Testing alternative models for the conservation of koalas in fragmented rural-urban landscapes

Predicting the various responses of different species to changes in landscape structure is a formidable challenge to landscape ecology. Based on expert knowledge and landscape ecological theory, we develop five competing a priori models for predicting the presence/absence of the Koala (Phascolarctos cinereus) in Noosa Shire, south-east Queensland (Australia). A priori predictions were nested within three levels of ecological organization: in situ (site level) habitat (< 1 ha), patch level (100 ha) and landscape level (100-1000 ha). To test the models, Koala surveys and habitat surveys (n = 245) were conducted across the habitat mosaic. After taking into account tree species preferences, the patch and landscape context, and the neighbourhood effect of adjacent present sites, we applied logistic regression and hierarchical partitioning analyses to rank the alternative models and the explanatory variables. The strongest support was for a multilevel model, with Koala presence best predicted by the proportion of the landscape occupied by high quality habitat, the neighbourhood effect, the mean nearest neighbour distance between forest patches, the density of forest patches and the density of sealed roads. When tested against independent data (n = 105) using a receiver operator characteristic curve, the multilevel model performed moderately well. The study is consistent with recent assertions that habitat loss is the major driver of population decline, however, landscape configuration and roads have an important effect that needs to be incorporated into Koala conservation strategies.

A GEOBIA methodology for fragmented agricultural landscapes

Very high resolution remotely sensed images are an important tool for monitoring fragmented agricultural landscapes, which allows farmers and policy makers to make better decisions regarding management practices. An object-based methodology is proposed for automatic generation of thematic maps of the available classes in the scene, which combines edge-based and superpixel processing for small agricultural parcels. The methodology employs superpixels instead of pixels as minimal processing units, and provides a link between them and meaningful objects (obtained by the edge-based method) in order to facilitate the analysis of parcels. Performance analysis on a scene dominated by agricultural small parcels indicates that the combination of both superpixel and edge-based methods achieves a classification accuracy slightly better than when those methods are performed separately and comparable to the accuracy of traditional object-based analysis, with automatic approach.

Linking ecological condition and the soundscape in fragmented Australian forests

Natural landscapes are increasingly subjected to anthropogenic pressure and fragmentation resulting in reduced ecological condition. In this study we examined the relationship between ecological condition and the soundscape in fragmented forest remnants of south-east Queensland, Australia. The region is noted for its high biodiversity value and increased pressure associated with habitat fragmentation and urbanisation. Ten sites defined by a distinct open eucalypt forest community dominated by spotted gum (Corymbia citriodora ssp. variegata) were stratified based on patch size and patch connectivity. Each site underwent a series of detailed vegetation condition and landscape assessments, together with bird surveys and acoustic analysis using relative soundscape power. Univariate and multivariate analyses indicated that the measurement of relative soundscape power reflects ecological condition and bird species richness, and is dependent on the extent of landscape fragmentation. We conclude that acoustic monitoring technologies provide a cost effective tool for measuring ecological condition, especially in conjunction with established field observations and recordings.

Landscape structure influences tree density patterns in fragmented woodlands in semi-arid eastern Australia.

Landscape and local-scale influences are important drivers of plant community structure. However, their relative contribution and the degree to which they interact remain unclear. We quantified the extent to which landscape structure, within-patch habitat and their confounding effects determine post-clearing tree densities and composition in agricultural landscapes in eastern subtropical Australia. Landscape structure (incorporating habitat fragmentation and loss) and within-patch (site) features were quantified for 60 remnant patches of Eucalyptus populnea (Myrtaceae) woodland. Tree density and species for three ecological maturity classes (regeneration, early maturity, late maturity) and local site features were assessed in one 100 × 10 m plot per patch. All but one landscape characteristic was determined within a 1.3-km radius of plots; Euclidean nearest neighbour distance was measured inside a 5-km radius. Variation in tree density and composition for each maturity class was partitioned into independent landscape, independent site and joint effects of landscape and site features using redundancy analysis. Independent site effects explained more variation in regeneration density and composition than pure landscape effects; significant predictors were the proportion of early and late maturity trees at a site, rainfall and the associated interaction. Conversely, landscape structure explained greater variation in early and late maturity tree density and composition than site predictors. Area of remnant native vegetation within a landscape and patch characteristics (area, shape, edge contrast) were significant predictors of early maturity tree density. However, 31% of the explained variation in early mature tree differences represented confounding influences of landscape and local variables. We suggest that within-patch characteristics are important in influencing semi-arid woodland tree regeneration. However, independent and confounding effects of landscape structure resulting from previous vegetation clearing may have exerted a greater historical influence on older cohorts and should be accounted for when examining woodland dynamics across a broader range of environments.

