Scale-dependent changes in the importance of larval supply and habitat to abundance of a reef fish

While there is great interest in the degree to which local interactions "scale-up" to predict regional patterns of abundance, few studies in marine systems have simultaneously examined patterns of abundance at both the large scale (tens of kilometers) typical of larval movement and the small scale (meters) typical of post-settlement interactions. We addressed this gap by monitoring larval supply, adult survivorship, and giant kelp (Macrocystis pyrifera, a primary habitat-forming species) abundance for 13 populations of kelp bass (Paralabrax clathratus) spread over approximately 200 km in the Santa Barbara Channel, California, USA. At the small, within-site scale, both recruitment and adult survivorship of kelp bass were density-dependent and positively related to kelp abundance. At the larger, among-site scale, the spatial pattern of adult kelp bass abundance was predicted well by the pattern of kelp bass larval supply, but there was a consistent negative spatial relationship between kelp abundance and kelp bass larval supply despite the positive effects of kelp on kelp bass at the smaller spatial scale. This large-scale negative relationship was likely a product of a channel-wide spatial mismatch between oceanographic conditions that favor kelp survival and those that concentrate and distribute fish larvae. These results generally support the recruit-adult hypothesis: kelp bass populations are limited by recruitment at low recruit densities but by density-dependent competition for food resources and/or predator refuges at high recruit densities. At the same time, spatial variation in kelp abundance produced substantial spatiotemporal heterogeneity in kelp bass demographics, which argues for a multispecies, metacommunity approach to predicting kelp bass dynamics.     

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 role of habitat disturbance and recovery in metapopulation persistence

Classical metapopulation theory assumes a static landscape. However, empirical evidence indicates many metapopulations are driven by habitat succession and disturbance. We develop a stochastic metapopulation model, incorporating habitat disturbance and recovery, coupled with patch colonization and extinction, to investigate the effect of habitat dynamics on persistence. We discover that habitat dynamics play a fundamental role in metapopulation dynamics. The mean number of suitable habitat patches is not adequate for characterizing the dynamics of the metapopulation. For a fixed mean number of suitable patches, we discover that the details of how disturbance affects patches and how patches recover influences metapopulation dynamics in a fundamental way. Moreover, metapopulation persistence is dependent not only on the average lifetime of a patch, but also on the variance in patch lifetime and the synchrony in patch dynamics that results from disturbance. Finally, there is an interaction between the habitat and metapopulation dynamics, for instance declining metapopulations react differently to habitat dynamics than expanding metapopulations. We close, emphasizing the importance of using performance measures appropriate to stochastic systems when evaluating their behavior, such as the probability distribution of the state of the metapopulation, conditional on it being extant (i.e., the quasistationary distribution).     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

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

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

Quantifying the importance of local niche-based and stochastic processes to tropical tree community assembly

Although niche-based and stochastic processes, including dispersal limitation and demographic stochasticity, can each contribute to community assembly, it is difficult to quantify the relative importance of each process in natural vegetation. Here, we extend Shipley's maxent model (Community Assembly by Trait Selection, CATS) for the prediction of relative abundances to incorporate both trait-based filtering and dispersal limitation from the larger landscape and develop a statistical decomposition of the proportions of the total information content of relative abundances in local communities that are attributable to trait-based filtering, dispersal limitation, and demographic stochasticity. We apply the method to tree communities in a mature, species-rich, tropical forest in French Guiana at 1-, 0.25- and 0.04-ha scales. Trait data consisted of species' means of 17 functional traits measured over both the entire meta-community and separately in each of nine 1-ha plots. Trait means calculated separately for each site always gave better predictions. There was clear evidence of trait-based filtering at all spatial scales. Trait-based filtering was the most important process at the 1-ha scale (34%), whereas demographic stochasticity was the most important at smaller scales (37-53%). Dispersal limitation from the meta-community was less important and approximately constant across scales (-9%), and there was also an unresolved association between site-specific traits and meta-community relative abundances. Our method allows one to quantify the relative importance of local niche-based and meta-community processes and demographic stochasticity during community assembly across spatial and temporal scales.     

