Robert J. Thomas

DIFFERENCES BETWEEN POPULATIONS OF REED WARBLERS IN DEFENCES AGAINST BROOD PARASITISM

Summary Two potential defences against brood parasitism by the cuckoo Cuculus canorus were compared experimentally between British populations of reed warblers Acrocephalus scirpaceus that are parasitised at different rates. (1) Rates of rejection of model cuckoo eggs were lower at two unparasitised populations which did not have resident cuckoos, than at a rarely parasitised population which had cuckoos nearby, and at a regularly parasitised population. (2) Reed warblers from an unparasitised population showed a slightly weaker response to taxidermic mounts of cuckoos and, unlike a parasitised population, did not differentiate between mounts of a cuckoo, sparrowhawk Accipiter nisus and jay Garrulus glandarius . Differences in exposure to real predators may explain the differences in responses to mounted predators between populations, as specie c aggressive responses to predators are likely to have been learned. Although evidence from dispersal and population turnover data suggests that there is likely to be gene e ow between reed warbler populations in Britain, the hypothesis that the population differences ree ect genotypic differences could not be ruled out. An alternative explanation of phenotypic plasticity in defences could also explain the population differences. Phenotypic plasticity in defences would be favoured in environments where the risk of parasitism e uctuates, if those defences are costly to unparasitised reed warblers.

Eye size in birds and the timing of song at dawn

Why do different species of birds start their dawn choruses at different times? We test the hypothesis that the times at which different species start singing at dawn are related to their visual capability at low light intensities. Birds with large eyes can achieve greater pupil diameters and hence, all other things being equal, greater visual sensitivity and resolution than birds with small eyes. We estimated the maximum pupil diameter of passerine birds by measuring the diameter of the exposed eye surface, and measured the times of the first songs at dawn of songbirds present in different bird communities, and the light intensities at these times. Using phylogenetic comparative analyses, we found that songbirds with large eyes started to sing at lower light intensities (and therefore earlier) than species with smaller eyes. These relationships were stronger when differences in body size were controlled for statistically, and were consistent between two phylogenies and when species were treated as independent data points. Our results therefore provide robust support for the hypothesis that visual capability at low light levels influences the times at which birds start to sing at dawn.

The costs of singing in nightingales

The costs of singing in birds are poorly understood. One potential type of cost is a metabolic cost of singing. Previous studies have measured short-term changes in oxygen consumption associated with bouts of vocalizations, with equivocal results. In this study, I used an alternative approach to measuring the metabolic cost of singing, by measuring overnight loss of body mass, in male common nightingales, Luscinia megarhynchos, singing at night at different rates. Nightingales were shown not to forage at night. They reached a higher mass at dusk prior to singing more at night, and lost more mass overnight when dusk mass and overnight song rate were high. These results show that singing at night is associated with increased overnight consumption of body reserves, which represents a significant metabolic cost of singing at night. However, the correlation between dusk mass and overnight song rate makes it impossible to determine whether these costs arise from the energetic costs of the singing itself, or from the metabolic costs of the additional body reserves laid down at dusk on nights when song rate was high. There are also likely to be costs associated with accumulating and carrying these extra body reserves during daylight, as well as other potential costs of singing such as an increased risk of predation. These results are consistent with those models of signalling in biology that predict or assume that honest signals are costly.

A biosynthetic classification of fungal and streptomycete fused-ring aromatic polyketides.

Polyketides are one of the largest and most structurally diverse classes of naturally occurring compounds, ranging from simple aromatic metabolites to complex macrocyclic lactones. Fungi and filamentous bacteria, particularly the actinomycetes, are major sources of polycyclic aromatic structures, which include many clinically important antibiotics and other useful metabolites. These fused‐ring polyketides are formed by the action of polyketide synthases (PKSs), which catalyse the assembly, folding and cross‐linking of poly‐β‐ketoacyl intermediates. In view of the taxonomic gulf between the eukaryotic fungi and prokaryotic bacteria, it is not surprising that they are rarely found to produce structurally identical fused‐ring metabolites. A review of [13C2]acetate incorporation data has revealed consistent differences in the reported cyclisation patterns, which require regiospecifically distinct cross‐linking of otherwise identical linear polyketide precursors. This observation provides the basis for a structural and biosynthetic classification of microbial fused‐ring polyketides, which has a number of useful ramifications.

PERSPECTIVE: THE EVOLUTION OF WARNING COLORATION IS NOT PARADOXICAL

Abstract Animals that are brightly colored have intrigued scientists since the time of Darwin, because it seems surprising that prey should have evolved to be clearly visible to predators. Often this self‐advertisement is explained by the prey being unprofitable in some way, with the conspicuous warning coloration helping to protect the prey because it signals to potential predators that the prey is unprofitable. However, such signals only work in this way once predators have learned to associate the conspicuous color with the unprofitability of the prey. The evolution of warning coloration is still widely considered to be a paradox, because it has traditionally been assumed that the very first brightly colored individuals would be at an immediate selective disadvantage because of their greater conspicuousness to predators that are naive to the meaning of the signal. As a result, it has been difficult to understand how a novel conspicuous color morph could ever avoid extinction for long enough for predators to become educated about the signal. Thus, the traditional view that the evolution of warning coloration is difficult to explain rests entirely on assumptions about the foraging behavior of predators. However, we review recent evidence from a range of studies of predator foraging decisions, which refute these established assumptions. These studies show that: (1) Many predators are so conservative in their food preferences that even very conspicuous novel prey morphs are not necessarily at a selective disadvantage. (2) The survival and spread of novel color morphs can be simulated in field and aviary experiments using real predators (birds) foraging on successive generations of artificial prey populations. This work demostrates that the foraging preferences of predators can regularly (though not always) result in the increase to fixation of a novel morph appearing in a population of familiar‐colored prey. Such fixation events occur even if both novel and familiar prey are fully palatable and despite the novel food being much more conspicuous than the familiar prey. These studies therefore provide strong empirical evidence that conspicuous coloration can evolve readily, and repeatedly, as a result of the conservative foraging decisions of predators.

