Do source-sink dynamics promote the spread of an invasive grass into a novel habitat?

Models of source-sink and other spatial patch dynamics have generated a number of ideas and predictions about species range expansion, the evolution of local adaptation, and the factors influencing population persistence, but relatively few empirical studies have applied these ideas due to the difficulty of measuring both patch-specific demography and movement rates. In this study, I used a combination of mark-recapture experiments, model fitting, and demographic approaches to ask how habitat-specific differences in population growth and dispersal affect spread of the invasive grass Aegilops triuncialis into serpentine environments. A. triuncialis germinated at lower rates but exhibited equivalent survival and greater growth in edge (extreme serpentine) than in core populations, even accounting for density differences between habitats. Estimated growth rates (lambda) for four of five edge subpopulations were strongly positive, ranging from lambda = 1.32 to 2.09 without propagule input from adjacent habitat. Local dispersal was best described by an exponential kernel, with a mean dispersal distance about twice as long on the edge (0.24-0.40 m) as in the core (0.18 m). Twenty-five percent of marked spikes in the edge were not relocated within the patch, suggesting greater rates of either seed predation or long-distance dispersal that reduced population growth. These results suggest that A. triuncialis can successfully spread into extreme serpentine habitats without sustained propagule input from adjacent populations. Further, asymmetric dispersal that may be both habitat- and density-dependent could slow growth rates on the edge. This pattern may also increase the importance of harsh edge patches as a source of long-distance dispersers.     

A multispecies test of source–sink indicators to prioritize habitat for declining populations

For species at risk of decline or extinction in source–sink systems, sources are an obvious target for habitat protection actions. However, the way in which source habitats are identified and prioritized can reduce the effectiveness of conservation actions. Although sources and sinks are conceptually defined using both demographic and movement criteria, simplifications are often required in systems with limited data. To assess the conservation outcomes of alternative source metrics and resulting prioritizations, we simulated population dynamics and extinction risk for 3 endangered species. Using empirically based habitat population models, we linked habitat maps with measured site‐ or habitat‐specific demographic conditions, movement abilities, and behaviors. We calculated source–sink metrics over a range of periods of data collection and prioritized consistently high‐output sources for conservation. We then tested whether prioritized patches identified the habitats that most affected persistence by removing them and measuring the population response. Conservation decisions based on different source–sink metrics and durations of data collection affected species persistence. Shorter time series obscured the ability of metrics to identify influential habitats, particularly in temporally variable and slowly declining populations. Data‐rich source–sink metrics that included both demography and movement information did not always identify the habitats with the greatest influence on extinction risk. In some declining populations, patch abundance better predicted influential habitats for short‐term regional persistence. Because source–sink metrics (i.e., births minus deaths; births and immigrations minus deaths and emigration) describe net population conditions and cancel out gross population counts, they may not adequately identify influential habitats in declining populations. For many nonequilibrium populations, new metrics that maintain the counts of individual births, deaths, and movement may provide additional insight into habitats that most influence persistence.     

Conservation of Species in Dynamic Landscapes: Divergent Fates of Silene tatarica Populations in Riparian Habitats

Abstract:  In transient environments, where local extinctions occur as a result of destruction or deterioration of the local habitat, the long-term persistence of a species requires successful colonizations at new, suitable sites. This kind of habitat tracking should be associated with the asynchronous dynamics of local populations, and it can be especially important for the conservation of rare plant species in riparian habitats. We determined spatiotemporal variation in the demography of the perennial Silene tatarica (L.) Pers. in 15 populations (1998–2003) located in periodically disturbed riparian habitats. The habitats differed according to their morphology (flat shores, slopes) and the amount of bare ground (open, intermediate, closed) along a successional gradient. We used elasticity and life-table response analyses and stochastic simulations to study the variation in population demography. Finite population growth rate was higher in intermediate habitats than in open and closed habitats. In stochastic simulations population size increased in most cases, but four populations were projected to become extinct within 12–70 years. The viability of local populations depended most on the survival and growth of juvenile individuals and on the fecundity of large fertile individuals. On a regional scale, the persistence of this species will require a viable network of local populations as protection against local extinctions caused by natural disturbances and succession. Accordingly, the long-term persistence of riparian species may depend on habitat changes; thus, their conservation requires maintenance of natural disturbance dynamics. Along regulated rivers, management activities such as the creation of open habitats for new colonization should be implemented. Similarly, these activities can be rather general requirements for the conservation of endangered species dependent on transient habitats along successional gradients.     

