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.

Estimating survival and movements using both live and dead recoveries : a case study of oystercatchers confronted with habitat change

1. Animals facing partial habitat loss can try to survive in the remaining habitat or emigrate. Effects on survival and movements should be studied simultaneously since survival rates may be underestimated if emigrants are not considered, and since emigrants may experience reduced survival.

2. We analysed movements and survival of adult wintering oystercatcher Haematopus ostralegus in response to the 1986–1987 partial closure of the Oosterschelde in the Dutch Delta. This reduced by one-third the tidal area of this major European wintering area for waders.

3. We developed a novel variant of a multistate capture–recapture model allowing simultaneous estimation of survival and movement between sites using a mixture of data (live recaptures and dead recoveries). We used a two-step process, first estimating movements between sites followed by site-specific survival rates.

4. Most birds were faithful to their ringing site. Winter survival was negatively affected by winter severity and was lowest among birds changing wintering site (i.e. moving outside of the Oosterschelde).

5. During mild winters, survival rates were very high, and similar to before the closure in both changed and unchanged sectors of the Oosterschelde. However, the combined effect of habitat loss with severe winters decreased the survival of birds from changed sectors and induced emigration.

6. The coastal engineering project coincided with three severe winters and high food stock, making assessment of its effects difficult. However, the habitat loss seems to have had less impact on adult survival and movements than did winter severity.

7. Synthesis and applications. Human-induced habitat change may result in population decline through costly emigration or reduced survival or reproduction of individuals that stay. Long-term monitoring of marked individuals helps to understand how populations respond to environmental change, but site-specific survival and movement rates should be integrated in the same model in order to maximize the information yield. Our modelling approach facilitates this because it allows the inclusion of recoveries from outside the study area.

The habitat value of Gorse Ulex europaeus L. and Hawthorn Crataegus monogyna Jacq. for birds in Quarry Hills Bushland Park, Victoria

Weeds are one of the primary threats to biodiversity; however, their impacts on wildlife can vary. This research investigated the habitat value of Gorse Ulex europaeus L. and Hawthorn Crataegus monogyna Jacq. and the impacts of its removal on birds in a bushland park in Victoria. The area search method was used to survey birds in vegetation dominated by these two weeds, in native vegetation and in areas where a weed removal program was undertaken; this included revegetated areas. The highest bird species richness and abundance was found in sites dominated by the weeds. At sites where the weed removal program was in the early stages, a much lower species richness and abundance occurred. The final stage of the weed removal program, where revegetated areas were older than five years, supported high richness and abundance of birds, but not as high as that of sites dominated by the weeds; nor was the composition the same. Thus, even after five years, revegetation may not provide for the bird community that was originally supported by weeds. This is an important weed management consideration in this park, and should be for weed removal projects elsewhere

Distributional overlap rather than habitat differentiation characterizes co-occurrence of bivalves in intertidal soft sediment systems

Diverse species assemblages are often associated with a diversity of habitat structures. Sedimentary systems seem to be no exception, as within sedimentary systems benthic species diversity within a sample point appears to correlate with sediment grain size complexity. However, it remains to be shown whether total benthic species diversity relates to a system’s sediment heterogeneity across multiple systems. In the present paper we examined whether bivalve diversity is associated with: (1) sediment heterogeneity across systems and (2) sediment grain size complexity within systems, at 9 temperate and tropical tidal flat systems. Although bivalve life-history strategies, like post-settlement habitat selection, might suggest that sediment heterogeneity should be important for bivalve species, bivalve diversity and sediment heterogeneity were not associated across systems. Interestingly, the association between total benthic diversity and sediment heterogeneity was also not significant, suggesting that changing species composition across systems does not account for the lack of a correlation between bivalve diversity and sediment heterogeneity. Instead of habitat differentiation, bivalve diversity within a sample point was highest in ‘complex’ fine-grained sediments and bivalve distributions showed a large degree of distributional overlap in all systems. The results of this study at both smaller and larger spatial scales suggest that coexistence between bivalve species in diverse tidal flats is not associated with increased sediment heterogeneity.

