Blowing in the wind? Nutrient enrichment of remnant woodlands in an agricultural landscape

Increasing fertiliser use in agricultural landscapes is likely to threaten the viability of remnant native vegetation in many parts of the world. Australia’s prime grain production landscapes have nutrient poor soils, which formerly supported semi-arid woodland. The ecological function and capacity for regeneration of these remnants may be particularly susceptible to nutrient enrichment. The key sources of nutrients are wind and water deposition from crop fertilisation, and manure and feed from sheep. We hypothesised that these sources would result in unequal deposition of nutrients within and among remnant vegetation patches. We surveyed soil nutrients (Total N, Available P and K, C:N ratio, and soil pH) in the edges and interiors of 60 remnant woodland patches of various sizes, and in adjacent cultivated paddocks. Nutrient load was negatively correlated with remnant size and patterns were particularly strong for available P. Small remnant patches (<3 ha) were accumulation zones for nutrients, with levels comparable or higher than within crop lands. The patterns are consistent with the hypothesis that small remnants are strongly enriched as a result of being used for livestock shelter. In larger remnants, the primary cause of enrichment is consistent with edge accumulation of nutrients due to wind and water movement. In large patches, remnant edges, particularly the windward edge, were elevated compared to interiors of large patches. In these semi-arid crop lands, current trends in intensification of cropping and a shift away from livestock may reduce the input of nutrients to small patches but increase the nutrient threat to larger remnants.     

Landscape context affects honeyeater communities and their foraging behaviour in Australia: implications for plant pollination

We investigated the species richness and composition of bird communities in mallee woodland remnants in a highly fragmented landscape, focusing specifically on honeyeaters and their foraging behaviour. We observed birds around flowering Eremophila glabra ssp. glabra plants in three replicated contexts: (1) the interior of large remnants, (2) linear remnants within ~3 km of a large remnant, and (3) linear remnants 5–7 km from a large remnant. We found species richness differed among elements, with an increase in the number of species that tolerate disturbed, open habitat and a decrease in the number of woodland-dependent species in linear elements. Honeyeater assemblages were similar in species richness and abundance among the elements, but differed in composition due to a higher number of large-sized honeyeater species in distant elements. Honeyeater movement patterns into a site and within a site were similar among the elements. Floral visitation varied among honeyeater species and was positively correlated with their abundance in the far element. Our results demonstrate that bird species respond differently to the spatial context of remnants in a fragmented landscape; however, the degree of isolation of linear remnants was not important. Linear remnants appear to be frequently used by honeyeaters, but the changes in community composition among the elements may influence the quality of pollination, which could have implications for plant reproduction.     

Edge geometry influences patch-level habitat use by an edge specialist in south-eastern Australia

We investigated patterns in habitat use by the noisy miner (Manorina melanocephala) along farmland-woodland edges of large patches of remnant vegetation (>300 ha) in the highly fragmented box-ironbark woodlands and forests of central Victoria, Australia. Noisy miners exclude small birds from their territories, and are considered a significant threat to woodland bird communities in the study region. Seventeen different characteristics of edge habitat were recorded, together with the detection or non-detection of noisy miners along 129 500-m segments of patch edge. Habitat characteristics ranged from patch-level factors related to patch-edge geometry to site-level floristic factors. Backward (stepwise) logistic regression analyses were used to identify habitat characteristics that were associated with the occupancy of a site by noisy miners. After accounting for the effects of spatial autocorrelation on the occurrence of noisy miners along edges, we identified projections of remnant vegetation from the patch edge into the agricultural matrix (e.g., corners of patches, peninsulas of vegetation) and clumps of trees in the agricultural matrix within 100 m of the edge as significant predictors of the occupancy of edges by noisy miners. This relationship was also confirmed in two other geographically and floristically distinct habitats within Victoria. The use of edges with projections by noisy miners may confer advantages in interspecific territorial defence. In light of these results, we advocate revegetation strategies that attempt to enclose projections within 100 m of the edge, with fencing placed out to this new boundary, to reduce the likelihood of colonisation and domination of an edge by noisy miners. Our study highlights the need for greater consideration to be given to the patterns in habitat use by aggressive edge specialists, particularly in relation to patch-edge geometry and other human-induced components of landscapes.     

