Pronounced inter-colony variation in the foraging ecology of Australasian gannets : influence of habitat differences

The predictability of prey due to oceanographic features can result in large aggregations of apex predators. Central place foragers, such as seabirds, are limited in their foraging duration and range during the breeding period, which can restrict their ability to reach such locations. Segregation by colony and sex can further restrict foraging range and may have direct implications for the foraging ecology of a species. In parts of its range, the Australasian gannet Morus serrator breeds in colonies of relatively close proximity and has recently been found to display sexual dimorphism. Two neighbouring breeding colonies in Bass Strait that experience divergent environmental conditions were investigated to determine whether these conditions or the sex of the individual are important variables influencing foraging behaviour in this species. GPS tracking and accelerometry were paired with stable isotope analysis to compare differences in foraging effort, habitat use and diet. Birds from Point Danger, a large colony located near a seasonally strong upwelling, travelled considerably further (77%) than birds from Pope's Eye, a small colony in a nutrient-poor embayment. However, within colonies no sexual differences in foraging effort were found. While the colonies did not overlap in foraging areas, a degree of sexual segregation was apparent within both colonies (Point Danger 46.3% overlap and Pope's Eye 73.7% overlap in home range). Furthermore, stable isotope analysis indicated birds from each colony fed at different trophic niches. This study reveals differences in habitat use and, consequently, dietary niches between and within neighbouring colonies.

Relationship between long-term environmental fluctuations and diving effort of female Australian fur seals

For predators foraging within spatially and temporally heterogeneous marine ecosystems, environmental fluctuations can alter prey availability. Using the proportion of time spent diving and foraging trip duration as proxies of foraging effort, a multi-year dataset was used to assess the response of 58 female Australian fur seals Arctocephalus pusillus doriferus to interannual environmental fluctuations. Multiple environmental indices (remotely sensed ocean colour data and numerical weather predictions) were assessed for their influence on inter-annual variations in the proportion of time spent diving and trip duration. Model averaging revealed strong evidence for relationships between 4 indices and the proportion of time spent diving. There was a positive relationship with effort and 2 yr-lagged spring sea-surface temperature, current winter zonal wind and southern oscillation index, while a negative relationship was found with 2 yr-lagged spring zonal wind. Additionally, a positive relationship was found between foraging trip duration and 1 yr-lagged spring surface chlorophyll a. These results suggest that environmental fluctuations may influence prey availability by affecting the survival and recruitment of prey at the larval and post-larval phases while also affecting current distribution of adult prey.

Time-in-area represents foraging activity in a wide-ranging pelagic forager

Successful Marine Spatial Planning depends upon the identification of areas with high importance for particular species, ecosystems or processes. For seabirds, advancements in biologging devices have enabled us to identify these areas through the detailed study of at-sea behaviour. However, in many cases, only positional data are available and the presence of local biological productivity and hence seabird foraging behaviour is inferred from these data alone, under the untested assumption that foraging activity is more likely to occur in areas where seabirds spend more time. We fitted GPS devices and accelerometers to northern gannets Morus bassanus and categorised the behaviour of individuals outside the breeding colony as plunge diving, surface foraging, floating and flying. We then used the locations of foraging events to test the efficiency of 2 approaches: time-in-area and kernel density (KD) analyses, which are widely employed to detect highly-used areas and interpret foraging behaviour from positional data. For KD analyses, the smoothing parameter (h) was calculated using the ad hoc method (KDad hoc), and KDh=9.1, where h = 9.1 km, to designate core foraging areas from location data. A high proportion of foraging events occurred in core foraging areas designated using KDad hoc, KDh=9.1, and time-in-area. Our findings demonstrate that foraging activity occurs in areas where seabirds spend more time, and that both KD analysis and the time-in-area approach are equally efficient methods for this type of analysis. However, the time-in-area approach is advantageous in its simplicity, and in its ability to provide the shapes commonly used in planning. Therefore, the time-in-area approach can be used as a simple way of using seabirds to identify ecologically important locations from both tracking and survey data.