Meso-scale hydrodynamic and reproductive asynchrony affects the source–sink metapopulation structure of the coastal polychaete Pectinaria koreni

The polychaete Pectinaria koreni exhibits a complex life cycle characterized by non-overlapping generations and widespread larval dispersal. To explore how “local” metapopulation genetic structure varies spatially and temporally during population turnover, we combined observations on demography, larval dispersal through hydrodynamic modelling and population genetics of successive age cohorts in the Baie de Seine (eastern English Channel, France). Mature adults (March), newly settled (July) and later-stage juveniles (September) were sampled in 2003 on the edge and in the main demes of the metapopulation. Demes displayed an asynchronous dynamics due to variations in habitat quality affecting reproductive timing (e.g. three distinct spawning events observed) and in local larval supply linked to temporal fluctuations of hydrodynamism. Two-source populations were identified among dense areas with the greatest larval retention and self-recruitment rates: one with a single recruitment event, stable temporal genetic variation and a strong spatial genetic re-homogenization during turnover, and the other with two recruitment events and significant allele frequency changes over time. Sink habitats displayed single recruitment event and experienced strong inter-generational (juveniles vs adults) genetic changes due to genetic drift associated with strong winter mortalities. Altogether, results suggested that adult spawning asynchrony and sweepstakes reproductive success, together with genetic drift, played a greater role than hydrodynamics itself in determining effective recruitment rates at some sites and generating genetic patchiness.     

The influence of disturbance events on survival and dispersal rates of Florida box turtles.

Disturbances have the potential to cause long-term effects to ecosystem structure and function, and they may affect individual species in different ways. Long-lived vertebrates such as turtles may be at risk from such events, inasmuch as their life histories preclude rapid recovery should extensive mortality occur. We applied capture-mark-recapture models to assess disturbance effects on a population of Florida box turtles (Terrapene carolina bauri) on Egmont Key, Florida, USA. Near the midpoint of the study, a series of physical disturbances affected the island, from salt water overwash associated with several tropical storms to extensive removal of nonindigenous vegetation. These disturbances allowed us to examine demographic responses of the turtle population and to determine if they affected dispersal throughout the island. Adult survival rates did not vary significantly either between sexes or among years of the study. Survival rates did not vary significantly between juvenile and adult turtles, or among years of the study. Furthermore, neither adult nor juvenile survival rates differed significantly between pre- and post-disturbance. However, dispersal rates varied significantly among the four major study sites, and dispersal rates were higher during the pre-disturbance sampling periods compared to post-disturbance. Our results suggest few long-term effects on the demography of the turtle population. Florida box turtles responded to tropical storms and vegetation control by moving to favorable habitats minimally affected by the disturbances and remaining there. As long as turtles and perhaps other long-lived vertebrates can disperse to non-disturbed habitat, and high levels of mortality do not occur in a population, a long life span may allow them to wait out the impact of disturbance with potentially little effect on long-term population processes.     

Effects of Habitat Loss and Fragmentation on the Behavior and Demography of Gray-Tailed Voles

We monitored the short term behavioral and demographic responses of gray-tailed voles (Microtus canicaudus) to the reduction and fragmentation of their habitat. Our objectives were (1) to test whether animals perished or moved into remaining fragments after 70% of their habitat was removed; and (2) to test the null hypothesis that the social structure and demography of animals would not differ between habitats consisting of one large continuous fragment (625 m²), a mosaic of 25 small fragments (each 25 m²) separated by 4 m of bare ground, and control, unmanipulated habitats (1850 m²). We conducted the experiment in 12, 0.2-ha enclosures planted with alfalfa with four replicates for each of two manipulated treatments and a control. A 70% reduction in habitat did not adversely affect adult survival, reproductive rate, juvenile recruitment, or population size. However, an influx of unrelated females into habitat fragments resulted in decreased juvenile recruitment in those fragments. Voles from cleared habitat moved into the remaining habitat and did not measurably affect the resident population. Similarly, the demography of voles did not differ significantly among the large-fragment, small-fragment, and control enclosures. Peak density estimates based on the amount of habitat in each enclosure were 545 animals per hectare in control, 1056 in large-fragment, and 2880 in small-fragment enclosures. Reduced movement of animals among the small fragments was the most obvious effect of habitat fragmentation. Six percent of females and 15% of males moved among small fragments within a week compared to approximately 60% moving comparable distances in large-fragment and control enclosures. Rates of juvenile dispersal and sexual maturation declined throughout the summer on all treatments, were associated with season and density, and were only marginally associated with habitat loss and fragmentation. We conclude that at the time of habitat removal and fragmentation, populations were small enough to accommodate a 70% reduction in habitat and still continue to increase in numbers. The social system of gray-tailed voles was sufficiently flexible to accommodate an influx of animals to withstand densities> 1000 voles per ha. The behavioral and demographic features of gray-tailed voles are similar to those reported for other small mammals, thus confirming the use of voles for ecological model systems in habitat fragmentation studies.     

