapopulation model for reef fish and uses it to evaluate the relative importance of connectivity versus demography and how this depends upon the level of local larval retention and the strength of density-dependent recruitment. Elasticity analyses indicated that patch contribution (source or sink) was more sensitive to demographic parameters (particularly survival) than connectivity and this effect was conserved even under strong levels of density-dependence and was generally strengthened as local retention increased. Variability in the relationship between parameter elasticity and local retention was shown to be dependent upon the magnitude of connectivity for an individual patch relative to a critical connectivity value. The proportion of larvae lost due to transport processes was an important parameter which directly affected the magnitude of this critical connectivity value. Patches with connectivity values less than the critical value contributed to the metapopulation largely via production (i.e., local demographics most important). As local retention increased, so did the importance of demographic parameters in these patches. Patches with connectivity values greater than the critical value contributed largely via dispersal of larvae and thus the importance of local demographics decreased as local retention increased.     

Finding regulation among seemingly unregulated populations: a practical framework for analyzing multivariate population time series for their interactions

The structure of ecological communities is often thought to be strongly influenced by population interactions. The interactions are often labeled as bottom-up and top-down control. Previous approaches to identify these processes often assume each population in the community is itself regulated. Therefore, each time series follows a stationary process. However, complex community structure and a lack of regulation in an individual population can result in inappropriate inferences based on traditional statistical approaches. Here, we introduce a statistical framework to analyze potentially non-stationary time series that are collectively regulated. We demonstrate the method with catch-per-unit-effort time series data of selected populations in the Gulf of Mexico. In the Gulf, we found that most of the time series data, which span 26 years, were non-stationary, thus individually unregulated. Species interaction patterns were location-dependent, but where brown shrimp interacted significantly with other species, we identified significant bottom-up forcing. On the other hand, we find almost no evidence of top-down forcing throughout the study areas.     

Grassland songbird survival and recruitment in agricultural landscapes: implications for source-sink demography

Population growth and decline are particularly sensitive to changes in three key life-history parameters: annual productivity, juvenile survival, and adult survival. However, for many species these parameters remain unknown. For example, although grassland songbirds are imperiled throughout North America, for this guild, only a small number of studies have assessed these parameters. From 2002 to 2006, in the agricultural landscape of the Champlain Valley of Vermont and New York, USA, we studied Savannah Sparrow (Passerculus sandwichensis) and Bobolink (Dolichonyx oryzivorus) demography on four grassland treatments: (1) early-hayed fields cut before 11 June and again in early- to mid-July; (2) middle-hayed fields cut once between 21 June and 10 July; (3) late-hayed fields cut after 1 August; and (4) rotationally grazed pastures. We assessed whether these treatments affected adult apparent survival (phi) and recruitment (f), how sensitive these parameters were to the presence of nonbreeders and local dispersal, and the populations' ability to persist in these four habitats. On average, birds using late-hayed fields had > 25% higher apparent survival than those on the more intensively managed early-hayed, middle-hayed, and grazed fields. Overall male phi was 35% higher than female phi, and Savannah Sparrow phi was 44% higher than Bobolink phi. Across all analyses and treatments, apparent survival estimates were 0.58-0.85 for male and 0.48-0.71 for female Savannah Sparrows, and 0.52-0.70 for male and 0.19-0.55 for female Bobolinks. For males of both species, potential nonbreeders decreased the precision of and lowered apparent survival estimates by 25%; female estimates showed little variation with the inclusion of nonbreeders. Inclusion of local dispersal observations increased apparent survival estimates and, in many cases, increased precision, though the effect was stronger for Savannah Sparrows than for Bobolinks, and also stronger for males than for females. High Savannah Sparrow apparent survival rates resulted in stable or near stable populations (lambda approximately 1), particularly in late-hayed and grazed fields, while low Bobolink apparent survival rates resulted in strongly declining populations (lambda < 1) in all treatments.     