Influence of intrinsic variation on foraging behaviour of adult female Australian fur seals

Phenotypic variation and individual experience can create behavioural and/or dietary variation within a population. This may reduce intra-specific competition, creating a buffer to environmental change. This study examined how intrinsic variation affects foraging behaviour of Australian fur seals. Foraging movements of 29 female Australian fur seals were recorded using FastLoc GPS and dive behaviour recorders. For each individual, body mass, flipper length and axis length were recorded, a tooth was sampled to determine age, and milk was collected for diet analysis. Clustering of fatty acid dietary analysis revealed 5 distinct groups in the population. Behaviour was described using 19 indices, which were then reduced to 7 principal components (>80% of the behavioural variation). Bayesian mixed effect models were developed to describe the relationship between these components and intrinsic variation. No association was found between diet and age or body shape; however, age had a negative relationship with component 1 (27% of variation). Older females spent less time at-sea and foraged nearer to the colony. Age had an effect on component 5 (7% of variation), which represented haul-outs and dive depth; older females made fewer visits to haul-out sites and dived deeper to the benthos. This suggests that as animals age they are able to utilise prior knowledge to exploit nearby foraging sites that younger animals are either unaware of, or have yet to gain the experience required to efficiently utilise. Mass had a negative effect on components representing the directedness of a foraging trip, suggesting heavier individuals were more likely to travel directly to a foraging site.

Exposure mediates transitions between bare and vegetated states in temperate mangrove ecosystems

The resilience of mangroves is dependent on their regeneration capacity. Patchy mid-19th century clearing dramatically affected this capacity, creating stable vegetated and unvegetated states in a fragmented temperate mangrove ecosystem. Mechanisms of mediation between states were tested by monitoring the survival and growth of planted mangrove seedlings and propagules on formerly forested bare mudflats and inside patches of existing forest. Survival (1 to 76%) and growth (-0.83 to 10.45 mm mo-1 increase in plant height) of seedlings was affected by (1) differing levels of exposure found at varying proximities to remnant forest and (2) differing inundation regimes both within and between sites that were randomly selected from locations that varied in aspect relative to prevailing winds. Increases in hydrodynamic energy within and between sites corresponded to a decrease in survival that was much more pronounced at locations that were exposed to prevailing winds. Growth rates were also generally lower at sites in exposed locations, but inundation regime was a more important determinant within sites, where growth was reduced at lower heights on the shore. Results suggest that stability of the bare mudflat state (caused by historical clearance of the mangrove forest) is dependent on level of exposure to hydrodynamic energy, and a return to a forested state is more likely where this exposure is lower. These results have implications for planning and implementing mangrove restoration projects and illustrate the role that physical factors can play in determining the resilience of disturbed temperate mangrove ecosystems.

Recovery pathways from small-scale disturbance in a temperate Australian seagrass

Recovery from disturbance is a key element of ecosystem persistence, and recovery can be influenced by large-scale regional differences and smaller local-scale variations in environmental conditions. Seagrass beds are an important yet threatened nearshore habitat and recover from disturbance by regrowth, vegetative extension and dispersive propagules. We described recovery pathways from small-scale disturbances in the seagrass Zostera nigricaulis in Port Phillip Bay, a large embayment in southeastern Australia, and tested whether these pathways differed between 5 regions with different hydrodynamic conditions and water quality, and between sites within those regions. Recovery pathways were broadly consistent. When aboveground biomass was removed, recovery, defined as the point at which disturbed areas converged with undisturbed controls, took from 2 to 8 mo, but when we removed above-and below-ground biomass, it took between 2 and 13 mo. There was no evidence of recovery resulting from sexual reproduction at any sites regardless of the presence of seeds in the sediment or flower production. We found no differences in recovery at the regional scale, but we found substantial differences between local sites. At some sites, rapid recovery occurred because seagrasses grew quickly, but at others, apparent recovery occurred because regrowth coincided with overall declines in cover of undisturbed areas. Recovery time was unrelated to seagrass canopy height, biomass, percentage cover, stem density, seed bank density, epiphyte cover or sediment organic matter in seagrass adjacent to disturbance experiments. This study highlights the importance of understanding fine-scale variation in local recovery mechanisms, which may override or obscure any regional signal.

