Setting Realistic Recovery Targets for Two Interacting Endangered Species,Sea Otter and Northern Abalone

Failure to account for interactions between endangered species may lead to unexpected population dynamics, inefficient management strategies, waste of scarce resources, and, at worst, increased extinction risk. The importance of species interactions is undisputed, yet recovery targets generally do not account for such interactions. This shortcoming is a consequence of species‐centered legislation, but also of uncertainty surrounding the dynamics of species interactions and the complexity of modeling such interactions. The northern sea otter (Enhydra lutris kenyoni) and one of its preferred prey, northern abalone (Haliotis kamtschatkana), are endangered species for which recovery strategies have been developed without consideration of their strong predator–prey interactions. Using simulation‐based optimization procedures from artificial intelligence, namely reinforcement learning and stochastic dynamic programming, we combined sea otter and northern abalone population models with functional‐response models and examined how different management actions affect population dynamics and the likelihood of achieving recovery targets for each species through time. Recovery targets for these interacting species were difficult to achieve simultaneously in the absence of management. Although sea otters were predicted to recover, achieving abalone recovery targets failed even when threats to abalone such as predation and poaching were reduced. A management strategy entailing a 50% reduction in the poaching of northern abalone was a minimum requirement to reach short‐term recovery goals for northern abalone when sea otters were present. Removing sea otters had a marginally positive effect on the abalone population but only when we assumed a functional response with strong predation pressure. Our optimization method could be applied more generally to any interacting threatened or invasive species for which there are multiple conservation objectives. Definición de Metas de Recuperación Realistas para Dos Especies en Peligro Interactuantes, Enhydra lutris y Haliotis kamtschatkana     

Evaluating mortality rates with a novel integrated framework for nonmonogamous species

The conservation of wildlife requires management based on quantitative evidence, and especially for large carnivores, unraveling cause‐specific mortalities and understanding their impact on population dynamics is crucial. Acquiring this knowledge is challenging because it is difficult to obtain robust long‐term data sets on endangered populations and, usually, data are collected through diverse sampling strategies. Integrated population models (IPMs) offer a way to integrate data generated through different processes. However, IPMs are female‐based models that cannot account for mate availability, and this feature limits their applicability to monogamous species only. We extended classical IPMs to a two‐sex framework that allows investigation of population dynamics and quantification of cause‐specific mortality rates in nonmonogamous species. We illustrated our approach by simultaneously modeling different types of data from a reintroduced, unhunted brown bear (Ursus arctos) population living in an area with a dense human population. In a population mainly driven by adult survival, we estimated that on average 11% of cubs and 61% of adults died from human‐related causes. Although the population is currently not at risk, adult survival and thus population dynamics are driven by anthropogenic mortality. Given the recent increase of human‐bear conflicts in the area, removal of individuals for management purposes and through poaching may increase, reversing the positive population growth rate. Our approach can be generalized to other species affected by cause‐specific mortality and will be useful to inform conservation decisions for other nonmonogamous species, such as most large carnivores, for which data are scarce and diverse and thus data integration is highly desirable.     

Accuracy of Short‐Term Demographic Data in Projecting Long‐Term Fate of Populations

Short‐term surveys are useful in conservation of species if they can be used to reliably predict the long‐term fate of populations. However, statistical evaluations of reliability are rare. We studied how well short‐term demographic data (1999–2002) of tartar catchfly (Silene tatarica), a perennial riparian plant, projected the fate and growth of 23 populations of this species up to the year 2010. Surveyed populations occurred along a river with natural flood dynamics and along a regulated river. Riparian plant populations are affected by flooding, which maintains unvegetated shores, while forest succession proceeds in areas with little flooding. Flooding is less severe along the regulated river, and vegetation overgrowth reduces abundance of tartar catchfly on unvegetated shores. We built matrix models to calculate population growth rates and estimated times to population extinction in natural and in regulated rivers, 13 and 10 populations, respectively. Models predicted population survival well (model predictions matched observed survival in 91% of populations) and accurately predicted abundance increases and decreases in 65% of populations. The observed and projected population growth rates differed significantly in all but 3 populations. In most cases, the model overestimated population growth. Model predictions did not improve when data from more years were used (1999–2006). In the regulated river, the poorest model predictions occurred in areas where cover of other plant species changed the fastest. Although vegetation cover increased in most populations, it decreased in 4 populations along the natural river. Our results highlight the need to combine disturbance and succession dynamics in demographic models and the importance of habitat management for species survival along regulated rivers. Precisión de Datos Demográficos de Corto Plazo en la Proyección del Destino de Poblaciones a Largo Plazo     

