Modeling ecological and evolutionary processes in spatially structured populations

Many species inhabit fragmented landscapes, resulting either from anthropogenic or from natural processes. The ecological and evolutionary dynamics of spatially structured populations are affected by a complex interplay between endogenous and exogenous factors. The metapopulation approach, simplifying the landscape to a discrete set of patches of breeding habitat surrounded by unsuitable matrix, has become a widely applied paradigm for the study of species inhabiting highly fragmented landscapes. In this thesis, I focus on the construction of biologically realistic models and their parameterization with empirical data, with the general objective of understanding how the interactions between individuals and their spatially structured environment affect ecological and evolutionary processes in fragmented landscapes. I study two hierarchically structured model systems, which are the Glanville fritillary butterfly in the Åland Islands, and a system of two interacting aphid species in the Tvärminne archipelago, both being located in South-Western Finland. The interesting and challenging feature of both study systems is that the population dynamics occur over multiple spatial scales that are linked by various processes. My main emphasis is in the development of mathematical and statistical methodologies. For the Glanville fritillary case study, I first build a Bayesian framework for the estimation of death rates and capture probabilities from mark-recapture data, with the novelty of accounting for variation among individuals in capture probabilities and survival. I then characterize the dispersal phase of the butterflies by deriving a mathematical approximation of a diffusion-based movement model applied to a network of patches. I use the movement model as a building block to construct an individual-based evolutionary model for the Glanville fritillary butterfly metapopulation. I parameterize the evolutionary model using a pattern-oriented approach, and use it to study how the landscape structure affects the evolution of dispersal. For the aphid case study, I develop a Bayesian model of hierarchical multi-scale metapopulation dynamics, where the observed extinction and colonization rates are decomposed into intrinsic rates operating specifically at each spatial scale. In summary, I show how analytical approaches, hierarchical Bayesian methods and individual-based simulations can be used individually or in combination to tackle complex problems from many different viewpoints. In particular, hierarchical Bayesian methods provide a useful tool for decomposing ecological complexity into more tractable components.

Flight metabolic rate in the Glanville fritillary butterfly

Dispersal is a highly important life history trait. In fragmented landscapes the long-term persistence of populations depends on dispersal. Evolution of dispersal is affected by costs and benefits and these may differ between different landscapes. This results in differences in the strength and direction of natural selection on dispersal in fragmented landscapes. Dispersal has been shown to be a nonrandom process that is associated with traits such as flight ability in insects. This thesis examines genetic and physiological traits affecting dispersal in the Glanville fritillary butterfly (Melitaea cinxia). Flight metabolic rate is a repeatable trait representing flight ability. Unlike in many vertebrates, resting metabolic rate cannot be used as a surrogate of maximum metabolic rate as no strong correlation between the two was found in the Glanville fritillary. Resting and flight metabolic rate are affected by environmental variables, most notably temperature. However, only flight metabolic rate has a strong genetic component. Molecular variation in the much-studied candidate locus phosphoglucose isomerase (Pgi), which encodes the glycolytic enzyme PGI, has an effect on carbohydrate metabolism in flight. This effect is temperature dependent: in low to moderate temperatures individuals with the heterozygous genotype at the single nucleotide polymorphism (SNP) AA111 have higher flight metabolic rate than the common homozygous genotype. At high temperatures the situation is reversed. This finding suggests that variation in enzyme properties is indeed translated to organismal performance. High-resolution data on individual female Glanville fritillaries moving freely in the field were recorded using harmonic radar. There was a strong positive correlation between flight metabolic rate and dispersal rate. Flight metabolic rate explained one third of the observed variation in the one-hour movement distance. A fine-scaled analysis of mobility showed that mobility peaked at intermediate ambient temperatures but the two common Pgi genotypes differed in their reaction norms to temperature. As with flight metabolic rate, heterozygotes at SNP AA111 were the most active genotype in low to moderate temperatures. The results show that molecular variation is associated with variation in dispersal rate through the link of flight physiology under the influence of environmental conditions. The evolutionary pressures for dispersal differ between males and females. The effect of flight metabolic rate on dispersal was examined in both sexes in field and laboratory conditions. The relationship between flight metabolic rate and dispersal rate in the field and flight duration in the laboratory were found to differ between the two sexes. In females the relationship was positive, but in males the longest distances and flight durations were recorded for individuals with low flight metabolic rate. These findings may reflect male investment in mate locating. Instead of dispersing, males with high flight metabolic rate may establish territories and follow a perching strategy when locating females and hence move less on the landscape level. Males with low metabolic rate may be forced to disperse due to low competitive success or may show adaptations to an alternative strategy: patrolling. In the light of life history trade-offs and the rate of living theory having high metabolic rate may carry a cost in the form of shortened lifespan. Experiments relating flight metabolic rate to longevity showed a clear correlation in the opposite direction: high flight metabolic rate was associated with long lifespan. This suggests that individuals with high metabolic rate do not pay an extra physiological cost for their high flight capacity, rather there are positive correlations between different measures of fitness. These results highlight the importance of condition.

