Conservation ecology and breeding biology of the white-browed treecreeper, Climacteris affinis

The White-browed Treecreeper Climacteris affinis is one of many woodland-dependent birds that are at risk from the encroachment of human-dominated land-uses into natural landscapes. The White-browed Treecreeper inhabits semi-arid woodlands in north-west Victoria, Australia, a vegetation community that has undergone extreme modification in the last century due to the expansion of agriculture in the region. Extant woodlands represent only 10% of the original woodland cover in the region, and are highly fragmented and disturbed in many districts. Thus, the survival of the White-browed Treecreeper may depend on active management. However, current knowledge of the ecology and biology of this species is virtually non-existent, and inadequate for informed and effective conservation actions. The aim of this thesis is to redress this situation and provide the ecological basis for sound conservation management of the species. The thesis consists of two parts: an investigation of habitat use at three spatial scales and a study of the social organization, nesting requirements, breeding behaviour and reproductive success of a population of White-browed Treecreepers. Fifty-six patches of remnant woodland in north-west Victoria were surveyed to determine the factors affecting the occurrence of the White-browed Treecreeper at the regional scale. It was detected in 16 patches, and was largely confined to two core districts - Yarrara and, Wyperfeld (Pine Plains). The floristic composition of the dominant tree species was an important determinant of patch occupancy, with the results providing quantitative support for the previously suspected affinity for Belah Casuarina pauper and Slender Cypress-pine Callitris gracilis — Buloke Allocasuarina luehmannii woodlands. However, the absence of the White-browed Treecreeper from several districts was due to factors other than a lack of appropriate habitat. Demographic isolation - the distance from the focal patch to the nearest population of the White-browed Treecreeper - was the most important variable in explaining variation in patch occupancy. Patches isolated from other treecreeper populations by more than 8.3 km in landscapes of non-preferred native vegetation, and 3 km in agricultural landscapes, were unlikely to support the White-browed Treecreeper. The impact of habitat loss and fragmentation on the capacity of individuals to move through the landscape (i.e. functional connectivity) is considered in relation to disruption to dispersal and migration, and the potential collapse of local metapopulations. Habitat use was then examined in a network of patches and linear strips of Belah woodland embedded in a predominantly cultivated landscape. A minimum area of 18.5 ha of Belah woodland was identified as the most important criterion for patch occupancy at the local scale. This landscape appeared to be permeable to movement by the White-browed Treecreeper, facilitated by the extensive network of linear habitat, and clusters of small to medium fragments. The third scale of habitat use investigated the frequency of use of 1-ha plots within tracts of occupied woodland. It is important to discriminate between habitat traits that operate at the population level, and those that act as proximate cues for habitat selection by individuals. Woodlands that have high tree density, extensive cover of low-stature shrubs, abundant lichen, a complex vertical structure, and relatively low cover of grass and herbs are likely to support larger populations of the White-browed Treecreeper. However, individuals appeared to be using tree dominance (positive) and tall shrub cover (negative) as proximate environmental stimuli for habitat selectivity. A relatively high cover of ground lichen, which probably reflects a ground layer with low disturbance and high structural complexity, was also a reliable indicator of habitat use. Predictive models were developed which could be used to plan vegetation management to enhance habitat for the White-browed Treecreeper. The results of the regional, landscape and patch-scale investigations emphasise that factors operating at multiple spatial scales influence the suitability of remnant vegetation as habitat for the White-browed Treecreeper. The White-browed Treecreeper is typical of many small Australian passerines in that it has high annual survival, small clutches, a long breeding season, multiple broods and relatively low reproductive rates. Reproductive effort is adjusted through the number of clutches laid rather than clutch size. They occupy relatively large, all-purpose territories throughout the year. However, unlike many group territorial birds, territory size was not related to the number of occupants. The White-browed Treecreeper nests in tree hollows. They select hollows with a southerly orientation where possible, and prefer hollows that were higher from the ground. At Yarrara, there was considerable spatial variation in hollow abundance that, in concert with territorial constraints, restricted the actual availability of hollows to less than the absolute abundance of hollows. Thus, the availability of suitable hollows may limit reproductive productivity in some territories, although the magnitude of this constraint on overall population growth is predicted to be small. However, lack of recruitment of hollow-bearing trees would increase the potential for hollow availability to limit population growth. This prospect is particularly relevant in grazed remnants and those outside the reserve system. Facultative cooperative breeding was confirmed, with groups formed through male philopatry. Consequently, natal dispersal is female-biased, although there was no skew in the sex ratio of the fledglings or the general adult population. Helpers were observed performing all activities associated with parenting except copulation and brooding. Cooperatively breeding groups enjoyed higher fledgling productivity than simple pairs, after statistically accounting for territory and parental quality. However, the difference reflected increased productivity in the 1999-breeding season only, when climatic conditions were more favourable than in 1998. Breeding commenced earlier in 1999, and all breeding units were more likely to attempt a second brood. However, only breeders with helpers were successful in fledging second brood young, and it was this difference that accounted for the overall discrepancy in productivity. The key mechanism for increased success in cooperative groups was a reduction hi the interval between first and second broods, facilitated by compensatory reductions in the level of care to the first brood. Thus, females with helpers probably achieved significant energetic savings during this period, which enabled them to re-lay sooner. Furthermore, they were able to recommence nesting when the fledglings from the first brood were younger because there were more adults to feed the dependent juveniles. The current utility, and possible evolutionary pathways, of cooperative breeding is examined from the perspective of both breeders and helpers. Breeders benefit through enhanced fledgling productivity in good breeding conditions and a reduction in the burden of parental care, which may impart significant energetic savings. Further, breeders may facilitate philopatry as a means for ensuring a minimum level of reproductive success. Helpers benefit through an increase in their inclusive fitness in the absence of opportunities for independent breeding (i.e. ecological constraints) and access to breeding vacancies in the natal or adjacent territories (i.e. benefits of philopatry). However, the majority of breeding unit-years comprised unassisted breeders, which suggests that pairs are selectively favoured under certain environmental or demographic conditions.

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.