Effects of an Invasive Plant on Population Dynamics in Toads

When populations decline in response to unfavorable environmental change, the dynamics of their population growth shift. In populations that normally exhibit high levels of variation in recruitment and abundance, as do many amphibians, declines may be difficult to identify from natural fluctuations in abundance. However, the onset of declines may be evident from changes in population growth rate in sufficiently long time series of population data. With data from 23 years of study of a population of Fowler's toad (Anaxyrus [ = Bufo] fowleri) at Long Point, Ontario (1989–2011), we sought to identify such a shift in dynamics. We tested for trends in abundance to detect a change point in population dynamics and then tested among competing population models to identify associated intrinsic and extrinsic factors. The most informative models of population growth included terms for toad abundance and the extent of an invasive marsh plant, the common reed (Phragmites australis), throughout the toads’ marshland breeding areas. Our results showed density‐dependent growth in the toad population from 1989 through 2002. After 2002, however, we found progressive population decline in the toads associated with the spread of common reeds and consequent loss of toad breeding habitat. This resulted in reduced recruitment and population growth despite the lack of significant loss of adult habitat. Our results underscore the value of using long‐term time series to identify shifts in population dynamics coincident with the advent of population decline. Efectos de una Planta Invasora sobre las Dinámica Poblacional de Sapos     

Individual based model of slug population and spatial dynamics

The slug, Deroceras reticulatum, is one of the most important pests of agricultural and horticultural crops in UK and Europe. In this paper, a spatially explicit individual based model (IbM) is developed to study the dynamics of a population of D. reticulatum. The IbM establishes a virtual field within which slug spatial dynamics and changes in abundance were simulated. The strong dependence of slug behaviour on environmental conditions is built into the model, which is based upon previous work on the environmental dependence of slug population dynamics. The simulation results show that the IbM described well changes in the slug population. The IbM proved capable of describing slug populations over 3.5 years, including the presence, magnitude and duration of D. reticulatum population crashes within this period. Moreover, the model was capable of reproducing slug population dynamics at two sites, with distinct weather and some 100 km apart, with minor changes in initialisation values but no change in model structure and parameter values. A study of field heterogeneity, which might simulate various field designs, indicated the importance of spatial structuring to slug population dynamics and the utility of the IbM for simulating a range of potential spatial management treatments for slug control to maximise crop yield. This IbM system performs well and is currently being used as part of an integrated approach to predict slug population dynamics and control in the UK.     

Population dynamics of Müllerian mimicry under interspecific competition

We ask what the effects of mutualism on population dynamics of two competitive species are. We model the population dynamics of mutualistic interactions with positive density- and frequency-dependences. We specifically assume the dynamics of Müllerian mimicry in butterflies, where the mortality of both species is reduced depending on the relative frequency of the other species. We assume that the two species are under Lotka–Volterra density-dependent competition. The equilibria are compared with the cases of competition alone. Unlike the traditional model of positive density-dependence, population explosion does not appear in the current dynamics, but the new equilibrium is simply achieved. It is because the effects of positive density- or frequency-dependence are restricted to parts of mortality. Both positive density- and frequency-dependences do promote coexistence of the mimetic species. However, the two models show a distinctive difference for coexistence. The effects of positive density-dependence are rather limited. In contrast, positive frequency-dependence always promotes coexistence, irrespective of environmental conditions. The results may imply that the evolutionary origin of Müllerian mimicry may depend on frequency-dependence (and density-dependence), but that its current population dynamics may depend solely on density-dependence. The role of frequency- and density-dependences on evolutionary dynamics is an open question.     

