ICES meets marine historical ecology: placing the history of fish and fisheries in current policy context

As a discipline, marine historical ecology (MHE) has contributed significantly to our understanding of the past state of the marine environment when levels of human impact were often very different from those today.What is less widely known is that insights from MHE have made headway into being applied within the context of present-day and long-term management and policy. This study draws attention to the applied value of MHE. We demonstrate that a broad knowledge base exists with potential for management application and advice, including the development of baselines and reference levels.Using a number of case studies from around the world,we showcase the value of historical ecology in understanding change and emphasize how it either has already informed management or has the potential to do so soon.We discuss these case studies in a context of the science–policy interface around six themes that are frequently targeted by current marine and maritime policies: climate change, biodiversity conservation, ecosystem structure, habitat integrity, food security, and human governance.We encourage science–policy bodies to actively engage with contributions from MHE, as well informed policy decisions need to be framed within the context of historical reference points and past resource or ecosystem changes.

Human dimensions of wildlife population management in Australasia – history, approaches and directions

It is now widely accepted that it is important to understand the ‘human dimensions’ of wildlife management issues in order to achieve management goals. This growing field of study was born in the 1960s and involves an examination of societal values, knowledge and behaviours associated with wildlife and wildlife management issues. This paper provides an overview of the history and directions in human dimensions research, focusing specifically on its application for wildlife population management in Australasia (in particular, Australia and New Zealand). It also provides a ‘toolkit’ of methods and approaches for those wishing to undertake, interpret or utilise human dimensions enquiries.

Ecology of epifaunal caridean shrimps in the Hopkins River estuary, and the role of estuaries in the life history of the atyid "Paratya Australiensis" Kemp, 1917 in south-eastern Australia

This study examined the factors affecting the distribution and abundance of epifaunal caridean shrimps in seagrass meadows of the Hopkins River estuary in south-western Victoria, Australia, and investigated the life history patterns of the freshwater Parana australiensis, found for the first time in estuaries. Adult and sub-adult shrimps were surveyed in seagrass meadows along the estuary over two years, and their planktonic larvae were surveyed in adjacent waters. Three species were collected. The marine Palaemon serenus occurred only near the mouth, summer to autumn, in high salinities. The marine/estuarine Macrobrachium intermedium occurred throughout the estuary. Adults were most abundant in late autumn, and least abundant in summer (unlike trends reported in marine meadows). Densities were higher and less variable in downstream meadows. P. australiensis occurred in the upper estuary all year, most abundantly in spring, due to migration from the river after peak discharge. Ovigerous females dominated, while males, showing less migration into the estuary, dominated above estuarine influence. Adults disappeared from the estuary in summer as salinity rose. Breeding period for P. australiensis was briefer in the estuary (September-December) than upstream (July-April). M. intermedium began breeding later in the upper estuary (November/December-March) than in the lower estuary (October-March), probably reflecting a physiological response to lower salinity, rather than an interaction with P. australiensis. No ovigerous P. serenus were found in the estuary. Larvae of P. australiensis and M intermedium occurred abundantly throughout the estuary, but P. serenus larvae did not. P. australiensis was an early coloniser to the plankton after peak discharge (November-December). Larvae concentrated in the deep saline layer at the head of the intruding salt wedge, thus probably maintaining longitudinal position. Diurnal vertical migrations were evident within the salt wedge, and in a deep pool above tidal influence. M. intermedium larvae occurred October-May in the lower estuary and November-April in the upper estuary, peaking in abundance one to two months after P. australiensis. They were associated with low surface flows and surface salinities greater than 10, over an anoxic deeper layer. All three species exhibited extended development of euryhaline larvae in the laboratory. Tolerances and optimal salinities of larvae of the three species reflected their distributions. M. intermedium was the most euryhaline species. P. australiensis larvae were tolerant of higher salinities than juveniles of adults: capable of developing in salinity of at least 15. Most P. australiensis juveniles recruited to the estuary November-December, after which numbers declined dramatically. After settlement, most recruits probably migrated upstream out of the estuary. Two cohorts of M. intermedium recruited to the estuary from larvae in summer (December and February), but some juveniles also migrated from adjacent coastal waters. Post-larval migration was at least as important a determinant of abundance as direct recruitment from estuarine, planktonic larvae in all three species. Distributions among seagrass meadows along the estuary were determined primarily by physico-chemical patterns driven by hydrological changes. Seasonal variations in salinity and temperature were strongly associated with seasonal variations in shrimp abundance. Salinity tolerances of adults of the three species reflected their distribution patterns. Biotic interactions were more important in determining distributions within meadows. P. australiensis, when abundant, were associated with seagrass biomass. M. intermedium were also, but when seagrass was sparsest and least extensive. The two species apparently partitioned the seagrass meadow according to depth in early summer. Laboratory experiments suggested P. australiensis was displaced from deeper water by M. intermedium. Preference for vegetative complexity and competition for position within meadows suggest the underlying importance of predation in regulating shrimp populations. A survey of south-eastern Australian estuaries found P. australiensis larvae abundant in all stable, open, well-developed, salt-wedge estuaries where adults were abundant. Adults were most abundant in low salinities among submerged leafy macrophytes. Reproductive traits of P. australiensis were compared in estuarine and fresh reaches of three rivers. Early in the breeding season, egg size was smaller, and (size-specific) egg number larger in estuaries than upstream. A trade-off between egg size and egg number resulted in no difference in total (size-specific) reproductive investment between locations. Reproductive investment tended to decrease at some locations over the breeding season, and this decrease was a result of decreased egg size in most cases. The decrease in reproductive investment probably reflected reduced food availability for the adult, while the reduced egg size was probably a response to improved conditions for larval development. In the Hopkins River, larger egg size at upstream sites was reflected in larger early stage larvae. Later stage larvae were larger in the estuary, suggesting more favourable conditions for larval development. Allozyme electrophoresis showed the P. australiensis populations in each of the three rivers to be distinct. Allozyme frequencies were not different within the Hopkins River, but upstream and estuarine locations in the Curdies and Gellibrand were different. Although some variation in reproductive traits within catchments may have been due to genotypic differences, trade-offs between egg size and number, and decreases in egg size over summer were probably due to plastic responses to environmental cues. It is proposed P. australiensis inhabits and reproduces in both estuarine and freshwater environments by plastic response to environmental conditions. Recruitment to estuaries is dependent on the presence of suitable adult, littoral habitat, and a stable salt wedge for larval retention. Estuaries are important recruitment sites for P. australiensis, potentially allowing an extra brood each year before riverine recruitment. Estuarine broods could constitute a large part of the total fecundity of P. australiensis females. Euryhaline larvae and estuarine recruitment of P. australiensis suggest marine transport of larvae between estuaries as a possible dispersal mechanism for Paratya species.