Conserving Slow-Growing, Long-Lived Tree Species: Input from the Demography of a Rare Understory Conifer, Taxus floridana

Abstract:  Although land preservation and promotion of successful regeneration are important conservation actions, their ability to increase population growth rates of slow-growing, long-lived trees is limited. We investigated the demography of Taxus floridana Nutt., a rare understory conifer, in three populations in different ravine forests spanning its entire geographic range along the Apalachicola River Bluffs in northern Florida (U.S.A.). We examined spatial and temporal patterns in demographic parameters and projected population growth rates by using four years of data on the recruitment and survival of seedlings and established stems, and on diameter growth from cross-sections of dead stems. All populations experienced a roughly 10-fold increase in seedling recruitment in 1996 compared with other years. The fates of seedlings and stems between 8 and 16 mm differed among populations. The fates of stems in two other size classes (the 2- to 4-mm class and the 4- to 8-mm class) differed among both populations and years. Individual stems in all populations exhibited similarly slow growth rates. Stochastic matrix models projected declines in all populations. Stochastic matrix analysis revealed the high elasticity of a measure of stochastic population growth rate to perturbations in the stasis of large reproductive stems for all populations. Additional analyses also indicated that occasional episodes of high recruitment do not greatly affect population growth rates. Conservation efforts directed at long-lived, slow-growing rare plants like Taxus floridana should both protect established reproductive individuals and further enhance survival of individuals in other life-history stages, such as juveniles, that often do not appear to contribute greatly to population growth rates.     

Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

Habitat complexity modifies post-settlement mortality and recruitment dynamics of a marine fish

For species that have an open population structure, local population size may be strongly influenced by a combination of propagule supply and post-settlement survival. While it is widely recognized that supply of larvae (or recruits) is variable and that variable recruitment may affect the relative contribution of pre- and post-settlement factors, less effort has been made to quantify how variation in the strength of post-settlement mortality (particularly density-dependent mortality) will affect the importance of processes that determine population size. In this study, I examined the effects of habitat complexity on mortality of blue rockfish (Sebastes mystinus) within nearshore reefs off central California. I first tested whether variation in habitat complexity (measured as three-dimensional complexity of rocky substrate) affected the magnitude of both density-independent and density-dependent mortality. I then used limitation analysis to quantify how variation in habitat complexity alters the relative influence of recruitment, density-independent mortality, and density-dependent mortality in determining local population size. Increased habitat complexity was associated with a reduction in both density-independent and density-dependent mortality. At low levels of habitat complexity, limitation analysis revealed that mortality was strong and recruitment had relatively little influence on population size. However, as habitat complexity increased, recruitment became more important. At the highest levels of habitat complexity, limitation by recruitment was substantial, although density-dependent mortality was ultimately the largest constraint on population size. In high-complexity habitats, population dynamics may strongly reflect variation in recruitment even though fluctuations may be dampened by density-dependent mortality. By affecting both density-independent and density-dependent mortality, variation in habitat complexity may result in qualitative changes in the dynamics of populations. These findings suggest that the relative importance of pre- vs. post-settlement factors may be determined by quantifiable habitat features, rather than ambient recruitment level alone. Because the magnitude of recruitment fluctuations can affect species coexistence and the persistence of populations, habitat-driven changes in population dynamics may have important consequences for both community structure and population viability.     