Who really ate the fruit? A novel approach to camera trapping for quantifying frugivory by ruminants

Tropical forest ruminants disperse several plants; yet, their effectiveness as seed dispersers is not systematically quantified. Information on frequency and extent of frugivory by ruminants is lacking. Techniques such as tree watches or fruit traps adapted from avian frugivore studies are not suitable to study terrestrial frugivores, and conventional camera traps provide little quantitative information. We used a novel time-delay camera-trap technique to assess the effectiveness of ruminants as seed dispersers for Phyllanthus emblica at Mudumalai, southern India. After being triggered by animal movement, cameras were programmed to take pictures every 2 min for the next 6 min, yielding a sequence of four pictures. Actual frugivores were differentiated from mere visitors, who did not consume fruit, by comparing the number of fruit remaining across the time-delay photograph sequence. During a 2-year study using this technique, we found that six terrestrial mammals consumed fallen P. emblica fruit. Additionally, seven mammals and one bird species visited fruiting trees but did not consume fallen fruit. Two ruminants, the Indian chevrotain Moschiola indica and chital Axis axis, were P. emblica's most frequent frugivores and they accounted for over 95% of fruit removal, while murid rodents accounted for less than 1%. Plants like P. emblica that are dispersed mainly by large mammalian frugivores are likely to have limited ability to migrate across fragmented landscapes in response to rapidly changing climates. We hope that more quantitative information on ruminant frugivory will become available with a wider application of our time-delay camera-trap technique.

Contrained linear spectral unmixing technique for regional land cover mapping using MODIS data

Over the last few decades, there has been a significant land cover (LC) change across the globe due to the increasing demand of the burgeoning population and urban sprawl. In order to take account of the change, there is a need for accurate and up- to-date LC maps. Mapping and monitoring of LC in India is being carried out at national level using multi-temporal IRS AWiFS data. Multispectral data such as IKONOS, Landsat- TM/ETM+, IRS-1C/D LISS-III/IV, AWiFS and SPOT-5, etc. have adequate spatial resolution (~ 1m to 56m) for LC mapping to generate 1:50,000 maps. However, for developing countries and those with large geographical extent, seasonal LC mapping is prohibitive with data from commercial sensors of limited spatial coverage. Superspectral data from the MODIS sensor are freely available, have better temporal (8 day composites) and spectral information. MODIS pixels typically contain a mixture of various LC types (due to coarse spatial resolution of 250, 500 and 1000 m), especially in more fragmented landscapes. In this context, linear spectral unmixing would be useful for mapping patchy land covers, such as those that characterise much of the Indian subcontinent. This work evaluates the existing unmixing technique for LC mapping using MODIS data, using end- members that are extracted through Pixel Purity Index (PPI), Scatter plot and N-dimensional visualisation. The abundance maps were generated for agriculture, built up, forest, plantations, waste land/others and water bodies. The assessment of the results using ground truth and a LISS-III classified map shows 86% overall accuracy, suggesting the potential for broad-scale applicability of the technique with superspectral data for natural resource planning and inventory applications.

