Drifting or walking? colonisation routes used by different instars and species of lotic, macroinvertebrate filter feeders

Many views of stream invertebrate populations centre on drift as the major route of larval dispersal, but few studies have presented unambiguous information about the role of drift. We present the results from an experiment designed to determine whether the major route of colonisation of substrata by hydropsychid larvae (commonly found in the drift) is by drifting directly onto substrata or by walking along the stream bottom. The experimental design contained four treatments: substrata open to drifters and walkers; fenced substrata open to drifters only; and two treatments open to drifters and walkers that provided forms of fence controls. Fifteen replicates of each treatment were set out at random locations within a riffle at each of three sites, with each site on a different river (the Little River, the Steavenson River and the Acheron River) in the Acheron River catchment. The experiment was run twice, once during autumn (April 1999) and once during early summer (January 2000). Both experiments were colonised by three species of hydropsychids, Asmicridea sp. AV1, and Smicrophylax sp. AV1 and AV2. We found that 2nd/3rd instars of Asmicridea sp. AV1 walked as well as drifted, whereas all others primarily drifted. No relation between numbers of recruits and water speed was found when substrata were open only to drifters, whereas substrata open also to walkers gained more recruits in faster flows. Additionally, larvae more frequently abandoned nets in slow than fast flows, indicating that drifting into unfavourable flow environments may result in mortality or redispersal of larvae. These findings demonstrate that, although drift is important, it is not necessarily the only method used by hydropsychids to colonise substrata. Larvae may have more capacity to choose substrata in fast flows when they colonise substrata by walking. Spot measures of hydropsychid distribution cannot distinguish between these explanations. The finding that walkers can sometimes comprise significant numbers of recruits raises the prospect that hydropsychids can be sourced locally and have not inevitably drifted in from upstream locations.

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.

Succession of small mammal communities after fire and reintroduction of the swamp antechinus 'Antechinus minimus'

This thesis is involved with changes that have occurred to small mammal populations following a major disturbance in the Anglesea region as a result of the 1983 Ash Wednesday fires. Fire, with its effects on spatial and temporal heterogeneity, was found to be an important factor in the maintenance of vegetation and small mammal community structure and diversity in the region. Successional changes in vegetation and small mammal communities were described by multivariate analyses, using data collected annually from 22 study sites. The use of factor analysis techniques, in reducing the annual capture data content, enabled long-term changes in the structure of mammal communities to be interpreted. The small mammal communities in the coastal heath and forest vegetation in the Anglesea region show evidence of a general resilience, (the degree and speed of recovery), to disturbance. Two phases of successional response to fire by mammal species have been proposed; a ‘re-establishment’ phase which occurs in the initial 5-6 years post-fire and is accompanied by rapid increase in species’ abundance, and a subsequent ‘maintenance’ phase accompanied by relatively minor changes in abundance. Habitat Suitability Indices were produced relating to these phases. Vertical density measures of understorey shrubs and herb layers showed significant relationships with small mammal species abundance at the study sites. Long term studies following major disturbances are needed to distinguish between short term recovery of plant and animal species and long term changes in these species. Studies extending over a number of years enable a better directional view of changes in small mammal communities than can be determined from . observations made over a short period. As a part of the investigation into temporal change, it was proposed to undertake trial reintroductions of the Swamp antechinus, Ant echinus minimus, a marsupial dasyurid species which was trapped in the area prior to the 1983 fire, but rarely subsequently. Other more commonly observed native small mammal species (e.g. Rattus fuscipes,R. lutreolus, Antechinus stuartii, Sminthopsis leucopus) had re-invaded the proposed reintroduction site after this fire. Failure of A. minimus to re-establish may have been due to spatial separation of the pre-fire populations coupled with the extensive area burnt in 1983, A source population of the species was located about 100km to the west and habitat utilization and interspecific and niche relationships between the species making the small mammal community explored. Discriminant analysis revealed some spatial separation of species within a habitat based on structural vegetation factors rather than floristic factors. Temporal separation of species was observed, asA. minimus were more active than Rattus species during daylight periods. There was evidence of micro-habitat selection by species, and structural vegetation factors were most commonly identified in statistical analyses as contributing towards selection by small mammal species. Following a theoretical modelling study three reintroduction trials were carried out near Anglesea during 1992-94. Individuals were subsequently radio tracked, and habitat relationships between the species in the small mammal community investigated. Although successful breeding of A, minimus occurred during the latter two trials, the subsequent fate of offspring was not determined. Invasive techniques required to adequately monitor young animals were considered potentially too damaging. Telemetry studies indicated a preference of A. minimus for short, wet heath vegetation. Structural vegetation factors were identified as being significant in discriminating between capture locations of species. Small scale and inexpensive trial reintroductions have yielded valuable additional data on this species and may be viewed as a useful tool in the conservation of other small native mammals.

Spatial ecology of E. australis

This research investigated links between the occurrence of endangered southern right whales and characteristics of their habitat. Habitat selection is based on physical environmental features, with important roles for social cues and site fidelity. This complex interaction of factors affects spatial recovery capacity and presents challenges for conservation management.

