Nestedness in fragmented landscapes: birds of the box-ironbark forests of south-eastern Australia

Nestedness in biota as a function of species richness – biota of depauperate assemblages being non-random subsets of richer biotas – has been widely documented in recent years (see Wright et al. 1998, Oecologia 113: 1–20). Ordering sites by richness maximizes nestedness indices; however, ordering by other criteria such as area or isolation may be more ecologically interpretable. We surveyed birds in true fragments (35 in all), and in "reference areas" in large extant forest blocks (30 locations), of the same range of areas (10, 20, 40, 80 ha). The avifauna was divided into "bush birds"– species dependent on forest and woodland, and "open country" species. We looked at nestedness in four data sets: "bush birds" in fragments and reference areas, and "all birds" in fragments and in reference areas. All data sets were significantly nested. Ordering by area in all cases was not significantly less nested than ordering by richness. Ordering by area in fragments was significantly greater than in reference areas, but the differences in standardized nestedness indices were small (<15%). We identified those birds that had distributions among fragments that conformed strongly with area, those that were more randomly distributed and some species that were more likely to occupy the smallest fragments. Among the latter was a hyperaggressive, invasive, colonial native species (noisy miner Manorina melanocephala). A suite of small, insectivorous birds were more likely to strongly conform with expected distributions in relation to area, which was consistent with observations of their vulnerability to the effects of the noisy miner in smaller fragments.

Disrupted fine-scale population processes in fragmented landscapes despite large-scale genetic connectivity for a widespread and common cooperative breeder : the superb fairy-wren (Malurus cyaneus)

Understanding how habitat fragmentation affects population processes (e.g. dispersal) at different spatial scales is of critical importance to conservation. We assessed the effects of habitat fragmentation on dispersal and regional and fine-scale population structure in a currently widespread and common cooperatively breeding bird species found across south-eastern Australia, the superb fairy-wren Malurus cyaneus. Despite its relative abundance and classification as an urban tolerant species, the superb fairy-wren has declined disproportionately from low tree-cover agricultural landscapes across the Box-Ironbark region of north-central Victoria, Australia. Loss of genetic connectivity and disruption to its complex social system may be associated with the decline of this species from apparently suitable habitat in landscapes with low levels of tree cover. To assess whether reduced structural connectivity has had negative consequences for genetic connectivity in the superb fairy-wren, we used a landscape-scale approach to compare patterns of genetic diversity and gene flow at large (landscape/regional) and fine (site-level) spatial scales. In addition, using genetic distances, for each sex, we tested landscape models of decreased dispersal through treeless areas (isolation-by-resistance) while controlling for the effect of isolation-by-distance. Landscape models indicated that larger-scale gene flow across the Box-Ironbark region was constrained by distance rather than by lack of structural connectivity. Nonetheless, a pattern of isolation-by-resistance for males (the less-dispersive sex) and lower genetic diversity and higher genetic similarity within sites in low-cover fragmented landscapes indicated disruption to fine-scale gene flow mechanisms and/or mating systems. Although loss of structural connectivity did not appear to impede gene flow at larger spatial scales, fragmentation appeared to affect fine-scale population processes (e.g. local gene flow mechanisms and/or mating systems) adversely and may contribute to the decline of superb fairy-wrens in fragmented landscapes in the Box-Ironbark region. © 2012 British Ecological Society.

Does reduced mobility through fragmented landscapes explain patch extinction patterns for three honeyeaters?

1.Habitat loss and associated fragmentation are major drivers of biodiversity decline, and understanding how they affect population processes (e.g. dispersal) is an important conservation goal. In a large-scale test employing 10 × 10 km units of replication, three species of Australian birds, the fuscous honeyeater, yellow-tufted honeyeater and white-plumed honeyeater, responded differently to fragmentation. The fuscous and yellow-tufted honeyeaters are ‘decliners’ that disappeared from suitable habitat in landscapes where levels of tree-cover fell below critical thresholds of 17 and 8%, respectively. The white-plumed honeyeater is a ‘tolerant’ species whose likelihood of occurrence in suitable habitat was independent of landscape-level tree-cover. 2.To determine whether the absence of the two decliner species in low tree-cover landscapes can be explained by reduced genetic connectivity, we looked for signatures of reduced mobility and gene flow in response to fragmentation across agricultural landscapes in the Box-Ironbark region of north-central Victoria, Australia. 3.We compared patterns of genetic diversity and population structure at the regional scale and across twelve 100 km2 landscapes with different tree-cover extents. We used genetic data to test landscape models predicting reduced dispersal through the agricultural matrix. We tested for evidence of sex-biased dispersal and sex-specific responses to fragmentation. 4.Reduced connectivity may have contributed to the disappearance of the yellow-tufted honey-eater from low tree-cover landscapes, as evidenced by male bias and increased relatedness among males in low tree-cover landscapes and signals of reduced gene flow and mobility through the agricultural matrix. We found no evidence for negative effects of fragmentation on gene flow in the other decliner, the fuscous honeyeater, suggesting that undetected pressures act on this species. As expected, there was no evidence for decreased movement through fragmented landscapes for the tolerant white-plumed honeyeater. 5.We demonstrated effects of habitat loss and fragmentation (stronger patterns of genetic differentiation, increased relatedness among males) on the yellow-tufted honeyeater above the threshold at which probability of occurrence dropped. Increasing extent and structural connectivity of habitat should be an appropriate management action for this species and other relatively sedentary woodland specialist species for which it can be taken as representative.