Importance of Habitat Quality and Landscape Connectivity for the Persistence of Endangered Natterjack Toads

Abstract: The natterjack toad (Bufo calamita) is endangered in several parts of its distribution, including Belgium, where it occurs mainly in artificial habitats. We parameterized a general model for natterjack population viability analysis (PVA) and tested its sensitivity to changes in the values of basic parameters. Then we assessed the relative efficiency of various conservation measures in 2 situations: a small isolated population and a system of 4 populations connected by rare dispersal movements. We based the population viability analysis on a stage‐structured model of natterjack population dynamics. We parameterized the model in the RAMAS GIS platform with vital rates obtained from our own field experience and from published studies. Simulated natterjack populations were highly sensitive to habitat quality (particularly pond drying), to dispersal from surrounding local populations, and to a lesser extent to values of fecundity and survival of terrestrial stages. Population trajectories were nearly insensitive to initial abundances, carrying capacities, and the frequency of extreme climatic conditions. The simulations showed that in habitats with highly ephemeral ponds, where premetamorphosis mortality was high, natterjack populations nearly always had a very high extinction risk. We also illustrated how low dispersal rates (<1 dispersing individual/generation) efficiently rescued declining local populations. Such source‐sink dynamics demonstrate that the identification and management of source populations should be a high priority.     

Evaluating the importance of demographic connectivity in a marine metapopulation

Recently researchers have gone to great lengths to measure marine metapopulation connectivity via tagging, genetic, and trace-elemental fingerprinting studies. These empirical estimates of larval dispersal are key to assessing the significance of metapopulation connectivity within a demographic context, but the life-history data required to do this are rarely available. To evaluate the demographic consequences of connectivity we constructed seasonal, size-structured metapopulation matrix models for two species of mytilid mussel in San Diego County, California, USA. The self-recruitment and larval exchange terms were produced from a time series of realized connectivities derived from trace-elemental fingerprinting of larval shells during spring and fall from 2003 to 2008. Both species exhibited a strong seasonal pattern of southward movement of recruits in spring and northward movement in fall. Growth and mortality terms were estimated using mark-recapture data from representative sites for each species and subpopulation, and literature estimates of juvenile mortality. Fecundity terms were estimated using county-wide settlement data from 2006-2008; these data reveal peak reproduction and recruitment in fall for Mytilus californianus, and spring for M. galloprovincialis. Elasticity and life-stage simulation analyses were employed to identify the season- and subpopulation-specific vital rates and connectivity terms to which the metapopulation growth rate (lambda) was most sensitive. For both species, metapopulation growth was most sensitive to proportional changes in adult fecundity, survival and growth of juvenile stages, and population connectivity, in order of importance, but relatively insensitive to adult growth or survival. The metapopulation concept was deemed appropriate for both Mytilus species as exchange between the subpopulations was necessary for subpopulation persistence. However, highest metapopulation growth occurred in years when a greater proportion of recruits was retained within the predominant source subpopulation. Despite differences in habitat and planktonic duration, both species exhibited similar overall metapopulation dynamics with respect to key life stages and processes. However, different peak reproductive periods in an environment of seasonal current reversals led to different regional (subpopulation) contributions to metapopulation maintenance; this result emphasizes the importance of connectivity analysis for spatial management of coastal resources.     

The transition from isolated patches to a metapopulation in the eastern collared lizard in response to prescribed fires

Habitat fragmentation often arises from human-induced alterations to the matrix that reduce or eliminate dispersal between habitat patches. Elimination of dispersal increases local extinction and decreases recolonization. These phenomena were observed in the eastern collared lizard (Crotaphytus collaris collaris), which lives in the mid-continental highland region of the Ozarks (Missouri, USA) on glades: habitats of exposed bedrock that form desert-like habitats imbedded in a woodland matrix. With the onset of woodland fire suppression, glade habitats degenerated and the woodland matrix was altered to create a strong barrier to dispersal. By 1980, lizard populations in the Ozarks were rapidly going extinct. In response to this decline, some glades were restored by clearing and burning. Starting in 1984, collared lizard populations were translocated onto these restored habitats. The translocated populations persisted but did not colonize nearby glades or disperse among one another. In 1994 prescribed woodland fires were initiated, which unleashed much dispersal and colonizing behavior. Dispersal was highly nonrandom by both intrinsic variables (age, gender) and extrinsic variables (overall demography, glade population sizes, glade areas, landscape features), resulting in different classes of lizards being dominant in creating demographic cohesiveness among glades, colonizing new glades on a mountain, and colonizing new mountain systems. A dramatic transition was documented from isolated fragments, to a nonequilibrium colonizing metapopulation, and finally to a stable metapopulation. This transition is characterized by the convergence of rates of extinction and recolonization and a major alteration of dispersal probabilities and pattern in going from the nonequilibrium to stable metapopulation states.     

