An Experimental Test of Matrix Permeability and Corridor Use by an Endemic Understory Bird

Abstract:  Because of widespread habitat fragmentation, maintenance of landscape connectivity has become a major focus of conservation planning, but empirical tests of animal movement in fragmented landscapes remain scarce. We conducted a translocation experiment to test the relative permeability of three landscape elements (open habitat, shrubby secondary vegetation, and wooded corridors) to movement by the Chucao Tapaculo ( Scelorchilus rubecula ), a forest understory bird endemic to South American temperate rainforest. Forty-one radio-tagged subjects were translocated (individually) to three landscape treatments consisting of small release patches that were either entirely surrounded by open habitat (pasture), entirely surrounded by dense shrubs, or linked to other patches by wooded corridors that were otherwise surrounded by open matrix. The number of days subjects remained in release patches before dispersal (a measure of habitat resistance) was significantly longer for patches surrounded by open habitat than for patches adjoining corridors or surrounded by dense shrubs. These results indicate that open habitat significantly constrains Chucao dispersal, in accord with expectation, but dispersal occurs equally well through wooded corridors and shrub-dominated matrix. Thus, corridor protection or restoration and management of vegetation in the matrix (to encourage animal movement) may be equally feasible alternatives for maintaining connectivity.     

Genes and song: genetic and social connections in fragmented habitat in a woodland bird with limited dispersal

Understanding the processes leading to population declines in fragmented landscapes is essential for successful conservation management. However, isolating the influence of disparate processes, and dispersal in particular, is challenging. The Grey Shrike-thrush, Colluricincla harmonica, is a sedentary woodland-dependent songbird, with learned vocalizations whose incidence in suitable habitat patches falls disproportionally with decline in tree cover in the landscape. Although it has been suggested that gaps in tree cover might act as barriers to its dispersal, the species remains in many remnants of native vegetation in agricultural landscapes, suggesting that it may have responded to habitat removal and fragmentation by maintaining or even increasing dispersal distances. We quantified population connectivity of the Grey Shrike-thrush in a system fragmented over more than 120 years using genetic (microsatellites) and acoustic (song types) data. First, we tested for population genetic and acoustic structure at regional and local scales in search of barriers to dispersal or gene flow and signals of local spatial structuring indicative of restricted dispersal or localized acoustic similarity. Then we tested for effects of habitat loss and fragmentation on genetic and acoustic connectivity by fitting alternative models of mobility (isolation-by-distance [the null model] and reduced and increased movement models) across treeless vs. treed areas. Birds within -5 km of each other had more similar genotypes and song types than those farther away, suggesting that dispersal and song matching are limited in the region. Despite restricted dispersal detected for females (but not males), populations appeared to be connected by gene flow and displayed some cultural (acoustic) connectivity across the region. Fragmentation did not appear to impact greatly the dispersal of the Grey Shrike-thrush: none of the mobility models fit the genetic distances of males, whereas for females, an isolation-by-distance model could not be rejected in favor of the models of reduced or increased movement through treeless gaps. However, dissimilarities of the song types were more consistent with the model of reduced cultural connectivity through treeless areas, suggesting that fragmentation impedes song type sharing in the Grey Shrike-thrush. Our paper demonstrates that habitat fragmentation hinders important population processes in an Australian woodland bird even though its dispersal is not detectably impacted.     

Consequences of ignoring dispersal variation in network models for landscape connectivity

Habitat loss and fragmentation can negatively influence population persistence and biodiversity, but the effects can be mitigated if species successfully disperse between isolated habitat patches. Network models are the primary tool for quantifying landscape connectivity, yet in practice, an overly simplistic view of species dispersal is applied. These models often ignore individual variation in dispersal ability under the assumption that all individuals move the same fixed distance with equal probability. We developed a modeling approach to address this problem. We incorporated dispersal kernels into network models to determine how individual variation in dispersal alters understanding of landscape-level connectivity and implemented our approach on a fragmented grassland landscape in Minnesota. Ignoring dispersal variation consistently overestimated a population's robustness to local extinctions and underestimated its robustness to local habitat loss. Furthermore, a simplified view of dispersal underestimated the amount of habitat substructure for small populations but overestimated habitat substructure for large populations. Our results demonstrate that considering biologically realistic dispersal alters understanding of landscape connectivity in ecological theory and conservation practice.     