Reconstructing the historic demography of an endangered seabird

Reducing extinction risk for threatened species requires determining which demographic parameters are depressed and causing population declines. Museum collections may constitute a unique, underutilized resource for measuring demographic changes over long time periods using age-ratio analysis. We reconstruct the historic demography of a U.S. federally endangered seabird, the Marbled Murrelet (Brachyramphus marmoratus), from specimens collected approximately 100 years ago for comparison with predictions from comparative analyses and with results from contemporary field studies using both age-ratio analysis and conventional demographic estimators. Reproduction in the late 1800s and early 1900s matched predictions from comparative analysis, but was 8-9 times greater than contemporary estimates, whereas adult survival was unchanged. Historic reproductive rates would support stable populations, but contemporary levels should result in population declines. Contemporary demographic estimates derived from age-ratio analysis were similar to estimates from conventional estimators. Using museum specimens to reconstruct historic demography provides a unique approach to identify causes of decline and to set demographic benchmarks for recovery of endangered species that meet most assumptions of age-ratio analysis.     

Quantifying the Indicator Power of an Indicator Species

Abstract:  Biodiversity indicator species are needed for classifying biotopes and sites for conservation, and a number of methods have been developed for determining indicator species for this purpose. Nevertheless, in addition to site classification, there is sometimes a need to define an indicator species that indicates the occurrence of another species. For example, when a species of interest (target species) is difficult to detect or identify, a reliable indicator species can function as a tool that saves time and money. We derived a method that provides a quantitative measure of the indicator power (IP) of an indicator species for the target species or any species assemblage. We calculated the measure of IP from a presence–absence matrix that covered several sites. The method provided a list of indicator species, the presence of which reliably indicated the presence of another species (e.g., a threatened or rare species in a given area). The IP of the species was highest when the number of shared occurrences between the indicator species and the target species was high and, simultaneously, when the indicator species and the target species occurred separately in only a few cases. The IP was also positively influenced by the number of sites with no occurrences of either the indicator or the target species. Our method can also be used to quantify different types of species occurrence indications. We refer to these types as presence–presence, presence–absence, absence–presence, and absence–absence indications. To clarify the use of the method, we examined the situation with red-listed polypores in White-backed Woodpecker (Dendrocopos leucotos) habitats in Fennoscandia and found some suitable indicator species. Our method provides a new, objective way to evaluate the IP of an indicator species.     

A state-dependent model for the optimal management of an invasive metapopulation.

Management of invasive species involves choosing between different management strategy options, but often the best strategy for a particular scenario is not obvious. We illustrate the use of optimization methods to determine the most efficient management strategy using one of the most devastating invasive forest pests in North America, the gypsy moth (Lymantria dispar), as a case study. The optimization approach involves the application of stochastic dynamic programming (SDP) to a metapopulation framework with different infestation patch sizes, with the goal of minimizing infestation spread. We use a novel "moving window" approach as a way to address a spatially explicit problem without being explicitly spatial. We examine results for two cases in order to develop general rules of thumb for management. We explore a model with limited parameter information and then assess how strategies change with specific parameterization for the gypsy moth. The model results in a complex but stable, state-dependent management strategy for a multiyear management program that is robust even under situations of uncertainty. The general rule of thumb for the basic model consists of three strategies: eradicating medium-density infestations, reducing large-density infestations, and reducing the colonization rate from the main infestation, depending on the state of the system. With specific gypsy moth parameterization, reducing colonization decreases in importance relative to the other two strategies. The application of this model to gypsy moth management emphasizes the importance of managing based on the state of the system, and if applied to a specific geographic area, has the potential to substantially improve the efficiency and cost-effectiveness of current gypsy moth eradication programs, helping to slow the spread of this pest. Additionally, the approach used for this particular invasive species can be extended to the optimization of management programs for the spread of other invasive and problem species exhibiting metapopulation dynamics.     