Risk spreading, connectivity, and optimal reserve spacing

Two important processes determining the dynamics of spatially structured populations are dispersal and the spatial covariance of demographic fluctuations. Spatially explicit approaches to conservation, such as reserve networks, must consider the tension between these two processes and reach a balance between distances near enough to maintain connectivity, but far enough to benefit from risk spreading. Here, we model this trade-off. We show how two measures of metapopulation persistence depend on the shape of the dispersal kernel and the shape of the distance decay in demographic covariance, and we consider the implications of this trade-off for reserve spacing. The relative rates of distance decay in dispersal and demographic covariance determine whether the long-run metapopulation growth rate, and quasi-extinction risk, peak for adjacent patches or intermediately spaced patches; two local maxima in metapopulation persistence are also possible. When dispersal itself fluctuates over time, the trade-off changes. Temporal variation in mean distance that propagules are dispersed (i.e., propagule advection) decreases metapopulation persistence and decreases the likelihood that persistence will peak for adjacent patches. Conversely, variation in diffusion (the extent of random spread around mean dispersal) increases metapopulation persistence overall and causes it to peak at shorter inter-patch distances. Thus, failure to consider temporal variation in dispersal processes increases the risk that reserve spacings will fail to meet the objective of ensuring metapopulation persistence. This study identifies two phenomena that receive relatively little attention in empirical work on reserve spacing, but that can qualitatively change the effectiveness of reserve spacing strategies: (1) the functional form of the distance decay in covariance among patch-specific demographic rates and (2) temporal variation in the shape of the dispersal kernel. The sensitivity of metapopulation recovery and persistence to how covariance of vital rates decreases with distance suggests that estimating the shape of this function is likely to be as important for effective reserve design as estimating connectivity. Similarly, because temporal variation in dispersal dynamics influences the effect of reserve spacing, approaches to reserve design that ignore such variation, and rely instead on long-term average dispersal patterns, are likely to lead to lower metapopulation viability than is actually achievable.     

Landscape Patterns of Florida Scrub Jay Habitat Use and Demographic Success

Remote sensing and geographical information systems are used to analyze landscapes important to species of conservation concern. The accuracy of the methods depends on how closely habitat mapping classes are linked to population demography. Habitat use by Florida Scrub Jays ( Aphelocoma c. coemlescens ) was quantified using circular plots. Habitat variation was mapped using high-resolution aerial photography on a site where all Florida Scrub Jays were color-banded. Nest site selection, nest success, yearling production, and breeder survival were measured within Florida Scrub Jay territories. Habitat use was lowest in areas without scrub oaks or areas within 136 m from forests. Open oak, dominated by scrub oaks and open sandy areas, had the highest use and nest success among habitats. Open oak occurred as narrow patches ( <20 m wide) in landscapes dominated by matrix habitat (palmetto-lyonia and swale marshes). Most wide patches (>50 m) of open oak were potential population sources, where reproduction exceeded mortality. Areas with patches of open oak of less than 1 ha were usually population sinks. Open oak occurred as less flammable patches in a landscape subject to frequent fires. Population sources varied temporally and spatially with fires and site potential to support scrub oaks (soils). Analyses of landscape patterns and dynamics indicated that habitat mapping should not only include patches of currently optimal habitat but should also include landscapes associated with open oak. The influences of landscape patterns on habitat use, reproductive success, survival, and territory size can be quantified at different scales starting with attributes associated with habitat patches, nest sites, and territories. Potential mapping errors occur, however, when habitats are used to quantify the areal extent of sources and sinks and similar population attributes important for species persistence.     