Linking nitrogen sources utilised by seagrass in a temperate marine embayment to patterns of seagrass change during drought

Reductions in the extent of seagrass Zostera nigricaulis coverage in Port Phillip Bay (PPB), Australia, between 2000 and 2011 coincided with a prolonged period of drought (1997 to 2009) characterized by decreases in freshwater and nutrient inputs. This led us to hypothesize that patterns of seagrass expansion and decline in PPB may be linked to nutrient availability. Seagrasses in PPB can make use of a range of different nitrogen (N) sources depending on their relative availability. Accordingly, there is a need to identify the origin of the N utilised by seagrasses in order to understand how changes in the availability of nutrients from various sources may influence seagrass growth. This study used stable isotope analysis to estimate the contribution of different sources of N to seagrass growth in different parts of PPB. Source modelling indicated that regional patterns of N source utilisation matched changes in seagrass extent from 2000 to 2011. Regions in which seagrass declined contained a similar array of sources, including significant contributions from the catchment area, whereas regions where seagrass areas remained unchanged were largely dependent on a single N source (either fixation/recycled or sewage-derived). We propose that reductions in N from the catchment during the drought may have contributed to the decline of seagrasses in regions where N from the catchment is an important source. This finding is likely to have implications for the growth, distribution and resilience of Z. nigricaulis seagrass in PPB as well as in other parts of its range in southern Australia.

Sexual segregation in habitat use is smaller than expected in a highly dimorphic marine predator, the southern sea lion

Sexual segregation in habitat use is widely reported in many taxa and can profoundly influence the distribution and behaviour of animals. However, our knowledge of the mechanisms driving sexual segregation is still in its infancy (particularly in marine taxa) and the influence of extrinsic factors in mediating the expression of sex differences in foraging behaviour is underdeveloped. Here, we combine data from biologging tags, with stable isotope analysis of vibrissae, to assess sexual segregation in southern sea lions (SSL) (Otaria flavescens) breeding at the Falkland Islands in the South Atlantic. We found evidence to support segregation, most notably in δ13C and δ15N values. However, in spite of extreme sexual size dimorphism and differing constraints related to female-only parental care, adult male and adult female SSL overlapped considerably in isotopic niches and foraging area, and shared similar foraging trip characteristics (such as distance and duration). This is in contrast to SSL breeding in Argentina, where prior studies report sexual differences in foraging locations and foraging trip characteristics. We posit that sexual segregation in SSL is influenced by habitat availability (defined here as the width of the Patagonian Shelf) and individual foraging preferences, rather than commonly invoked individual-based limiting factors per se.

Utilization of organic matter by invertebrates along an estuarine gradient in an intermittently open estuary

In intermittently open estuaries, the sources of organic matter sustaining benthic invertebrates are likely to vary seasonally, particularly between periods of connection and disconnection with the ocean and higher and lower freshwater flows. This study investigated the contribution of allochthonous and autochthonous primary production to the diet of representative invertebrate species using stable isotope analysis (SIA) during the austral summer and winter (2008, 2009) in an intermittently open estuary on the south-eastern coast of Australia. As the study was conducted towards the end of a prolonged period of drought, a reduced influence of freshwater/terrestrial organic matter was expected. Sampling was conducted along an estuarine gradient, including upper, middle and lower reaches and showed that the majority of assimilated organic matter was derived from autochthonous estuarine food sources. Additionally, there was an input of allochthonous organic matter, which varied along the length of the estuary, indicated by distinct longitudinal trends in carbon and nitrogen stable isotope signatures along the estuarine gradient. Marine seaweed contributed to invertebrate diets in the lower reaches of the estuary, while freshwater/terrestrial organic matter had increased influence in the upper reaches. Suspension-feeding invertebrates derived large parts of their diet from freshwater/terrestrial material, despite flows being greatly reduced in comparison with non-drought years.