Quantifying the importance of regional and local filters for community trait structure in tropical and temperate zones

The influence of regional and local processes on community structure is a major focus of ecology. Classically, ecologists have used local-regional richness regressions to evaluate the role of local and regional processes in determining community structure, an approach that has numerous flaws. Here, we implemented a novel trait-based approach that treats local and regional influences as a continuum, rather than a dichotomy. Using hylid frogs (Hylidae), we compared trait dispersion among members of local species assemblages to the trait dispersion in the regional assemblage from which they were drawn. Similarly, we compared trait dispersion in the regional assemblages to dispersion in the continental species pool. We estimated the contributions of local and regional filters, and we compared their strength in temperate and tropical zones. We found that regional and local filters explained 80% of the total variation among local assemblages in community body size dispersion. Overall, regional filters reduced trait dispersion, and local filters increased it, a pattern driven by particularly strong antagonistic effects in temperate zones that reduced the realized total variation by more than 40%. In contrast, local and regional filters acted in concert in tropical regions. Patterns within the tropics did not differ from the random expectation based on a null model, but within the temperate zone, local community filtering was stronger than expected by chance. Furthermore, in temperate regions, antagonistic regional and local filtering masked from 76% to 90% of the total variation in trait dispersion. Together, these results suggest that there are fundamental differences in the scale and identity of the processes determining community structure in temperate and tropical regions.     

Using individual‐condition measures to predict the long‐term importance of habitat extent for population persistence

Habitat loss and fragmentation are causing widespread population declines, but identifying how and when to intervene remains challenging. Predicting where extirpations are likely to occur and implementing management actions before losses result may be more cost‐effective than trying to reestablish lost populations. Early indicators of pressure on populations could be used to make such predictions. Previous work conducted in 2009 and 2010 identified that the presence of Eastern Yellow Robins (Eopsaltria australis) in 42 sites in a fragmented region of eastern Australia was unrelated to woodland extent within 500 m of a site, but the robins’ heterophil:lymphocyte (H:L) ratios (an indicator of chronic stress) were elevated at sites with low levels of surrounding woodland. We resurveyed these 42 sites in 2013 and 2014 for robin presence to determine whether the H:L ratios obtained in 2009 and 2010 predicted the locations of extirpations and whether the previous pattern in H:L ratios was an early sign that woodland extent would become an important predictor of occupancy. We also surveyed for robins at 43 additional sites to determine whether current occupancy could be better predicted by landscape context at a larger scale, relevant to dispersal movements. At the original 42 sites, H:L ratios and extirpations were not related, although only 4 extirpations were observed. Woodland extent within 500 m had become a strong predictor of occupancy. Taken together, these results provide mixed evidence as to whether patterns of individual condition can reveal habitat relationships that become evident as local shifts in occupancy occur but that are not revealed by a single snapshot of species distribution. Across all 85 sites, woodland extent at scales relevant to dispersal (5 km) was not related to occurrence. We recommend that conservation actions focus on regenerating areas of habitat large enough to support robin territories rather than increasing connectivity within the landscape.     

Seasonal changes in movements and habitat preferences of the scalloped hammerhead shark (Sphyrna lewini) while refuging near an oceanic island

