Mimicry and the eye of the beholder

Recent experiments (Dittrich et al. (Proc. R. Soc. Lond. B 251, 195 (1993))) suggest that pigeon perception of wasp mimicry by hoverflies is similar to that of humans and of computer-based image matching. However, the relations are nonlinear and may explain why some species are abundant despite their being poor mimics to the human eye. We suggest that these discrepancies between pigeon and human categorization may lie in the differences between avian and primate colour vision. As pigeon categorization and computer image analysis were both assessed by using colour slides designed for human vision, they lacked the natural colour information available to wild birds, in particular that from ultraviolet (uv) wavelengths.

Tetrachromacy, oil droplets and bird plumage colours

There is a growing body of data on avian eyes, including measurements of visual pigment and oil droplet spectral absorption, and of receptor densities and their distributions across the retina. These data are sufficient to predict psychophysical colour discrimination thresholds for light-adapted eyes, and hence provide a basis for relating eye design to visual needs. We examine the advantages of coloured oil droplets, UV vision and tetrachromacy for discriminating a diverse set of avian plumage spectra under natural illumination. Discriminability is enhanced both by tetrachromacy and coloured oil droplets. Oil droplets may also improve colour constancy. Comparison of the performance of a pigeon's eye, where the shortest wavelength receptor peak is at 410 nm, with that of the passerine Leiothrix, where the ultraviolet-sensitive peak is at 365 nm, generally shows a small advantage to the latter, but this advantage depends critically on the noise level in the sensitivity mechanism and on the set of spectra being viewed.

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.

Estimating the contribution of carnivorous waterbirds to nutrient loading in freshwater habitats

1. We estimated nitrogen (N) and phosphorus (P) loading into wetlands by carnivorous waterbirds with alternative physiological models using a food-intake and an excreta-production approach. The models were applied for non-breeding and breeding Dutch inland carnivorous waterbird populations to quantify their contribution to nutrient loading on a landscape scale.

2. Model predictions based on food intake exceeded those based on excretion by 59–62% for N and by 2–36% for P, depending on dietary assumptions. Uncertainty analysis indicated that the intake model was most affected by errors in energy requirement, while the excretion model was dependent on faecal nutrient composition.

3. Per capita loading rate of non-breeders increased with body mass from 0.3–0.8 g N day⁻¹ and 0.15 g P day⁻¹ in little gulls Larus minutus to 4.5–11.5 g N day⁻¹ and 2.1–3.2 g P day⁻¹ in great cormorants Phalacrocorax carbo. For breeding birds, the estimated nutrient loading by a family unit over the entire breeding period ranged between 17.6–443.0 g N and 8.6 g P for little tern Sterna albifrons to 619.6–1755.6 g N and 316.2–498.1 g P for great cormorants.

4. We distinguished between external (i.e. importing) and internal (i.e. recycling) nutrient loading by carnivorous waterbirds. For the Netherlands, average external-loading estimates ranged between 38.1–91.5 tonnes N and 16.7–18.2 tonnes P per year, whilst internal-loading estimates ranged between 53.1–140.5 tonnes N and 25.2–39.2 tonnes P and per year. The average contribution of breeding birds was estimated to be 17% and 32% for external and internal loading respectively. Most important species were black-headed gull Larus ridibundus and mew gull Larus canus for external loading, and great cormorant and grey heron Ardea cinerea for internal loading.

5. On a landscape scale, loading by carnivorous waterbirds was of minor importance for freshwater habitats in the Netherlands with 0.26–0.65 kg N ha⁻¹ a⁻¹ and 0.12–0.16 kg P ha⁻¹ a⁻¹. However, on a local scale, breeding colonies may be responsible for significant P loading.

Habitat use throughout migration : linking individual consistency, prior breeding success and future breeding potential

1. Habitat use can influence individual performance in a wide range of animals, either immediately or through carry-over effects in subsequent seasons. Given that many animal species also show consistent individual differences in reproductive success, it seems plausible that individuals may have consistent patterns of habitat use representing individual specializations, with concomitant fitness consequences.

2. Stable-carbon isotope ratios from a range of tissues were used to discern individual consistency in habitat use along a terrestrial–aquatic gradient in a long-distance migrant, the Bewick’s swan (Cygnus columbianus bewickii). These individual specialisations represented <15% of the isotopic breadth of the population for the majority of individuals and were seen to persist throughout autumn migration and overwintering until aquatic habitats were no longer available.

3. Individual foraging specialisations were then used to demonstrate two consecutive carry-over effects associated with macroscale habitat segregation: consequences of breeding season processes for autumn habitat use; and consequences of autumn habitat use for future reproductive success. Adults that were successful breeders in the year of capture used terrestrial habitats significantly more than adults that were not successful, revealing a substantial cost of reproduction and extended parental care. Use of aquatic habitats during autumn was, however, associated with increased body condition prior to spring migration; and increased subsequent breeding success in adults that had been unsuccessful the year before. Yet adults that were successful breeders in the year of capture remained the most likely to be successful the following year, despite their use of terrestrial habitats.