Using gene-history and expression analyses to assess the involvement of LGI genes in human disorders

Mutations in the leucine-rich, glioma-inactivated 1 gene, LGI1, cause autosomal-dominant lateral temporal lobe epilepsy via unknown mechanisms. LGI1 belongs to a subfamily of leucine-rich repeat genes comprising four members (LGI1–LGI4) in mammals. In this study, both comparative developmental as well as molecular evolutionary methods were applied to investigate the evolution of the LGI gene family and, subsequently, of the functional importance of its different gene members. Our phylogenetic studies suggest that LGI genes evolved early in the vertebrate lineage. Genetic and expression analyses of all five zebrafish lgi genes revealed duplications of lgi1 and lgi2, each resulting in two paralogous gene copies with mostly nonoverlapping expression patterns. Furthermore, all vertebrate LGI1 orthologs experience high levels of purifying selection that argue for an essential role of this gene in neural development or function. The approach of combining expression and selection data used here exemplarily demonstrates that in poorly characterized gene families a framework of evolutionary and expression analyses can identify those genes that are functionally most important and are therefore prime candidates for human disorders.

Cancer: an emergent property of disturbed resource-rich environments? Ecology meets personalized medicine

For an increasing number of biologists, cancer is viewed as a dynamic system governed by evolutionary and ecological principles. Throughout most of human history, cancer was an uncommon cause of death and it is generally accepted that common components of modern culture, including increased physiological stresses and caloric intake, favor cancer development. However, the precise mechanisms for this linkage are not well understood. Here, we examine the roles of ecological and physiological disturbances and resource availability on the emergence of cancer in multicellular organisms. We argue that proliferation of 'profiteering phenotypes' is often an emergent property of disturbed, resource-rich environments at all scales of biological organization. We review the evidence for this phenomenon, explore it within the context of malignancy, and discuss how this ecological framework may offer a theoretical background for novel strategies of cancer prevention. This work provides a compelling argument that the traditional separation between medicine and evolutionary ecology remains a fundamental limitation that needs to be overcome if complex processes, such as oncogenesis, are to be completely understood.