Incidence of competitors and landscape structure as predictors of woodland-dependent birds

Globally, modification of landscapes for agriculture has had a strong influence on the distribution and abundance of biota. In particular, woodland-dependent birds are under threat across agricultural landscapes in Britain, North America and Australia, with their decline and extirpation attributed to the loss and fragmentation of habitat. Other native species have become over-abundant in response to anthropogenic landscape change and have strong interactive effects on avian assemblage structure. In eastern Australia, the hyper-aggressive noisy miner (Manorina melanocephala) often dominates woodlands in agricultural landscapes through interspecific competition, resulting in declines of species richness of woodland-dependent birds. We aimed to determine the relative influence and importance of interspecific competition, in situ habitat structure and landscape structure for woodland-dependent bird species at the landscape level. We recorded species-specific landscape incidence of woodland-dependent birds in 24 agricultural-woodland mosaics (25 km²) in southern Queensland, Australia. We selected extensively cleared landscapes (10–23 % woodland cover) where fragmentation effects are expected to be greatest. We applied generalised linear models and hierarchical partitioning to quantify the relative importance of the landscape-level incidence of the noisy miner, mistletoe abundance, shrub cover, woodland extent, woodland subdivision and land-use intensity for the incidence of 46 species of woodland birds at the landscape-scale. The landscape-level incidence of the noisy miner was the most important explanatory variable across the assemblage. Both in situ habitat structure and landscape structure were of secondary importance to interspecific aggression, although previous research suggests that the increasing incidence of the noisy miner in fragmented agricultural landscapes is itself a consequence of anthropogenic changes to landscape structure. Species’ responses to fragmentation varied from positive to negative, but complex habitat structure had a consistently positive effect, suggesting in situ restoration of degraded habitats could be a conservation priority. Landscape wide conservation of woodland-dependent bird populations in agricultural landscapes may be more effective if direct management of noisy miner populations is employed, given the strong negative influence of this species on the incidence of woodland-dependent birds among landscapes.     

How does a transforming landscape influence bird breeding success?

Context

The conversion of agricultural landscapes to tree plantations is a major form of landscape transformation worldwide, but its effects on biodiversity, particularly key population processes like reproductive success, are poorly understood.

Objectives

We compared bird breeding success between woodland remnants surrounded by maturing stands of plantation Radiata Pine and a matched set of woodland remnants in semi-cleared grazing land.

Methods

Our study was conducted in the Nanangroe region in south-eastern New South Wales, Australia. Using repeated field measurements, we quantified bird breeding success in 23 woodland remnants; 13 surrounded by Radiata Pine plantations and 10 on farms where remnants were surrounded by semi-cleared grazing land. We matched the attributes of native remnant patches between two types of matrix.

Results

We found that: (1) rates of nesting success of smaller-bodied birds in woodland remnants surrounded by grazing land were significantly higher than in woodland remnants surrounded by pine plantations; and (2) taxa with domed nests were more successful at nesting than species that constructed open cup/bowl nests in woodland remnants within farmlands.

Conclusions

Our findings suggest that bird breeding success in remnant woodland patches is significantly diminished as a result of the conversion of semi-cleared grazing land to pine plantations.

     

Patterns of exotic plant invasions in fragmented urban and rural grasslands across continents

Linear native grassland remnants in fragmented landscapes are usually at a great risk of exotic species invasion from their edges. Changes in species distribution near habitat edges are extensively studied in ecology as knowledge about edge responses is important to understand the development of patterns and processes in landscapes. However, elucidating robust general principles for edge effects has been difficult as species responses to habitat edges are highly variable and dependent on a large number of attributes which affect the function and structure of edges and therefore the distance that edge effects penetrate into fragmented natural vegetation. The objective of this study was to investigate the generality of exotic species invasion patterns from edges in native grassland patches surrounded by urban and rural landscapes. This was done by comparing the results of research from Victoria, Australia with a similar study from North-West Province, South Africa. Despite their occurrence on different continents, the grasslands are floristically and structurally similar and are dominated by the same grass species. Invasion patterns were quantified using two spatial statistics methods; block kriging and spatially constrained clustering. Two distinct patterns of exotic species invasion were identified in native grassland remnants in South Africa and Australia, namely exotic species invasion from the edge where the cover of exotic species increased with increasing proximity to the edge and a pattern that suggests that gap phase vegetation dynamics may also drive exotic species invasion at urban grasslands. Although urbanization and weed invasions are complex processes similar patterns of exotic species invasion in urban grasslands were found in two different continents suggesting that general patterns may occur. Implications of this for the conservation of native grasslands in contrasting landscapes are discussed.     