Ecological dynamics of extinct species in empty habitat networks. 1. The role of habitat pattern and quantity, stochasticity and dispersal

We examined a remnant host plant (Primula veris L.) habitat network that was last inhabited by the rare butterfly Hamearis lucina L. in north Wales in 1943, to assess the relative contribution of several spatial parameters to its regional extinction. We first examined relationships between P. veris characteristics and H. lucina eggs in surviving H. lucina populations, and used these to predict the suitability and potential carrying capacity of the habitat network in north Wales. This resulted in an estimate of roughly 4500 eggs (ca 227 adults). We developed a discrete space, discrete time metapopulation model to evaluate the relative contribution of dispersal distance, habitat and environmental stochasticity as possible causes of extinction. We simulated the potential persistence of the butterfly in the current network as well as in three artificial (historical and present) habitat networks that differed in the quantity (current and X3) and fragmentation of the habitat (current and aggregated). We identified that reduced habitat quantity and increased isolation would have increased the probability of regional extinction, in conjunction with environmental stochasticity and H. lucina's dispersal distance. This general trend did not change in a qualitative manner when we modified the ability of dispersing females to stay in, and find suitable habitats (by changing the size of the grid cells used in the model). Contrary to most metapopulation model predictions, system persistence declined with increasing migration rate, suggesting that the mortality of migrating individuals in fragmented landscapes may pose significant risks to system-wide persistence. Based on model predictions for the present landscape we argue that a major programme of habitat restoration would be required for a re-established metapopulation to persist for > 100 years.

The investigation of edge effects within ragmented forest islands in Short Hills Provincial Park

It has been well documented, within the field of landscape ecology, that terrestrial fragmentation contributes to increased heterogeneity at the landscape level. It has also been observed that elevated areas of edge habitat occur within fragmented landscapes. Spatial and temporal edge effects were investigated in four areas designated as Nature Reserve Zones within Short Hills Provincial Park, near St. Catharines, Ontario. Random sampling along exposed edges was performed on trees and saplings, at 5 and 25 ill edge depths, using the point-centred quarter method. Diameter at breast height (dbh) and distance from point measurements were used to establish relative density, dominance, frequency and importance value. One-way analyses of variance were used on dbh measurements of tree species and Chi-Square contingency tables were used on size class distributions of saplings species to determine significant differences between 5 and 25 metres. Qualitative comparisons of importance values were also used to determine differences between 5 and 25 metres as well as between trees and saplings. These statistical and qualitative comparisons suggest that a significant overall spatial edge effect is currently exhibited by fragmented wooded islands within the park. The major species of the park, Acersaccharuln, may be exhibiting a temporal edge effect. The heterogeneous nature of the park may be of importance in understanding this area as a complex, ecological system. It is possible that the remaining forest tracts of the park have been affected, and continue to be affected by previous disturbances. Based on these findings, recommendations are made to the Ontario Ministry of Natural Resources concerning the management of Short Hills Provincial Park in accordance with their 1990 proposed Management Plan.

Identifying time lags in the restoration of grassland butterfly communities: a multi-site assessment

Although grasslands are crucial habitats for European butterflies, large-scale declines in quality and area have devastated many species. Grassland restoration can contribute to the recovery of butterfly populations, although there is a paucity of information on the long-term effects of management. Using eight UK data sets (9-21 years), we investigate changes in restoration success for (1) arable reversion sites, were grassland was established on bare ground using seed mixtures, and (2) grassland enhancement sites, where degraded grasslands are restored by scrub removal followed by the re-instigation of cutting/grazing. We also assessed the importance of individual butterfly traits and ecological characteristics in determining colonisation times. Consistent increases in restoration success over time were seen for arable reversion sites, with the most rapid rates of increase in restoration success seen over the first 10 years. For grasslands enhancement there were no consistent increases in restoration success over time. Butterfly colonisation times were fastest for species with widespread host plants or where host plants established well during restoration. Low mobility butterfly species took longer to colonise. We show that arable reversion is an effective tool for the management of butterfly communities. We suggest that as restoration takes time to achieve, its use as a mitigation tool against future environmental change (i.e. by decreasing isolation in fragmented landscapes) needs to take into account such time lags.

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.

Can agroforest woodlots work as stepping stones for birds in the Atlantic forest region?

