Time and energy constraints in demanding phases of the annual cycle : an example of time limitation in refuelling migratory swans

The daily metabolizable energy intake of an animal is potentially limited by either the available feeding time or by its capacity to process energy. Animals are generally considered not to be time-limited but rather to be energy-processing-limited. This is concluded from the common observation that an animal's feeding time per day increases with a decrease in food density. We argue that such changes in feeding time are in theory also expected when no constraints are operating. Thus, a study of the constraints on energy intakes of free-living animals should be performed during demanding phases of the year. As an example, we collected data on time and energy budgets of Bewick's swan (Cygnus columbianus bewickii) refuelling during migration on fennel pondweed (Potamogeton pectinatus) tubers in two years differing two-fold in tuber biomass density. As predicted by time limitation, the feeding time (defined as the time with the head submerged) did not change in response to a change in food biomass density, both within and between years (averaging 12.2 h d⁻¹). Contrary to energy-processing limitation, and again in line with time limitation, the daily metabolizable energy intake varied, being greater in the year with high than in the year with low food densities. We conclude that more studies are needed of animals operating under demanding conditions before it can be assessed whether free-living animals are generally energy-processing- or time-limited.

Challenges within the annual cycle of long-distance migratory waders along the East Asian-Australasian flyway

 The thesis focuses on the conservation of migratory shorebirds, particularly in this part of the world. Loss of habitats is one of the greatest threats and prioritizing the conservation of these habitats is of great importance to the future of these species.