Mallee Emu-wren (Stipiturus mallee) : multi-scale habitat requirements and population structure

This multi-disciplinary investigation found that: i) in Triodia-mallee the Mallee Emu-wren requires vegetation greater than 16-18 years since last burned, with high coverage of mature growth-phase Triodia scariosa (spinifex) and, ii) the species is panmictic with relatively low genetic diversity and evidence of genetic drift and bottlenecks.

Blue whale habitat selection and within-season distribution in a regional upwelling system off southern Australia

Blue whales Balaenoptera musculus aggregate to feed in a regional upwelling system during November–May between the Great Australian Bight (GAB) and Bass Strait. We analysed sightings from aerial surveys over 6 upwelling seasons (2001–02 to 2006–07) to assess within-season patterns of blue whale habitat selection, distribution, and relative abundance. Habitat variables were modelled using a general linear model (GLM) that ranked sea surface temperature (SST) and sea surface chlorophyll (SSC) of equal importance, followed by depth, distance to shore, SSC gradient, distance to shelf break, and SST gradient. Further discrimination by hierarchical partitioning indicated that SST accounted for 84.4% of variation in blue whale presence explained by the model, and that probability of sightings increased with increasing SST. The large study area was resolved into 3 zones showing diversity of habitat from the shallow narrow shelf and associated surface upwelling of the central zone, to the relatively deep upper slope waters, broad shelf and variable upwelling of the western zone, and the intermediate features of the eastern zone. Density kernel estimation showed a trend in distribution from the west during November–December, spreading south-eastward along the shelf throughout the central and eastern zones during January–April, with the central zone most consistently utilised. Encounter rates in central and eastern zones peaked in February, coinciding with peak upwelling intensity and primary productivity. Blue whales avoided inshore upwelling centres, selecting SST ~1°C cooler than remotely sensed ambient SST. Whales selected significantly higher SSC in the central and eastern zones than the western zone, where relative abundance was extremely variable. Most animals departed from the feeding ground by late April.

Understanding fish-habitat associations using video observations and sonar imaging

This thesis was motivated by the increasing role of species distribution models in managing marine fishes and their habitats. These models provided new information about fish-habitat associations. However, models are potentially influenced by fish behaviour towards underwater video systems.

Foraging trip strategies and habitat use during late pup rearing by lactating Australian fur seals

Australian fur seals (Arctocephalus pusillus doriferus) are the most conspicuous and abundant marine mammal in shelf waters of south-eastern Australia. To successfully rear offspring, the females must encounter sufficient prey on each foraging trip out of a central place for periods up to11 months each year. We investigated foraging trip strategies and habitat use by the females in three winter–spring periods, 2001–03, from four colonies that span the species’ latitudinal range and contribute 80% of pup production. Trip durations of 37 females averaged 6.1±0.5 (s.e.) days, although >90% of the seal’s time at sea was spent <150 km travel (<2 days) away. Most females exhibited strong fidelities to individually preferred hotspots Females from colonies adjacent to productive shelf-edge waters generally had shorter trips, had smaller ranges, foraged closer to colonies and exhibited less diversity in trip strategies than did those from colonies more distant from a shelf-edge. From a management perspective, there was minimal overlap (<1%) between where females foraged and a system of marine reserves established in 2007, suggesting that habitats visited by lactating Australian fur seals currently receive minimal legislative protection

Comparing the seasonal survival of resident and migratory oystercatchers : carry-over effects of habitat quality and weather conditions