Movements and habitat preferences of sharks relative to a central location are widely documented for many species; however, the reasons for such behaviors are currently unknown. Do movements vary spatially or temporally or between individuals? Do sharks have seasonal habitat and environmental preferences or simply perform movements at random at any time of the year? To help understand requirements for the designation of critical habitats for an endangered top predator and to develop zoning and management plans for key habitats, we examined vertical and horizontal movements, and determined habitat and environmental preferences of scalloped hammerhead sharks (Sphyrna lewini). We tracked seven hammerheads for 19–96 h at Wolf Island (1.38ºN, 91.82ºW) between 2007 and 2009 using ultrasonic transmitters with depth and temperature sensors, and we profiled temperature through the water column. Movements of individual hammerheads fell in two classes: constrained (remaining near the island) and dispersive (moving offshore to pelagic environments). The central activity space or kernel off the southeast side of Wolf Island was small and common to most, but the area varied among individuals (mean ± SE 0.25 ± 0.2 km²), not exceeding 0.6 km² for any of the sharks, and not changing significantly between seasons. In general, hammerheads showed preference for the up-current habitat on the eastern side of Wolf Island in both the warm and cold seasons. However, the depth of sharks varied with season, apparently in response to seasonal changes in the vertical structure of temperature. Hammerheads performed frequent vertical excursions above the thermocline during offshore movements and, in general, were observed to prefer temperatures of 23–26 °C found above the thermocline. At times, though individuals moved into the thermocline and made brief dives below it. Our results provided evidence that hammerheads (1) are highly selective of location (i.e., habitat on up-current side of island) and depth (i.e., top of the thermocline) while refuging, where they may carry out essential activities such as cleaning and thermoregulation, and (2) perform exploratory vertical movements by diving the width of the mixed layer and occasionally diving below the thermocline while moving offshore, most likely for foraging.     

Quantifying species recovery and conservation success to develop an IUCN Green List of Species

Stopping declines in biodiversity is critically important, but it is only a first step toward achieving more ambitious conservation goals. The absence of an objective and practical definition of species recovery that is applicable across taxonomic groups leads to inconsistent targets in recovery plans and frustrates reporting and maximization of conservation impact. We devised a framework for comprehensively assessing species recovery and conservation success. We propose a definition of a fully recovered species that emphasizes viability, ecological functionality, and representation; and use counterfactual approaches to quantify degree of recovery. This allowed us to calculate a set of 4 conservation metrics that demonstrate impacts of conservation efforts to date (conservation legacy); identify dependence of a species on conservation actions (conservation dependence); quantify expected gains resulting from conservation action in the medium term (conservation gain); and specify requirements to achieve maximum plausible recovery over the long term (recovery potential). These metrics can incentivize the establishment and achievement of ambitious conservation targets. We illustrate their use by applying the framework to a vertebrate, an invertebrate, and a woody and an herbaceous plant. Our approach is a preliminary framework for an International Union for Conservation of Nature (IUCN) Green List of Species, which was mandated by a resolution of IUCN members in 2012. Although there are several challenges in applying our proposed framework to a wide range of species, we believe its further development, implementation, and integration with the IUCN Red List of Threatened Species will help catalyze a positive and ambitious vision for conservation that will drive sustained conservation action.     

Evaluating the effects of foraging habitat restoration on shorebird reproduction: the importance of performance criteria and comparative design

Coastal development and engineering projects preclude ecosystem processes that provide habitat for beach nesting birds. Management for coastal species may depend on actions that attempt to restore important habitat features and mitigate disturbance effects. However, species response to restoration or other management actions may be difficult to predict or measure. At Jones Beach State Park, on Long Island, New York, a 0.49 ha restoration project provided moist substrate foraging habitat for breeding Piping Plovers (Charadrius melodus) from 2002 to 2005. We examined whether foraging habitat restoration affected Piping Plover breeding population size, productivity, and fledgling production within 300 m of the restoration site. We found a positive relationship between habitat restoration and the number of fledglings produced per year. However, foraging habitat restoration did not significantly increase the number of Plover pairs breeding at Jones Beach. Our ability to evaluate restoration effects on Plovers depended on: 1) use of multiple performance criteria; 2) a design that allowed comparison of pre- and post-restoration data; and 3) a spatial control that allowed comparison of similar areas that were near and far from the restoration site. Despite the small size of the restoration project, there were measurable benefits to Plovers, indicating that foraging habitat restoration may be an effective tool for species recovery.     

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.     