Is the matrix important to butterflies in fragmented landscapes?

The quality and extent of the 'matrix' in terrestrial fragmented landscapes may influence the persistence and behaviour of patch-associated fauna. Butterflies are a popular target group for fragmentation studies and represent an ideal assemblage to explore the impact and role of the matrix in patchy landscapes. To date, there has been no attempt to synthesise available research and assess the extent to which the matrix is included in studies of fragmented butterfly populations. Addressing this issue is important for improved understanding of habitat use in fragmented landscapes, and for the successful management and conservation of butterfly biodiversity. Our systematic review of 100 empirical research papers spans 50 years, and identifies how (and indeed if) the matrix is recognised in studies of butterfly populations in fragmented landscapes. We found that it was significantly more likely for studies not to include the matrix in their experimental design. This is of particular concern given 60 % of papers that excluded the matrix in their research did so in systems where the matrix was expected to contain resources of value for patch-associated species (as it was either a heterogeneous landscape or had similar structure to patches). Of the papers that did consider the matrix, 80 % (n = 24) reported a negative effect of the matrix on butterfly species and/or communities. Matrix effects may influence the survival and persistence of faunal groups in a world increasingly dominated by fragmented habitats. Our review suggests that future research should clearly define the matrix, and incorporate it in appropriate experimental designs.

Species conservation in fragmented landscapes: implications for the Grey-Crowned Babbler

The Grey-crowned Babbler is a bird species under threat in Victoria and this research describes how they are using the limited and fragmented habitat available and the dire affects habitat fragmentation is having on their genetic processes

Conceptual domain of the matrix in fragmented landscapes

In extensively modified landscapes, how the matrix is managed determines many conservation outcomes. Recent publications revise popular conceptions of a homogeneous and static matrix, yet we still lack an adequate conceptual model of the matrix. Here, we identify three core effects that influence patch-dependent species, through impacts associated with movement and dispersal, resource availability, and the abiotic environment. These core effects are modified by five 'dimensions': spatial and temporal variation in matrix quality; spatial scale; temporal scale of matrix variation; and adaptation. The conceptual domain of the matrix, defined as three core effects and their interaction with these five dimensions, provides a much-needed framework to underpin management of fragmented landscapes and highlights new research priorities.

Small mammals in a fragmented landscape

Small mammals displayed contrasting patterns of occurence in forest fragments. Population studies revealed that key processes responsible for survival in habitat patches include demographic flexibility and the movement of individuals between patches. Combining pattern- and process-based approaches provides a more complete understanding of fauna in modified landscapes.

Genes and song : genetic and social connections in fragmented habitat in a woodland bird with limited dispersal

Understanding the processes leading to population declines in fragmented landscapes is essential for successful conservation management. However, isolating the influence of disparate processes, and dispersal in particular, is challenging. The Grey Shrike-thrush, Colluricincla harmonica, is a sedentary woodland-dependent songbird, with learned vocalizations whose incidence in suitable habitat patches falls disproportionally with decline in tree cover in the landscape. Although it has been suggested that gaps in tree cover might act as barriers to its dispersal, the species remains in many remnants of native vegetation in agricultural landscapes, suggesting that it may have responded to habitat removal and fragmentation by maintaining or even increasing dispersal distances. We quantified population connectivity of the Grey Shrike-thrush in a system fragmented over more than 120 years using genetic (microsatellites) and acoustic (song types) data. First, we tested for population genetic and acoustic structure at regional and local scales in search of barriers to dispersal or gene flow and signals of local spatial structuring indicative of restricted dispersal or localized acoustic similarity. Then we tested for effects of habitat loss and fragmentation on genetic and acoustic connectivity by fitting alternative models of mobility (isolation-by-distance [the null model] and reduced and increased movement models) across treeless vs. treed areas. Birds within 5 km of each other had more similar genotypes and song types than those farther away, suggesting that dispersal and song matching are limited in the region. Despite restricted dispersal detected for females (but not males), populations appeared to be connected by gene flow and displayed some cultural (acoustic) connectivity across the region. Fragmentation did not appear to impact greatly the dispersal of the Grey Shrike-thrush: none of the mobility models fit the genetic distances of males, whereas for females, an isolation-by-distance model could not be rejected in favor of the models of reduced or increased movement through treeless gaps. However, dissimilarities of the song types were more consistent with the model of reduced cultural connectivity through treeless areas, suggesting that fragmentation impedes song type sharing in the Grey Shrike-thrush. Our paper demonstrates that habitat fragmentation hinders important population processes in an Australian woodland bird even though its dispersal is not detectably impacted.

