Individual preferences for sound tool design in a parrot.

The rarity of tool manufacture in wild parrots is surprising because they share key life-history traits with advanced tool-using species, including large brains, complex sociality and prolonged parental care. When it does occur, tool manufacture in parrots tends to be innovative, spontaneous and individually variable, but most cases have been in captivity. In the wild, only palm cockatoos (Probosciger aterrimus) have been observed using tools regularly. However, they are unusual because they use tools to enhance their displays rather than for foraging or self-maintenance. Males in northern Australia make two types of tool from sticks and seed pods, which they tap rhythmically against a tree during display. We analysed 256 sound tools retrieved from 70 display trees. Drumsticks (89% of tools) were used more often than seed pod tools; most males manufactured only drumsticks, but some made both types. Individual males differed significantly in the design of their drumsticks including the length, width and mass but we found no evidence that neighbours copied each other. We discuss the highly individualized preferences for sound tool design in context of the behavioural predispositions behind the rarity of tool manufacture in wild parrots.

Can an introduced predator select for adaptive sex allocation?

Most species produce equal numbers of sons and daughters, and sex differences in survival after parental care do not usually affect this pattern. Temporary overproduction of the scarcer sex can be adaptive when generations overlap, the sexes differ in life-history expectations, and parents can anticipate future mating opportunities. However, an alternative strategy of maximizing the competitiveness of the more abundant sex in these circumstances remains unexplored. We develop theory showing how mothers can maximize reproductive value when future mate competition will be high by producing more sons in the advantageous early hatching positions within their broods. Our model for optimal birth order was supported by long-term data of offspring sex in a parrot facing catastrophic female mortality caused by introduced predators. Swift parrots (Lathamus discolor) suffer high female mortality due to introduced sugar gliders (Petaurus breviceps) creating fluctuating male-biased adult sex ratios. Offspring hatched early within broods fledged in better condition, and in support of our model were more likely to be male in years with higher adult female mortality. We found a highly significant rank-order correlation between observed and predicted birth sex ratios. Our study shows the potential for mothers to maximize reproductive value via strategic biases in offspring sex depending on the advantages conferred by birth order and the predictability of future mate competition. Our long-term data support the predictions and appear to suggest that sex allocation strategies may evolve surprisingly quickly when anthropogenic pressures on populations are severe.

Egg size investment in superb fairy-wrens: helper effects are modulated by climate.

Natural populations might exhibit resilience to changing climatic conditions if they already show adaptive flexibility in their reproductive strategies. In cooperative breeders, theory predicts that mothers with helpers should provide less care when environmental conditions are favourable, but maintain high investment when conditions are challenging. Here, we test for evidence of climate-mediated flexibility in maternal investment in the cooperatively breeding superb fairy-wren Malurus cyaneus We focus on egg size because in this species egg size influences offspring size, and females reduce egg investment when there are helpers at the nest. We report that females lay larger eggs during dry, hot conditions. However, the effect of temperature is modulated by the presence of helpers: the average egg size of females with helpers is reduced during cooler conditions but increased during hot conditions relative to females without helpers. This appears to reflect plasticity in egg investment rather than among female differences. Analysis of maternal survival suggests that helped females are better able to withstand the costs of breeding in hot conditions than females without helpers. Our study suggests that females can use multiple, independent cues to modulate egg investment flexibly in a variable environment.

Brood parasitism and the evolution of cooperative breeding in birds.

The global distribution of cooperatively breeding birds is highly uneven, with hotspots in Australasia and sub-Saharan Africa. The ecological drivers of this distribution remain enigmatic yet could yield insights into the evolution and persistence of cooperative breeding. We report that the global distributions of avian obligate brood parasites and cooperatively breeding passerines are tightly correlated and that the uneven phylogenetic distribution of cooperative breeding is associated with the uneven targeting of hosts by brood parasites. With a long-term field study, we show that brood parasites can acquire superior care for their young by targeting cooperative breeders. Conversely, host defenses against brood parasites are strengthened by helpers at the nest. Reciprocally selected interactions between brood parasites and cooperative breeders may therefore explain the close association between these two breeding systems.