Effects of forest patch size on avian diversity

The effects of landscape patchiness on the diversity of birds of the Georgia Piedmont were investigated during 1993. Birds were sampled along line transects within relatively large (10–13.25 ha) and small (less than 3.25 ha) forest patches located within nonforest agricultural landscapes. Patterns of habitat use in these patches were compared to those in contiguous forest patches larger than 13.25 ha. Analysis of variance revealed significant differences in diversity between large and small woodlots and between contiguous and fragmented landscapes, especially in terms of the numbers of edge and interior species and winter-resident, summer-resident, and year-round birds observed.     

Edge and area effects on avian assemblages and insectivory in fragmented native forests

Disentangling the confounded effects of edge and area in fragmented landscapes is a recurrent challenge for landscape ecologists, requiring the use of appropriate study designs. Here, we examined the effects of forest fragment area and plot location at forest edges versus interiors on native and exotic bird assemblages on Banks Peninsula (South Island, New Zealand). We also experimentally measured with plasticine models how forest fragment area and edge versus interior location influenced the intensity of avian insectivory. Bird assemblages were sampled by conducting 15?min point-counts at paired edge and interior plots in 13 forest fragments of increasing size (0.5?C141?ha). Avian insectivory was measured as the rate of insectivorous bird attacks on plasticine models mimicking larvae of a native polyphagous moth. We found significant effects of edge, but not of forest patch area, on species richness, abundance and composition of bird assemblages. Exotic birds were more abundant at forest edges, while neither edge nor area effects were noticeable for native bird richness and abundance. Model predation rates increased with forest fragmentation, both because of higher insectivory in smaller forest patches and at forest edges. Avian predation significantly increased with insectivorous bird richness and foraging bird abundance. We suggest that the coexistence of native and exotic birds in New Zealand mosaic landscapes enhances functional diversity and trait complementation within predatory bird assemblages. This coexistence results in increased avian insectivory in small forest fragments through additive edge and area effects.     

Habitat amount and quality,not patch size,determine persistence of a woodland-dependent mammal in an agricultural landscape

Context

The classical theory of island biogeography explains loss of species in fragmented landscapes as an effect of remnant patch size and isolation. Recently this has been challenged by the habitat amount and habitat continuum hypotheses, according to which persistence in modified landscapes is related to total habitat amount rather than habitat configuration or the ability of species to use all habitats to varying degrees. Distinguishing between these theories is essential for effective conservation planning in modified landscapes.

Objective

Identify which factors of habitat type, amount and configuration predict the persistence of a keystone woodland specialist, the eastern bettong Bettongia gaimardi, in a fragmented landscape.

Method

In the Midlands region of Tasmania we carried out camera surveys at 62 sites in summer and winter. We included habitat and landscape features to model whether habitat amount or patch size and isolation influenced the presence of the eastern bettong, and to measure effects of habitat quality.

Results

Habitat amount within a 1 km buffer was a better predictor of occupancy than patch size and isolation. Occupancy was also affected by habitat quality, indicated by density of regenerating stems.

Conclusion

Our results support the habitat amount hypothesis as a better predictor of presence. For a species that is able to cross the matrix between remnant patches and utilise multiple patches, the island biogeography concept does not explain habitat use in fragmented landscapes. Our results emphasize the value of small remnant patches for conservation of the eastern bettong, provided those patches are in good condition.

     

Patterns of bird predation on reptiles in small woodland remnant edges in peri-urban north-western Sydney, Australia

The predator–prey relationship plays an integral role in community structure. In the presence of habitat fragmentation, the dynamic interaction among co-existing species may be disrupted. In this paper we investigated the interaction between small skinks resident in open woodland remnants and the predatory birds that cross-forage between the remnants and the surrounding peri-urban matrix. Skinks were found in significantly fewer numbers in the edge of remnants compared to their core. In contrast, predatory birds were in largest numbers at the edge compared to the core of remnants. We found that there was a strong negative correlation between skink numbers and predatory birds (individually and combined) consistent with higher predation pressure in the edge compared to the core of remnants. Strike rates on decoys that mimicked skinks were also higher in the edge compared to core habitats, consistent with higher predation rates in this edge habitat.     