Importance of dispersal routes that minimize open‐ocean movement to the genetic structure of island populations

Islands present a unique scenario in conservation biology, offering refuge yet imposing limitations on insular populations. The Kimberley region of northwestern Australia has more than 2500 islands that have recently come into focus as substantial conservation resources. It is therefore of great interest for managers to understand the driving forces of genetic structure of species within these island archipelagos. We used the ubiquitous bar‐shouldered skink (Ctenotus inornatus) as a model species to represent the influence of landscape factors on genetic structure across the Kimberley islands. On 41 islands and 4 mainland locations in a remote area of Australia, we genotyped individuals across 18 nuclear (microsatellite) markers. Measures of genetic differentiation and diversity were used in two complementary analyses. We used circuit theory and Mantel tests to examine the influence of the landscape matrix on population connectivity and linear regression and model selection based on Akaike's information criterion to investigate landscape controls on genetic diversity. Genetic differentiation between islands was best predicted with circuit‐theory models that accounted for the large difference in resistance to dispersal between land and ocean. In contrast, straight‐line distances were unrelated to either resistance distances or genetic differentiation. Instead, connectivity was determined by island‐hopping routes that allow organisms to minimize the distance of difficult ocean passages. Island populations of C. inornatus retained varying degrees of genetic diversity (N_A = 1.83 – 7.39), but it was greatest on islands closer to the mainland, in terms of resistance‐distance units. In contrast, genetic diversity was unrelated to island size. Our results highlight the potential for islands to contribute to both theoretical and applied conservation, provide strong evidence of the driving forces of population structure within undisturbed landscapes, and identify the islands most valuable for conservation based on their contributions to gene flow and genetic diversity.     

Effects of Habitat Fragmentation on Effective Dispersal of Florida Scrub‐Jays

Abstract: Studies comparing dispersal in fragmented versus unfragmented landscapes show that habitat fragmentation alters the dispersal behavior of many species. We used two complementary approaches to explore Florida Scrub‐Jay (Aphelocoma c?rulescens) dispersal in relation to landscape fragmentation. First, we compared dispersal distances of color‐marked individuals in intensively monitored continuous and fragmented landscapes. Second, we estimated effective dispersal relative to the degree of fragmentation (as inferred from two landscape indexes: proportion of study site covered with Florida Scrub‐Jay habitat and mean distance to nearest habitat patch within each study site) by comparing genetic isolation‐by‐distance regressions among 13 study sites having a range of landscape structures. Among color‐banded individuals, dispersal distances were greater in fragmented versus continuous landscapes, a result consistent with other studies. Nevertheless, genetic analyses revealed that effective dispersal decreases as the proportion of habitat in the landscape decreases. These results suggest that although individual Florida Scrub‐Jays may disperse farther as fragmentation increases, those that do so are less successful as breeders than those that disperse short distances. Our study highlights the importance of combining observational data with genetic inferences when evaluating the complex biological and life‐history implications of dispersal.     

Combining Measures of Dispersal to Identify Conservation Strategies in Fragmented Landscapes