Consumption of crops by feral pigs (Sus scrofa) in a fragmented agricultural landscape

Feral pigs (Sus scrofa) consume and damage crops and impact the environment through predation, competition and habitat disturbance, although supporting dietary data are lacking in agricultural landscapes. This study was undertaken to determine the relative importance of food items in the diet of feral pigs in a fragmented agricultural landscape, particularly to assist in predicting the breadth of likely impacts. Diet composition was assessed from the stomach contents of 196 feral pigs from agricultural properties in southern Queensland. Feral pigs were herbivorous, with plant matter comprising >99% of biomass consumed. Crops were consumed more frequently than non-crop species, and comprised >60% of dietary biomass, indicating a clear potential for direct economic losses. Consumption of pasture and forage species also suggests potential competition for pasture with domestic stock. There is little evidence of direct predation on native fauna, but feral pig feeding activities may impact environmental values. Seasonal differences in consumption of crop, pasture or animal food groups probably reflect the changing availability of food items. We recommend that future dietary studies examine food availability to determine any dietary preferences to assist in determining the foods most susceptible to damage. The outcomes of this study are important for developing techniques for monitoring the impacts of feral pigs, essential for developing management options to reduce feral pig damage on agricultural lands.

Wild Ungulate Decision-Making and the Role of Tiny Refuges in Human-Dominated Landscapes

Wildlife conservation in human-dominated landscapes requires that we understand how animals, when making habitat-use decisions, obtain diverse and dynamically occurring resources while avoiding risks, induced by both natural predators and anthropogenic threats. Little is known about the underlying processes that enable wild animals to persist in densely populated human-dominated landscapes, particularly in developing countries. In a complex, semi-arid, fragmented, human-dominated agricultural landscape, we analyzed the habitat-use of blackbuck, a large herbivore endemic to the Indian sub-continent. We hypothesized that blackbuck would show flexible habitat-use behaviour and be risk averse when resource quality in the landscape is high, and less sensitive to risk otherwise. Overall, blackbuck appeared to be strongly influenced by human activity and they offset risks by using small protected patches (similar to 3 km(2)) when they could afford to do so. Blackbuck habitat use varied dynamically corresponding with seasonally-changing levels of resources and risks, with protected habitats registering maximum use. The findings show that human activities can strongly influence and perhaps limit ungulate habitat-use and behaviour, but spatial heterogeneity in risk, particularly the presence of refuges, can allow ungulates to persist in landscapes with high human and livestock densities.

Effects of species turnover on reserve site selection in a fragmented landscape

Changes in species composition is an important process in many ecosystems but rarely considered in systematic reserve site selection. To test the influence of temporal variability in species composition on the establishment of a reserve network, we compared network configurations based on species data of small mammals and frogs sampled during two consecutive years in a fragmented Atlantic Forest landscape (SE Brazil). Site selection with simulated annealing was carried out with the datasets of each single year and after merging the datasets of both years. Site selection resulted in remarkably divergent network configurations. Differences are reflected in both the identity of the selected fragments and in the amount of flexibility and irreplaceability in network configuration. Networks selected when data for both years were merged did not include all sites that were irreplaceable in one of the 2 years. Results of species number estimation revealed that significant changes in the composition of the species community occurred. Hence, temporal variability of community composition should be routinely tested and considered in systematic reserve site selection in dynamic systems.

Are corridors, fragment size and forest structure important for the conservation of leaf-litter lizards in a fragmented landscape?

To investigate the implications of forest fragmentation for conservation of leaf-litter lizards the importance of fragment size, corridors and forest structure was examined in 20 forest fragments and six localities within a continuous forest in the Atlantic Plateau of Sao Paulo state, Brazil. The fragments were 2-276 ha in area and had different degrees of connectivity depending on the presence or absence of corridors. Two species of lizards were dominant, Ecpleopus gaudichaudii and Enyalius perditus. Variation in forest structure among sites was important only in explaining the abundance of E. perditus. Regardless of variation in forest structure, lizard species composition, total lizard abundance, number of species and abundance of E. perditus were sensitive to fragmentation per se but not to fragment size or corridor linkage. The inhospitable matrix surrounding fragments is probably what determines the presence and abundance of E. perditus and the high er lizard richness in continuous forests. These conditions may have prevented lizard species from recolonizing the forest fragments. Our results emphasize that the conservation of this leaf-litter fauna depends on the maintenance of large tracts of continuous forests and not on the size of fragments or on the presence of forest connections. Strategies for conservation of leaf-litter lizards in such highly fragmented Atlantic Forest landscapes should consider the enlargement of landscape connectivity between fragments and continuous forest, allowing the latter areas to act as a source of individuals for fragments.