Ecological value of riparian zones to birds in forest landscapes

Riparian zones are a characteristic component of many landscapes throughout the world and increasingly are valued as key areas for biodiversity conservation. Their importance for bird communities has been well recognised in semi-arid environments and in modified landscapes where there is a marked contrast between riparian and adjacent non-riparian vegetation. The value of riparian zones in largely intact landscapes with continuous vegetation cover is less well understood. This research examined the importance of riparian habitats for avifauna conservation by investigating the ecological interactions contributing to the pattern of bird assemblages in riparian and adjacent non-riparian habitats. Specifically, the focus is on the bird assemblages of riparian zones and those of adjacent non-riparian vegetation types and the influence that associated differences in resource availabilities, habitat structure and conditions have on observed patterns. This study was conducted in the foothill forests of the Victorian Highlands, south-east Australia. Mixed-species eucalypt (genus Eucalyptus) forests dominate the vegetation of this region. Site selection was based on the occurrence of suitable riparian habitat interspersed within extensive, relatively undisturbed (i.e. no recent timber harvesting or fire events) forest mosaics. A series of 30 paired riparian and non-riparian sites were established among six stream systems in three forest areas (Bunyip State Park, Kinglake National Park and Marysville State Forest). Riparian sites were positioned alongside the stream and the non-riparian partner site was positioned on a facing slope at a distance of approximately 750 m. Bird surveys were carried out during 29 visits to each site between July 2001 and December 2002. Riparian sites were floristically distinct from non-riparian sites and had a more complex vegetation structure, including a mid-storey tree layer mostly absent from non-riparian sites, extensive fine litter and coarse woody debris, and dense ground-layer vegetation (e.g. sedges and ground ferns). The characteristic features of non-riparian habitats included a relatively dense canopy cover, a ground layer dominated by grasses and fine litter, and a high density of canopy-forming trees in the smaller size-classes. Riparian zones supported a significantly greater species richness, abundance and diversity of birds when compared to non-riparian habitats. The composition of bird assemblages differed significantly between riparian and non-riparian habitats, with riparian assemblages displaying a higher level of similarity among sites. The strongest contributors to observed dissimilarities between habitat types included species that occurred exclusively in either habitat type or species with large contrasts in abundance between habitat types. Much of the avifauna (36%) of the study area is composed of species that are common and widespread in south-east Australia (i.e. forest generalists). Riparian habitats were characterised by a suite of species more typical of wetter forest types in south-east Australia and many of these species had a restricted distribution in the forest mosaic. Some species (7%) occurred exclusively in riparian habitats (i.e. riparian selective species) while others (43%) were strongly linked to these habitats (i.e. riparian associated species). A smaller proportion of species occurred exclusively (2%) in non-riparian habitats (i.e. non-riparian selective species) or were strongly linked to these habitats (10%; i.e. non-riparian associated species). To examine the seasonal dynamics of assemblages, the variation through time in species richness, abundance and composition was compared between riparian and non-riparian sites. Riparian assemblages supported greater richness and abundance, and displayed less variation in these parameters, than non-riparian assemblages at all times. The species composition of riparian assemblages was distinct from non-riparian assemblages throughout the annual cycle. An influx of seasonal migrants elevated species richness and abundance in the forest landscape during spring and summer. The large-scale movement pattern (e.g. coastal migrant, inland migrant) adopted by migrating species was associated with their preference for riparian or non-riparian habitats in the landscape. Species which migrate north-south along the east coast of mainland Australia (i.e. coastal migrants) used riparian zones disproportionately; eight of eleven species were riparian associated species. Species which migrate north-south through inland Australia (i.e. inland migrants) were mostly associated with non-riparian habitats. The significant differences in the dynamics of community structure between riparian and non-riparian assemblages shows that there is a disproportionate use of riparian zones across the landscape and that they provide higher quality habitat for birds throughout the annual cycle. To examine the ecological mechanisms by which riparian assemblages are richer and support more individual birds, the number of ecological groups (foraging, nest-type and body mass groups) represented, and the species richness of these groups, was compared between riparian and non-riparian assemblages. The structurally complex vegetation and distinctive habitat features (e.g. aquatic environments, damp sheltered litter) provided in the riparian zone, resulted in the consistent addition of ecological groups to riparian assemblages (e.g. sheltered ground – invertebrates foraging group) compared with non-riparian assemblages. Greater species richness was accommodated in most foraging, nest-type and body mass groups in riparian than non-riparian assemblages. Riparian zones facilitated greater richness within ecological groups by providing conditions (i.e. more types of resources and greater abundance of resources) that promoted ecological segregation between ecologically similar species. For a set of commonly observed species, significant differences in their use of structural features, substrates and heights were registered between riparian and non-riparian habitats. The availability and dynamics of resources in riparian and non-riparian habitats were examined to determine if there is differential availability of particular resources, or in their temporal availability, throughout the annual cycle. Riparian zones supported more abundant and temporally reliable eucalypt flowering (i.e. nectar) than non-riparian habitats throughout the annual cycle. Riparian zones also supported an extensive loose bark resource (an important microhabitat for invertebrates) including more peeling bark and hanging bark throughout the year than at non-riparian sites. The productivity of eucalypts differed between habitat types, being higher in riparian zones at most times for all eucalypts combined, and for some species (e.g. Narrow-leaved Peppermint Eucalyptus radiata). Non-riparian habitats provided an abundant nectar resource (i.e. shrub flowering) at particular periods in the annual cycle. Birds showed clear relationships with the availability of specific food (i.e. nectar) and foraging resources (i.e. loose bark). The demonstration of a greater abundance of resources and higher primary productivity in riparian zones is consistent with the hypothesis that these linear strips that occupy only a small proportion of the landscape have a disproportionately high value for birds. Riparian zones in continuous eucalypt forest provide high quality habitats that contribute to the diversity of habitats and resources available to birds in the forest mosaic, with positive benefits for the landscape-level species pool. Despite riparian and non-riparian habitat supporting distinct assemblages of birds, strong linkages are maintained along the riparian-upslope gradient. Clearly, the maintenance of diverse and sustainable assemblages of birds in forest landscapes depends on complementary management of both riparian and non-riparian vegetation.

Modelling behavioural and fitness consequences of disturbance for geese along their spring flyway

1. For migratory birds the implications of environmental change may be difficult to predict because they use multiple sites during their annual cycle. Moreover, the migrants’ use of these sites may be interdependent. Along the flyway of the Svalbard pink-footed goose Anser brachyrhynchus population, Norwegian farmers use organized scaring to minimize goose use of their grasslands in spring. We assessed the consequences of this practice for regional site use of pink-footed geese along their spring migration route.