Abstract: Understanding the way in which habitat fragmentation disrupts animal dispersal is key to identifying effective and efficient conservation strategies. To differentiate the potential effectiveness of 2 frequently used strategies for increasing the connectivity of populations in fragmented landscapes—corridors and stepping stones—we combined 3 complimentary methods: behavioral studies at habitat edges, mark‐recapture, and genetic analyses. Each of these methods addresses different steps in the dispersal process that a single intensive study could not address. We applied the 3 methods to the case study of Atrytonopsis new species 1, a rare butterfly endemic to a partially urbanized stretch of barrier islands in North Carolina (U.S.A.). Results of behavioral analyses showed the butterfly flew into urban and forested areas, but not over open beach; mark‐recapture showed that the butterfly dispersed successfully through short stretches of urban areas (<500 m); and genetic studies showed that longer stretches of forest (>5 km) were a dispersal barrier, but shorter stretches of urban areas (≤5 km) were not. Although results from all 3 methods indicated natural features in the landscape, not urbanization, were barriers to dispersal, when we combined the results we could determine where barriers might arise: forests restricted dispersal for the butterfly only when there were long stretches with no habitat. Therefore, urban areas have the potential to become a dispersal barrier if their extent increases, a finding that may have gone unnoticed if we had used a single approach. Protection of stepping stones should be sufficient to maintain connectivity for Atrytonopsis new species 1 at current levels of urbanization. Our research highlights how the use of complementary approaches for studying animal dispersal in fragmented landscapes can help identify conservation strategies.     

The Paradox of Forest Fragmentation Genetics

Abstract:  Theory predicts widespread loss of genetic diversity from drift and inbreeding in trees subjected to habitat fragmentation, yet empirical support of this theory is scarce. We argue that population genetics theory may be misapplied in light of ecological realities that, when recognized, require scrutiny of underlying evolutionary assumptions. One ecological reality is that fragment boundaries often do not represent boundaries for mating populations of trees that benefit from long-distance pollination, sometimes abetted by long-distance seed dispersal. Where fragments do not delineate populations, genetic theory of small populations does not apply. Even in spatially isolated populations, where genetic theory may eventually apply, evolutionary arguments assume that samples from fragmented populations represent trees that have had sufficient time to experience drift, inbreeding, and ultimately inbreeding depression, an unwarranted assumption where stands in fragments are living relicts of largely unrelated predisturbance populations. Genetic degradation may not be as important as ecological degradation for many decades following habitat fragmentation.     

Dispersal depression with habitat fragmentation in the bog fritillary butterfly

Habitat fragmentation is expected to impose strong selective pressures on dispersal rates. However, evolutionary responses of dispersal are not self-evident, since various selection pressures act in opposite directions. Here we disentangled the components of dispersal behavior in a metapopulation context using the Virtual Migration model, and we linked their variation to habitat fragmentation in the specialist butterfly Proclossiana eunomia. Our study provided a nearly unique opportunity to study how habitat fragmentation modifies dispersal at the landscape scale, as opposed to microlandscapes or simulation studies. Indeed, we studied the same species in four landscapes with various habitat fragmentation levels, in which large amounts of field data were collected and analyzed using similar methodologies. We showed the existence of quantitative variations in dispersal behavior correlated with increased fragmentation. Dispersal propensity from habitat patches (for a given patch size), and mortality during dispersal (for a given patch connectivity) were lower in more fragmented landscapes. We suggest that these were the consequences of two different evolutionary responses of dispersal behavior at the individual level: (1) when fragmentation increased, the reluctance of individuals to cross habitat patch boundaries also increased; (2) when individuals dispersed, they flew straighter in the matrix, which is the best strategy to improve dispersal success. Such evolutionary responses could generate complex nonlinear patterns of dispersal changes at the metapopulation level according to habitat fragmentation. Due to the small size and increased isolation of habitat patches in fragmented landscapes, overall emigration rate and mortality during dispersal remained high. As a consequence, successful dispersal at the metapopulation scale remained limited. Therefore, to what extent the selection of individuals with a lower dispersal propensity and a higher survival during dispersal is able to limit detrimental effects of habitat fragmentation on dispersal success is unknown, and any conclusion that metapopulations would compensate for them is flawed.     