Two tests of a stochastic dynamic programming model of daily singing routines in birds.

Many hypotheses have been put forward to account for the dawn chorus in birds. Few of these, however, are able to account for variation in song output over the whole day, or for differences in daily singing routines between species, individuals, seasons and environmental conditions. One hypothesis that does offer a more general explanation is based on a stochastic dynamic programming (SDP) model of daily singing routines. This model relates the relative costs and benefits of feeding and singing at different times of day to the size of a birds fat reserves and calculates the optimal daily routines of singing and foraging that will maximize the amount that the bird can sing while avoiding starvation. The use of SDP models in behavioural ecology has become well established, but they remain largely untested empirically. I tested two predictions of the SDP model of daily routines of singing, using free-living European robins Erithacus rubecula. The results supported both predictions: (1) food supplementation causing unpredictable short-term increases in foraging success increased subsequent song output; and (2) changes in ambient temperature were positively associated with changes in subsequent song output. Copyright 1999 The Association for the Study of Animal Behaviour.

Body mass regulation and the daily singing routines of European robins

Stochastic dynamic programming (SDP) models of daily singing and foraging routines in birds relate an individuals fat reserves to the relative costs and benefits of singing and foraging at different times of day. Two central predictions of such models are that: (1) overnight loss of fat reserves is higher on colder nights, and (2) birds sing more at dawn when their fat reserves are high. We tested these predictions in free-living European robins, Erithacus rubecula, by examining the relationships between ambient temperature, body mass change (an index of changes in fat reserves) and song rate. In support of prediction (1), overnight mass loss was positively associated with overnight temperature. However, robins also put on more mass at dusk if the night ahead was going to be cold, which tended to buffer the effects of overnight temperature on dawn body mass. In support of prediction (2), robins sang more at dawn on days when their dawn body mass was high, although this association was detectable only after controlling for foraging intensity during the dawn chorus. We analyse differences between the results of this and related studies and discuss the implications of these results for existing SDP models of daily singing routines.

Strategic diel regulation of body mass in European robins

Stochastic dynamic programming (SDP) is a computational technique that has been used to model daily routines of foraging in small birds. A diurnal bird must build up its fat reserves towards dusk in order to avoid starvation during the night, when it cannot feed. However, as well as the benefits of avoiding starvation, storing fat imposes costs such as an increased predation risk and higher flight and metabolic costs. There is therefore an optimal level of fat reserves for a bird to reach at dusk in order to survive overnight without being left with excessive fat reserves at dawn. I tested a prediction common to all SDP models of daily foraging routines, that a bird will attempt to reach this level at dusk, regardless of its fat reserves the previous dawn. I provided supplementary food to manipulate the fat reserves at dawn of free-living European robins, Erithacus rubecula. Diurnal changes in body mass (a reliable estimate of fat reserves) were then monitored remotely. Robins provided with an ad libitum food supply reached almost exactly the same body mass at dusk, regardless of their body mass at dawn, supporting the prediction that birds attempt to reach a target level of reserves at dusk. Copyright 2000 The Association for the Study of Animal Behaviour.

The trade-off between singing and mass gain in a daytime-singing bird, the European robin

The functions of bird song are well described, but empirical studies examining the costs of singing are scarce. Two potential costs are a metabolic cost of singing, and lost feeding opportunities, but such energetic costs will only be biologically important if they have a significant effect on the birds body reserves. Overnight loss of reserves has previously been found to increase with increasing song rates in nocturnally singing common nightingales Luscinia megarhynchos. However, it is not clear how such costs compare with those incurred by daytime-singing birds, which may forfeit foraging opportunities when they sing. In this paper we investigated the effect of variation in song rate on the body reserves of a typical daytimesinging bird, the European robin Erithacus rubecula, singing at different rates in three different circumstances: (i) Natural variation in song rate of free-living robins. (ii) Manipulations of the song rate of free-living robins using playbacks of conspecific song. (iii) Manipulations of the song rate of aviary-housed robins using playbacks of conspecific song. In all three parts of our study, birds gained less mass when they sang more. Our analyses suggested that this might have been due primarily to a reduction in food intake rate, rather than to the metabolic cost of the singing itself or a concurrent increase in locomotor costs. These results together demonstrate that the costs of singing, as measured by their overall net effects on body reserves, can have a significant impact on the energetic state of daytime-singing birds.

The effect of variability in the food supply on the daily singing routines of European robins: a test of a stochastic dynamic programming model

A stochastic dynamic programming (SDP) model offers a general explanation of daily singing routines in birds, but remains almost untested empirically. I examined a central prediction of the SDP model, that a more variable food supply decreases the birds song output at dawn, relative to its song output at dusk. I provided supplementary food to make the food supply more or less variable over 2-week periods in the territories of free-living European robins Erithacus rubecula. Robins sang relatively less at dawn than at dusk after weeks in which their supplementary food supply was variable, and more at dawn than at dusk after weeks in which their food supplementation was constant. These results provide strong support for the prediction of the SDP model. Copyright 1999 The Association for the Study of Animal Behaviour.