2. We used dynamic programming to find the sequence of migratory decisions that maximizes the fitness of female geese during spring migration, assuming scaring impinges on both food-intake rates and predation risk. The parameterization of the model was based on data gathered from individually marked pink-footed geese between 1991 and 2003.

3. The effect of scaring in terms of fitness and site use was most noticeable regarding food-intake rate. Scaring resulted in a redistribution of geese along the flyway. Furthermore, the outcomes of the modelling exercises were highly dependent on whether or not the geese were omniscient or naive: at moderate scaring levels naive geese were predicted to succumb.

4. On a qualitative basis there was good correspondence between the predictions from the model and the empirical evidence gathered to date.

5. Synthesis and applications. Besides highlighting the importance of learning and changing behaviour in an adaptive fashion, our modelling exercise indicated the potential vulnerability of the geese to abrupt environmental change. In addition, the exercise emphasized the interdependence of site use along the migratory flyway. The model supports the necessity for an integrated flyway management approach. In Norway, discussion is ongoing about the future management of the spring conflict between farming interests and geese. Farmers in north and mid-Norway have announced that they will expand the scaring campaign if a long-term solution, including a compensation scheme, is not forthcoming. If scaring on such a large scale is implemented abruptly, it may have severe consequences for the population: management of both the scaring intensity and its geographical extent is urgently required.

On the growth and reproductive biology of asp, Aspius vorax, population from the middle reaches of Euphrates river

Endemic asp, Aspius vorax, from the middle section of the Euphrates River flowing through eastern Syria were studied to determine the main characteristics of their population structure, morphological parameters and reproductive biology. Samples ranged between 0+ and 4+ years of age and were dominated by 2+ years old group. Total length (TL) ranged between 19 and 70 cm corresponding with 46 to 2824.5 g weight, respectively. Fish growth has isometric pattern and the overall sex ratio was unbiased. Seasonal changes in the condition factor were related with the water temperature as well as the spawning season. Annual cycle of gonadosomatic index (GSI) readings indicated that spawning season occur around March when fish longer than 36 cm can mate. Average pre-spawning GSI was greater in individuals older than 2 years. Meanwhile, female fecundity was highly related to TL and weight. These findings did not always concur with previous observations from other asp populations, mainly in southern and northern Mesopotamia. Our results highlighted basic biological aspects of the local population and indicated differences between populations which can assist in fisheries management, conservation and commercial culture of the investigated species.

The performance of a student evaluation of teaching system

Student evaluation of teaching (SET) is now commonplace in many universities internationally. While much effort has been devoted to examining the statistical validity of SET instruments, there has been limited examination of the methodological and consequential validity (together referred to as ‘utility’) of the ways in which SET data are used. This paper examines the SET system at Deakin University from the perspective of utility. It draws on publicly available SET results for an entire annual cycle of unit offerings. Consideration is given to the representativeness of the data produced, and to the utility of the data reported, by the system. While this investigation focuses on the SET system currently employed at Deakin University, it offers both an analysis methodology and conclusions that can be applied more generally.

GeoLight - processing and analysing light-based geolocator data in R

1. Determining global position by light measurements (‘geolocation’) has revolutionised the methods used to track migratory birds throughout their annual cycle.2. To date, there is no standard way of analysing geolocator data, making communication of analyses cumbersome and hampering the reproducibility of results.3. We have, therefore, developed the R package GeoLight, which provides basic functions for all steps of determining global positioning and a new approach in analysing movement pattern.4. Here, we briefly introduce and discuss the major functions of this package using example movement data of European hoopoe (Upupa epops).