4. Our results uniquely demonstrate not only individual foraging specializations throughout the migration period, but also that processes during breeding and autumn migration, mediated by individual consistency, may play a fundamental role in the population dynamics of long-distance migrants. These findings, therefore, highlight the importance of long-term consistency to our understanding of habitat function, interindividual differences in fitness, population dynamics and the evolution of migratory strategies.

Building genetic networks using relatedness information : a novel approach for the estimation of dispersal and characterization of group structure in social animals

Natal dispersal is an important life history trait driving variation in individual fitness, and therefore, a proper understanding of the factors underlying dispersal behaviour is critical to many fields including population dynamics, behavioural ecology and conservation biology. However, individual dispersal patterns remain difficult to quantify despite many years of research using direct and indirect methods. Here, we quantify dispersal in a single intensively studied population of the cooperatively breeding chestnut-crowned babbler (Pomatostomus ruficeps) using genetic networks created from the combination of pairwise relatedness data and social networking methods and compare this to dispersal estimates from re-sighting data. This novel approach not only identifies movements between social groups within our study sites but also provides an estimation of immigration rates of individuals originating outside the study site. Both genetic and re-sighting data indicated that dispersal was strongly female biased, but the magnitude of dispersal estimates was much greater using genetic data. This suggests that many previous studies relying on mark–recapture data may have significantly underestimated dispersal. An analysis of spatial genetic structure within the sampled population also supports the idea that females are more dispersive, with females having no structure beyond the bounds of their own social group, while male genetic structure expands for 750 m from their social group. Although the genetic network approach we have used is an excellent tool for visualizing the social and genetic microstructure of social animals and identifying dispersers, our results also indicate the importance of applying them in parallel with behavioural and life history data.

Kin selection, not group augmentation, predicts helping in an obligate cooperatively breeding bird

Kin selection theory has been the central model for understanding the evolution of cooperative breeding, where non-breeders help bear the cost of rearing young. Recently, the dominance of this idea has been questioned; particularly in obligate cooperative breeders where breeding without help is uncommon and seldom successful. In such systems, the direct benefits gained through augmenting current group size have been hypothesized to provide a tractable alternative (or addition) to kin selection. However, clear empirical tests of the opposing predictions are lacking. Here, we provide convincing evidence to suggest that kin selection and not group augmentation accounts for decisions of whether, where and how often to help in an obligate cooperative breeder, the chestnut-crowned babbler (Pomatostomus ruficeps). We found no evidence that group members base helping decisions on the size of breeding units available in their social group, despite both correlational and experimental data showing substantial variation in the degree to which helpers affect productivity in units of different size. By contrast, 98 per cent of group members with kin present helped, 100 per cent directed their care towards the most related brood in the social group, and those rearing half/full-sibs helped approximately three times harder than those rearing less/non-related broods. We conclude that kin selection plays a central role in the maintenance of cooperative breeding in this species, despite the apparent importance of living in large groups.

The role of objective and subjective evaluation in the production and marketing of goats for meat

Objective and subjective evaluations of goats for meat production are related to important determinants of production and profitability. The most important attributes in assessment of goats for market are: live weight; body condition score; and the age of goats. As goats grow, their carcass and body organs increase in weight in proportion to the empty body weight. For farmers and field workers the linear regression approach for estimating carcass weight by measuring live weight is the most suitable as it accounts for 88 to 97% of the variation in carcass, offal and boneless meat weight. Live weight scales or heart girth tapes should be used and the risks and errors associated with these methods are summarized. The proportion of a live goat that is the carcass, known as dressing percentage, increases from 35% to about 50% as goats grow. The usefulness and errors associated with dressing percentage in field estimation are discussed. A valuable subjective method for estimating the nutritional status of goats is the use of body condition scoring as it accounts for 60 to 67% of the variation in live weight change, carcass weight and fat reserves of goats. A method for body condition scoring and a similar fat scoring system are explained. Body condition score is also associated with mortality risk and reproductive performance of goats. The number of permanent incisors in the lower jaw of goats is a method of estimating the age of goats but is biased by differences in live weights of goats. The value and role of ultrasound scanning the carcasses of goats is summarized. For the marketing of kid meat no permanent incisors should have erupted. Other useful practices for the successful marketing of goat meat are discussed including: knowing market specifications and chemical withholding periods; animal health; prevention of bruising; identification of goats; size of consignments; timeliness; provision of paperwork. A checklist is provided. The use of subjective and objective assessment techniques in evaluating goats for meat production will provide the best results. Where only subjective assessment techniques are available they will provide satisfactory performance provided the skills have been learnt and are applied.