Host behaviour and physiology underpin individual variation in avian influenza virus infection in migratory Bewick's swans

Individual variation in infection modulates both the dynamics of pathogens and their impact on host populations. It is therefore crucial to identify differential patterns of infection and understand the mechanisms responsible. Yet our understanding of infection heterogeneity in wildlife is limited, even for important zoonotic host-pathogen systems, owing to the intractability of host status prior to infection. Using novel applications of stable isotope ecology and eco-immunology, we distinguish antecedent behavioural and physiological traits associated with avian influenza virus (AIV) infection in free-living Bewick's swans (Cygnus columbianus bewickii). Swans infected with AIV exhibited higher serum δ13C (-25.3 ± 0.4) than their non-infected counterparts (-26.3±0.2). Thus, individuals preferentially foraging in aquatic rather than terrestrial habitats experienced a higher risk of infection, suggesting that the abiotic requirements of AIV give rise to heterogeneity in pathogen exposure. Juveniles were more likely to be infected (30.8% compared with 11.3% for adults), shed approximately 15-fold higher quantity of virus and exhibited a lower specific immune response than adults. Together, these results demonstrate the potential for heterogeneity in infection to have a profound influence on the dynamics of pathogens, with concomitant impacts on host habitat selection and fitness.

Early microhabitat use by age 0 year brook charr Salvelinus fontinalis in lakes

The early habitat use of age 0 year brook charr Salvelinus fontinalis in three north temperate lakes which differ in terms of shoreline physical habitat is described. In the two lakes, which contained abundant shoreline woody debris and inundated vegetation, brook charr were observed in extremely close proximity with these habitat features, near shore and near the surface. Fish were absent from open areas away from shore unless in close proximity with fallen floating logs near the surface, extending offshore. In a third lake that had no woody debris or inundated shoreline vegetation, brook charr were observed exclusively in close proximity with the shoreline itself, and near the surface. In all three lakes, fish were most closely associated with the shoreline and with woody debris and inundated vegetation (when present) shortly after emergence, and significantly farther from shore and deeper in the water column thereafter.

Satellite tracking reveals unusual diving characteristics for a marine reptile, the olive ridley turtle : Lepidochelys olivacea

The movements, diving behaviour and thermal environment occupied by 4 adult female olive ridley turtles Lepidochelys olivacea in northern Australia were determined through satellite telemetry. Patterns of behaviour recorded were rather unusual compared to other sea turtles in that dives were mainly deep, largely benthic and exceptionally long (>2 h) in some cases, characteristics typical of over-wintering turtles in colder environments. One individual occupied shallow coastal foraging zones, while the others foraged far from land (probably on the seabed) in relatively deep water (>100 m). Individuals performed long dives (frequently >100 min), but from the short post-dive intervals we suggest that these dives were mainly aerobic. Maximum dive depth recorded was 200 ± 20 m (mean maximum depths ranged from 20.1 to 46.7 m across individuals; n = 17328 dives in total; depths ≥3 m were considered ‘dives’) and the maximum duration was 200 ± 20 min (mean durations ranged from 24.5 to 48.0 min across individuals). Temperature profiles indicate that turtles experienced temperatures ranging from 23 to 29°C at the surface, with the lowest temperature recorded (18.7°C) at a depth of 98 m. Only 6.9% of the dives were in water <20°C. From time-allocation at depth (TAD) scores, we demonstrated that many dives reaching the known or inferred sea bottom were U-shaped, but there was no apparent diel signal in dive depth. This suggests that many benthic dives were not associated exclusively with resting behaviour and likely had a foraging component as well. The ability to perform long benthic dives allows this species to exploit deeper benthic environments in addition to the shallow coastal areas more generally occupied by adult hard-shelled sea turtles (e.g. green and hawksbill turtles). Deep benthic dives also occur in certain marine mammals (e.g. narwhals) and sea birds (e.g. rockhopper penguins) and therefore seem to be a general foraging strategy exploited by animals that can perform long dives.

Ontogeny of long distance migration

The movements of some long-distance migrants are driven by innate compass headings that they follow on their first migrations (e.g., some birds and insects), while the movements of other first-time migrants are learned by following more experienced conspecifics (e.g., baleen whales). However, the overall roles of innate, learned, and social behaviors in driving migration goals in many taxa are poorly understood. To look for evidence of whether migration routes are innate or learned for sea turtles, here for 42 sites around the world we compare the migration routes of >400 satellite-tracked adults of multiple species of sea turtle with ∼45 000 Lagrangian hatchling turtle drift scenarios. In so doing, we show that the migration routes of adult turtles are strongly related to hatchling drift patterns, implying that adult migration goals are learned through their past experiences dispersing with ocean currents. The diverse migration destinations of adults consistently reflected the diversity in sites they would have encountered as drifting hatchlings. Our findings reveal how a simple mechanism, juvenile passive drift, can explain the ontogeny of some of the longest migrations in the animal kingdom and ensure that adults find suitable foraging sites.

