Influence of habitat quality, population size, patch size, and connectivity on patch-occupancy dynamics of the middle spotted woodpecker

Despite extensive research on the effects of habitat fragmentation, the ecological mechanisms underlying colonization and extinction processes are poorly known, but knowledge of these mechanisms is essential to understanding the distribution and persistence of populations in fragmented habitats. We examined these mechanisms through multiseason occupancy models that elucidated patch-occupancy dynamics of Middle Spotted Woodpeckers (Dendrocopos medius) in northwestern Spain. The number of occupied patches was relatively stable from 2000 to 2010 (15-24% of 101 patches occupied every year) because extinction was balanced by recolonization. Larger and higher quality patches (i.e., higher density of oaks >37 cm dbh [diameter at breast height]) were more likely to be occupied. Habitat quality (i.e., density of large oaks) explained more variation in patch colonization and extinction than did patch size and connectivity, which were both weakly associated with probabilities of turnover. Patches of higher quality were more likely to be colonized than patches of lower quality. Populations in high-quality patches were less likely to become extinct. In addition, extinction in a patch was strongly associated with local population size but not with patch size, which means the latter may not be a good surrogate of population size in assessments of extinction probability. Our results suggest that habitat quality may be a primary driver of patch-occupancy dynamics and may increase the accuracy of models of population survival. We encourage comparisons of competing models that assess occupancy, colonization, and extinction probabilities in a single analytical framework (e.g., dynamic occupancy models) so as to shed light on the association of habitat quality and patch geometry with colonization and extinction processes in different settings and species.     

Effectiveness of Corridors Relative to Enlargement of Habitat Patches

Abstract:  The establishment of biological corridors between two otherwise isolated habitat patches is a common yet contentious strategy for conserving populations in fragmented landscapes. We compared the effectiveness of corridors with the effectiveness of an alternate conservation strategy, the enlargement of existing habitat patches. We used a spatially explicit population model that simulated population size in two kinds of patches. One patch had a corridor that connected it to a larger "source" patch and the other patch was unconnected and enlarged at the periphery by an area the same size as the corridor. Patch isolation, corridor width, patch size, and the probability that individuals would cross the border from habitat to matrix were varied independently. In general, population size was greater in enlarged patches than in connected patches when patches were relatively large and isolated. Corridor width and the probability of crossing the border from habitat to matrix did not affect the relative benefit of corridors versus patch enlargement. Although biological corridors may mitigate potential effects of inbreeding depression at long time scales, our results suggest that they are not always the best method of conserving fragmented populations.     

Obstruction of biodiversity conservation by minimum patch size criteria

Minimum patch size criteria for habitat protection reflect the conservation principle that a single large (SL) patch of habitat has higher biodiversity than several small (SS) patches of the same total area (SL > SS). Nonetheless, this principle is often incorrect, and biodiversity conservation requires placing more emphasis on protection of large numbers of small patches (SS > SL). We used a global database reporting the abundances of species across hundreds of patches to assess the SL > SS principle in systems where small patches are much smaller than the typical minimum patch size criteria applied for biodiversity conservation (i.e., ∼85% of patches <100 ha). The 76 metacommunities we examined included 4401 species in 1190 patches. From each metacommunity, we resampled species–area accumulation curves to evaluate how biodiversity responded to habitat existing as a few large patches or as many small patches. Counter to the SL > SS principle and consistent with previous syntheses, species richness accumulated more rapidly when adding several small patches (45.2% SS > SL vs. 19.9% SL > SS) to reach the same cumulative area, even for the very small patches in our data set. Responses of taxa to habitat fragmentation differed, which suggests that when a given total area of habitat is to be protected, overall biodiversity conservation will be most effective if that habitat is composed of as many small patches as possible, plus a few large ones. Because minimum patch size criteria often require larger patches than the small patches we examined, our results suggest that such criteria hinder efforts to protect biodiversity.     

