The need for biological realism in habitat modeling: a reinterpretation of Zharikov et al. (2006)

Zharikov et al. (2006: Landscape Ecology 21:107–120) modeled the nest-site habitat use of marbled murrelets (Brachyramphus marmoratus) in Desolation Sound (DS) and Clayoquot Sound (CS), British Columbia. They compared known nest sites, located with radio-telemetry, with randomly-located points within the same areas. Their conclusions suggest that murrelets tended to nest in disproportionately smaller fragments within the more disturbed DS landscape; streams, steeper slopes, and lower elevations were selected in both landscapes; murrelets nested closer to recent clearcuts than would be expected in the DS landscape; and survivorship of nestlings was greater in areas with recent clearcuts and was positively correlated with recent habitat fragmentation. These conclusions are contrary to current management guidelines in British Columbia, and therefore require close scrutiny. Our detailed examination reveals flaws in their use of data, application of modeling, and most seriously, interpretation of the results. Problems include: conceptual errors in the interpretation of models; inappropriate spatial resolution; confusing use and interpretation of fragmentation and patch size data; overemphasis of statistically significant but biologically trivial results; and ignoring some contradictory studies. We conclude that it would be risky to apply the results from Zharikov et al. in the selection of murrelet nesting habitat for management purposes in British Columbia. Our review identifies issues that may arise in other ecological modeling studies and stresses the need for biological realism in addition to statistical rigour.     

Habitat Selection and Breeding Success in a Forest-nesting Alcid, the Marbled Murrelet, in Two Landscapes with Different Degrees of Forest Fragmentation

We studied habitat selection and breeding success in marked populations of a protected seabird (family Alcidae), the marbled murrelet (Brachyramphus marmoratus), in a relatively intact and a heavily logged old-growth forest landscape in south-western Canada. Murrelets used old-growth fragments either proportionately to their size frequency distribution (intact) or they tended to nest in disproportionately smaller fragments (logged). Multiple regression modelling showed that murrelet distribution could be explained by proximity of nests to landscape features producing biotic and abiotic edge effects. Streams, steeper slopes and lower elevations were selected in both landscapes, probably due to good nesting habitat conditions and easier access to nest sites. In the logged landscape, the murrelets nested closer to recent clearcuts than would be expected. Proximity to the ocean was favoured in the intact area. The models of habitat selection had satisfactory discriminatory ability in both landscapes. Breeding success (probability of nest survival to the middle of the chick rearing period), inferred from nest attendance patterns by radio-tagged parents, was modelled in the logged landscape. Survivorship was greater in areas with recent clearcuts and lower in areas with much regrowth, i.e. it was positively correlated with recent habitat fragmentation. We conclude that marbled murrelets can successfully breed in old-growth forests fragmented by logging.     

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.     

Regeneration Strategies,Disturbance and Plant Interactions as Organizers of Vegetation Spatial Patterns in a Pine Forest

To determine how vegetation pattern in early successional forests may be related to plant traits and types of disturbance, we measured percent cover of individual taxa annually in a South Carolina Pinus elliottii forest, starting one year before, and ending four years after harvest and tree girdling disturbances were applied. The 17 most important taxa surveyed were grouped into four regeneration strategies chosen a priori, and the spatial patterns of these groups and of the soil were investigated using global variability, semivariograms and kriged maps. We also examined spatial correlations across years, across taxa, and between species and soil disturbance. Seed bank taxa represented by Dichanthelium spp. increased rapidly and formed large patches, and then quickly declined. Taxa that regenerate by newly dispersed seeds, represented by Rhus copallina and Rubus spp. occurred at first in a few patches, and became widespread later. Stump sprouters, represented by Quercus spp. and Myrica cerifera, had rapid increases in cover, but their spatial patterns were largely determined by their pre-disturbance patterns. Prunus serotina, which relies on both sprouting and dispersed seed, had moderate cover and a random distribution. Within-species temporal correlation of spatial pattern was lower in girdled than in harvested plots, and was not clearly related to regeneration strategy. Forest floor disturbance was patchy and affected the pattern of Dichanthelium spp. in the harvested plots. Negative correlations between herbs and woody plants in harvested plots reflected the role of biotic (i.e., successional) filters on vegetation pattern. Surprisingly, no spatial correlations were detected between the nitrogen fixer, Myrica cerifera and other taxa in this N-limited system. In comparing the spatial and temporal patterns, we found kriged maps more informative than analysis of semivariograms alone. The maps and correlation statistics demonstrated that regeneration traits, spatial patterns of soil disturbances, and interactions among taxa influence dynamics of the spatial patterns of the plants. We also demonstrated that disturbance types affected the importance and interactions among these three factors, and caused different spatial patterns of the plant taxa.     