Population dynamics of splitnose rockfish (Sebastes diploproa) in the Northeast Pacific Ocean

We developed an age-structured population model of splitnose rockfish, Sebastes diploproa, in the Northeast Pacific Ocean. Splitnose rockfish is a bycatch species that co-occurs with several commercially important species that are currently declared overfished. Bycatch species are typically not the focus of stock assessment efforts because of their limited economic importance, but they may suffer the same population declines as species with which they co-occur. To examine the dynamics of splitnose rockfish for the first time, we analyzed data from three groundfish fisheries and four research surveys conducted in the Northeast Pacific Ocean. To develop a model, we used Stock Synthesis, a statistical framework for the construction of a population dynamics models utilizing both fishery-dependent and fishery-independent data. In the model, we reconstructed the total catch of the species back to 1900, estimated the dynamics of the stock spawning output and recruitment and evaluated biomass depletion relative to the stock's unfished state, as well as sources of uncertainty in model outputs. The results indicate that the splitnose rockfish is currently not overfished even though it has experienced several periods of abrupt decline in its biomass. Revisiting age data from earlier years, monitoring fishery discard, and investigating the spatial dynamics of splitnose rockfish is important to further improve the understanding of this species’ population dynamics, and decrease uncertainty in model results.     

Integrating Landscape and Metapopulation Modeling Approaches: Viability of the Sharp-Tailed Grouse in a Dynamic Landscape

Abstract:  The lack of management experience at the landscape scale and the limited feasibility of experiments at this scale have increased the use of scenario modeling to analyze the effects of different management actions on focal species. However, current modeling approaches are poorly suited for the analysis of viability in dynamic landscapes. Demographic (e.g., metapopulation) models of species living in these landscapes do not incorporate the variability in spatial patterns of early successional habitats, and landscape models have not been linked to population viability models. We link a landscape model to a metapopulation model and demonstrate the use of this model by analyzing the effect of forest management options on the viability of the Sharp-tailed Grouse (  Tympanuchus phasianellus ) in the Pine Barrens region of northwestern Wisconsin (U.S.A.). This approach allows viability analysis based on landscape dynamics brought about by processes such as succession, disturbances, and silviculture. The landscape component of the model (LANDIS) predicts forest landscape dynamics in the form of a time series of raster maps. We combined these maps into a time series of patch structures, which formed the dynamic spatial structure of the metapopulation component (RAMAS). Our results showed that the viability of Sharp-tailed Grouse was sensitive to landscape dynamics and demographic variables such as fecundity and mortality. Ignoring the landscape dynamics gave overly optimistic results, and results based only on landscape dynamics (ignoring demography) lead to a different ranking of the management options than the ranking based on the more realistic model incorporating both landscape and demographic dynamics. Thus, models of species in dynamic landscapes must consider habitat and population dynamics simultaneously.     

Demographic Models and Reality in Reintroductions: Persian Fallow Deer in Israel

Abstract:  Because most reintroduced species are rare, data on their dynamics are scarce. Consequently, reintroduction programs often rely on data from other species or captive populations to project the performance of the reintroduced population in the wild. We compared the reproductive success and survival of a Persian fallow deer ( Dama mesopotamica ) population reintroduced in Israel over the first 5 years of the project with the survival and reproduction parameters estimated while planning the reintroduction. In addition, we compared the actual growth of the wild population with the growth originally projected by a computer model in the original reintroduction program. We monitored 74 radio-collared individuals (57 females and 17 males) released semiannually 1996–2001. Survival during the first year after release was lower than later years (0.90 and 0.82 versus 0.95 and 0.88, for females and males, respectively). Such an impact was not anticipated in the original plan, but overall survival was higher than originally projected. As assumed in the reintroduction program, reproductive success improved significantly with time since release and overall, was higher than expected. The mean number of animals released annually was lower than planned. Overall, the growth of the reintroduced population was slower than projected, but the deviation was close to confidence limits and the pattern similar. After 5 years it appears that the original time frame of 8–10 years for project completion can be met or at worst will cause a 1-year delay. Over the short term of 5 years, projection models in reintroduction programs are useful tools for assessing the sustained use of the breeding core, depicting the dynamics of the population in the wild, providing a relatively accurate time frame for the successful completion of the project, and assessing project success.     