Role of Patch Size, Disease, and Movement in Rapid Extinction of Bighorn Sheep

Abstract: The controversy (  Berger 1990, 1999 ; Wehausen 1999 ) over rapid extinction in bighorn sheep ( Ovis canadensis ) has focused on population size alone as a correlate to persistence time. We report on the persistence and population performance of 24 translocated populations of bighorn sheep. Persistence in these sheep was strongly correlated with larger patch sizes, greater distance to domestic sheep, higher population growth rates, and migratory movements, as well as to larger population sizes. Persistence was also positively correlated with larger average home-range size ( p = 0.058, n = 10 translocated populations) and home-range size of rams ( p = 0.087, n = 8 translocated populations). Greater home-range size and dispersal rates of bighorn sheep were positively correlated to larger patches. We conclude that patch size and thus habitat carrying capacity, not population size per se, is the primary correlate to both population performance and persistence. Because habitat carrying capacity defines the upper limit to population size, clearly the amount of suitable habitat in a patch is ultimately linked to population size. Larger populations (250+ animals) were more likely to recover rapidly to their pre-epizootic survey number following an epizootic ( p = 0.019), although the proportion of the population dying in the epizootic also influenced the probability of recovery ( p = 0.001). Expensive management efforts to restore or increase bighorn sheep populations should focus on large habitat patches located ≥23 km from domestic sheep, and less effort should be expended on populations in isolated, small patches of habitat.     

Inferring Metapopulation Dynamics from Patch-Level Incidence of Florida Scrub Plants

Spatial structure and dynamics of multiple populations may explain species distribution patterns in patchy communities with heterogeneous disturbance regimes, especially when species have poor dispersal. The endemic-rich Florida (U.S.A.) rosemary scrub occupies about 4% of the west portion of Archbold Biological Station and occurs scattered within a matrix of less xeric vegetation. Longer fire-return times and higher frequency of open patches in rosemary scrub provide favorable habitat for many plant species. Occupancy of 123 species of vascular plants and ground lichens in 89 patches was determined by repeated site surveys. About two-thirds of the species occurring at more than 14 patches had a significant logistic regression of presence on time-since-fire, patch size, patch isolation, or their interactions. Species with presence related to the interaction between patch isolation and patch size were primarily herbs and small shrubs specializing in rosemary scrub. These results suggest the importance of spatial characteristics of the landscape for population turnover of these species. An incidence-based metapopulation model was used to predict extinction and colonization probabilities of those species with presence in rosemary scrub patches related to the studied spatial variables. This is the first attempt to apply incidence-based metapopulation models to plants. The results showed stronger effects of patch size and patch isolation on extinction probabilities of herbs than on those of woody species. Because of their effect on spatial heterogeneity and habitat availability, fire suppression and habitat destruction may decrease persistence probabilities for these rosemary scrub specialists, many of which are endangered species.     

Estimating Extinction Risk with Metapopulation Models of Large‐Scale Fragmentation

Habitat loss is the principal threat to species. How much habitat remains—and how quickly it is shrinking—are implicitly included in the way the International Union for Conservation of Nature determines a species’ risk of extinction. Many endangered species have habitats that are also fragmented to different extents. Thus, ideally, fragmentation should be quantified in a standard way in risk assessments. Although mapping fragmentation from satellite imagery is easy, efficient techniques for relating maps of remaining habitat to extinction risk are few. Purely spatial metrics from landscape ecology are hard to interpret and do not address extinction directly. Spatially explicit metapopulation models link fragmentation to extinction risk, but standard models work only at small scales. Counterintuitively, these models predict that a species in a large, contiguous habitat will fare worse than one in 2 tiny patches. This occurs because although the species in the large, contiguous habitat has a low probability of extinction, recolonization cannot occur if there are no other patches to provide colonists for a rescue effect. For 4 ecologically comparable bird species of the North Central American highland forests, we devised metapopulation models with area‐weighted self‐colonization terms; this reflected repopulation of a patch from a remnant of individuals that survived an adverse event. Use of this term gives extra weight to a patch in its own rescue effect. Species assigned least risk status were comparable in long‐term extinction risk with those ranked as threatened. This finding suggests that fragmentation has had a substantial negative effect on them that is not accounted for in their Red List category. Estimación del Riesgo de Extinción Mediante Modelos Metapoblacionales de Fragmentación a Gran Escala     