Modeling fire and landform influences on the distribution of old-growth pinyon-juniper woodland

Expansion of Pinus and Juniperus species into shrub steppe in semi-arid regions of the western United States has been widely documented and attributed in part to fire exclusion. If decreased fire frequency has been an important cause of woodland expansion, one would expect to find age structures dominated by younger trees on more fire-prone sites, with old-growth pinyon-juniper woodland limited to sites with lower fire risk. We compared current old-growth distribution with spatial models for fire risk in a 19-km² watershed in central Nevada, USA. Multiple GIS models were developed to represent fire susceptibility, according to abiotic factors representing fuels and topographic barriers to fire spread. We also developed cellular automata models to generate fire susceptibility surfaces that additionally account for neighborhood effects. Rule-based GIS models failed to predict old-growth distribution better than random models. Cellular automata models incorporating spatial heterogeneity of site productivity predicted old-growth distribution better than random models but with low accuracy, ranging from 58% agreement at the single-pixel (0.09-ha) scale to 80% agreement for 20-pixel neighborhoods. The best statistical model for predicting old-growth occurrence included the negative effect of topographic convergence index (local wetness), and the positive effects of solar insolation and proximity to rock outcrops. Results support the hypothesis that old-growth woodlands in the Great Basin are more likely to occur on sites with low fire risk. However, weak relationships suggest that old-growth woodlands have not been confined to fire-safe sites. Conservation efforts should consider the landscape context of old-growth woodlands across a broad landscape, with an emphasis on conserving landscape variability in tree age structure.     

Old-growth forest landscape transitions from pre-European settlement to present

We conducted a multi-temporal spatial analysis of forest cover for a 9600 ha landscape in northern Wisconsin, U.S.A., using data from pre-European settlement (1860s), post-settlement (1931), and current (1989) periods. Using GIS we have shown forest landscape changes and trajectories that have been generally described in aggregate for the norther Great Lake States region. We created the pre-European settlement map from the witness tree data of the original federal General Land Office survey notes. The 1931 cover was produced from the Wisconsin Land Economic Inventory, and the 1989 cover map was based on color infrared photography. We used GIS to analyze 1) land area occupied by different forest types at different dates, 2) temporal transitions between dates and their driving proceses, and 3) successional trajectories with landforms and spatial associations of forest types. Over the 120 year period, forest cover has changed from a landscape dominated by old-growth hemlock (Tsuga canadensis) and hardwood forests (Acer saccharum, Betula alleghaniensis) to largely second-growth hardwoods and conifers. The former dominant hemlock is largely eliminated from the landscape. From 1860 to 1931, large-scale disturbances associated with logging were the dominant processes on the landscape. Early successional forest types covered much of the landscape by the 1930s. From 1931 to 1989, succession was the dominant process driving forest transitions as forest types succeeded to a diverse group of upland hardwood and conifer forest types. If successional trajectories continue, a more homogeneous landscape may develop comprised of both a northern hardwood type dominated by sugar maple, and a boreal conifer/hardwood forest.     

Is bird incidence in Atlantic forest fragments influenced by landscape patterns at multiple scales?

The degree to which habitat fragmentation affects bird incidence is species specific and may depend on varying spatial scales. Selecting the correct scale of measurement is essential to appropriately assess the effects of habitat fragmentation on bird occurrence. Our objective was to determine which spatial scale of landscape measurement best describes the incidence of three bird species (Pyriglena leucoptera, Xiphorhynchus fuscus and Chiroxiphia caudata) in the fragmented Brazilian Atlantic forest and test if multi-scalar models perform better than single-scalar ones. Bird incidence was assessed in 80 forest fragments. The surrounding landscape structure was described with four indices measured at four spatial scales (400-, 600-, 800- and 1,000-m buffers around the sample points). The explanatory power of each scale in predicting bird incidence was assessed using logistic regression, bootstrapped with 1,000 repetitions. The best results varied between species (1,000-m radius for P. leucoptera; 800-m for X. fuscus and 600-m for C. caudata), probably due to their distinct feeding habits and foraging strategies. Multi-scale models always resulted in better predictions than single-scale models, suggesting that different aspects of the landscape structure are related to different ecological processes influencing bird incidence. In particular, our results suggest that local extinction and (re)colonisation processes might simultaneously act at different scales. Thus, single-scale models may not be good enough to properly describe complex pattern–process relationships. Selecting variables at multiple ecologically relevant scales is a reasonable procedure to optimise the accuracy of species incidence models.     