Developing Metapopulation Connectivity Criteria from Genetic and Habitat Data to Recover the Endangered Mexican Wolf

Restoring connectivity between fragmented populations is an important tool for alleviating genetic threats to endangered species. Yet recovery plans typically lack quantitative criteria for ensuring such population connectivity. We demonstrate how models that integrate habitat, genetic, and demographic data can be used to develop connectivity criteria for the endangered Mexican wolf (Canis lupus baileyi), which is currently being restored to the wild from a captive population descended from 7 founders. We used population viability analysis that incorporated pedigree data to evaluate the relation between connectivity and persistence for a restored Mexican wolf metapopulation of 3 populations of equal size. Decreasing dispersal rates greatly increased extinction risk for small populations (<150–200), especially as dispersal rates dropped below 0.5 genetically effective migrants per generation. We compared observed migration rates in the Northern Rocky Mountains (NRM) wolf metapopulation to 2 habitat‐based effective distance metrics, least‐cost and resistance distance. We then used effective distance between potential primary core populations in a restored Mexican wolf metapopulation to evaluate potential dispersal rates. Although potential connectivity was lower in the Mexican wolf versus the NRM wolf metapopulation, a connectivity rate of >0.5 genetically effective migrants per generation may be achievable via natural dispersal under current landscape conditions. When sufficient data are available, these methods allow planners to move beyond general aspirational connectivity goals or rules of thumb to develop objective and measurable connectivity criteria that more effectively support species recovery. The shift from simple connectivity rules of thumb to species‐specific analyses parallels the previous shift from general minimum‐viable‐population thresholds to detailed viability modeling in endangered species recovery planning. Desarrollo de Criterios de Conectividad Metapoblacional a Partir de Datos Genéticos y de Hábitat para Recuperar al Lobo Mexicano en Peligro de Extinción     

Effect of Rotational Shepherding on Demographic and Genetic Connectivity of Calcareous Grassland Plants

Response to habitat fragmentation may not be generalized among species, in particular for plant communities with a variety of dispersal traits. Calcareous grasslands are one of the most species‐rich habitats in Central Europe, but abandonment of traditional management has caused a dramatic decline of calcareous grassland species. In the Southern Franconian Alb in Germany, reintroduction of rotational shepherding in previously abandoned grasslands has restored species diversity, and it has been suggested that sheep support seed dispersal among grasslands. We tested the effect of rotational shepherding on demographic and genetic connectivity of calcareous grassland specialist plants and whether the response of plant populations to shepherding was limited to species dispersed by animals (zoochory). Specifically, we tested competing dispersal models and source and focal patch properties to explain landscape connectivity with patch‐occupancy data of 31 species. We fitted the same connectivity models to patch occupancy and nuclear microsatellite data for the herb Dianthus carthusianorum (Carthusian pink). For 27 species, patch connectivity was explained by dispersal by rotational shepherding regardless of adaptations to zoochory, whereas population size (16% species) and patch area (0% species) of source patches were not important predictors of patch occupancy in most species. [Correction made after online publication, February 25, 2014: Population size and patch area percentages were mistakenly inverted, and have now been fixed.] Microsite diversity of focal patches significantly increased the model variance explained by patch occupancy in 90% of the species. For D. carthusianorum, patch connectivity through rotational shepherding explained both patch occupancy and population genetic diversity. Our results suggest shepherding provides dispersal for multiple plant species regardless of their dispersal adaptations and thus offers a useful approach to restore plant diversity in fragmented calcareous grasslands. Efectos del Pastoreo Rotacional sobre la Conectividad Genética y Demográfica de Plantas de Pastizales Calcáreos     

Complex effects of habitat amount and fragmentation on functional connectivity and inbreeding in a giant panda population

The relationships between habitat amount and fragmentation level and functional connectivity and inbreeding remain unclear. Thus, we used genetic algorithms to optimize the transformation of habitat area and fragmentation variables into resistance surfaces to predict genetic structure and examined habitat area and fragmentation effects on inbreeding through a moving window and spatial autoregressive modeling approach. We applied these approaches to a wild giant panda population. The amount of habitat and its level of fragmentation had nonlinear effects on functional connectivity (gene flow) and inbreeding. Functional connectivity was highest when approximately 80% of the surrounding landscape was habitat. Although the relationship between habitat amount and inbreeding was also nonlinear, inbreeding increased as habitat increased until about 20% of the local landscape contained habitat, after which inbreeding decreased as habitat increased. Because habitat fragmentation also had nonlinear relationships with functional connectivity and inbreeding, we suggest these important responses cannot be effectively managed by minimizing or maximizing habitat or fragmentation. Our work offers insights for prioritization of protected areas.     