Geometrical Factors and Metapopulation Dynamics of the Bush Cricket, Metrioptera bicolor Philippi (Orthoptera: Tettigoniidae)

This paper presents a metapopulation study of the bush cricket, Metrioptera bicolor , living in a recently fragmented landscape. The species inhabits grass and heathland patches of varying area and isolation. Analyses are made of how these geometrical factors affect local population size and density, distribution pattern, and the probability of local extinction and colonization. The proportion of available patches occupied varied between 72 and 79% during 1985–1990. Unoccupied patches were smaller and more isolated than those that were occupied. Patches where populations became extinct during this period were smaller than those with persisting populations. Since local population size was well correlated with patch area, it was clear that stochastic extinctions only occurred in small populations. Critical patch size for population extinction was approximately half a hectare. Colonized patches were less isolated than those that had not been colonized. Critical inter-patch distance for colonization was about 100 meters. The turnover was restricted to an identifiable share of the available patches. Only 33% of the patches were so small that extinction due to stochastic causes could be considered highly probable. This metapopulation will therefore most likely persist over a considerable period in its present spatial structure. There are apparent threats of further fragmentation, however, and nothing is known about the likelihood of large-scale extinctions resulting from extremely unfavorable weather conditions. Nevertheless, our results show that it is appropriate to include geometrical factors in metapopulation models.     

Local Population Dynamics in Metapopulation Models: Implications for Conservation

Abstract: Habitat fragmentation and the division of populations into spatially separated units have led to the increasing use of metapopulation models to characterize these populations. One prominent model that has served as a heuristic tool was introduced by Levins and is based on a collection of simplifying assumptions that exclude information on the dynamics and spatial distribution of local populations. Levins's and similar models predict the proportion of occupied habitat patches at equilibrium and the conditions needed to avoid total extinction. There are many obvious concerns about using such models, including how realistic alterations might change the predictions and whether occupancy has any relationship to population-level processes. Although many of the assumptions of these simple models are known to be unrealistic, we do not know how the assumptions affect model predictions. We simulated a metapopulation, and our results show that assumptions such as homogeneity of habitat patches, random migration among patches, equivalent extinction probabilities in all patches, and a large number of patches can lead to large overestimations of habitat occupancy. But when we explicitly modeled the underlying population dynamics within each patch, we found (1) that there was a strong correlation between proportion of occupied patches and total metapopulation size and (2) that the distribution of individuals among patches was relatively insensitive to model assumptions. Thus, our results show that although realistic modifications will change model predictions for occupancy, occupancy and population trends will be correlated. These correlations between occupancy and population size suggest that occupancy models may have some utility in conservation applications.     

Avian Survival Rates and the Extinction Process on Barro Colorado Island, Panama

Abstract: Selective extinction following isolation of habitat patches may be due to biogeographical (e.g., island size or isolation) and ecological (species natural histories, interspecifc interactions) factors, or their interactions. Among the demographic and life history attributes commonly associated with high extinction probability are small populations, large size of individuals, and population variability. Long-term capture-recapture data from forest habitat in central Panama permit an examination of the association between mainland survival rates and extinction on a nearby land-bridge island Species of birds that no longer occur on Barro Colorado Island (BCI), Panama, have, on average, lower survival rates on the adjacent mainland than species that have persisted on BCI. Moreover, of the species that no longer occur on BCI, those with lower mainland survival rates generally disappeared earlier from the island. My analysis provides little evidence of a relationship between extinction and population size. Recolonization of BCI from the adjacent mainland by the forest undergrowth species studied here is unlikely. Reduced reproductive success on BCI combined with naturally low adult survival rates seems to be responsible for these BCI extinctions. High nest predation and/or altered landscape dynamics are probable agents in the low reproductive success. The methods used here could be employed in other circumstances to identify fragmentation-sensitive species.     