Demographic Side Effects of Selective Hunting in Ungulates and Carnivores

Abstract:  Selective harvesting regimes are often implemented because age and sex classes contribute differently to population dynamics and hunters show preferences associated with body size and trophy value. We reviewed the literature on how such cropping regimes affect the demography of the remaining population (here termed demographic side effects ). First, we examined the implications of removing a large proportion of a specific age or sex class. Such harvesting strategies often bias the population sex ratio toward females and reduce the mean age of males, which may consequently delay birth dates, reduce birth synchrony, delay body mass development, and alter offspring sex ratios. Second, we reviewed the side effects associated with the selective removal of relatively few specific individuals, often large trophy males. Such selective harvesting can destabilize social structures and the dominance hierarchy and may cause loss of social knowledge, sexually selected infanticide, habitat changes among reproductive females, and changes in offspring sex ratio. A common feature of many of the reported mechanisms is that they ultimately depress recruitment and in some extreme cases even cause total reproductive collapse. These effects could act additively and destabilize the dynamics of populations, thus having a stronger effect on population growth rate than first anticipated. Although more experimental than observational studies reported demographic side effects, we argue that this may reflect the quite subtle mechanisms involved, which are unlikely to be detected in observational studies without rigorous monitoring regimes. We call for more detailed studies of hunted populations with marked individuals that address how the expression of these effects varies across mating systems, habitats, and with population density. Theoretical models investigating how strongly these effects influence population growth rates are also required.     

Demographic Responses by Birds to Forest Fragmentation

Abstract:  Despite intensive recent research on the effects of habitat loss and fragmentation on bird populations, our understanding of underlying demographic causes of population declines is limited. We reviewed avian demography in relation to habitat fragmentation. Then, through a meta-analysis, we compared specific demographic responses by forest birds to habitat fragmentation, providing a general perspective of factors that make some species and populations more vulnerable to fragmentation than others. We obtained data from the scientific literature on dispersal, survival, fecundity, and nesting success of birds. Birds were divided into subgroups on the basis of region, nest site, biogeographical history, and migration strategy. Species most sensitive to fragmentation were ground- or open-nesters nesting in shrubs or trees. Residents were equally sensitive to fragmentation in the Nearctic and Palearctic regions, but Nearctic migrants were more sensitive than Palearctic migrants. Old World species were less sensitive than New World species, which was predicted based on the history of forest fragmentation on these two continents. Pairing success was the variable most associated with fragmentation, suggesting an important role of dispersal. Fledgling number or condition, timing of nesting, and clutch size were not associated with sensitivity to fragmentation, suggesting that negative fragmentation effects on birds do not generally result from diminished food resources with increasing level of fragmentation. Future studies on demographic responses of birds to habitat fragmentation would be more effective if based on a combination of measures that can distinguish among the demographic mechanisms underlying population changes related to habitat fragmentation.     

Patch Size and Connectivity Thresholds for Butterfly Habitat Restoration

Abstract:  Recovery of endangered species in highly fragmented habitats often requires habitat restoration. Selection of restoration sites typically involves too many options and too much uncertainty to reach a decision based on existing reserve design methods. The Fender's blue butterfly (  Icaricia icarioides fenderi ) survives in small, isolated patches of remnant prairie in Oregon's Willamette Valley—a habitat for which <0.5% of the original remains. Recovery of this species will require considerable habitat restoration. We investigated the potential of biologically based rules of thumb and more complex models to serve as tools in making land acquisitions. Based on Fender's blue dispersal behavior and demography, we have estimated that restored patches should be <1 km from existing habitat and at least 2 ha. We compared these rules to the results of two modeling approaches: an incidence function model and a spatially explicit simulation of demography and dispersal behavior. Not surprisingly, the simple rules and complex models all conclude that large (>2 ha) connected (<1 km) patches have the highest restoration value. The dispersal model, however, suggests that small, connected patches have more restoration value than large, isolated patches, whereas the incidence function model suggests that size and connectivity are equally important. These differences stem from model assumptions. We used incidence functions to predict long-term, stochastic, steady-state conditions and dispersal simulations to predict short-term (25-year) colonization dynamics. To apply our results in the context of selecting restoration sites on the ground, we recommend selecting nearby sites when short-term colonization dynamics are expected to be an important aspect of a species' biology.     