Habitat edge effects decrease litter accumulation and increase litter decomposition in coastal salt marshes

Context

Habitat fragmentation is known to be one of the leading causes of species extinctions, however few studies have explored how habitat fragmentation impacts ecosystem functioning and carbon cycling, especially in wetland ecosystems.

Objectives

We aimed to determine how habitat fragmentation, defined by habitat area and distance from habitat edge, impacts the above-ground carbon cycling and nutrient stoichiometry of a foundation species in a coastal salt marsh.

Methods

We conducted our research in a salt marsh in the Mid-Atlantic United States, where the foundation grass species Spartina patens is being replaced by a more flood-tolerant grass, leading to highly fragmented habitat patches. We quantified decomposition rates, live biomass, and litter accumulation of S. patens at patch edges and interiors. Additionally, we measured relevant characteristics (e.g., habitat area, elevation, microclimate) of S. patens patches.

Results

Habitat edge effects, and not habitat area effects, had distinct impacts on ecosystem functioning. Habitat edges had less litter accumulation, faster decomposition rates, a warmer and drier microclimate, and lower elevations than patch interiors. Patches with low elevation edges had the fastest decomposition rates, while interiors of patches at any elevation had the slowest decomposition rates. Notably, these impacts were not driven by changes in primary production.

Conclusion

Habitat fragmentation impacts the above-ground carbon cycling of S. patens in coastal wetlands by altering litter decomposition, but not primary production, through habitat edge effects. Future research should investigate whether this pattern scales across broader landscapes and if it is observable in other wetland ecosystems.

     

Large termitaria act as refugia for tall trees,deadwood and cavity-using birds in a miombo woodland

Landscape heterogeneity can play an important role in providing refugia and sustaining biodiversity in disturbed landscapes. Large Macrotermes (Isoptera) termite mounds in miombo woodlands form nutrient rich islands that sustain a different suite of woody plant species relative to the woodland matrix. We investigated the role of termitaria in providing habitat for cavity-using birds in miombo woodlands that had been greatly impacted by elephants and fire, by comparing the availability of habitat favored by cavity-using birds (tall trees, trees with deadwood, and cavities) on and off mounds, and then testing its effect on species richness and abundance of cavity-using birds. We surveyed 48 termitaria paired with 48 woodland matrix sites in the breeding season; and 54 matrix-termitarium pairs in the non-breeding season in Chizarira National Park, Zimbabwe. Generalized linear mixed-effects models showed that termitaria harboured significantly higher densities (ha⁻¹) of habitat components considered important for cavity nesting birds. Density of trees >6 m in height and incidence of trees with deadwood was nearly 10 times greater on mounds than in the matrix, and the density of cavities was nine times higher on mounds compared to the matrix. A model selection procedure showed that termitaria provided refugia for cavity-using birds and contributed to the resilience of bird communities through high on-mound densities of trees with deadwood. Large termitaria thus appear to play an important role in maintaining functionally important components of the avifauna in heavily impacted Miombo woodlands.     

Markov models of land cover dynamics in a southern Great Plains grassland region

Grassland regions of the southern Great Plains are fragmented by agricultural activity and many habitat remnants have experienced encroachment by juniper (Juniperus virginiana L.). Recently, many cropland areas have been converted to monoculture grassland (pastures) and enrolled into the Conservation Reserve Program (CRP). Our objectives were to develop spatial and temporal Markov models to characterize land cover dynamics relative to juniper expansion and CRP using aerial photography from 1965, 1981, and 1995. We used landscapes surrounding three Breeding Bird Survey routes with varying levels of juniper encroachment in Oklahoma as study areas. As expected, land cover changes from 1965 to 1995 included increases in juniper woodland, mixed juniper-deciduous woodland, and pastures from CRP activity. Markov models revealed that juniper had a low likelihood of self-replacement in early stages of encroachment. In all areas, relatively little native grassland was lost to juniper encroachment, but other native habitat types such as deciduous woodland were heavily impacted. Transition probabilities for land cover dynamics varied significantly both spatially and temporally. Projections of these raw transition matrices produced widely varying models of future land cover conditions. By modifying the matrices to account for recent and potential socio-political and ecological changes occurring in this region, a number of more plausible land cover scenarios were produced than those resulting from simple projections of raw transition matrices.     