Reptiles in restored agricultural landscapes: the value of linear strips, patches and habitat condition

Habitat restoration, including revegetation of linear strips and enlargement of remnant patches, may benefit native fauna in highly fragmented landscapes. Such restoration has occurred around the world, even though the relative importance of strips and patches of vegetation remains controversial. Using reptile communities from south-eastern Australia, we assessed the conservation value of revegetation in strips and alongside remnant patches compared with remnant vegetation and cleared roadsides. We also examined the distance that reptiles occurred from remnant patches into linear vegetation. We found that reptile species richness and counts did not substantially differ between revegetated, remnant and cleared habitats, or between linear strip and patch treatments. This may indicate that species sensitive to land clearing have already been lost from the landscape. These results imply that if specialist species have already been lost, we may be unable to measure the effects of agriculture on biodiversity. Furthermore, revegetation with the expectation that fauna will recolonize may be unrealistic and translocations may be necessary. Unexpectedly, we recorded higher species richness and counts of rare reptile species in remnant linear strips as distance from remnant patches increased. Ground-layer attributes were important for increasing reptile species richness and counts and in structuring reptile communities, explaining approximately three times as much variation as remnant shape or vegetation type (remnant, revegetated, cleared). Management agencies should protect and effectively manage remnant linear strips if rarer reptiles are to be retained, paying particular attention to ground-layer attributes. The decision to include ground layers in future revegetation activities will be more important than the shape of restored areas.

Spatial dynamics of the knob-tailed gecko Nephrurus stellatus in a fragmented agricultural landscape

In fragmented landscapes, a species' dispersal ability and response to habitat condition are key determinants of persistence. To understand the relative importance of dispersal and condition for survival of Nephrurus stellatus (Gekkonidae) in southern Australia, we surveyed 92 woodland remnants three times. This gecko favours early post-fire succession conditions so may be at risk of extinction in the long-unburnt agricultural landscape. Using N-mixture models, we compared the influence of four measures of isolation, patch area and two habitat variables on the abundance and occurrence of N. stellatus, while taking into account detection probability. Patch occupancy was high, despite the long-term absence of fire from most remnants. Distance to the nearest occupied site was the most informative measure of patch isolation, exhibiting a negative relationship with occupancy. Distance to a nearby conservation park had little influence, suggesting that mainland-island metapopulation dynamics are not important. Abundance and occurrence were positively related to %-cover of spinifex (Triodia), indicating that niche-related factors may also contribute to spatial dynamics. Patterns of patch occupancy imply that N. stellatus has a sequence of spatial dynamics across an isolation gradient, with patchy populations and source-sink dynamics when patches are within 300 m, metapopulations at intermediate isolation, and declining populations when patches are separated by >1-2 km. Considering the conservation needs of the community, habitat condition and connectivity may need to be improved before fire can be reintroduced to the landscape. We speculate that fire may interact with habitat degradation and isolation, increasing the risk of local extinctions.

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.

Habitat fragmentation and landscape change

This chapter begins by summarizing the conceptual approaches used to understand conservation in fragmented landscapes. We then examine the biophysical aspects of landscape change, and how such change affects species and communities, posing two main questions: (i) what are the implications for the patterns of occurrence of species and communities?; and (ii) how does landscape change affect processes that influence the distribution and viability of species and communities. The chapter concludes by identifying the kinds of actions that will enhance the conservation of biota in fragmented landscapes.

Fine-scale effects of habitat loss and fragmentation despite large-scale gene flow for some regionally declining woodland bird species

Habitat loss and associated fragmentation effects are well-recognised threats to biodiversity. Loss of functional connectivity (mobility, gene flow and demographic continuity) could result in population decline in altered habitat, because smaller, isolated populations are more vulnerable to extinction. We tested whether substantial habitat reduction plus fragmentation is associated with reduced gene flow in three 'decliner' woodland-dependent bird species (eastern yellow robin, weebill and spotted pardalote) identified in earlier work to have declined disproportionately in heavily fragmented landscapes in the Box-Ironbark forest region in north-central Victoria, Australia. For these three decliners, and one 'tolerant' species (striated pardalote), we compared patterns of genetic diversity, relatedness, effective population size, sex-ratios and genic (allele frequency) differentiation among landscapes of different total tree cover, identified population subdivision at the regional scale, and explored fine-scale genotypic (individual-based genetic signature) structure. Unexpectedly high genetic connectivity across the study region was detected for 'decliner' and 'tolerant' species. Power analysis simulations suggest that moderate reductions in gene flow should have been detectable. However, there was evidence of local negative effects of reduced habitat extent and structural connectivity: slightly lower effective population sizes, lower genetic diversity, higher within-site relatedness and altered sex-ratios (for weebill and eastern yellow robin) in 10 x 10 km 'landscapes' with low vegetation cover. We conclude that reduced structural connectivity in the Box-Ironbark ecosystem may still allow sufficient gene flow to avoid the harmful effects of inbreeding in our study species. Although there may still be negative consequences of fragmentation for demographic connectivity, the high genetic connectivity of mobile bird species in this system suggests that reconnecting isolated habitat patches may be less important than increasing habitat extent and/or quality if these need to be traded off.