Pre-EuroAmerican settlement forests in Redwood National Park,California, USA: a reconstruction using line summaries in historic land surveys

Extensive logging in the twentieth century destroyed much of the coniferous forests in the lower Redwood Creek basin of Redwood National Park. Restoration of cutover lands requires the identification of historical, pre-logging reference conditions. Field notes from the original Public Land Surveys were used to reconstruct the pre-EuroAmerican settlement forests. Most reconstructive studies based on historic surveys rely on bearing tree evidence over large areas to determine vegetation patterns over several hundreds to thousands of square kilometers. Due to the small size of the study area (approximately 200 km²), bearing tree evidence could not accurately reconstruct the vegetation at this scale. Instead, lists of the overstory and understory vegetation for each surveyed mile (line summaries) were employed. Analysis of line summaries evidence identified the historical importance, geographical range, and environmental influences on woody species and vegetation communities. Topography, especially elevation, and soil texture were significantly correlated with plot-scale ordination scores derived from non-metric multidimensional scaling. The influence of topography and distance to ocean coast on the historical distribution of dominant woody species concurs with findings from present-day field studies of local and regional old-growth forest. A comparison with present-day vegetation maps revealed that coast redwood (Sequoia sempervirens), Douglas fir (Pseudotsuga menziesii), Sitka spruce (Picea sitchensis), and red alder (Alnus rubra) experienced the most substantive changes in the vegetation as a result of twentieth century land use activities.     

The Importance of Spatial Scale in Habitat Models: Capercaillie in the Swiss Alps

The role of scale in ecology is widely recognized as being of vital importance for understanding ecological patterns and processes. The capercaillie (Tetrao urogallus) is a forest grouse species with large spatial requirements and highly specialized habitat preferences. Habitat models at the forest stand scale can only partly explain capercaillie occurrence, and some studies at the landscape scale have emphasized the role of large-scale effects. We hypothesized that both the ability of single variables and multivariate models to explain capercaillie occurrence would vary with the spatial scale of the analysis. To test this hypothesis, we varied the grain size of our analysis from 1 to just over 1100 hectares and built univariate and multivariate habitat suitability models for capercaillie in the Swiss Alps. The variance explained by the univariate models was found to vary among the predictors and with spatial scale. Within the multivariate models, the best single-scale model (using all predictor variables at the same scale) worked at a scale equivalent to a small annual home range. The multi-scale model, in which each predictor variable was entered at the scale at which it had performed best in the univariate model, did slightly better than the best single-scale model. Our results confirm that habitat variables should be included at different spatial scales when species-habitat relationships are investigated.     

The influence of forest harvesting on landscape spatial patterns and old-growth-forest fragmentation in southeast British Columbia

Habitat fragmentation is considered one of the major conservation issues of recent decades. We tested predictions of landscape patterns in a 352,253-ha managed forest area in southeast British Columbia. We did this by focussing on forest fragmentation concerns among old-growth, harvest, and wildfire patches in 44 delineated landscapes using patch indices as measures of landscape pattern. We found no significant association between amount of harvesting and 15 old-growth patch indices. Comparisons among patch types revealed that amounts and spatial patterns of harvest patches differed little from amounts and spatial patterns of old-growth patches in control landscapes. Variability indices revealed similar variability between harvest patches and old-growth patches, and more variability between harvest patches and wildfire patches. Little of the evidence gathered in this study supported predictions of fragmentation of old-growth spatial patterns, or predicted differences between harvest spatial patterns and more naturally occurring spatial patterns. We suggest these results could be due to the relatively small amounts of harvesting and old-growth forest in these landscapes, and therefore habitat amount may be a more important factor than spatial configuration of patches in these landscapes.     