Value of Semi-Open Corridors for Simultaneously Connecting Open and Wooded Habitats: a Case Study with Ground Beetles

Abstract:  To counteract habitat fragmentation, the connectivity of a landscape should be enhanced. Corridors are thought to facilitate movement between disconnected patches of habitat, and linear strips of habitat connecting isolated patches are a popular type of corridor. On the other hand, the creation of new corridors can lead to fragmentation of the surrounding habitat. For example, heathland corridors connect patches of heathland and alternatively hedgerows connect patches of woodland. Nevertheless, these corridors themselves also break up previously connected patches of their surrounding habitat and in so doing fragment another type of habitat (heathland corridors fragment woodlands and woodland strips or hedgerows fragment heathlands). To overcome this challenge we propose the use of semi-open habitats (a mixture of heathland and woodland vegetation) as conservation corridors to enable dispersal of both stenotopic heathland and woodland species. We used two semi-open corridors with a mosaic of heathland and woody vegetation to investigate the efficiency of semi-open corridors for species dispersal and to assess whether these corridors might be a suitable approach for nature conservation. We conducted a mark-recapture study on three stenotopic flightless carabid beetles of heathlands and woodlands and took an inventory of all the carabid species in two semi-open corridors. Both methodological approaches showed simultaneous immigration of woodland and heathland species in the semi-open corridor. Detrended correspondence analysis showed a clear separation of the given habitats and affirmed that semi-open corridors are a good strategy for connecting woodlands and heathlands. The best means of creating and preserving semi-open corridors is probably through extensive grazing .     

The One-Migrant-per-Generation Rule in Conservation and Management

In the face of continuing habitat fragmentation and isolation, the optimal level of connectivity between populations has become a central issue in conservation biology. A common rule of thumb holds that one migrant per generation into a subpopulation is sufficient to minimize the loss of polymorphism and heterozygosity within subpopulations while allowing for divergence in allele frequencies among subpopulations. The one-migrant-per-generation rule is based on numerous simplifying assumptions that may not hold in natural populations. We examine the conceptual and theoretical basis of the rule and consider both genetic and nongenetic factors that influence the desired level of connectivity among subpopulations. We conclude that one migrant per generation is a desirable minimum, but it may be inadequate for many natural populations. We suggest that a minimum of 1 and a maximum of 10 migrants per generation would be an appropriate general rule of thumb for genetic purposes, bearing in mind that factors other than genetics may further influence the ideal level of connectivity.     

Effects of habitat fragmentation on the genetic structure and connectivity of the black-lipped pearl oyster Pinctada margaritifera populations in French Polynesia

Habitat destruction leading to increased fragmentation is detrimental to species by reducing population size and genetic diversity and by restraining population connectivity. However, little is known about the effects of naturally fragmented habitats on wild populations, especially when it comes to marine benthic invertebrates with long pelagic larval duration. In this framework, we investigated the connectivity and genetic diversity variation among nine wild populations of the black-lipped pearl oyster, Pinctada margaritifera, throughout French Polynesia using ten microsatellite DNA markers. Despite the naturally fragmented habitat (South Pacific oceanic islands), we found high values of genetic diversity and population admixture, indicating connectivity at small and large spatial scales within sampled sites of the Tuamotu, and between the Society and Tuamotu Archipelagos. In the meantime, habitat geomorphology increased genetic drift in populations occurring in small, closed lagoons. Significant genetic structure not correlated to geographic distance was detected mainly between closed and open lagoons. The Marquesas Islands hosted the most divergent populations, likely a result of vicariance. Our results also highlight that migration patterns among lagoons are not symmetrical. Altogether, the general pattern of gene flow, nonsymmetrical migration rates among populations, absence of isolation by distance and absence of recent extinction events found in our study strongly suggest that P. margaritifera populations of French Polynesia follow an asymmetrical island model of dispersal.     