Context-dependent correlation between resting metabolic rate and daily energy expenditure in wild chipmunks

Several empirical studies have shown that variation in daily energy expenditure (DEE) and resting metabolic rate (RMR) is influenced by environmental and individual factors, but whether these shared influences are responsible for, or independent of, relationships between DEE and RMR remains unknown. The objectives of this study were to (i) simultaneously evaluate the effects of environmental and individual variables on DEE and RMR in free-ranging eastern chipmunks (Tamias striatus) and (ii) quantify the correlation between DEE and RMR before and after controlling for common sources of variation. We found that the influence of individual factors on DEE and RMR is most often shared, whereas the influence of environmental factors tends to be distinct. Both raw and mass-adjusted DEE and RMR were significantly correlated, but this correlation vanished after accounting for the shared effect of reproduction on both traits. However, within reproductive individuals, DEE and RMR remained positively correlated after accounting for all other significant covariates. The ratio of DEE to RMR was significantly higher during reproduction than at other times of the year and was negatively correlated with ambient temperature. DEE and RMR appear to be inherently correlated during reproduction, but this correlation does not persist during other, less energy-demanding periods of the annual cycle.

Prolonged and flexible primary moult overlaps extensively with breeding in beach-nesting Hooded Plovers Thinornis rubricollis

We present the first report of complete overlap of breeding and moult in a shorebird. In southeastern Australia, Hooded Plovers Thinornis rubricollis spend their entire lives on oceanic beaches, where they exhibit biparental care. Population moult encompassed the 6-month breeding season. Moult timing was estimated using the Underhill-Zucchini method for Type 2 data with a power transformation to accommodate sexual differences in rates of moult progression in the early and late stages of moult. Average moult durations were long in females (170.3 ± 14.2 days), and even longer in males (210.3 ± 13.5 days). Breeding status was known for most birds in our samples, and many active breeders (especially males) were also growing primaries. Females delayed the onset of primary moult but were able to increase the speed of moult and continue breeding, completing moult at about the same time as males. The mechanism by which this was achieved appeared to be flexibility in moult sequence. All moult formulae fell on one of two linked moult sequences, one faster than the other. The slower sequence had fewer feathers growing concurrently and also had formulae indicating suspended moults. Switching between sequences via common formulae is possible at many points during the moult cycle, and three of 12 recaptures were confirmed to have switched sequences in the same moult season. Hooded Plovers thus have a prolonged primary moult with the flexibility to change their rate of moult; this may facilitate high levels of replacement clutches that are associated with passive nest defence and low reproductive success. © 2014 British Ornithologists' Union.

Regional differences in the reproductive parameters of the sparsely-spotted stingaree, Urolophus paucimaculatus, from South-Eastern Australia

Whether spatial variation occurs in the life-history traits of chondrichthyan species is important to fisheries modelling and assessments. A study on the reproductive parameters of Urolophus paucimaculatus from four separate regions across south-eastern Australia found regional differences in maximum total length (TL), size-at-maturity, size-at-maternity and litter sizes. Inshore embayments (Port Phillip Bay (PPB) and Corner Inlet (CI)) appear to allow for larger TLs (females and males) than do offshore areas (Lakes Entrance (LE) and Western Bass Strait (WBS)). Size-at-maturity and size-at-maternity decreased across longitude from west (PPB) to east (LE) and seasonality of parturition and ovulation occurred earlier in PPB (August-October) than in LE (September-December). Maximum litter size correlated with maximum TL (six in PPB, five in each of CI and LE, and four in WBS). There was uncertainty in classifying females for maternal condition because the reproductive cycle appears to range from a continuous annual cycle to a non-continuous biennial cycle. Much of the uncertainty arises from the ambiguity of observation of non-pregnant mature females, which have either aborted through capture and handling, or are in a 'resting year' between pregnancies. Most likely, the majority are reproducing annually with an unknown proportion of females non-continuous and resting between pregnancies.