Incorporating anthropogenic effects into trophic ecology: predator-prey interactions in a human-dominated landscape

Apex predators perform important functions that regulate ecosystems worldwide. However, little is known about how ecosystem regulation by predators is influenced by human activities. In particular, how important are top-down effects of predators relative to direct and indirect human-mediated bottom-up and top-down processes? Combining data on species' occurrence from camera traps and hunting records, we aimed to quantify the relative effects of top-down and bottom-up processes in shaping predator and prey distributions in a human-dominated landscape in Transylvania, Romania. By global standards this system is diverse, including apex predators (brown bear and wolf), mesopredators (red fox) and large herbivores (roe and red deer). Humans and free-ranging dogs represent additional predators in the system. Using structural equation modelling, we found that apex predators suppress lower trophic levels, especially herbivores. However, direct and indirect top-down effects of humans affected the ecosystem more strongly, influencing species at all trophic levels. Our study highlights the need to explicitly embed humans and their influences within trophic cascade theory. This will greatly expand our understanding of species interactions in human-modified landscapes, which compose the majority of the Earth's terrestrial surface.

Life history and the ecology of stress: how do glucocorticoid hormones influence life-history variation in animals?

Glucocorticoids hormones (GCs) are intuitively important for mediation of age-dependent vertebrate life-history transitions through their effects on ontogeny alongside underpinning variation in life-history traits and trade-offs in vertebrates. These concepts largely derive from the ability of GCs to alter energy allocation, physiology and behaviour that influences key life-history traits involving age-specific life-history transitions, reproduction and survival. Studies across vertebrates have shown that the neuroendocrine stress axis plays a role in the developmental processes that lead up to age-specific early life-history transitions. While environmental sensitivity of the stress axis allows for it to modulate the timing of these transitions within species, little is known as to how variation in stress axis function has been adapted to produce interspecific variation in the timing of life-history transitions. Our assessment of the literature confirms that of previous reviews that there is only equivocal evidence for correlative or direct functional relationships between GCs and variation in reproduction and survival. We conclude that the relationships between GCs and life-history traits are complex and general patterns cannot be easily discerned with current research approaches and experimental designs. We identify several future research directions including: (i) integration of proximate and ultimate measures, including longitudinal studies that measure effects of GCs on more than one life-history trait or in multiple environmental contexts, to test explicit hypotheses about how GCs and life-history variation are related and (ii) the measurement of additional factors that modulate the effects of GCs on life-history traits (e.g. GC receptors and binding protein levels) to better infer neurendocrine stress axis actions. Conceptual models of HPA/I axis actions, such as allostatic load and reactive scope, to some extent explicitly predict the role of GCs in a life-history context, but are descriptive in nature. We propose that GC effects on life-history transitions, survival probabilities and fecundity can be modelled in existing quantitative demographic frameworks to improve our understanding of how GC variation influences life-history evolution and GC-mediated effects on population dynamics Lay Summary Functional Ecology

Urbanisation and the ecology of powerful owls ninox strenua in outer Melbourne, Victoria

Historically, the Powerful Owl (Ninox strenua) has been seen as a southeastern Australian species restricted to, or most numerous in, dense gullies of tall open forests in hilly or mountainous areas of the coast and Great Divide. However, recent research has revealed that Powerful Owls may breed numerously and successfully in a wider range of habitats than previously believed, including the forests and woodlands within the metropolitan areas of some major cities.Here we report on the breeding of a number of pairs of Powerful Owls in the Yarra Valley, Victoria. Study sites ranged from relatively undisturbed, wet sclerophyll forest 80 km from central Melbourne, through dry sclerophyll, eucalypt-dominated open forest with some disturbance, to a highly disturbed urban parkland only 18 km from central Melbourne. We found that Powerful Owls breed successfully in some urban areas, but are limited in the amount of human disturbance they can tolerate near their nesting hollow. In the most heavily utilized section of the urban parkland, all breeding attempts were unsuccessful and in one year the young were apparently eaten by one of the parents. This followed construction of a timber boardwalk under the nest tree during the breeding season. The Powerful Owls subsequently moved to a more secluded nesting hollow and raised two young. Recommendations for management of Powerful Owls in urban areas are discussed in the context
of these results.