Temporal patterns of populations in a warming world: a modelling framework

In this paper, we present an approach for describing the environmentally induced temporal pattern of structured populations by partial integro-differential equations. Populations are structured according to size or stage. Growth, energy allocation and stage transitions are affected by environmental conditions of which temperature, photoperiod, water depth and food supply were taken into account. The resulting modelling framework was applied to describe, analyse and predict alterations in populations with continuous development, populations with distinct state structures and interacting populations. Our exemplary applications consider populations of freshwater Amphipoda, Isopoda and Odonata. The model was capable of simulating life cycle alterations in dependence on temperature in interaction with other environmental factors: (1) population dynamics, (2) seasonal regulation, (3) water depth-dependent dispersal, (4) intraguild predation and (5) consumer-resource dynamics.     

Comparing geographic boundaries in songbird demography data with vegetation boundaries: a new approach to evaluating habitat quality

To plan for the habitat needs of forest songbirds of conservation concern, managers need to understand how spatial heterogeneity in forest conditions influences habitat quality. I used difference boundary detection (wombling) and spatially constrained clustering to delineate boundaries in various combinations of four forest vegetation variables (understory height, understory density, percent deciduous vs. conifer understory, and percent canopy closure) in two Michigan northern hardwood forests. My goal was to identify vegetation boundaries that corresponded with boundaries in an understory-dependent songbird’s distribution, and with boundaries in demographic measures for this songbird that indicate habitat quality (e.g., occupancy by older vs. yearling males, reproductive success). Both forests were actively-managed, mature stands: The first site (78 ha) was heavily deer-browsed (HB), with many browse-resistant conifers in the understory, and the second (62 ha) was less-browsed (LB), with deciduous-dominated understory. I compared the vegetation difference and cluster boundaries to difference boundaries based on 6 years of distribution and demographic data for black-throated blue warblers (Dendroica caerulescens). At the HB site, warbler boundaries overlapped strongly with vegetation boundaries that included all four variables, and clustering effectively divided the habitat into areas with different warbler occupancy and demographic characteristics. At the LB site, warbler distribution showed high overlap with difference and cluster boundaries based on just the height and density of understory vegetation, and cluster boundaries again effectively partitioned the study area into sites that varied in habitat quality. Thus, geographic boundary analysis is likely to be a useful tool for identifying key vegetation variables for management, and for delineating clusters (habitat patches) within sites that capture differences in habitat quality.

The tunicate Salpa thompsoni ecology in the Southern Ocean. I. Distribution, biomass, demography and feeding ecophysiology

Distribution, density, and feeding dynamics of the pelagic tunicate Salpa thompsoni have been investigated during the expedition ANTARKTIS XVIII/5b to the Eastern Bellingshausen Sea on board RV Polarstern in April 2001. This expedition was the German contribution to the field campaign of the Southern Ocean Global Ocean Ecosystems Dynamics Study (SO-GLOBEC). Salps were found at 31% of all RMT-8 and Bongo stations. Their densities in the RMT-8 samples were low and did not exceed 4.8 ind m⁻² and 7.4 mg C m⁻². However, maximum salp densities sampled with the Bongo net reached 56 ind m⁻² and 341 mg C m⁻². A bimodal salp length frequency distribution was recorded over the shelf, and suggested two recent budding events. This was also confirmed by the developmental stage composition of solitary forms. Ingestion rates of aggregate forms increased from 2.8 to 13.9 μg (pig) ind⁻¹ day⁻¹ or from 0.25 to 2.38 mg C ind⁻¹ day⁻¹ in salps from 10 to 40 mm oral-atrial length, accounting for 25–75% of body carbon per day. Faecal pellet production rates were on average 0.08 pellet ind⁻¹ h⁻¹ with a pronounced diel pattern. Daily individual egestion rates in 13 and 30 mm aggregates ranged from 0.6 to 4.8 μg (pig) day⁻¹ or from 164 to 239 μg C day⁻¹. Assimilation efficiency ranged from 73 to 90% and from 65 to 76% in 13 and 30 mm aggregates, respectively. S. thompsoni exhibited similar ingestion and egestion rates previously estimated for low Antarctic (~50°S) habitats. It has been suggested that the salp population was able to develop in the Eastern Bellingshausen Sea due to an intrusion into the area of the warm Upper Circumpolar Deep Water