Methodological, temporal and spatial factors affecting modeled occupancy of resident birds in the perennially cultivated landscape of Uttar Pradesh, India

Biodiversity persistence in non-woody tropical farmlands is poorly explored, and multi-species assessments with robust landscape-scale designs are sparse. Modeled species occupancy in agricultural mosaics is affected by multiple factors including survey methods (convenience-based versus systematic), landscape-scale agriculture-related variables, and extent of remnant habitat. Changes in seasonal crops can additionally alter landscape and habitat conditions thereby influencing species occupancy. We investigated how these factors affect modeled occupancy of 56 resident bird species using a landscape-scale multi-season occupancy framework across 24 intensively cultivated and human-dominated districts in Uttar Pradesh state, north India. Convenience-based roadside observations provided considerable differences in occupancy estimates and associations with remnant habitat and intensity of cultivation relative to systematic transect counts, and appeared to bias results to roadside conditions. Modeled occupancy of only open-area species improved with increasing intensity of cultivation, while remnant habitat improved modeled occupancy of scrubland, wetland and woodland species. Strong seasonal differences in occupancy were apparent for most species across all habitat guilds. Further habitat loss will be most detrimental to resident scrubland, wetland and woodland species. Uttar Pradesh’s agricultural landscape has a high conservation value, but will require a landscape-level approach to maintain the observed high species richness. Obtaining ecological information from unexplored landscapes using robust landscape-scale surveys offers substantial advantages to understand factors affecting species occupancy, and is necessary for efficient conservation planning.     

Predator activity and predation on songbird nests on forest-field edges in east-central Illinois

We measured the activity of mammalian predators, numbers of singing male songbirds, and predation rates on nests of songbirds (152 natural, open-cup nests and 380 artificial nests) on 38 250 m transects located along various types of forest-field edges in a wildlife management area in east-central Illinois. We then related these variables to each other and to measures of the vegetative structure of our transects that we anticipated might influence predator activity or predation rates on nests of birds characteristic of edge and shrubland habitats. Mammalian predators, particularly raccoons (Procyon lotor), were abundant in the wildlife area and present on all transects surveyed. We did not find significant relationships among the variables we measured. Rather, rates of nest predation were consistently high (>70%) and generally evenly distributed around our study site. Medium-sized, generalist mammalian predators in the midwestern United States reach their highest population densities in fragmented landscapes with abundant edge habitat, particularly agricultural edges. Areas of natural habitat in these landscapes dominated by agriculture may concentrate predators and act as ecological traps for nesting birds because they attract high densities of breeding birds that are subjected to high rates of nest predation.     

Movement across woodland edges suggests plantations and farmland are barriers to dispersal

Landscape Ecology - The behavioural response of animals to edges between habitat patches and the matrix can influence population dynamics and species persistence in fragmented landscapes. We aimed...     

Landscape perception by forest understory birds in the Atlantic Rainforest: black-and-white versus shades of grey

Even among forest specialists, species-specific responses to anthropogenic forest fragmentation may vary considerably. Some appear to be confined to forest interiors, and perceive a fragmented landscape as a mosaic of suitable fragments and hostile matrix. Others, however, are able to make use of matrix habitats and perceive the landscape in shades of grey rather than black-and-white. We analysed data of 42 Chiroxiphia caudata (Blue Manakin), 10 Pyriglena leucoptera (White-shouldered Fire-eye) and 19 Sclerurus scansor (Rufous-breasted Leaftosser) radio-tracked in the Atlantic Rainforest of Brazil between 2003 and 2005. We illustrate how habitat preferences may determine how species respond to or perceive the landscape structure. We compared available with used habitat to develop a species-specific preference index for each of six habitat classes. All three species preferred old forest, but relative use of other classes differed significantly. S. scansor perceived great contrast between old forest and matrix, whereas the other two species perceived greater habitat continuity. For conservation planning, our study offers three important messages: (1) some forest specialist species are able to persist in highly fragmented landscapes; (2) some forest species may be able to make use of different anthropogenic habitat types to various degrees; whereas (3) others are restricted to the remaining forest fragments. Our study suggests species most confined to forest interiors to be considered as potential umbrella species for landscape-scale conservation planning.     