A dispersal-constrained habitat suitability model for predicting invasion of alpine vegetation

Developing tools to predict the location of new biological invasions is essential if exotic species are to be controlled before they become widespread. Currently, alpine areas in Australia are largely free of exotic plant species but face increasing pressure from invasive species due to global warming and intensified human use. To predict the potential spread of highly invasive orange hawkweed (Hieracium aurantiacum) from existing founder populations on the Bogong High Plains in southern Australia, we developed an expert-based, spatially explicit, dispersal-constrained, habitat suitability model. The model combines a habitat suitability index, developed from disturbance, site wetness, and vegetation community parameters, with a phenomenological dispersal kernel that uses wind direction and observed dispersal distances. After generating risk maps that defined the relative suitability of H. aurantiacum establishment across the study area, we intensively searched several locations to evaluate the model. The highest relative suitability for H. aurantiacum establishment was southeast from the initial infestations. Native tussock grasslands and disturbed areas had high suitability for H. aurantiacum establishment. Extensive field searches failed to detect new populations. Time-step evaluation using the location of populations known in 1998-2000, accurately assigned high relative suitability for locations where H. aurantiacum had established post-2003 (AUC [area under curve] = 0.855 +/- 0.035). This suggests our model has good predictive power and will improve the ability to detect populations and prioritize areas for ongoing monitoring.     

Differences in gypsum plant communities associated with habitat fragmentation and livestock grazing.

The negative consequences of habitat fragmentation for plant communities have been documented in many regions of the world. In some fragmented habitats, livestock grazing has been proposed to be a dispersal mechanism reducing isolation between fragments. In others, grazing acts together with fragmentation in a way that increases habitat degradation. Iberian gypsum plant communities have been grazed and fragmented by agricultural practices for centuries. Although their conservation is considered a priority by the European Community, the effects of fragmentation on gypsum plant communities and the possible role of livestock grazing remain unknown. In addition, a substantial proportion of plant species growing in gypsum environments are gypsum specialists. They could be particularly affected by fragmentation, as was found for other habitat specialists (i.e., serpentine and calcareous specialists). In this study (1) we investigated the effect of fragmentation and grazing on gypsum plant community composition (species and life-forms), and (2) we tested to see if gypsum specialists were differently affected by fragmentation and grazing than habitat generalists. A vegetation survey was conducted in the largest gypsum outcrop of Europe (Middle Ebro Valley, northeast Spain). Fragmented and continuous sites in grazed and ungrazed areas were compared. Measurements related to species and composition of life-forms were contrasted first for the whole gypsum plant community and then specifically for the gypsum specialists. In the whole community, our results showed lower plant species diversity in fragmented sites, mainly due to the larger dominance of species more tolerant to fragmented habitat conditions. With livestock grazing, the plant species richness and the similarity in plant species composition between remnants was larger, suggesting that animals were acting as dispersal agents between fragments. As expected, gypsum specialists were less abundant in fragmented areas, and grazing led to the disappearance of the rare gypsum specialist Campanula fastigiata. According to our results, conservation strategies for gypsum plant communities in human-dominated landscapes should consider that fragmentation and grazing modify plant community composition affecting gypsum specialists in particular.     