The demographic consequences of the cost of reproduction in ungulates

The cost of reproduction can generate covariation between demographic rates that can potentially influence demography and population dynamics in long-lived iteroparous species. However, there has been relatively little work linking the survival cost of reproduction and population dynamics. The apparent scarcity of information on this important link is potentially due to covariation between vital rates, which can substantially influence fluctuations in population size. In this paper we examine the opportunity for survival costs of reproduction to leave a dynamic signature using a simulation model based broadly on an ungulate life history. We find that an increase in the cost delays the onset of reproduction and reduces reproductive rates of young, but not of prime-age, females. Accordingly, the number of offspring produced declines and the interval between reproductive events increases among young females experiencing high cost. These effects are translated to an age structure skewed toward young ages and reduced population density. These results suggest that, by delaying reproduction when conditions deteriorate, females protect their survival during the critical first three years of life, after which the negative effect of reproduction on survival declines. Unless conditions for reproduction are severe, it is not profitable to delay reproduction beyond age 3 years due to the high risk of death before having a chance to reproduce. We also demonstrate that lack of adjustment of reproductive strategies to elevated levels of the cost of reproduction, for example due to rapid changes in environmental conditions, results in lower average density and longevity compared to females that have sufficient time to adjust to changes in the cost. This suggests that even moderate costs of reproduction may have a major negative effect on population dynamics of ungulates.     

Two-way coupling versus one-way forcing of plankton and fish models to predict ecosystem changes in the Benguela

‘End-to-end’ models have been adopted in an attempt to capture more of the processes that influence the ecology of marine ecosystems and to make system wide predictions of the effects of fishing and climate change. Here, we develop an end-to-end model by coupling existing models that describe the dynamics of low (ROMS–N₂P₂Z₂D₂) and high trophic levels (OSMOSE). ROMS–N₂P₂Z₂D₂ is a biogeochemical model representing phytoplankton and zooplankton seasonal dynamics forced by hydrodynamics in the Benguela upwelling ecosystem. OSMOSE is an individual-based model representing the dynamics of several species of fish, linked through opportunistic and size-based trophic interactions. The models are coupled through a two-way size-based predation process. Plankton provides prey for fish, and the effects of predation by fish on the plankton are described by a plankton mortality term that is variable in space and time. Using the end-to-end model, we compare the effects of two-way coupling versus one-way forcing of the fish model with the plankton biomass field. The fish-induced mortality on plankton is temporally variable, in part explained by seasonal changes in fish biomass. Inclusion of two-way feedback affects the seasonal dynamics of plankton groups and usually reduces the amplitude of variation in abundance (top-down effect). Forcing and coupling lead to different predicted food web structures owing to changes in the dominant food chain which is supported by plankton (bottom-up effect). Our comparisons of one-way forcing and two-way coupling show how feedbacks may affect abundance, food web structure and food web function and emphasise the need to critically examine the consequences of different model architectures when seeking to predict the effects of fishing and climate change.     

The relationship between ecology and the incidence of cooperative breeding in Australian birds

Summary Australia has many cooperatively breeding species of birds. These tend to occur in eucalypt and semi-arid woodlands rather than in rainforests or deserts. They tend to be insectivores that pursue rather than sit and wait for their prey, and tend to forage on the ground rather than above it. We propose that environments where resources do not show marked seasonal fluctuations are those in which cooperative breeding is most likely to evolve. Under these conditions birds might experience difficulty acquiring the extra food necessary to breed, especially if inexperienced. When adult survival was high, young and inexperienced birds could delay breeding. Unpredictable environments may also favor cooperative breeding, but our data do not strongly support this. Groupliving would be favored further if young birds are particularly vulnerable to predators when alone. They should therefore remain in the family group and delay their dispersal unitl a suitable breeding vacancy becomes available. These hypotheses are not mutually exclusive, but are complementary. Both may be required to ensure that at least some year-old birds do not breed and also do not disperse. We believe that they give rise to predictions, which can be tested in future field studies.     