Carnivore mammals in a fragmented landscape in northeast of Sao Paulo State, Brazil

Sao Paulo is the most developed state in Brazil and little of its native vegetation remains. In Luiz Antonio and Santa Rita do Passa Quatro municipalities, only small fragments of cerrado (Brazilian savanna) physiognomies (cerrado, cerrado sensu stricto) and of semideciduous forest have been left, surrounded by eucalyptus silviculture and sugar-cane agriculture. However, that vegetation mosaic still shelters large mammals, including several carnivore species. To detect the carnivores present in such a mosaic area (50,000 ha), and to find out how they use the landscape, we recorded them through 21 camera traps and 21 track plots, during 18 months. Species richness, diversity and relative frequency were evaluated according to the habitat. Ten species were recorded, some of them locally threatened to extinction (Puma concolor, Leopardus pardalis, Chrysocyon brachyurus). Species diversity did not significantly differ among fragments, and although most species preferred one or another habitat, the carnivore community as a whole explored all the study area regardless of the vegetation cover;eucalyptus plantations were as used by the carnivores as the native fragments. Therefore, it seems possible to maintain such animals in agricultural landscapes, where some large native fragments are left and the matrix is permeable to native fauna.

Occurrence of small mammals in a fragmented landscape: the role of vegetation heterogeneity

Most studies of habitat use by small mammals rely on data from trapping grids. Such studies pertain to habitat use by individuals, which may not reflect population-level requirements. To meet the challenge of landscape change, it is important to understand habitat use by populations across large geographic areas. We surveyed small mammals in 48 forest remnants across a 300 km2 study area, to investigate the influence of vegetation heterogeneity on regional distributions. Information-theoretic techniques were used to evaluate models of vegetation associations. Richness of native mammals was influenced by vegetation condition: disturbed sites supported fewer species. Models for individual species showed the agile antechinus, Antechinus agilis, to prefer structurally diverse forest vegetation, the long-nosed potoroo, Potorous tridactylus, to favour mesic shrub communities, the bush rat, Rattus fuscipes, to prefer complex low cover regardless of composition, the swamp rat, Rattus lutreolus, to favour reduced canopy cover, and the house mouse, Mus domesticus, to prefer disturbed vegetation. To satisfy the needs of all native species, a mosaic of natural vegetation is required. Degradation and simplification of forest vegetation have detrimental consequences. These results highlight the need to consider habitat quality, together with more traditional biogeographic variables, when investigating factors influencing patch occupancy by native fauna in modified landscapes.