Effect of matrix habitat on the spread of flea beetle introductions for biological control of leafy spurge

Matrix habitats are known to influence the movement patterns of a variety of species but it is less well known whether these effects have strong implications for spatial population dynamics, including the spread of biological introductions. Using a spatially explicit simulation model parameterized with empirical data, we examine how grass and shrub matrix habitats, each offering different resistance to dispersal, influence the spread and impact of a biocontrol agent, Aphthona lacertosa, on the invasive weed, leafy spurge. Model predictions indicate that differential responses to matrix habitat have little effect on the agent’s spread over the study landscape and this is supported by statistical models fit to observed A. lacertosa incidence on the same landscape. Subsequent experimentation with the simulation model suggested that A. lacertosa colonization rates were largely unaffected by increases in amount of the more restrictive shrub matrix. However, simulations of an hypothetical species with greater overall dispersal ability but reduced dispersal rate through shrub matrix showed that colonization rates were noticeably reduced when the percentage of shrub matrix on the landscape approached 50%. Combined these results suggest that some tailoring of release strategies may be required to accommodate the unique dispersal capabilities of different biocontrol agents on particular release landscapes, but for A. lacertosa there appears to be little effect of matrix habitat structure on rates of spread.     

Interaction of food resources and landscape structure in determining the probability of patch use by carnivores in fragmented landscapes

Studies on the distribution of mammalian carnivores in fragmented landscapes have focused mainly on structural aspects such as patch and landscape features; similarly, habitat connectivity is usually associated with landscape structure. The influence of food resources on carnivore patch use and the important effect on habitat connectivity have been overlooked. The aim of this study is to evaluate the relative importance of food resources on patch use patterns and to test if food availability can overcome structural constraints on patch use. We carried out a patch-use survey of two carnivores: the beech marten (Martes foina) and the badger (Meles meles) in a sample of 39 woodland patches in a fragmented landscape in central Italy. We used the logistic model to investigate the relative effects on carnivore distribution of patch, patch neighbourhood and landscape scale variables as well as the relative abundance of food resources. Our results show how carnivore movements in fragmented landscapes are determined not only by patch/landscape structure but also by the relative abundance of food resources. The important take-home message of our research is that, within certain structural limits (e.g. within certain limits of patch isolation), by modifying the relative amount of resources and their distribution, it is possible to increase suitability in smaller/relatively isolated patches. Conversely, however, there are certain thresholds above which an increase in resources will not achieve high probability of presence. Our findings have important and generalizable consequences for highly fragmented landscapes in areas where it may not be possible to increase patch sizes and/or reduce isolation so, for instance, forest regimes that will increase resource availability could be implemented. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Prediction of the spatial distribution and relative abundance of ground-dwelling mammals using remote sensing imagery and simulation models

We present an approach that allows current, retrospective and future relative abundances of mammal species to be predicted across landscapes. A spatial generalized regression model of species relative abundance based on habitat quality and time since disturbance was combined with coverages of the spatial distribution of habitat quality derived from a simulation model which predicts the historical and future spatial arrangement of forest habitat. The strength of this approach is that the input habitat data can be derived as part of a standard forest inventory mapping program with the addition of high spatial resolution remote sensing imagery. Furthermore, it operates at the scale used for wildlife management in Australia, which makes it widely applicable. To demonstrate the approach we use data collected over 20 years on the long-nosed potoroo (Potorous tridactylus) and the large wallabies (red-necked wallaby, Macropus rufogriseus, and swamp wallaby, Wallabia bicolor) and their habitats following wildfire. Results indicate the relative abundance of the potoroo has increased, from initially sparse numbers of less than 0.5 % of plot-night occurrences to close to 3% approximately twenty years after a major fire event. The large wallabies by contrast decreased in relative abundance from about 20% since the major fire event. Presently the relative abundance of large wallabies was modelled at 2% of plot-nights with tracks which was very low. Predictions of future relative abundance without additional disturbance were low, with the region likely to be unsuitable for the species in the next 5 years. These models offer tools for investigating the current and historical abundances of key species which can provide data to forest managers for wildlife management thereby translating current scientific understanding into tools suitable for every-day use by forest managers. This revised version was published online in July 2006 with corrections to the Cover Date.     