Flocking and feeding in the fiddler crab (UCA tangeri): prey availability as risk-taking behaviour

For a full understanding of prey availability, it is necessary to study risk-taking behaviour of the prey. Fiddler crabs are ideally suited for such a study, as they have to leave their safe burrow to feed on the surface of the intertidal flats during low tide, thereby exposing themselves to avian predators. A study in an intertidal area along the coast of Mauritania showed that small crabs always stayed in the vicinity of their burrow, but large crabs wandered in large flocks (also referred to as droves) to feed on sea-grass beds downshore. Transplanting downshore feeding substrate to the burrowing zone of the small crabs proved that they too preferred to feed on it. Since small crabs can be preyed upon by more species of birds, this suggests that the decision not to leave the burrowing zone might be related to the risk of being fed upon by birds. We calculated predation risk from measurements on the density and feeding activity of the crabs, as well as the feeding density, the intake rate and the size selection of the avian predators. Per hour on the surface, crabs in a flock were more at risk than crabs feeding near their burrow. Thus, though flocking crabs may have benefited from ‘swamping the predator’ by emerging in maximum numbers during some tides only, this did not reduce their risk of predation below that of non-flocking crabs. Furthermore we found that irrespective of activity, large crabs suffered a higher mortality per tide from avian predators than small crabs. This suggests that large crabs could not sufficiently reduce their foraging time to compensate for the increased risk while foraging in a flock, even though they probably experienced better feeding conditions than small crabs staying near their burrow. The greater energy demands of large crabs were reflected in a greater surface area grazed. Thus, with increasing size a fiddler crab has to feed further away from its burrow and so may derive less protection from staying near to it. It seems that growing big does not reduce the risk of predation for fiddler crabs, as it does in many other species with indeterminate growth. As in such species, the most probable advantage of growing big is increased mating success. Ultimately, therefore, prey availability must be understood from the life-history decisions of the prey species.

Habitat selection and energetics of the fiddler crab (Uca tangeri)

We tried to unravel the possible links between the skewed predation risk in Uca tangeri (where large individuals are more at risk from avian predators) and size-dependent changes in the physiology and habitat choice of this fiddler crab species. Over a transect running from low to high in the tidal zone of a beach in Mauritania, the temperature profile at various depths in the substrate, the water-table level of seep water, salt concentration of seep water, depth of the aerobic level, operative temperatures on the surface, and size distribution of crabs were assessed. In addition, resting metabolic rates, Q10 and thermal and starvation tolerances were estimated. Going from low to high in the tidal zone, crab size and burrow depth increased. At the preferred burrowing depth, microclimatological conditions appeared to be equally favourable at all sites. At the surface, conditions were more favourable low in the tidal zone, where also food availability is sufficient to enable small crabs to forage in the vicinity of their burrows. Large crabs have higher energy requirements and are thereby forced to forage in flocks low in the tidal zone where food is probably more abundant. Low in the tidal zone, digging deeply is impossible as the aerobic layer is rather thin. Large crabs prefer living high in the tidal zone as (1) deep burrows ensure better protection against predators, (2) more time is available for digging holes and (3) the substrate is better suited for reproduction. Energy reserves in late summer ensured an average of 34 days of survival. It is argued that the allotment of energy to growth must be considerable even in reproducing animals; the rewards of growth being the disproportional increase in reproductive output with size.

Patterns of connectivity and population structure of the southern calamary Sepioteuthis australis in southern Australia

The southern calamary, Sepioteuthis australis, is a commercially and recreationally important inshore cephalopod endemic to southern Australia and New Zealand. Typical of other cephalopods, S. australis has a short life span, form nearshore spawning aggregations and undergo direct development. Such life history traits may restrict connectivity between spawning grounds creating highly structured and genetically differentiated populations that are susceptible to population crashes. Here we use seven polymorphic microsatellite markers to assess connectivity and population structure of S. australis across a large part of its geographic range in Australia. Little genetic differentiation was found between sampling locations. Overall, FST was low (0.005, 95% CI≤<0.001-0.011) and we detected no significant genetic differentiation between any of the locations sampled. There was no strong relationship between genetic and geographical distance, and our neighbour joining analysis did not show clustering of clades based on geographical locations. Similarly, network analysis showed strong connectivity amongst most locations, in particular, Tasmania appears to be well connected with several other locations and may act as an important source population. High levels of gene flow and connectivity between S. australis sampling sites across Australia are important for this short-lived species, ensuring resilience against spatial and temporal mortality fluctuations.