Extinction Rates in Archipelagoes: Implications for Populations in Fragmented Habitats

To study the effect of habitat fragmentation on population viability, I used extinction rates on islands in archipelagoes and estimated the relative probability of extinction per species on single large islands and sets of smaller islands with the same total area. Data on lizards, birds, and mammals on oceanic islands and mammals on mountaintops and in nature reserves yield similar results. Species are likely to go extinct on all the small islands before they go extinct on the single, large island. In the short term, the analysis indicates that extinction probabilities may be lower on a set of small islands. This is perhaps an artifact due to underestimation of extinction rates on small islands and/or the necessity of pooling species in a focal taxon to obtain estimates of extinction rates (which may obscure area thresholds and underestimate the slope and curvature of extinction rates as a function of area). Ultimately, cumulative extinction probabilities are higher for a set of small islands than for single large islands. Mean and median times to extinction tend to be shorter in the fragmented systems, in some cases much shorter. Thus, to minimize extinction rates in isolated habitat remnants and nature reserve systems, the degree of fragmentation should be minimized     

Extinction debt of forest plants persists for more than a century following habitat fragmentation

Following habitat fragmentation individual habitat patches may lose species over time as they pay off their "extinction debt." Species with relatively low rates of population extinction and colonization ("slow" species) may maintain extinction debts for particularly prolonged periods, but few data are available to test this prediction. We analyzed two unusually detailed data sets on forest plant distributions and land-use history from Lincolnshire, United Kingdom, and Vlaams-Brabant, Belgium, to test for an extinction debt in relation to species-specific extinction and colonization rates. Logistic regression models predicting the presence-absence of 36 plant species were first parameterized using data from Lincolnshire, where forest cover has been relatively low (approximately 5-8%) for the past 1000 years. Consistent with extinction debt theory, for relatively slow species (but not fast species) these models systematically underpredicted levels of patch occupancy in Vlaams-Brabant, where forest cover was reduced from approximately 25% to <10% between 1775 and 1900 (it is presently 6.5%). As a consequence, the ability of the Lincolnshire models to predict patch occupancy in Vlaams-Brabant was worse for slow than for fast species. Thus, more than a century after forest fragmentation reached its current level an extinction debt persists for species with low rates of population turnover.     

Populations in small, ephemeral habitat patches may drive dynamics in a Daphnia magna metapopulation

Migration is the key process to understand the dynamics and persistence of a metapopulation. Many metapopulation models assume a positive correlation between habitat patch size or stability and the number of emigrants. However, few empirical data exist, and habitat patch size and habitat stability may affect dispersal differently than they affect local persistence. Here, we studied the production of the migration stage (i.e., resting eggs called ephippia) of the cladoceran Daphnia magna in a metapopulation consisting of 530 rock pool habitat patches over 25 years. Earlier, the functioning of this metapopulation was explained with a Levins-type metapopulation model or with a mainland-island metapopulation model, based on local extinction and colonization data or time series data, respectively. We used pool volume, hydroperiod length, and number of desiccation events to calculate per-pool production of ephippia (i.e., migration stages). We estimated that populations in small and ephemeral habitat patches produced more than half of the 250 000 to 1 million ephippia that were produced in the metapopulation as a whole per year between 1982 and 2006. Furthermore, these small populations contributed approximately 90% of the ephippia exposed during desiccation events, while the contribution of the long-lived populations in large pools was minimal. We term this an "inverse mainland-island" type metapopulation and propose that populations in small, ephemeral habitat patches may also be the driving force for metapopulation dynamics in other systems.     

Linking edge effects and patch size effects: Importance of matrix nest predators

The edge effect is usually considered to be the proximate cause of area sensitivity in forest birds. We tested if birds nesting in large patches are less vulnerable to the edge effect using a simple model that assumes an increase in patch size reduces the probability of a matrix predator moving to the core areas of forest and that larger perimeter/area ratios result in a higher number of matrix predators per unit of area. The probability of a nest being successful decreased asymptotically with an increase in either the patch penetration distance of predators or predator density, but those effects were reduced when patch size was increased. Large patches have a lower probability of being affected by an Allee effect and they can function as sink habitats only if penetration distance and predator density are largely increased. However, the transition from an Allee effect to a sink condition occurs with a small increase in penetration distance and predator density. Since birds nesting in large patches are less vulnerable to an increase in matrix predator populations, persistence of bird populations may be possible by increasing the size of habitat patches that can act as source populations.     