Events happening in one season can affect life-history traits at (the) subsequent season(s) by carry-over effects. Wintering conditions are known to affect breeding success, but few studies have investigated carry-over effects on survival. The Eurasian oystercatcher Haematopus ostralegus is a coastal wader with sedentary populations at temperate sites and migratory populations in northern breeding grounds of Europe. We pooled continental European ringing-recovery datasets from 1975 to 2000 to estimate winter and summer survival rates of migrant and resident populations and to investigate long-term effects of winter habitat changes. During mild climatic periods, adults of both migratory and resident populations exhibited survival rates 2% lower in summer than in winter. Severe winters reduced survival rates (down to 25% reduction) and were often followed by a decline in survival during the following summer, via short-term carry-over effects. Habitat changes in the Dutch wintering grounds caused a reduction in food stocks, leading to reduced survival rates, particularly in young birds. Therefore, wintering habitat changes resulted in long-term (>10 years) 8.7 and 9.4% decrease in adult annual survival of migrant and resident populations respectively. Studying the impact of carry-over effects is crucial for understanding the life history of migratory birds and the development of conservation measures.

Habitat use throughout migration : linking individual consistency, prior breeding success and future breeding potential

1. Habitat use can influence individual performance in a wide range of animals, either immediately or through carry-over effects in subsequent seasons. Given that many animal species also show consistent individual differences in reproductive success, it seems plausible that individuals may have consistent patterns of habitat use representing individual specializations, with concomitant fitness consequences.

2. Stable-carbon isotope ratios from a range of tissues were used to discern individual consistency in habitat use along a terrestrial–aquatic gradient in a long-distance migrant, the Bewick’s swan (Cygnus columbianus bewickii). These individual specialisations represented <15% of the isotopic breadth of the population for the majority of individuals and were seen to persist throughout autumn migration and overwintering until aquatic habitats were no longer available.

3. Individual foraging specialisations were then used to demonstrate two consecutive carry-over effects associated with macroscale habitat segregation: consequences of breeding season processes for autumn habitat use; and consequences of autumn habitat use for future reproductive success. Adults that were successful breeders in the year of capture used terrestrial habitats significantly more than adults that were not successful, revealing a substantial cost of reproduction and extended parental care. Use of aquatic habitats during autumn was, however, associated with increased body condition prior to spring migration; and increased subsequent breeding success in adults that had been unsuccessful the year before. Yet adults that were successful breeders in the year of capture remained the most likely to be successful the following year, despite their use of terrestrial habitats.

4. Our results uniquely demonstrate not only individual foraging specializations throughout the migration period, but also that processes during breeding and autumn migration, mediated by individual consistency, may play a fundamental role in the population dynamics of long-distance migrants. These findings, therefore, highlight the importance of long-term consistency to our understanding of habitat function, interindividual differences in fitness, population dynamics and the evolution of migratory strategies.

Predicting landscape-genetic consequences of habitat loss, fragmentation and mobility for multiple species of woodland birds

Inference concerning the impact of habitat fragmentation on dispersal and gene flow is a key theme in landscape genetics. Recently, the ability of established approaches to identify reliably the differential effects of landscape structure (e.g. land-cover composition, remnant vegetation configuration and extent) on the mobility of organisms has been questioned. More explicit methods of predicting and testing for such effects must move beyond post hoc explanations for single landscapes and species. Here, we document a process for making a priori predictions, using existing spatial and ecological data and expert opinion, of the effects of landscape structure on genetic structure of multiple species across replicated landscape blocks. We compare the results of two common methods for estimating the influence of landscape structure on effective distance: least-cost path analysis and isolation-by-resistance. We present a series of alternative models of genetic connectivity in the study area, represented by different landscape resistance surfaces for calculating effective distance, and identify appropriate null models. The process is applied to ten species of sympatric woodland-dependant birds. For each species, we rank a priori the expectation of fit of genetic response to the models according to the expected response of birds to loss of structural connectivity and landscape-scale tree-cover. These rankings (our hypotheses) are presented for testing with empirical genetic data in a subsequent contribution. We propose that this replicated landscape, multi-species approach offers a robust method for identifying the likely effects of landscape fragmentation on dispersal.

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.