Ecosystems and human well-being

■ Human well-being has several key components: the basic material needs for a good life, freedom and choice, health, good social relations, and personal security. Well-being exists on a continuum with poverty, which has been defined as"pronounced deprivation in well-being."
■ How well-being and ill-being, or poverty, are expressed and experienced is context- and situation-dependent, reflecting local social and personal factors such as geography, ecology, age, gender,and culture.These concepts are complex and value-laden.
■ Ecosystems are essential for human well-being through their provisioning, regulating, cultural, and supporting services. Evidence in recent decades of escalating human impacts. on ecological systems worldwide raises concerns about the consequences of ecosystem changes for human well-being.
■ Human well-being can be enhanced through sustainable human interaction with ecosystems with the support of appropriate instruments, institutions, organizations, and technology. creation of these through participation and transparency may contribute to people's freedoms and choices and to increased economic, social,and ecological security.
■ Some believe that the problems from the depletion and degradation of ecological capital can be largely overcome by the substitution of physical and human capital. Others believe that there are more significant limits to such substitutions.The scope for substitutions varies by socioeconomic status.
■ We identify direct and indirect pathways between ecosystem change and human well-being,whether it be positive or negative.lndirect effects are characterized by more complex webs of causation, involving social, economic, and political threads. Threshold points exist beyond which rapid changes to human well-being can occur.
■ Indigent poorly resourced, and otherwise disadvantaged communities are generally the most vulnerable to adverse ecosystem change. Spirals, both positive and negative, can occur for any population, but the poor are more vulnerable.      
■ Functioning institutions are vital to enable equitable access to ecosystem services. lnstitutions sometimes fail or remain undeveloped because of powerful individuals or groups. Bodies that mediate the distribution of goods and services may also be appropriated for the benefit of powerful minorities.
■ For poor people, the greatest gains in well-being will occur through more equitable and secure access to ecosystem services. In the long run, the rich can contribute greatly to human well-being by reducing their substantial impacts on ecosystems and by facilitating greater access to ecosystem services by the poor.
■ We argue ecological security warrants recognition as a sixth freedom of equal weight with participative freedom, economic   facilities, social opportunities, transparency guarantees, and protective security.

Intraspecific phenotypic variability in the black-eared miner (Manorina melanotis); human-facilitated introgression and the consequences for an endangered taxon

Prior to this study the circumscription of the endangered black-eared miner (Manorina melanotis) and the common yellow-throated miner (Manorina flavigula) has been clouded by the existence of hybrid individuals. We examined the intra- and inter-specific phenotypic variation of the two taxa. All available museum specimens (n=138) and a sample of live individuals (n=83) were examined. Cluster analysis revealed a continuum of phenotypic traits now exists between the two taxa. However, further analysis revealed the black-eared miner and yellow-throated miner were separable on phenotypic characters prior to extensive modification of mallee habitat after 1950, suggesting the black-eared miner should be afforded full species status [contrary to Schodde and Mason, 1999. (Schodde, R., Mason, I.J., 1999. The Directory of Australian Birds: Passerines. CSIRO Wildlife and Ecology, Canberra]. Our study highlights the need to carefully examine, not only intraspecific phenoptyic variation within a taxon, but to also consider how such variation may be affected by hybridisation facilitated by human disturbance of habitat.

Switching on the chronometer of history : communism, postcommunism and heritage

The destruction of monuments accompanying the fall of Communism ignited debates about preservation of manifestations of a hated regime. While heritage professionals called for their preservation as ‘historical documents’, many monuments were destroyed or removed. Yampolsky sees anti-Communist iconoclasm as a rejection of the totalitarianism of time embodied in Communist monuments. These ‘intentional monuments’ were intended to ‘negate the march of time and oppose to it the permanence of human action’. They demonstrated the alleged end of history in a classless utopia.

Iconoclastic acts against these monuments involved the crossing of ‘the invisible boundaries of the sacral zone surrounding monuments, switching on the chronometer of history’. In doing so, iconoclasts provide the conditions for reassertion of heritage practices: heritage requires a sense of the flow of time, a difference between past, present and future.