Metacommunity dynamics over 16 years in a pyrogenic shrubland

Metacommunity theory allows predictions about the dynamics of potentially interacting species' assemblages that are linked by dispersal, but strong empirical tests of the theory are rare. We analyzed the metacommunity dynamics of Florida rosemary scrub, a patchily distributed pyrogenic community, to test predictions about turnover rates, community nestedness, and responses to patch size, arrangement, and quality. We collected occurrence data for 45 plant species from 88 rosemary scrub patches in 1989 and 2005 and used growth form, mechanism of regeneration after fire, and degree of habitat specialization to categorize species by life history. We tested whether patch size, fire history, and structural connectivity (a measure of proximity and size of surrounding patches) could be used to predict apparent extinctions and colonizations. In addition, we tested the accuracy of incidence-function models built with the patch survey data from 1989. After fire local extinction rates were higher for herbs than woody plants, higher for species that regenerated only from seed than species able to resprout, and higher for generalist than specialist species. Fewer rosemary specialists and a higher proportion of habitat generalists were extirpated on recently burned patches than on patches not burned between 1989 and 2005. Nestedness was highest for specialists among all life-history groups. Estimated model parameters from 1989 predicted the observed (1989-2005) extinction rates and the number of patches with persistent populations of individual species. These results indicate that species with different life-history strategies within the same metacommunity can have substantially different responses to patch configuration and quality. Real metacommunities may not conform to certain assumptions of simple models, but incidence-function models that consider only patch size, configuration, and quality can have significant predictive accuracy.     

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.     

How does the spatial structure of habitat loss affect the eco-epidemic dynamics?

Habitat loss is considered as one of the primary causes of species extinction, especially for a species that also suffers from an epidemic disease. Little attention has been paid to the combined effect of habitat loss and epidemic transmission on the species spatiotemporal dynamics. Here, a spatial model of the parasite–host/prey–predator eco-epidemiological system with habitat loss was studied. Habitat patches in the model, instead of undergoing a random loss, were spatially clustered by different degrees. Not only the quantity of habitat loss but also its clustering degree was shown to affect the equilibrium of the system. The infection rate and the probability of successful predation were keys to determine the spatial patterns of species. The epidemic disease is more likely to break out if only a small amount of suitable patches were lost. Counter-intuitively, infected preys are more sensitive to habitat loss than predators if the lost patches are highly clustered. This result is new to eco-epidemiology and implies a possibility of using spatial arrangement of suitable (or unsuitable) patches to control the spread of epidemics in the ecological system.     

Harvesting creates ecological traps: consequences of invisible mortality risks in predator-prey metacommunities

Models of two-patch predator-prey metacommunities are used to explore how the global predator population changes in response to additional mortality in one of the patches. This could describe the dynamics of a predator in an environment that includes a refuge area where that predator is protected and a spatially distinct ("risky") area where it is harvested. The predator's movement is based on its perceived fitness in the two patches, but the risk from the additional mortality is potentially undetectable; this often occurs when the mortality is from human harvesting or from a novel type of top predator. Increases in undetected mortality in the risky area can produce an abrupt collapse of either the refuge population or of the entire predator population when the mortality rate exceeds a threshold level. This is due to the attraction of the risky patch, which has abundant prey due to its high predator mortality. Extinction of the refuge predator population does not occur when the refuge patch has a higher maximum per capita predator growth rate than the exploited patch because the refuge is then more attractive when the predator is rare. The possibility of abrupt extinction of one or both patches from high densities in response to a small increase in harvest is often associated with alternative states. In such cases, large reductions in mortality may be needed to avoid extinction in a collapsing predator population, or to reestablish an extinct population. Our analysis provides a potential explanation for sudden collapses of harvested populations, and it argues for more consideration of adaptive movement in designing protected areas.     