Post-storm surveys reveal large-scale spatial patterns and influences of site factors,forest structure and diversity in endemic bark-beetle populations

The storm that struck France on december 26^th and 28^th 1999 felled 140 million m³ of timber and had a high economic, social and landscape impact. This event offered the opportunity to study large-scale patterns in populations of forest insect pests that would benefit from the abundant breeding material. A large-scale survey was carried out in France in 2000 to sample the most frequently observed species developing on spruce (Ips typographus, Pityogene schalcographus) and pine (Tomicus piniperda, Ips sexdentatus) in 898 locations distributed throughout wind-damaged areas. The local abundance of each species scored on a 0 to 5 scale was analysed using geostatistical estimators to explore the extent and intensity of spatial autocorrelation, and was related to site, stand, and neighbourhood landscape metrics of the forest cover (in particular the interconnection with broadleaf forest patches) found within dispersal distance. All species but I. sexdentatus, which was much less abundant, displayed large-scale spatial dependence and regional variations in abundance. Lower infestation levels per tree (windfalls and standing trees) were observed in stands with a high proportion of wind-damaged trees, which was interpreted as the result of beetles distributing themselves among the available breeding material. More infestations were observed in wind-broken trees as compared to wind-felled trees. More importantly, populations showed significant relationships with the structure of coniferous stands (in particular with the number of coniferous patches). T. piniperda population levels were negatively correlated to the amount of coniferous edge shared with broadleaf forest patches, possibly because of the disruptive effect of non-host volatiles on host-finding processes at the landscape-scale. The differences observed between species regarding patterns and relationships to site, stand, and forest cover characteristics are discussed in relation to the ecological characteristics of each species.     

Clone identification and genetic relationship among vine cacti from the genera Hylocereus and Selenicereus based on RAPD analysis

Clones of Hylocereus and of Selenicereus species were distinguished from each other by the banding pattern generated by one to nine 10-mer oligonucleotide primers in the random amplified polymorphic DNA (RAPD) reaction. RAPD analysis was also applied to estimate the genetic relationship among five Hylocereus and nine Selenicereus species. A dendrogram was constructed based on a data matrix of 173 polymorphic bands originated by nine primers. Two groups were identified, one consisting of Hylocereus species and the other consisting of Selenicereus species. These results are consistent with the accepted taxonomic classification of the genera studied. The principal coordinate analysis (PCO), i.e. the plot drawn on the basis of the RAPD data, clearly distinguished between three groups, namely, Hylocereus species, S. megalanthus and the rest of the Selenicereus species studied. PCO thus strongly support the notion that the tetraploid S. megalanthus is an exception among the Selenicereus group. The RAPD results support our hypothesis regarding the allopolyploid (rather than autopolyploid) origin of this species.     

Dispersion of kangaroo rat mounds at multiple scales in New Mexico,USA

Burrowing mammals create disturbances that increase the ecological heterogeneity of landscapes. In desert systems, banner-tailed kangaroo rats (Dipodomys spectabilis) construct large mounds that greatly influence the spatial patterning of soils, plants, and animals. The overall effects of the patches generated by D. spectabilis depend on the dispersion patterns of the mounds; these patterns may be sensitive to scale and landscape position. We examined the distribution of D. spectabilis mounds across multiple scales in four 40-ha grassland plots in New Mexico, USA. We used Ripley's K-function for our point-pattern analysis. The dispersion patterns of mounds were generally scale-sensitive but depended somewhat on plot-level densities, which were related to topographic position and grazing history. Mound spacing was either regular or random at small scales (0–50 m), random or aggregated at intermediate scales (50–300 m), and aggregated at large scales (300–3000 m). This scale-dependency of pattern reflected spatial domains in which different biotic (territoriality, dispersal, grazing) and abiotic (soil texture and drainage) factors exerted strong influences. How other organisms perceive the spatial patterning of mounds will depend on the extent of their movements. Patches may appear regular to one species but aggregated to another. The dispersion of D. spectabilis mounds also has implications for the spatial structuring of desert rodent communities. D. spectabilis excludes smaller species of kangaroo rats from areas around their mounds; they create spatial heterogeneity in behavioral dominance that may influence the distribution of subordinate species at multiple scales.     