Modelling the effect of in-stream and overland dispersal on gene flow in river networks

Modelling gene flow across natural landscapes is a current challenge of population genetics. Models are essential to make clear predictions about conditions that cause genetic differentiation or maintain connectivity between populations. River networks are a special case of landscape matrix. They represent stretches of habitat connected according to a branching pattern where dispersal is usually limited to upstream or downstream movements. Because of their peculiar topology, and the increasing concern about conservation issues in hydrosystems, there has been a recent revival of interest in modelling dispersal in river networks. Network complexity has been shown to influence global population differentiation. However, geometric characteristics are likely to interact with the way individuals move across space. Studies have focused on in-stream movements. None of the work published so far took into consideration the ability of many species to disperse overland between branches of the same network though. We predicted that the relative contribution of these two dispersal modalities (in-stream and overland) would affect the overall genetic structure. We simulated dispersal in synthetic river networks using an individual-based model. We tested the effect of dispersal modalities, i.e. the ratio of overland/in-stream dispersal, and two geometric parameters, bifurcation angle between branches and network complexity. Data revealed that if geometrical parameters affected population differentiation, dispersal parameters had the strongest effect. Interestingly, we observed a quadratic relationship between p the proportion of overland dispersers and population differentiation. We interpret this U-shape pattern as a balance between isolation by distance caused by in-stream movements at low values of p and intense migrant exchanges within the same branching unit at high values of p. Our study is the first attempt to model out-of-network movements. It clearly shows that both geometric and dispersal parameters interact. Both should be taken into consideration in order to refine predictions about dispersal and gene flow in river network.     

How do dispersal costs and habitat selection influence realized population connectivity?

Despite the importance of dispersal for population connectivity, dispersal is often costly to the individual. A major impediment to understanding connectivity has been a lack of data combining the movement of individuals and their survival to reproduction in the new habitat (realized connectivity). Although mortality often occurs during dispersal (an immediate cost), in many organisms costs are paid after dispersal (deferred costs). It is unclear how such deferred costs influence the mismatch between dispersal and realized connectivity. Through a series of experiments in the field and laboratory, we estimated both direct and indirect deferred costs in a marine bryozoan (Bugula neritina). We then used the empirical data to parameterize a theoretical model in order to formalize predictions about how dispersal costs influence realized connectivity. Individuals were more likely to colonize poor-quality habitat after prolonged dispersal durations. Individuals that colonized poor-quality habitat performed poorly after colonization because of some property of the habitat (an indirect deferred cost) rather than from prolonged dispersal per se (a direct deferred cost). Our theoretical model predicted that indirect deferred costs could result in nonlinear mismatches between spatial patterns of potential and realized connectivity. The deferred costs of dispersal are likely to be crucial for determining how well patterns of dispersal reflect realized connectivity. Ignoring these deferred costs could lead to inaccurate predictions of spatial population dynamics.     

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.     

Dispersal ecology of deadwood organisms and connectivity conservation

Limited knowledge of dispersal for most organisms hampers effective connectivity conservation in fragmented landscapes. In forest ecosystems, deadwood‐dependent organisms (i.e., saproxylics) are negatively affected by forest management and degradation globally. We reviewed empirically established dispersal ecology of saproxylic insects and fungi. We focused on direct studies (e.g., mark‐recapture, radiotelemetry), field experiments, and population genetic analyses. We found 2 somewhat opposite results. Based on direct methods and experiments, dispersal is limited to within a few kilometers, whereas genetic studies showed little genetic structure over tens of kilometers, which indicates long‐distance dispersal. The extent of direct dispersal studies and field experiments was small and thus these studies could not have detected long‐distance dispersal. Particularly for fungi, more studies at management‐relevant scales (1–10 km) are needed. Genetic researchers used outdated markers, investigated few loci, and faced the inherent difficulties of inferring dispersal from genetic population structure. Although there were systematic and species‐specific differences in dispersal ability (fungi are better dispersers than insects), it seems that for both groups colonization and establishment, not dispersal per se, are limiting their occurrence at management‐relevant scales. Because most studies were on forest landscapes in Europe, particularly the boreal region, more data are needed from nonforested landscapes in which fragmentation effects are likely to be more pronounced. Given the potential for long‐distance dispersal and the logical necessity of habitat area being a more fundamental landscape attribute than the spatial arrangement of habitat patches (i.e., connectivity sensu strict), retaining high‐quality deadwood habitat is more important for saproxylic insects and fungi than explicit connectivity conservation in many cases.