Spatial scale of local breeding habitat quality and adjustment of breeding decisions

Experimental studies provide evidence that, in spatially and temporally heterogeneous environments, individuals track variation in breeding habitat quality to adjust breeding decisions to local conditions. However, most experiments consider environmental variation at one spatial scale only, while the ability to detect the influence of a factor depends on the scale of analysis. We show that different breeding decisions by adults are based on information about habitat quality at different spatial scales. We manipulated (increased or decreased) local breeding habitat quality through food availability and parasite prevalence at a small (territory) and a large (patch) scale simultaneously in a wild population of Great Tits (Parus major). Females laid earlier in high-quality large-scale patches, but laying date did not depend on small-scale territory quality. Conversely, offspring sex ratio was higher (i.e., biased toward males) in high-quality, small-scale territories but did not depend on large-scale patch quality. Clutch size and territory occupancy probability did not depend on our experimental manipulation of habitat quality, but territories located at the edge of patches were more likely to be occupied than central territories. These results suggest that integrating different decisions taken by breeders according to environmental variation at different spatial scales is required to understand patterns of breeding strategy adjustment.     

Effects of Patch Size on Birds in Old-Growth Montane Forests

Following habitat alteration or fragmentation, competition, parasitism, and predation from species that live in the new habitats may reduce the survival and reproductive success of species living in the original habitats. Negative influences from species that live outside the remnant patches are expected to be greater in small rather than in large remnant patches because more "external" species are expected to move through the centers of small remnant patches. We surveyed birds within remnant patches of old-growth montane forests on Vancouver Island, Canada, (1) to evaluate whether the richness and abundance of non-old-growth bird species were larger at the center of small rather than large patches and (2) to evaluate whether the opposite was true of old-growth bird species. More non-old-growth bird species were present at the center of small remnant patches of old growth than in large old-growth patches. We found no relationship, however, between patch size and richness or abundance of old-growth bird species at the center of remnant patches of old growth. This was true for old-growth species with open, cup-shaped nests and cavity nests. Old-growth birds may have been affected less in our study area than in other areas because they evolved within heterogeneous montane forests and interacted with non-old-growth species throughout their evolutionary histories or because the contrast between old-growth forests and logged areas was less than that between the forests and agricultural/urban areas that were surveyed in other studies.     

Effects of temperature and precipitation on grassland bird nesting success as mediated by patch size

Grassland birds are declining faster than any other bird guild across North America. Shrinking ranges and population declines are attributed to widespread habitat loss and increasingly fragmented landscapes of agriculture and other land uses that are misaligned with grassland bird conservation. Concurrent with habitat loss and degradation, temperate grasslands have been disproportionally affected by climate change relative to most other terrestrial biomes. Distributions of grassland birds often correlate with gradients in climate, but few researchers have explored the consequences of weather on the demography of grassland birds inhabiting a range of grassland fragments. To do so, we modeled the effects of temperature and precipitation on nesting success rates of 12 grassland bird species inhabiting a range of grassland patches across North America (21,000 nests from 81 individual studies). Higher amounts of precipitation in the preceding year were associated with higher nesting success, but wetter conditions during the active breeding season reduced nesting success. Extremely cold or hot conditions during the early breeding season were associated with lower rates of nesting success. The direct and indirect influence of temperature and precipitation on nesting success was moderated by grassland patch size. The positive effects of precipitation in the preceding year on nesting success were strongest in relatively small grassland patches and had little effect in large patches. Conversely, warm temperatures reduced nesting success in small grassland patches but increased nesting success in large patches. Mechanisms underlying these differences may be patch‐size‐induced variation in microclimates and predator activity. Although the exact cause is unclear, large grassland patches, the most common metric of grassland conservation, appears to moderate the effects of weather on grassland‐bird demography and could be an effective component of climate‐change adaptation.