Study of long-distance movement of Plum pox virus (Sharka) as an alternative resistance-evaluation method in Prunus

During the breeding programs for Plum pox virus (PPV, Sharka) resistance in Prunus, the evaluation of the new releases through symptoms observation on leaves has been contradictory and represents one of the main handicaps in these programs. In order to increase the accuracy of this traditional evaluation method, we here analyze an alternative method based on the study of the ability of a genotype to allow the long-distance movement of the virus through its vascular vessels. Two different plant models have been assayed: (a) in Model I, the inoculation was performed in the ‘GF305’ rootstock with a later grafting of the genotype under evaluation and a scion of healthy control ‘GF305’, to evaluate the long-distance movement through the studied genotype from the rootstock to the scion (xylem transport), and (b) in Model II, the inoculation with ‘GF305’ diseased scions was performed by grafting these diseased scions onto the studied genotypes, which were grafted onto healthy ‘GF305’ peach seedlings, to evaluate the long-distance movement through the studied genotype from the scion to the rootstock (phloem transport). The results show that, regardless of the presence of symptoms, susceptible genotypes allowed the movement of the virus through their vascular vessels in both directions studied. However, the resistant apricot ‘Stark Early Orange’ did not allow this movement. We propose the study of the ability of a genotype to allow the long-distance movement of the virus as an alternative and more accurate method for the evaluation of PPV resistance. However, this protocol is much more tedious than the traditional one and could be used mainly in the evaluation of a reduced number of more interesting genotypes.     

Evaluating effects of low quality habitats on regional population growth in <Emphasis Type="Italic">Peromyscus leucopus</Emphasis>: Insights from field-parameterized spatial matrix models

Due to complex population dynamics and source–sink metapopulation processes, animal fitness sometimes varies across landscapes in ways that cannot be deduced from simple density patterns. In this study, we examine spatial patterns in fitness using a combination of intensive field-based analyses of demography and migration and spatial matrix models of white-footed mouse (Peromyscus leucopus) population dynamics. We interpret asymptotic population growth rates from these spatial models as fitness-based measures of habitat-quality and use elasticity analysis to further explore model behavior and the roles of migration. In addition, we compare population growth rates at the spatial scale of single habitats and the landscape-level scale at which these habitats are assembled. To this end, we employ emerging techniques in multi-scale estimation of demography and movement and recently described vec-permutation methods for spatial matrix notation and analysis. Our findings indicate that the loss of low quality habitats or reductions in movement from these habitats into higher quality areas could negatively affect landscape-level population fitness.     

The effects of timber harvesting on the structure and composition of adjacent old-growth coast redwood forest,California, USA

Data collected across timber harvest boundaries on nine sites within the Redwood National and State Park management area in California, USA, were used to estimate the effective size of old-growth coast redwood preserves. Fourteen variables related to stand structure and composition, wildlife habitat, and physical environment were significantly correlated to distance from the timber harvest boundary using multiple regression analysis. A maximum depth of edge influence of 200 m was determined for variables exhibiting a significant correlation to the distance from the harvest edge. A spatial analysis using ArcView indicated that 53% of the old growth preserved within the study area was influenced by edge conditions, leaving 47% as effective old-growth.This revised version was published online in May 2005 with corrections to the Cover Date.     

Site fidelity of female caribou at multiple spatial scales

Studies of site fidelity have been hampered by arbitrary designations of spatial scale and the lack of null models for comparison. We generated null expectations of fidelity at different scales from the distribution of radio-tracked animals in a population. We applied the models to space use of satellite-tracked caribou (Rangifer tarandus caribou), the most vagile nonvolant terrestrial animal, from populations representing sedentary and migratory ecotypes. We compared distances between consecutive-year locations of adult females to expectations based on the total range and seasonal range of each population. At the scale of the total range, sedentary and migratory caribou displayed remarkably similar philopatry, despite a 30-fold difference in size of their population ranges, from time of calving (late May) to breeding (late October). The most intense fidelity occurred during post-calving when, on average, sedentary and migratory females returned to as near as 6.7 km and 123 km, respectively, of locations occupied the previous year. At the scale of the seasonal range, the ecotypes differed. Sedentary caribou still displayed fidelity from calving to breeding; migratory caribou exhibited fidelity only during late autumn. For migratory, but not sedentary caribou, inter-year distances during winter were negatively correlated with age, implying that older females were more philopatric. We conclude that reproductive activities delimit the season of fidelity of female caribou of both ecotypes, and that scale-dependent ecotypic differences in fidelity may reflect different factors of population limitation. A spatially-explicit approach to site fidelity is essential for synthesizing patterns across studies.