Trophic status drives interannual variability in nesting numbers of marine turtles

Large annual fluctuations are seen in breeding numbers in many populations of non–annual breeders. We examined the interannual variation in nesting numbers of populations of green (Chelonia mydas) (n = 16 populations), loggerhead (Caretta caretta) (n =10 populations), leatherback (Dermochelys coriacea) (n = 9 populations) and hawksbill turtles (Eretmochelys imbricata) (n = 10 populations). Interannual variation was greatest in the green turtle. When comparing green and loggerhead turtles nesting in Cyprus we found that green turtles were more likely to change the interval between laying seasons and showed greater variation in the number of clutches laid in a season. We suggest that these differences are driven by the varying trophic statuses of the different species. Green turtles are herbivorous, feeding on sea grasses and macro–algae, and this primary production will be more tightly coupled with prevailing environmental conditions than the carnivorous diet of the loggerhead turtle.

Whale watching disrupts feeding activities of minke whales on a feeding ground

Human disturbances of wildlife, such as tourism, can alter the activities of targeted individuals. Repeated behavioural disruptions can have long-term consequences for individual vital rates (survival and reproduction). To manage these sub-lethal impacts, we need to understand how activity disruptions can influence bioenergetics and ultimately individual vital rates. Empirical studies of the mechanistic links between whale-watching boat exposure and behavioural variation and vital rates are currently lacking for baleen whales (mysticetes). We compared minke whale Balaenoptera acutorostrata behaviour on a feeding ground in the presence and absence of whale-watching boats. Effects on activity states were inferred from changes in movement metric data as well as the occurrence of surface feeding events. Linear mixed effects models and generalised estimation equations were used to investigate the effect of whale-watching boat interactions. Measurement errors were quantified, and their effects on model parameter estimates were investigated using resampling methods. Minke whales responded to whale-watching boats by performing shorter dives and increased sinuous movement. A reduction in the probability of observing longer inter-breath intervals during sinuous movement showed that whale-watching boat interactions reduced foraging activity. Further, the probability of observing surface feeding events also decreased during interactions with whale-watching boats. This indicates that whalewatching boats disrupted the feeding activities of minke whales. Since minke whales are capital breeders, a decrease in feeding success on the feeding grounds due to whale-watching boats could lead to a decrease in energy available for foetus development and nursing on the breeding grounds. Such impact could therefore alter the calving success of this species.

Nest visit synchrony is high and correlates with reproductive success in the wild Zebra finch Taeniopygia guttata

Nest visit synchrony, whereby adults coordinate their visits to the nest, has been documented in several species of cooperative breeders. Visit synchrony may reduce nest predation rate or sibling competition, or instead follow from synchronisation of other behaviours, such as foraging. However, nest visit synchrony has rarely been considered in species with bi-parental care, even though it could conceivably bring similar fitness benefits to that seen in cooperative breeders. In addition, in species with bi-parental care, we might expect nest visit synchrony to reflect the quality of the pair or the overall coordination of breeding activity between partners. Here, we tested whether nest visit synchrony occurs in a classic avian model for the study of bi-parental care, the zebra finch Taeniopygia guttata. We found that in the wild, both zebra finch parents visited the nest very infrequently during nestling provisioning, with only one visit per hour, and that nest visits were highly synchronised with parents visiting the nest together on 78% of the visits. In addition, we found that nest visit synchrony was correlated with hatching rate, brood size at hatching and the number of offspring in the nest a few days prior to fledging. Our results suggest that, while more work is required to understand the benefits of nest visit synchrony in this species, considering behavioural synchrony and cooperation between mated partners may offer new insight into the study of parental investment, including in species with bi-parental care.

Seasonal- and sex-specific correlations between dispersal and exploratory behaviour in the great tit

While the importance of personality in explaining individual dispersal strategies is increasingly recognized, limited information is still available on how patterns of personality-dependent dispersal may develop, hampering our understanding of the ecological significance of behavioural dispersal syndromes. Here, we examine the relative importance of personality at different stages of dispersal in the great tit, by analysing the sex-specific relationship between exploratory behaviour (EB; quantified in a novel environment) and dispersal distances in different seasons over the course of the first year of life (summer, autumn, winter, and until the first breeding attempt). In females, we found that EB was an important predictor of dispersal distances in summer and autumn, but only a weak to moderate correlation remained for females captured in winter or for natal dispersal distances based on first breeding records. We obtained a contrasting pattern at the population level, whereby male (but not female) immigrants captured in summer and autumn had higher EB scores than locally born birds, while this was not the case in birds captured in winter and those recruited as breeders into the population. In addition to providing further evidence for the existence of a behaviour dispersal syndrome in birds, our results show that correlations between EB and dispersal appear strongest at the early stages of the dispersal process, rather than being developed gradually. These findings show the importance of analysing the effect of phenotypic attributes on dispersal across different stages of the dispersal phenomenon and in each sex separately.