Tidal influences on the habitat use of Indo-Pacific humpback dolphins in an estuary

This paper offers the first study of diurnal variations in the use of an estuarine habitat by Indo-Pacific humpback dolphins. Passive acoustic data loggers were deployed in the Xin Huwei River Estuary, Western Taiwan, from July 2009 to December 2010, to collect biosonar clicks. Acoustic encounter rates of humpback dolphins on the riverside of the estuary changed significantly over the four tidal phases, instead of the two diurnal phases based on the recordings from 268 days. Among the tidal phases, the encounter rates were lowest during ebb tides. Additionally, circling movements associated with the hunt for epipelagic fish significantly changed in temporal and spatial presence over the four tidal phases, matching the overall pattern of encounter rate changes in the focal estuary. Our findings suggest that the occurrence pattern and habitat utilization of humpback dolphins are likely to be influenced by the tidal-driven activity of their epipelagic prey.     

Reservoir‐host amplification of disease impact in an endangered amphibian

Emerging wildlife pathogens are an increasing threat to biodiversity. One of the most serious wildlife diseases is chytridiomycosis, caused by the fungal pathogen, Batrachochytrium dendrobatidis (Bd), which has been documented in over 500 amphibian species. Amphibians vary greatly in their susceptibility to Bd; some species tolerate infection, whereas others experience rapid mortality. Reservoir hosts—species that carry infection while maintaining high abundance but are rarely killed by disease—can increase extinction risk in highly susceptible, sympatric species. However, whether reservoir hosts amplify Bd in declining amphibian species has not been examined. We investigated the role of reservoir hosts in the decline of the threatened northern corroboree frog (Pseudophryne pengilleyi) in an amphibian community in southeastern Australia. In the laboratory, we characterized the response of a potential reservoir host, the (nondeclining) common eastern froglet (Crinia signifera), to Bd infection. In the field, we conducted frog abundance surveys and Bd sampling for both P. pengilleyi and C. signifera. We built multinomial logistic regression models to test whether Crinia signifera and environmental factors were associated with P. pengilleyi decline. C. signifera was a reservoir host for Bd. In the laboratory, many individuals maintained intense infections (>1000 zoospore equivalents) over 12 weeks without mortality, and 79% of individuals sampled in the wild also carried infections. The presence of C. signifera at a site was strongly associated with increased Bd prevalence in sympatric P. pengilleyi. Consistent with disease amplification by a reservoir host, P. pengilleyi declined at sites with high C. signifera abundance. Our results suggest that when reservoir hosts are present, population declines of susceptible species may continue long after the initial emergence of Bd, highlighting an urgent need to assess extinction risk in remnant populations of other declined amphibian species.     

Quantifying dispersal and establishment limitation in a population of an epiphytic lichen

Dispersal is a process critical for the dynamics and persistence of metapopulations, but it is difficult to quantify. It has been suggested that the old-forest lichen Lobaria pulmonaria is limited by insufficient dispersal ability. We analyzed 240 DNA extracts derived from snow samples by a L. pulmonaria-specific real-time PCR (polymerase chain reaction) assay of the ITS (internal transcribed spacer) region allowing for the discrimination among propagules originating from a single, isolated source tree or propagules originating from other locations. Samples that were detected as positives by real-time PCR were additionally genotyped for five L. pulmonaria microsatellite loci. Both molecular approaches demonstrated substantial dispersal from other than local sources. In a landscape approach, we additionally analyzed 240 snow samples with real-time PCR of ITS and detected propagules not only in forests where L. pulmonaria was present, but also in large unforested pasture areas and in forest patches where L. pulmonaria was not found. Monitoring of soredia of L. pulmonaria transplanted to maple bark after two vegetation periods showed high variance in growth among forest stands, but no significant differences among different transplantation treatments. Hence, it is probably not dispersal limitation that hinders colonization in the old-forest lichen L. pulmonaria, but ecological constraints at the stand level that can result in establishment limitation. Our study exemplifies that care has to be taken to adequately separate the effects of dispersal limitation from a limitation of establishment.