Human threats to sandy beaches: a meta-analysis of ghost crabs illustrates global anthropogenic impacts

Beach and coastal dune systems are increasingly subjected to a broad range of anthropogenic pressures that on many shorelines require significant conservation and mitigation interventions. But these interventions require reliable data on the severity and frequency of adverse ecological impacts. Such evidence is often obtained by measuring the response of 'indicator species'.Ghost crabs are the largest invertebrates inhabiting tropical and subtropical sandy shores and are frequently used to assess human impacts on ocean beaches. Here we present the first global meta-analysis of these impacts, and analyse the design properties and metrics of studies using ghost-crabs in their assessment. This was complemented by a gap analysis to identify thematic areas of anthropogenic pressures on sandy beach ecosystems that are under-represented in the published literature.Our meta-analysis demonstrates a broad geographic reach, encompassing studies on shores of the Pacific, Indian, and Atlantic Oceans, as well as the South China Sea. It also reveals what are, arguably, two major limitations: i) the near-universal use of proxies (i.e. burrow counts to estimate abundance) at the cost of directly measuring biological traits and bio-markers in the organism itself; and ii) descriptive or correlative study designs that rarely extend beyond a simple 'compare and contrast approach', and hence fail to identify the mechanistic cause(s) of observed contrasts.Evidence for a historically narrow range of assessed pressures (i.e., chiefly urbanisation, vehicles, beach nourishment, and recreation) is juxtaposed with rich opportunities for the broader integration of ghost crabs as a model taxon in studies of disturbance and impact assessments on ocean beaches. Tangible advances will most likely occur where ghost crabs provide foci for experiments that test specific hypotheses associated with effects of chemical, light and acoustic pollution, as well as the consequences of climate change (e.g. species range shifts).

Examining the functional role of current area closures used for the conservation of an overexploited and highly mobile fishery species

Protecting essential habitats through the implementation of area closures has been recognized as a useful management tool for rebuilding overfished populations and minimizing habitat degradation. School shark (Galeorhinus galeus) have suffered significant stock declines in Australia; however, recent stock assessments suggest the population may have stabilized and the protection of closed nursery areas has been identified as a key management strategy to rebuilding their numbers. Young-of-The-year (YOY) and juvenile G. galeus were acoustically tagged and monitored to determine ontogenetic differences in residency and seasonal use of an important protected nursery area (Shark Refuge Area or SRA) in southeastern Tasmania. BothYOYand juvenile G. galeus showed a distinct seasonal pattern of occurrence in the SRAwith most departing the area during winter and only a small proportion of YOY (33%) and no juveniles returning the following spring, suggesting areas outside the SRA may also be important during these early life-history stages. While these behaviors confirm SRAs continue to function as essential habitat during G. galeus early life history, evidence of YOY and juveniles emigrating from these areas within their first 1-2 years and the fact that few YOY return suggest that these areas may only afford protection for a more limited amount of time than previously thought. Determining the importance of neighbouring coastal waters and maintaining the use of traditional fisheries management tools are therefore required to ensure effective conservation of G. galeus during early life history.

Identifying knowledge gaps in seagrass research and management: An Australian perspective.

Seagrass species form important marine and estuarine habitats providing valuable ecosystem services and functions. Coastal zones that are increasingly impacted by anthropogenic development have experienced substantial declines in seagrass abundance around the world. Australia, which has some of the world's largest seagrass meadows and is home to over half of the known species, is not immune to these losses. In 1999 a review of seagrass ecosystems knowledge was conducted in Australia and strategic research priorities were developed to provide research direction for future studies and management. Subsequent rapid evolution of seagrass research and scientific methods has led to more than 70% of peer reviewed seagrass literature being produced since that time. A workshop was held as part of the Australian Marine Sciences Association conference in July 2015 in Geelong, Victoria, to update and redefine strategic priorities in seagrass research. Participants identified 40 research questions from 10 research fields (taxonomy and systematics, physiology, population biology, sediment biogeochemistry and microbiology, ecosystem function, faunal habitats, threats, rehabilitation and restoration, mapping and monitoring, management tools) as priorities for future research on Australian seagrasses. Progress in research will rely on advances in areas such as remote sensing, genomic tools, microsensors, computer modeling, and statistical analyses. A more interdisciplinary approach will be needed to facilitate greater understanding of the complex interactions among seagrasses and their environment.