A Behaviorally Explicit Demographic Model Integrating Habitat Selection and Population Dynamics in California Sea Lions

Abstract: Although there has been a call for the integration of behavioral ecology and conservation biology, there are few tools currently available to achieve this integration. Explicitly including information about behavioral strategies in population viability analyses may enhance the ability of conservation biologists to understand and estimate patterns of extinction risk. Nevertheless, most behavioral‐based PVA approaches require detailed individual‐based data that are rarely available for imperiled species. We present a mechanistic approach that incorporates spatial and demographic consequences of behavioral strategies into population models used for conservation. We developed a stage‐structured matrix model that includes the costs and benefits of movement associated with 2 habitat‐selection strategies (philopatry and direct assessment). Using a life table for California sea lions (Zalophus californianus), we explored the sensitivity of model predictions to the inclusion of these behavioral parameters. Including behavioral information dramatically changed predicted population sizes, model dynamics, and the expected distribution of individuals among sites. Estimated population sizes projected in 100 years diverged up to 1 order of magnitude among scenarios that assumed different movement behavior. Scenarios also exhibited different model dynamics that ranged from stable equilibria to cycles or extinction. These results suggest that inclusion of behavioral data in viability models may improve estimates of extinction risk for imperiled species. Our approach provides a simple method for incorporating spatial and demographic consequences of behavioral strategies into population models and may be easily extended to other species and behaviors to understand the mechanisms of population dynamics for imperiled populations.     

Using Modeling to Improve Monitoring of Structured Populations: Are We Collecting the Right Data?

Abstract:  Population monitoring is central to most demographic studies and conservation efforts, but it may not always be directed at the most appropriate life stage. We used stochastic simulation modeling to evaluate the effectiveness of a monitoring program for a well-studied population of Eastern Imperial Eagles ( Aquila heliaca ) in Kazakhstan. Specifically, we asked whether the most appropriate data were being collected to understand system state and population dynamics. Our models were parameterized with data collected over the course of 25 years of study of this population. We used the models to conduct simulation experiments to evaluate relationships between monitored or potentially monitored parameters and the demographic variables of interest—population size ( N ) and population growth (λ). Static analyses showed that traditional territory-based monitoring was a poor indicator of eagle population size and growth and that monitoring survivorship would provide more information about these parameters. Nevertheless, these same traditionally monitored territory-based parameters had greater power to detect long-term changes in population size than did survivorship or population structure. Regardless of the taxa considered, threats can have immediate impacts on population size and growth or longer-term impacts on population dynamics. Prudently designed monitoring programs for any species will detect the demographic effects of both types of threats.     

The impact of parasite manipulation and predator foraging behavior on predator-prey communities

Parasites are known to directly affect their hosts at both the individual and population level. However, little is known about their more subtle, indirect effects and how these may affect population and community dynamics. In particular, trophically transmitted parasites may manipulate the behavior of intermediate hosts, fundamentally altering the pattern of contact between these individuals and their predators. Here, we develop a suite of population dynamic models to explore the impact of such behavioral modifications on the dynamics and structure of the predator-prey community. We show that, although such manipulations do not directly affect the persistence of the predator and prey populations, they can greatly alter the quantitative dynamics of the community, potentially resulting in high amplitude oscillations in abundance. We show that the precise impact of host manipulation depends greatly on the predator's functional response, which describes the predator's foraging efficiency under changing prey availabilities. Even if the parasite is rarely observed within the prey population, such manipulations extend beyond the direct impact on the intermediate host to affect the foraging success of the predator, with profound implications for the structure and stability of the predator-prey community.     

Sensitivity of plant-pollinator-herbivore communities to changes in phenology

For many species in seasonal environments, warmer springs associated with anthropogenic climate change are causing phenological changes. Within ecological communities, the timing of interactions among species may be altered if the species experience asymmetrical phenological shifts. We present a model that examines the consequences of changes in the relative timing of herbivory and pollination in a community of herbivores and pollinators that share a host plant. Our model suggests that phenological shifts can alter the abundances of these species and, in some cases, their population dynamics. If historical patterns of interactions in a community change and herbivores become active before pollinators, the community could see a reduction in pollinators and an increase in herbivores, while if pollinators become active before herbivores, there could be a loss of stable coexistence. Previous studies have warned of the potential for climate change to cause large phenological mismatches whereby species that depend on one another become so separated in time that they can no longer interact. Our results suggest that climate change-induced phenological shifts can have a major impact on communities even in cases where complete phenological mismatches do not occur.     