Social constraint and an absence of sex-biased dispersal drive fine-scale genetic structure in white-winged choughs.

This study used eight polymorphic microsatellite loci to examine the relative effects of social organization and dispersal on fine-scale genetic structure in an obligately cooperative breeding bird, the white-winged chough (Corcorax melanorhamphos). Using both individual-level and population-level analyses, it was found that the majority of chough groups consisted of close relatives and there was significant differentiation among groups (F(ST) = 0.124). However, spatial autocorrelation analysis revealed strong spatial genetic structure among groups up to 2 km apart, indicating above average relatedness among neighbours. Multiple analyses showed a unique lack of sex-biased dispersal. As such, choughs may offer a model species for the study of the evolution of sex-biased dispersal in cooperatively breeding birds. These findings suggest that genetic structure in white-winged choughs reflects the interplay between social barriers to dispersal resulting in large family groups that can remain stable over long periods of times, and short dispersal distances which lead to above average relatedness among neighbouring groups.

Cryptic gentes revealed in pallid cuckoos Cuculus pallidus using reflectance spectrophotometry.

Many cuckoo species lay eggs that match those of their hosts, which can significantly reduce rejection of their eggs by the host species. However, egg mimicry is problematic for generalist cuckoos that parasitize several host species with different egg types. Some generalist cuckoos have overcome this problem by evolving several host-specific races (gentes), each with its own, host-specific egg type. It is unknown how generalist cuckoos lacking gentes are able to avoid egg rejection by hosts. Here we use reflectance spectrophotometry (300-700 nm) on museum egg collections to test for host-specific egg types in an Australian generalist cuckoo reported to have a single egg type. We show that the colour of pallid cuckoo (Cuculus pallidus) eggs differed between four host species, and that their eggs closely mimicked the eggs of the host they parasitized. These results reveal that pallid cuckoos have host-specific egg types that have not been detected by human observation, and indicate that gentes could be more common than previously realized.

Coalitions of relatives and reproductive skew in cooperatively breeding white-winged choughs.

We used DNA fingerprinting to examine reproductive skew in cooperatively breeding white-winged choughs, Corcorax melanorhamphos, which live in groups of up to 20 individuals. Before a severe drought, groups that had been stable for multiple years were characterized by long-term monogamy involving a single breeding pair (high skew). After the drought, new groups formed from the amalgamation of multiple individuals and coalitions of relatives. At most one member of each faction succeeded in breeding, such that skew was dependent on the number of unrelated factions, and not group size. In the new groups, dominant males and females with supporting relatives were always successful. Whereas most females without support also gained breeding positions, many males without family support failed to breed. Thus subordinates gain indirect fitness by first helping related males to secure a breeding position, and then helping to raise their young. Our study demonstrates the advantage of operating in coalitions, and suggests that the acquisition of future allies may be a major benefit of helping behaviour in this species.

The cost of helping.

Cooperative breeding in mammals, birds and fish has provided evolutionary biologists with a rich framework for studying the causes and consequences of group-based reproduction. Helping behaviour is especially enigmatic because it often entails an individual sacrificing personal reproduction while assisting others in their breeding attempts. The decision to help others to reproduce is affected by immediate and future costs analogous to those of direct reproduction, but these components of the equation have usually been neglected. Recent research suggests that the type of benefit sought could determine the extent of help given.

Group territoriality in two populations of African lions

Lionesses, Panthera leoin the Serengeti ecosystem, Tanzania, assess the odds of winning group-territorial contests by counting the number of opponents they hear roaring. They will approach intruders aggressively only if they outnumber them. Here the lionesses in the Serengeti are compared with another population living nearby but in strikingly different ecological circumstances. The lions of Ngorongoro Crater live at much higher densities owing to year-round availability of non-migratory prey species, but also suffer higher mortality from fighting. Playback experiments showed that lionesses in the crater differ from those in the Serengeti by approaching 'intruders' more quickly when the odds of winning are low. This increased aggression is interpreted in terms of the greater difficulty of holding a territory at high population density.