Ecology and conservation of insectivorous bats in rural landscapes

Throughout the world, the increasing use of land for agriculture has been associated with extensive loss and fragmentation of natural habitats and, frequently, the degradation of remaining habitats. The effects of such habitat changes have been well studied for some faunal groups, but little is known of their consequences for bats. The aim of this study was to investigate the ecology and conservation of an assemblage of insectivorous bats in a rural landscape, with particular focus on their foraging and roosting requirements. This increased knowledge will, hopefully, assist the formulation of policy and management decisions to ensure the long-term survival of bats in these altered environments. The distribution and abundance of insectivorous bats in the Northern Plains of Victoria was investigated to determine the impacts of land-use change and to identify factors influencing the distribution of bats in rural landscapes. Thirteen species of insectivorous bats were recorded across the region by sampling at 184 sites. Two species were rare, but the remaining 11 species were widespread and occurred in all types of remnant wooded vegetation, ranging from large blocks (≥200 ha) to small isolated remnants (≤5 ha) and scattered trees in cleared farm paddocks. There was no significant difference between remnant types in the relative abundance of bat species, in species richness, or in the composition of bat assemblages at study sites. In a subsequent study, no difference in the activity levels of bats was found between remnants with different tree densities, ranging from densely-vegetated blocks to single paddock trees. However, sites in open paddocks devoid of trees differed significantly from all types of wooded remnants and had significantly lower levels of bat activity and a different species composition. In highly cleared and modified landscapes, all native vegetation has value to bats, even the smallest remnant, roadside and single paddock tree. Roost sites are a key habitat requirement for bats and may be a limiting resource in highly modified environments. Two species, the lesser long-eared bat Nyctophilus geoffroyi and Gould's wattled bat Chalinolobus gouldii, were investigated as a basis for understanding the capacity of bats to survive in agricultural landscapes. These species have different wing morphologies, which may be influential in how they use the landscape, and anecdotal evidence suggested differences in their roosting ecology. Roosting ecology was examined using radio-tracking to locate 376 roosts in two study areas with contrasting tree cover in northern Victoria. Both species were highly selective in the location of their roosts in the landscape, in roost-site selection and in roosting behaviour, and responded differently to differing levels of availability of roosts. The Barmah-Picola study area incorporated remnant vegetation in farmland and an adjacent extensive floodplain forest (Barmah forest). Male N. geojfroyi roosted predominantly within 3 km of their foraging areas in remnants in farmland. However, most female N. geoffroyi, and both sexes of C. gouldii, roosted in Barmah forest up to 12 km from their foraging areas in farmland remnants. These distances were greater than previously recorded for these species and further than predicted by wing morphology. In contrast, in the second study area (Naring) where only small remnants of wooded vegetation remain in farmland, individuals of both species moved significantly shorter distances between roost sites and foraging areas. There were marked inter- and intra-specific differences in the roosts selected. C. gouldii used similar types of roosts in both areas - predominantly dead spouts in large, live trees. N. geoffroyi used a broader range of roost types, especially in the farmland environment. Roosts were typically under bark and in fissures, with males in particular also using anthropogenic structures. A strong preference was shown by both sexes for roosts in dead trees, and entrance dimensions of roosts were consistently narrow (2.5 cm). In Barmah forest, maternity roosts used by N. geoffroyi were predominantly in narrow fissures in large-diameter, dead trees, while at Naring maternity roosts were also found under bark, in buildings, and in small-diameter, live and dead trees. The number of roost trees that are required for an individual or colony is influenced by the frequency with which bats move between roosts, the proportion of roosts that are re-used, the distance between consecutive roosts, and the size of roosting colonies. Both species roosted in small colonies and regularly shifted roost sites within a discrete roost area. These behavioural traits suggest that a high density of roost sites is required. There were marked differences in these aspects of behaviour between individuals roosting in Barmah forest and in the fragmented rural landscape. At Naring, N. geqffroyi remained in roosts for longer periods and moved greater distances between consecutive roosts than in Barmah forest. In contrast, C. gouldii used a smaller pool of roosts in the farmland environment by re-using roosts more frequently. Within Barmah forest, there is an extensive area of forest but the density of hollow-bearing trees is reduced due to timber harvesting and silvicultural practices. Individuals were selective in the location of their roosting areas, with both species selecting parts of the forest that contained higher densities of their preferred roost trees than was generally available in the forest. In contrast, in farmland at Naring, where there were small pockets of remnant vegetation with high densities of potential roost sites surrounded by cleared paddocks with few roosting opportunities, little selection was shown. This suggests that in Barmah forest the density of trees with potential roosts is lower than optimal, while in farmland roosting resources may be adequate in woodland remnants, but limiting at the landscape scale since more than 95% of the landscape now provides no roosting opportunities. Insectivorous bats appear to be less severely affected than some other faunal groups by habitat fragmentation and land-use change. A highly developed capacity for flight, the spatial scale at which they move and their ability to cross open areas means that they can regularly move among multiple landscape elements, rather than depend on single remnants for all their resources. In addition, bats forage and roost mainly at elevated levels in trees and so are less sensitive to degradation of wooded habitats at ground level. Although seemingly resilient to habitat fragmentation, insectivorous bats are fundamentally dependent on trees for roosting and foraging, and so are vulnerable to habitat loss and ongoing rural tree decline. Protection of the remaining large old trees and measures to ensure regeneration to provide ongoing replacement of hollow-bearing trees through time are critical to ensure the long-term conservation of bats in rural landscapes.