Information processing in models for density-dependent emigration: A comparison

Density-dependent emigration has been recognized as a fitness enhancing strategy. Yet, especially in the modelling literature there is no consensus about how density-dependent emigration should quantitatively be incorporated into metapopulation models. In this paper we compare the performance of five different dispersal strategies (defined by the functional link between density and emigration probability). Four of these strategies are based on published functional relationships between local population density and emigration probability, one assumes density-independent dispersal. We use individual-based simulations of time-discrete metapopulation dynamics and conduct evolution experiments for a broad range of values for dispersal mortality and environmental stochasticity. For each set of these conditions we analyze the evolution of emigration rates in ‘monoculture experiments’ (with only one type of dispersal strategy used by all individuals in the metapopulation) as well as in selection experiments that allow a pair-wise comparison of the performance of each functional type. We find that a single-parameter ‘asymptotic threshold’ strategy - derived from the marginal value theorem - with a decelerating increase of emigration rate with increasing population density, out-competes any other strategy, i.e. density-independent emigration, a ‘linear threshold’ strategy and a flexible three-parameter strategy. Only when environmental conditions select for extremely high emigration probabilities (close to one), strategies may perform approximately equally. A simple threshold strategy derived for the case of continuous population growth performs even worse than the density-independent strategy. As the functional type of the dispersal function implemented in metapopulation models may severely affect predictions concerning the survival of populations, range expansion, or community changes we clearly recommend to carefully select adequate functions to model density-dependent dispersal.     

Use of stochastic patch occupancy models in the California red-legged frog for Bayesian inference regarding past events and future persistence

Assessing causes of population decline is critically important to management of threatened species. Stochastic patch occupancy models (SPOMs) are popular tools for examining spatial and temporal dynamics of populations when presence–absence data in multiple habitat patches are available. We developed a Bayesian Markov chain method that extends existing SPOMs by focusing on past environmental changes that may have altered occupancy patterns prior to the beginning of data collection. Using occupancy data from 3 creeks, we applied the method to assess 2 hypothesized causes of population decline—in situ die-off and residual impact of past source population loss—in the California red-legged frog. Despite having no data for the 20–30 years between the hypothetical event leading to population decline and the first data collected, we were able to discriminate among hypotheses, finding evidence that in situ die-off increased in 2 of the creeks. Although the creeks had comparable numbers of occupied segments, owing to different extinction–colonization dynamics, our model predicted an 8-fold difference in persistence probabilities of their populations to 2030. Adding a source population led to a greater predicted persistence probability than did decreasing the in situ die-off, emphasizing that reversing the deleterious impacts of a disturbance may not be the most efficient management strategy. We expect our method will be useful for studying dynamics and evaluating management strategies of many species.     

Running to stand still: temperature effects on species richness, species turnover, and functional community dynamics

The information on temperature-mediated changes in biodiversity in local assemblages is scarce and mainly addresses the change in species richness. However, warming may have more consistent effects on species turnover than on the number of species. Moreover, very few studies extended the analysis of changes in biodiversity and species composition to questions of associated ecosystem functions such as primary production. Here, we synthesize 4 case studies employing microalgal microcosms within the Aquashift priority program to ask (1) do warming-related shifts in species richness correspond to changes in the rate of biomass production, (2) do similar relationships prevail for evenness, and (3) do warming-related shifts in species turnover stabilize or destabilize biomass production? Two of the four cases are previously unpublished, and for a third case, the link between diversity and functional consequences of temperature was not analyzed before. We found accelerated loss of species with warming in all cases. Biomass production was lower with lower species richness in most cases but increased with lower evenness. Most importantly, the relation between functional and compositional stability was different between cases: More rapid extinction resulted in more variable biomass in 2 cases conducted with a limited species pool, indicating that compositional destabilization relates to functional variability. By contrast, the only experiment with a large species pool (30 species) allowed previously rare species to become dominant in the community and showed more stable biomass at high turnover, indicating that compensatory dynamics (turnover) can promote functional stability. These 4 independent experiments highlight the need to consider both compositional and functional consequences of altered temperature regimes.