Having restarted the chronometer of history, a society is forced to assess where it stands in relation to its past. Will it continue on a path of ‘wilful forgetting’, or seek to confront the past? The danger of wilful forgetting is the creation of nostalgia. Alternatively, preservation of places of memory helps processing of the past required for movement into the future. ‘One need only consider the way in which Berliners tore down the hated Berlin Wall in the aftermath of 1989’, Fulbrook writes, ‘to understand the desire to rid the landscape of a hated excrescence, a symbol of a rejected political past. But…for those who come after, the effort of historical imagination is all the greater for lack of a topography of experience’.

Heritage preservation can produce a ‘topography of experience’, through which the experience of Communism is examined. Reassertion of a humanistic historical time through heritage practices reveals the arrogant futility of utopian projects seeking to bring history to an end.

Bob Brown: ecology, economy, equality and eternity

"Against a backdrop of advancing neoliberalism and globalisation, this timely book examines nine prominent Australians from diverse backgrounds - ʻglobal citizensʾ who have each enhanced public life through promoting universal values and human rights. The book charts over 50 years of campaigning, and espouses perennial causes such as peace, social justice, ecological sustainability and gender and racial equality. Ultimately, this inspiring volume sends a message of hope for Australian society and provides a benchmark for all proponents of change."--Publisher description.

Community development and ecology : exploring the links

This paper will explore connections between the concepts of community development and ecology. Initially the tendency was to think there should be a total melding of the principles and practices of community development with those of an ecological understanding but on reflection this has not and indeed is not necessarily the case. The relative epistemological positioning of two different groups, one strongly associating with social justice and the need for people to be at the centre of our economic, environmental and social understanding; and the other clearly seeing the plant and ecology/environment being paramount. While there are a myriad of connections the focus of much community development has been around human welfare based on principles of social, political and economic justice. This has at times been to the detriment of ecological sustainability. Conversely ecology and particularly aspects of deep ecology have focussed on the 'other than human' aspects of the planet and at times seemed almost 'anti 'human and overlooking the need to work with the social almost entirely. This paper briefly outlines the historical separation of the social from the ecological then goes on to explore alternative understandings that bring together principles of community development and ecology. Three examples are used to highlight the principles and practices that are being used across diverse contexts but all informed by common norms and values that are consistent with both community development and ecology. Concepts such as subsidiarity, participation and empowerment that form the basis of community development praxis are critical to the development of local sustainability. The combination of these aspects is evidenced in the three examples. Each is very clearly located in the local context and is built on sound ecological and community development understandings but each is also well aware that the need for a broader perspective is imperative to achieving global goals.

Knowledge and education for a sustainable and socially just world : the ecology of innovation and creativity for humanity

Sustainability is a critical aim of Malaysian public policy and an important aim in education. Nonetheless, what sustainability means as it relates to education and the relationship between education and a sustainable future is unclear. In this paper I shall investigate the role that Universities in Malaysia play in shifting the practice and culture of innovation and creativity towards more sustainable values and outcomes. Sustainable education is based on ensuring that the capacities of students and the broader society are reengaged and empowered through connecting education to the needs and aspirations of civil society and moving away from neoliberal ideas of education as a practice of consumption towards, sustainable values of advancing human dignity.

Creativity and innovation within such an educational framework are goals and practices deeply connected and embedded within sustainable commitments to social justice, the public good, as well as individual growth and development which provide a critical legitimizing principle for university research and teaching. One of the key theoretical influences in making this argument will draw from the arguments of Amartya Sen whose theorization of capability may provide us with a way of thinking about social growth and development that is not possessively individualistic but rather socially concerned. I will discuss this in reference to the approach of University Sains Malaysia which provides an example of a public University seeking to engage sustainability and tie educational creativity and innovation back to the common good and a sustainable future.

The philosophical aim of this paper is to show how universities can pursue creativity and innovation as socially useful practices for advancing humane and sustainable values throughout Malaysian society and avoid the fusion of creativity with possessive individualism, consummerization and social irresponsibility. In this respect this paper addresses directly the theme of the conference: ‘Thinking Minds: Nurturing the Design of a Better Future'. '

To realise our national aspirations, a concerted effort is needed to increase our nation’s competitiveness, productivity and innovativeness. Attributes such as desire for knowledge, innovative thinking, creativity and competitiveness must be imbued within our people. The inculcation of moral values, progressiveness and performance-based cultures must also be instilled if we are to nurture successful individuals of the highest quality. This will determine our success as a knowledge-based economy.’ (Badawi 2007)

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.