In fragmented landscapes, agroforest woodlots can potentially act as stepping stones facilitating movement between forest fragments. We assessed the influence of agroforest woodlots on bird distribution and diversity in the Atlantic forest region (SE Brazil), and also tested which categories of species can use different types of connection elements, and whether this use is influenced by the distance to large forest patches. We studied two fragmented landscapes, with and without stepping stones linking large fragments, and one forested landscape. Using a point count, a bird survey was undertaken in the fragmented landscapes in five different elements: large remnants (> 400 ha), agroforest woodlots (0.4-1.1 ha), small patches (0.5-7 ha), riparian corridor, and pasture areas (the main matrix). Generalist and open-area species were commonly observed in the agroforest system or other connection elements, whereas only a few forest species were present in these connections. For the latter species, the distance of woodlots to large patches was essential to determine their richness and abundance. Based on our results and data from literature, we suggest that there is an optimal relationship between the permeability of the matrix and the efficiency of stepping stones, which occurs at intermediate degrees of matrix resistance, and is species-dependent. Because the presence of agroforest system favors a higher richness of generalist species, they appeared to be more advantageous for conservation than the monoculture system; for this reason, they should be considered as a management alternative, particularly when the matrix permeability requirement is met.

Movements of neotropical understory passerines affected by anthropogenic forest edges in the Brazilian Atlantic rainforest

Edge effects are suggested to have great impact on the persistence of species in fragmented landscapes. We tested edge avoidance by forest understory passerines in the Brazilian Atlantic Rainforest and also compared their mobility and movement patterns in contiguous and fragmented landscapes to assess whether movements would increase in the fragmented landscape. Between 2003 and 2005, 96 Chiroxiphia caudata, 38 Pyriglena leucoptera and 27 Sclerurus scansor were radio-tracked. The most strictly forest species C. caudata and S scansor avoided forest edges while P leucoptera showed affinities for the edge Both sensitive species showed larger mean step length and maximal observed daily distance in the fragmented forest versus the unfragmented forest. P. leucoptera did not show any significant difference. There were no significant differences in proportional daily home range use for any of the three species. Our results suggested that fragmentation and the consequent increase in edge areas do influence movement behavior of sensitive forest understory birds that avoided the use of edges and increased the speed and distance they covered daily. For the most restricted forest species, it would be advisable to protect larger patches of forest instead of many small or medium fragments connected by narrow corridors. However, by comparing our data with that obtained earlier, we concluded that movement behavior of resident birds differs from that of dispersing birds and might not allow to infer functional connectivity or landscape-scale sensitivity to fragmentation; a fact that should be taken into consideration when suggesting conservation strategies. (c) 2008 Elsevier Ltd. All rights reserved.

Time-lag in biological responses to landscape changes in a highly dynamic Atlantic forest region

Time-lagged responses of biological variables to landscape modifications are widely recognized, but rarely considered in ecological studies. In order to test for the existence of time-lags in the response of trees, small mammals, birds and frogs to changes in fragment area and connectivity, we studied a fragmented and highly dynamic landscape in the Atlantic forest region. We also investigated the biological correlates associated with differential responses among taxonomic groups. Species richness and abundance for four taxonomic groups were measured in 21 secondary forest fragments during the same period (2000-2002), following a standardized protocol. Data analyses were based on power regressions and model selection procedures. The model inputs included present (2000) and past (1962, 1981) fragment areas and connectivity, as well as observed changes in these parameters. Although past landscape structure was particularly relevant for trees, all taxonomic groups (except small mammals) were affected by landscape dynamics, exhibiting a time-lagged response. Furthermore, fragment area was more important for species groups with lower dispersal capacity, while species with higher dispersal ability had stronger responses to connectivity measures. Although these secondary forest fragments still maintain a large fraction of their original biodiversity, the delay in biological response combined with high rates of deforestation and fast forest regeneration imply in a reduction in the average age of the forest. This also indicates that future species losses are likely, especially those that are more strictly-forest dwellers. Conservation actions should be implemented to reduce species extinction, to maintain old-growth forests and to favour the regeneration process. Our results demonstrate that landscape history can strongly affect the present distribution pattern of species in fragmented landscapes, and should be considered in conservation planning. (C) 2009 Elsevier Ltd. All rights reserved.