Development of cooperative territoriality in juvenile lions.

African lions, Panthera leo, engage in many cooperative activities including hunting, care of young, and group territoriality, but the contribution of juvenile lions to these activities has never been documented. Here we present experimental evidence that juvenile lionesses make a gradual transition to group-territorial defence between weaning (8 months) and sexual maturity (42 months). When challenged by simulated intruders played from a loud-speaker, juvenile females (but not males) become progressively more likely to join the adult females in territorial defence with age, and their behaviour is affected by both the number of defending adults and the number of intruders. We interpret the ability of juveniles to assess relative numbers as an adaptation for assessing the risk of territorial conflict according to their own fighting ability, and the ability of their pride of successfully defend the territory. The difference between the sexes reflects the greater value of the natal territory to philopatric females. Adult females display a variety of strategies when defending the territory, including unconditional and conditional forms of cooperation. We show here that individuals display the rudiments of these strategies as juveniles.

Complex cooperative strategies in group-territorial African lions.

Female lions (Panthera leo) showed persistent individual differences in the extent to which they participated in group-territorial conflict. When intergroup encounters were simulated by playback of aggressive vocalizations, some individuals consistently led the approach to the recorded intruder, whereas others lagged behind and avoided the risks of fighting. The lead females recognized that certain companions were laggards but failed to punish them, which suggests that cooperation is not maintained by reciprocity. Modification of the "odds" in these encounters revealed that some females joined the group response when they were most needed, whereas other lagged even farther behind. The complexity of these responses emphasizes the great diversity of individual behavior in this species and the inadequacy of current theory to explain cooperation in large groups.

Avian cooperative breeding: Old hypotheses and new directions.

In cooperatively breeding birds, individuals that appear capable of reproducing on their own may instead assist others with their breeding efforts. Research into avian cooperative breeding has attempted to reconcile the apparent altruism of this behaviour with maximization of inclusive fitness. Most explanations of cooperative breeding have suggested that philopatry is enforced by ecological constraints, such as a shortage of resources critical to breeding. Non-dispersers may then benefit both directly and indirectly from contributing at the nest. Recent research has shown that such benefits may be sufficient to promote philopatry, without the need for ecological constraints, and emphasizes that consideration of both costs and benefits of philopatry is essential for a comprehensive approach to the problem. The growing body of data from long-term studies of different species should combine with an improved phylogenetic perspective on cooperative breeding, to provide a useful base for future comparative analyses and experimentation.

Computer-aided hybridoma screening. Production of monoclonal antibodies against membrane fractions from mammary tumour tissue.

Monoclonal antibodies were raised against membrane fractions from mammary tumour tissue. 746 hybridomas were obtained against the so-called membrane fraction '2a' and 1393 hybridomas against the so-called 'top-region'. These hybridomas were tested with an immunodot assay, cell-binding radioimmunoassay and an immunocytological screening assay using several antigens. In the evaluation of these data we used the common database software (dBASE II) on a personal computer. To facilitate computer handling, even for the non-specialist, we programmed our individual menu-driven software for computer-aided hybridoma screening.

Screening of hybridoma supernatants raised against membrane fractions from breast cancer using an immunodot assay.

Membranes were prepared from mammary tumors and fractionized using gel filtration and gradient polyacrylamide gel electrophoresis. To obtain monoclonal antibodies against tumor-associated antigens, mice were immunized against membrane fraction 2a. After hybridization, we obtained 746 hybridoma cell lines. We performed the immunodot assay for screening. Every supernatant was tested against 15 antigens. Eight antigens were purified fractions from tumor membrane preparation, and five were crude membrane preparations from benign and malignant breast cell lines. Additionally, the Thomsen-Friedenreich antigen of erythrocytes was tested. We selected 83 hybridomas for further characterization.