Use of forest fragments by blue-winged macaws (Primolius maracana) within a fragmented landscape

Parrots are the most threatened group of birds in the world, mainly because of the reduction and fragmentation of their natural habitats. However, few studies have investigated the dynamics of parrot populations in disturbed landscapes on a broad scale. In this paper, we studied the ecological interactions of the vulnerable blue-winged macaw (Primolius maracana) in a fragmented landscape surrounding a large protected park in southeastern Brazil. We sampled 36 forest fragments that varied in size, characteristics, degree of isolation and type of surrounding matrix in order to assess the importance of habitat features on the maintenance of these birds. Blue-winged macaws were recorded in 70% of the satellite remnants that were sampled, which included large and small blocks of forest. These areas were used as sites for feeding, nesting or overnight rests, and also provided connectivity for birds' displacements. However, the frequency of macaw visits varied among the remnants, and this was related to habitat features such as patch size, human use of surrounding land, and the proximity to the protected park, to urban areas and to the birds' roosting areas. In general, landscape-scale parameters explained more of the variation in the frequency of visits by macaws than did patch-scale parameters. These results demonstrate the importance of landscape mosaics for the survival of blue-winged macaws.

Environmental suitability of a highly fragmented and heterogeneous landscape for forest bird species in south-eastern Brazil

Assessment of the suitability of anthropogenic landscapes for wildlife species is crucial for setting priorities for biodiversity conservation. This study aimed to analyse the environmental suitability of a highly fragmented region of the Brazilian Atlantic Forest, one of the world's 25 recognized biodiversity hotspots, for forest bird species. Eight forest bird species were selected for the analyses, based on point counts (n = 122) conducted in April-September 2006 and January-March 2009. Six additional variables (landscape diversity, distance from forest and streams, aspect, elevation and slope) were modelled in Maxent for (1) actual and (2) simulated land cover, based on the forest expansion required by existing Brazilian forest legislation. Models were evaluated by bootstrap or jackknife methods and their performance was assessed by AUC, omission error, binomial probability or p value. All predictive models were statistically significant, with high AUC values and low omission errors. A small proportion of the actual landscape (24.41 +/- 6.31%) was suitable for forest bird species. The simulated landscapes lead to an increase of c. 30% in total suitable areas. In average, models predicted a small increase (23.69 +/- 6.95%) in the area of suitable native forest for bird species. Being close to forest increased the environmental suitability of landscapes for all bird species; landscape diversity was also a significant factor for some species. In conclusion, this study demonstrates that species distribution modelling (SDM) successfully predicted bird distribution across a heterogeneous landscape at fine spatial resolution, as all models were biologically relevant and statistically significant. The use of landscape variables as predictors contributed significantly to the results, particularly for species distributions over small extents and at fine scales. This is the first study to evaluate the environmental suitability of the remaining Brazilian Atlantic Forest for bird species in an agricultural landscape, and provides important additional data for regional environmental planning.

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.

Large unburnt areas, not small unburnt patches, are needed to conserve avian diversity in fire-prone landscapes

Mitigating the impacts of large-scale fires on biodiversity is becoming increasingly important as their frequency increases. In response, fire managers have engaged with the concept that retaining small unburnt residual areas of vegetation within extensively burnt landscapes may facilitate biodiversity conservation. However, it remains uncertain how the size and isolation of these unburnt residuals influence faunal distributions, persistence and recovery following fire. We used a replicated observation study to test bird responses to the size and isolation of unburnt residuals in a mallee woodland area recently burnt by a 28 000 ha wildfire in southern Australia. The scale of our study provided a rare opportunity to consider the responses of large mobile organisms to fire-induced habitat fragmentation. Within five replicated spatial blocks, we crossed two levels of isolation with large (5-7 ha) and small (1-3 ha) unburnt patches and matrix sites burnt 5 years previously. We compared these site types to six continuous (non-fragmented) unburnt sites. We surveyed each site on eight occasions. Most birds occurred more frequently in unburnt habitat beyond the extent of the fire. Bird responses to the availability and spatial distribution of unburnt remnants within the fire were largely influenced by their ability to use the recently burnt matrix. Occurrence of five species was higher in unburnt residuals when more of the landscape within 500 m was burnt. A fire refuge effect may be likely for two competitive species that occurred more frequently in unburnt residuals than in the burnt matrix or continuous unburnt habitat. For the weebill, recolonization following fire was likely to occur gradually over time from ex situ sources. Synthesis and applications. To maintain avian diversity in fire-prone landscapes, our results suggest a need to shift management focus from creating networks of small unburnt patches towards preserving large, intact areas of habitat. However, five species common to the burnt matrix preferentially selected residual patches when unburnt resources were locally scarce. Therefore, to benefit birds, land managers should limit the extent of applied burns and use narrow burns. When planning large burns, practitioners should consider that a number of species will remain absent from the landscape for several decades.