Importance of interhabitat gaps and stepping-stones for lesser woodcreepers (Xiphorhynchus fuscus) in the Atlantic forest, Brazil

Translocation experiments showed that a woodcreeper bird species is able to move between isolated forest fragments, but this ability is limited by increasing interpatch distances. Larger distances (> 100 m) were overcome by using small stepping-stones (isolated trees), which enhance connectivity and are useful for the species conservation in fragmented landscapes.

Passive Restoration in Biodiversity Hotspots: Consequences for an Atlantic Rainforest Lizard Taxocene

Long-term conservation in biodiversity hotspots depends on the recovery of communities in secondary forest fragments. In most cases, however, recovery strategies for these areas are based only on passive restoration. It is therefore necessary to determine the efficiency of such strategies. In this study, we assess the efficiency of passive restoration on a 567-ha 28-yr-old fragment of Atlantic Rainforest in Northeastern Brazil. We measured richness, composition, abundance and biomass of a lizard taxocene and also vegetation structure and availability of several microhabitat descriptors in 18 plots of this secondary forest. We then compared them with measures in 29 plots from two neighboring reference sites. Species richness, abundance, biomass and microhabitat descriptors availability inside the secondary fragment did not differ from reference sites. However, composition and vegetation structure showed small differences. Some forest specialist lizards, which should be a focus of conservation efforts in fragmented landscapes of the Atlantic Rainforest, were not found in the fragment and data indicate that this was not due to sampling or a lack of suitable habitat or microhabitat. In the presence of preserved source sites, passive restoration may be a cheap and effective way to recover lizard taxocenes of the Atlantic Rainforest. Some of the species may need to be re-introduced to accelerate the full recovery of original composition of lizard taxocenes in secondary Atlantic Rainforests.

Distribution and habitat requirements of the Yellow-Footed Antechinus Antechinus flavipes at multiple scales: a review

This review synthesises present knowledge of the distribution patterns and habitat requirements of the Yellow-footed Antechinus Antechinus flavipes. Factors influencing the distribution of A. flavipes are examined at several spatial scales ranging from the broad climatic conditions prevalent over the species’ entire range to the characteristics of nest sites used by individual animals. Analysis of the literature suggests that: 1) at the broad-scale, A. flavipes distribution is largely determined by warm, dry climatic conditions, the distribution of dry forests and woodlands and competition with closely related species; 2) at the landscape-scale the determinants of A. flavipes distribution are largely unknown, although initial investigations suggest some tolerance of fragmented landscapes; and 3) at a local-scale the distribution of A. flavipes is largely determined by the presence of large diameter trees, tree hollows, coarse woody debris, rocky crevices and leaf-litter. Directions for future research are suggested throughout the review.