An experimental study of habitat selection by birds in a coffee plantation

Unique components of tropical habitats, such as abundant vascular epiphytes, influence the distribution of species and can contribute to the high diversity of many animal groups in the tropics. However, the role of such features in habitat selection and demography of individual species has not been established. Understanding the mechanisms of habitat selection requires both experimental manipulation of habitat structure and detailed estimation of the behavioral and demographic response of animals, e.g., changes in movement patterns and survival probabilities. Such studies have not been conducted in natural tropical forest, perhaps because of high habitat heterogeneity, high species diversity, and low abundances of potential target species. Agroforestry systems support a less diverse flora, with greater spatial homogeneity which, in turn, harbors lower overall species diversity with greater numerical dominance of common species, than natural forests. Furthermore, agroforestry systems are already extensively managed and lend themselves easily to larger scale habitat manipulations than protected natural forest. Thus, agroforestry systems provide a good model environment for beginning to understand processes underlying habitat selection in tropical forest animals. Here, we use multistate, capture-recapture models to investigate how the experimental removal of epiphytes affected monthly movement and survival probabilities of two resident bird species (Common Bush-Tanager [Chlorospingus ophthalmicus] and Golden-crowned Warbler [Basileuterus culicivorus]) in a Mexican shade coffee plantation. We established two paired plots of epiphyte removal and control. We found that Bush-Tanagers were at least five times more likely to emigrate from plots where epiphytes were removed compared to control plots. Habitat-specific movement patterns were not detected in the warbler. However, unlike the Golden-crowned Warbler, Common Bush-Tanagers depend upon epiphytes for nest sites and (seasonally) for foraging. These dispersal patterns imply that active habitat selection based on the presence or absence of epiphytes occurs in C. ophthalmicus on our study area. Survival rates did not vary with habitat in either species. Interestingly, in both species, survival was higher in the nonbreeding season, when birds were in mixed-species flocks. Movement by Common Bush-Tanagers into areas with epiphytes occurred mostly during the breeding season, when mortality-driven opportunity was greatest.     

Using demography and movement behavior to predict range expansion of the southern sea otter

In addition to forecasting population growth, basic demographic data combined with movement data provide a means for predicting rates of range expansion. Quantitative models of range expansion have rarely been applied to large vertebrates, although such tools could be useful for restoration and management of many threatened but recovering populations. Using the southern sea otter (Enhydra lutris nereis) as a case study, we utilized integro-difference equations in combination with a stage-structured projection matrix that incorporated spatial variation in dispersal and demography to make forecasts of population recovery and range recolonization. In addition to these basic predictions, we emphasize how to make these modeling predictions useful in a management context through the inclusion of parameter uncertainty and sensitivity analysis. Our models resulted in hind-cast (1989-2003) predictions of net population growth and range expansion that closely matched observed patterns. We next made projections of future range expansion and population growth, incorporating uncertainty in all model parameters, and explored the sensitivity of model predictions to variation in spatially explicit survival and dispersal rates. The predicted rate of southward range expansion (median = 5.2 km/yr) was sensitive to both dispersal and survival rates; elasticity analysis indicated that changes in adult survival would have the greatest potential effect on the rate of range expansion, while perturbation analysis showed that variation in subadult dispersal contributed most to variance in model predictions. Variation in survival and dispersal of females at the south end of the range contributed most of the variance in predicted southward range expansion. Our approach provides guidance for the acquisition of further data and a means of forecasting the consequence of specific management actions. Similar methods could aid in the management of other recovering populations.     

A model for behavioral regulation of metapopulation dynamics

Habitat fragmentation can decrease local population persistence by reducing connectivity, which is a function of dispersal of individuals among habitat fragments. Dispersal is often treated as diffusion in population models, even though for many species it is a result of a series of behavioral decisions. We developed a metapopulation model to explore the potential importance of dispersal behaviors in driving metapopulation dynamics. We incorporated types of behavior that affect dispersal—colonization inhibiting, colonization enhancing, extinction inhibiting, extinction enhancing, rescue enhancing, rescue inhibiting—into Levins’ (1969) metapopulation model and projected occupancy rates for a variety of parameter values. Examples from the literature of behaviors associated with each of these parameters are provided. Our model simplifies into previously published metapopulation models that incorporate only a single behavior, and we present a density-dependent rescue function that leads to multiple non-zero equilibria. We found a variety of behavioral effects on metapopulations. Rescue enhancement fills patches faster than does colonization enhancement or extinction inhibition, and declines in patch occupancy are moderate with extinction enhancement, but colonization inhibition causes metapopulation extinction. We also found that with colonization and extinction inhibitions, equilibrium patch occupancy is inversely related to patch turnover rate. With density-dependent rescue, persistence depends not only on the strength of the strong rescue effect, but also on having a sufficient initial fraction of patches occupied; the stronger the rescue effect, the lower this fraction can be. This study suggests that dispersal behavior can have strong influences on metapopulation dynamics. It confirms the importance of understanding the relationship between landscape structure and dispersal behavior in understanding population persistence.