Migratory animals couple biodiversity and ecosystem functioning worldwide

Animal migrations span the globe, involving immense numbers of individuals from a wide range of taxa. Migrants transport nutrients, energy, and other organisms as they forage and are preyed upon throughout their journeys. These highly predictable, pulsed movements across large spatial scales render migration a potentially powerful yet underappreciated dimension of biodiversity that is intimately embedded within resident communities. We review examples from across the animal kingdom to distill fundamental processes by which migratory animals influence communities and ecosystems, demonstrating that they can uniquely alter energy flow, food-web topology and stability, trophic cascades, and the structure of metacommunities. Given the potential for migration to alter ecological networks worldwide, we suggest an integrative framework through which community dynamics and ecosystem functioning may explicitly consider animal migrations.

Determinants of individual foraging specialization in large marine vertebrates, the Antarctic and subantarctic fur seals

The degree of individual specialization in resource use differs widely among wild populations where individuals range from fully generalized to highly specialized. This interindividual variation has profound implications in many ecological and evolutionary processes. A recent review proposed four main ecological causes of individual specialization: interspecific and intraspecific competition, ecological opportunity and predation. Using the isotopic signature of subsampled whiskers, we investigated to what degree three of these factors (interspecific and intraspecific competition and ecological opportunity) affect the population niche width and the level of individual foraging specialization in two fur seal species, the Antarctic and subantarctic fur seals (Arctocephalus gazella and Arctocephalus tropicalis), over several years. Population niche width was greater when the two seal species bred in allopatry (low interspecific competition) than in sympatry or when seals bred in high-density stabilized colonies (high intraspecific competition). In agreement with the niche variation hypothesis (NVH), higher population niche width was associated with higher interindividual niche variation. However, in contrast to the NVH, all Antarctic females increased their niche width during the interbreeding period when they had potential access to a wider diversity of foraging grounds and associated prey (high ecological opportunities), suggesting they all dispersed to a similar productive area. The degree of individual specialization varied among populations and within the annual cycle. Highest levels of interindividual variation were found in a context of lower interspecific or higher intraspecific competition. Contrasted results were found concerning the effect of ecological opportunity. Depending on seal species, females exhibited either a greater or lower degree of individual specialization during the interbreeding period, reflecting species-specific biological constraints during that period. These results suggest a significant impact of ecological interactions on the population niche width and degree of individual specialization. Such variation at the individual level may be an important factor in the species plasticity with significant consequences on how it may respond to environmental variability.

Hampered performance of migratory swans: intra- and inter-seasonal effects of avian influenza virus

The extent to which animal migrations shape parasite transmission networks is critically dependent on a migrant's ability to tolerate infection and migrate successfully. Yet, sub-lethal effects of parasites can be intensified through periods of increased physiological stress. Long-distance migrants may, therefore, be especially susceptible to negative effects of parasitic infection. Although a handful of studies have investigated the short-term, transmission-relevant behaviors of wild birds infected with low-pathogenic avian influenza viruses (LPAIV), the ecological consequences of LPAIV for the hosts themselves remain largely unknown. Here, we assessed the potential effects of naturally-acquired LPAIV infections in Bewick's swans, a long-distance migratory species that experiences relatively low incidence of LPAIV infection during early winter. We monitored both foraging and movement behavior in the winter of infection, as well as subsequent breeding behavior and inter-annual resighting probability over 3 years. Incorporating data on infection history we hypothesized that any effects would be most apparent in naïve individuals experiencing their first LPAIV infection. Indeed, significant effects of infection were only seen in birds that were infected but lacked antibodies indicative of prior infection. Swans that were infected but had survived a previous infection were indistinguishable from uninfected birds in each of the ecological performance metrics. Despite showing reduced foraging rates, individuals in the naïve-infected category had similar accumulated body stores to re-infected and uninfected individuals prior to departure on spring migration, possibly as a result of having higher scaled mass at the time of infection. And yet individuals in the naïve-infected category were unlikely to be resighted 1 year after infection, with 6 out of 7 individuals that never resighted again compared to 20 out of 63 uninfected individuals and 5 out of 12 individuals in the re-infected category. Collectively, our findings indicate that acute and superficially harmless infection with LPAIV may have indirect effects on individual performance and recruitment in migratory Bewick's swans. Our results also highlight the potential for infection history to play an important role in shaping ecological constraints throughout the annual cycle.