Use of space by the yellow-footed antechinus, Antechinus flavipes, in a fragmented landscape in South Australia

An understanding of how individual species are able to persist and move within fragmented landscapes is critical for elucidating the effects of fragmentation and aiding in the management of species. Here, we studied movement behaviour of the dasyurid Antechinus flavipes in a heavily fragmented landscape using trapping and radiotracking. We assessed the ability of animals to move within and amongst small (<6 ha) remnants and make use of the matrix, and investigated how females used the available space within remnants. Seventeen between-remnant movements were detected from 428 recaptures, ranging in length from 30 to 720 m and averaging 352 m. Most were by adult males during the breeding season, with 40% more than 500 m. Landscape types traversed would have included exotic pine plantations, open grazed areas and roads. Between-site movements of juveniles were only detected on three occasions. However, few young males were captured as adults, suggesting high dispersal rates and considerable matrix use. Conversely, despite high female recapture rates, again only three between-site movements were recorded. Radiotracking further indicated that females confined foraging to remnants, with occasional forays to isolated trees in paddocks. Female home range areas were similar for remnants and forest (0.04–0.66 ha). A. flavipes is clearly able to persist in very small patches of native vegetation in the landscape studied here. Its long-term persistence appears dependent on the ability of females to maintain a presence in the small remnants, and of unrelated males to move between remnants to breed with resident females. This study illustrates the importance of recognising the occurrence of metapopulations in fragmented landscapes for conservation management.     

Biodiversity functions of urban cemeteries: Evidence from one of the largest Jewish cemeteries in Europe

As the world becomes more urbanized, urban cemeteries may become increasingly valuable for biodiversity conservation as cemeteries are ubiquitous elements of the green infrastructure in cities worldwide. By implementing a multi-taxon approach at different spatial extents, we analyzed habitat functions of a large urban cemetery in Berlin (Weiÿensee Jewish Cemetery) and explored related environmental variables. This cemetery is an outstanding cultural heritage site but it also stands for old urban cemeteries that have progressed to urban woodland, an ecosystem type that exists in many regional and religious contexts. The cemetery provided a habitat for 604 species; species of conservation concern comprised 1.6⿿100% of total species among different groups of taxa (in decreasing order: bats, birds, lichens, bryophytes, carabids, vascular plants, spiders). Species richness and species composition at the plot level were significantly related to differences in management intensity and resulting vegetation structures but differed among taxonomic groups. In vascular plants, carabids and spiders, the species composition varied significantly with habitat age, and there was a set of characteristic species for different age classes in each species group. Our results thus support the use of differentiated management approaches to maintain habitat heterogeneity by allowing wilderness development in some parts of a cemetery while keeping others more open. Since these aims can be combined with efforts to preserve outstanding grave architectures and allow access to visitors, our study indicates ways of reconciling conflicting aims of heritage preservation and biodiversity conservation, a promising perspective for biodiversity conservation in culturally shaped urban landscapes. We conclude that cemeteries provide important cultural ecosystem services within the urban green infrastructure.     

Multi-scale landscape and seascape patterns associated with marbled murrelet nesting areas on the U.S. west coast

Habitat for wide-ranging species should be addressed at multiple scales to fully understand factors that limit populations. The marbled murrelet (Brachyramphus marmoratus), a threatened seabird, forages on the ocean and nests inland in large trees. We developed statistical relationships between murrelet use (occupancy and abundance) and habitat variables quantified across many spatial scales (statewide to local) and two time periods in California and southern Oregon, USA. We also addressed (1) if old-growth forest fragmentation was negatively associated with murrelet use, and (2) if some nesting areas are more important than others due to their proximity to high quality marine habitat. Most landscapes used for nesting were restricted to low elevation areas with frequent fog. Birds were most abundant in unfragmented old-growth forests located within a matrix of mature second-growth forest. Murrelets were less likely to occupy old-growth habitat if it was isolated (> 5 km) from other nesting murrelets. We found a time lag in response to fragmentation, where at least a few years were required before birds abandoned fragmented forests. Compared to landscapes with little tono murrelet use, landscapes with many murrelets were closer to the ocean's bays, river mouths, sandy shores, submarine canyons, and marine waters with consistently high primary productivity. Within local landscapes (≤ 800ha), inland factors limited bird abundance, but at the broadest landscape scale studied (3200 ha), proximity to marine habitat was most limiting. Management should focus on protecting or creating large, contiguous old-growth forest stands, especially in low-elevation areas near productive marine habitat. This revised version was published online in July 2006 with corrections to the Cover Date.