Ecology of arboreal marsupials in a network of remnant linear habitats

Linear strips of vegetation set within a less-hospitable matrix are common features of landscapes throughout the world. Depending on location, form and function, these linear landscape elements include hedgerows, fencerows, shelterbelts, roadside or streamside strips and wildlife corridors. In many anthropogenically-modified landscapes, linear strips are important components for conservation because they provide a large proportion of the remaining wooded or shrubby habitat for fauna. They may also function to provide connectivity across the landscape. In some districts, the linear strips form an interconnected network of habitat. The spatial configuration of remnant habitat (size, shape and arrangement) may influence habitat suitability, and hence survival, of many species of plant and animal in modified landscapes. Near Euroa in south-eastern Australia, the clearing and fragmentation of temperate woodlands for agriculture has been extensive and, at present, less than 5% tree cover remains, most of which (83%) occurs as linear strips along roads and streams. The remainder of the woodland occurs as relatively small patches and single isolated trees scattered across the landscape. As an assemblage, arboreal marsupials are woodland dependent and vary in their sensitivity to habitat loss and fragmentation. This thesis focusses on determining the conservation status of arboreal marsupials in the linear network and understanding how they utilise the landscape mosaic. Specifically, the topics examined in this thesis are: (1) the composition of the arboreal marsupial assemblage in linear and non-linear woodland remnants; (2) the status and habitat preferences of species of arboreal marsupial within linear remnants; and (3) the ecology of a population of the Squirrel Glider Petaurus norfolcensis in the linear network, focusing on population dynamics, spatial organisation, and use of den trees. The arboreal marsupial fauna in the linear network was diverse, and comprised seven out of eight species known to occur in the district. The species detected within the strips were P. norfolcensis, the Sugar Glider Petaurus breviceps, Common Brushtail Possum Trichosums vulpecula, Common Ringtail Possum Pseudocheirus peregrinus, Brush-tailed Phascogale Phascogale tapoatafa, Koala Phascolarctos cinereus and Yellow-footed Antechinus Antechinus flavipes. The species not detected was the Feathertail Glider Acrabates pygmaeus. Survey sites in linear remnants (strips of woodland along roads and streams) supported a similar richness and density of arboreal mammals to sites in non-linear remnants (large patches or continuous tracts of woodland nearby). Furthermore, the combined abundance of all species of arboreal marsupials was significantly greater in sites in the linear remnants than in the non-linear remnants. This initial phase of the study provided no evidence that linear woodland remnants support a degraded or impoverished arboreal marsupial fauna in comparison with the nonlinear remnants surveyed. Intensive trapping of arboreal marsupials within a 15 km linear network between February 1997 and June 1998 showed that all species of arboreal marsupial (except A. pygmaeus) were present within the linear strips. Further analyses related trap-based abundance estimates to measures of habitat quality and landscape structure. Width of the linear habitat was significantly positively correlated with the combined abundance of all arboreal marsupials, as well as with the abundance of P. norfolcensis and T. vulpecula. The abundance of T. vulpecula was also significantly positively correlated with variation in overstorey species composition, Acacia density and the number of hollow-bearing trees. The abundance of P. norfolcensis was positively correlated with Acacia density and canopy width, and negatively correlated with distance to the nearest intersection with another linear remnant. No significant variables were identified to explain the abundance of P. tapoatafa, and there were insufficient captures of the remaining species to investigate habitat preferences. Petaurus norfolcensis were resident within the linear network and their density (0.95 -1.54 ha-1) was equal to the maximum densities recorded for this species in continuous forest elsewhere in south-eastern Australia. Rates of reproduction were also similar to those in continuous forest, with births occurring between May and December, a mean natality rate of 1.9, and a mean litter size of 1.7. Sex ratios never differed significantly from parity. Overall, the population dynamics of P. norfolcensis were comparable with published results for the species in contiguous forest, clearly suggesting that the linear remnants currently support a self-sustaining, viable population. Fifty-one P. norfolcensis were fitted with radio transmitters and tracked intermittently between December 1997 and November 1998. Home ranges were small (1.3 - 2.8 ha), narrow (20 - 40 m) and elongated (322 - 839 m). Home ranges were mostly confined to the linear remnants, although 80% of gliders also utilised small clumps of adjacent woodland within farm paddocks for foraging or denning. Home range size was significantly larger at intersections between two or more linear remnants than within straight sections of linear remnants. Intersections appeared to be important sites for social interaction because the overlap of home ranges of members of adjacent social groups was significantly greater at intersections than straight sections. Intersections provided the only opportunity for members of three or more social groups to interact, while still maintaining their territories. The 51 gliders were radiotracked to 143 different hollow-bearing trees on 2081 occasions. On average, gliders used 5.3 den trees during the study (range 1-15), and changed den trees every 4.9 days. The number of den trees used by each glider is likely to be conservative because the cumulative number of den trees continued to increase over the full duration of the study. When gliders shifted between den trees, the mean distance between consecutive den sites was 247 m. Den trees were located throughout a glider's home range, thereby reducing the need to return to a central den site and potentially minimising energy expenditure. Dens were usually located in large trees (mean diameter 88.5 cm) and were selected significantly more often than expected based on their occurrence within the landscape. The overall conclusion of this thesis is that the linear network I studied provides high quality habitat for resident populations of arboreal marsupials. Important factors influencing the suitability of the linear remnants appear to be the high level of network connectivity, the location on soils of high nutrient status, the high density of large trees and an acacia understorey. In highly fragmented landscapes, linear habitats as part of the remaining woodland mosaic have the potential to be an integral component in the conservation of woodland-dependent fauna. The habitat value of linear strips of vegetation should not be underestimated.