Winter foraging site fidelity of king penguins breeding at the Falkland Islands

Foraging site fidelity has profound consequences for individual fitness, population processes and the effectiveness of species conservation measures. Accordingly, quantifying site fidelity has become increasingly important in animal movement and habitat selection studies. To assess foraging site fidelity in king penguins (Aptenodytes patagonicus) breeding at the Falkland Islands (51.48°S, 57.83°W), we measured overlap in time spent in foraging areas (at a 0.1° × 0.1° grid resolution) between successive foraging trips and foraging route consistency during the crèche period. In total, 30 complete foraging trips from seven king penguins were recorded between April and October 2010. King penguins predominantly foraged on the highly productive Patagonian slope, to the north of the Falkland Islands [median foraging trip distance 213 km (SD = 215 km) and duration 12.8 days (SD = 14.7 days)]. Overlap in time spent in an area on consecutive foraging trips ranged between 2 and 73 % (mean 27 %, SD = 22 %). Bearing during the outbound portion of foraging trips was typically highly repeatable for individual birds, but foraging trip duration and distance were not. Travel during the outbound phase of foraging trips was consistent with the direction of the northward-flowing Falkland Current that may act as a directional cue or facilitate rapid transit to foraging areas. Flexibility in foraging trip distances and durations may be a response to changes in resource availability and changes in the energetic requirements of adults and chicks over an extended breeding cycle.

Towards a cohesive, holistic view of top predation: a definition, synthesis and perspective

Research on the ecology of top predators - upper trophic level consumers that are relatively free from predation once they reach adult size - has provided regular contributions to general ecology and is a rapidly expanding and increasingly experimental, multidisciplinary and technological endeavour. Yet, an exponentially expanding literature coupled with rapid disintegration into specialized, disconnected subfields for study (e.g. vertebrate predators versus invertebrate predators, community ecology versus biological control etc.) increasingly means that we are losing a coherent, integrated understating of the role and importance of these species in ecosystems. This process of canalization is likely to hinder sharing of scientific discovery and continued progress, especially as there is a growing need to understand the generality of the top-down forcing, as demonstrated for some members of this group. Here, we propose ways to facilitate synthesis by promoting changes in mentality and awareness among specialists through increased debate and collaboration, conceptual reviews and a series of exemplary case studies. The strategy will rely on the collective contribution by all scientists in the field and will strive to consolidate and formalise top-order predation as a holistic, cohesive, cross-taxonomical field of research studying the ecology, evolution and behaviour of apex predators and their capability to exert top-down forcing on lower trophic levels. © 2014 The Authors.

Diving deeper into individual foraging specializations of a large marine predator, the southern sea lion

Despite global declines in the abundance of marine predators, knowledge of foraging ecology, necessary to predict the ecological consequences of large changes in marine predator abundance, remains enigmatic for many species. Given that populations suffering severe declines are of conservation concern, we examined the foraging ecology of southern sea lions (SSL) (Otaria flavescens)-one of the least studied otariids (fur seal and sea lions)-which have declined by over 90 % at the Falkland Islands since the 1930s. Using a combination of biologging devices and stable isotope analysis of vibrissae, we redress major gaps in the knowledge of SSL ecology and quantify patterns of individual specialization. Specifically, we revealed two discrete foraging strategies, these being inshore (coastal) and offshore (outer Patagonian Shelf). The majority of adult female SSL (72 % or n = 21 of 29 SSL) foraged offshore. Adult female SSL that foraged offshore travelled further (92 ± 20 vs. 10 ± 4 km) and dived deeper (75 ± 23 vs. 21 ± 8 m) when compared to those that foraged inshore. Stable isotope analysis revealed long-term fidelity (years) to these discrete foraging habitats. In addition, we found further specialization within the offshore group, with adult female SSL separated into two clusters on the basis of benthic or mixed (benthic and pelagic) dive behavior (benthic dive proportion was 76 ± 9 vs. 51 ± 8 %, respectively). We suggest that foraging specialization in depleted populations such as SSL breeding at the Falkland Islands, are influenced by foraging site fidelity, and could be independent of intraspecific competition. Finally, the behavioral differences we describe are crucial to understanding population-level dynamics, impediments to population recovery, and threats to population persistence.

Animal behaviour and cancer

Scientists are increasingly coming to realize that oncogenic phenomena are both frequent and detrimental for animals, and must therefore be taken into account when studying the biology of wildlife species and ecosystem functioning. Here, we argue that several behaviours that are routine in an individual's life can be associated with cancer risks, or conversely prevent/cure malignancies and/or alleviate their detrimental consequences for fitness. Although such behaviours are theoretically expected to be targets for natural selection, little attention has been devoted to explore how they influence animal behaviour. This essay provides a summary of these issues as well as an overview of the possibilities offered by this research topic, including possible applications for cancer prevention and treatments in humans.

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.

Growth and reproductive dynamics of the South American red shrimp, Pleoticus muelleri (Crustacea: Solenoceridae), from the southeastern coast of Brazil

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)