Roger Jovani

Resource unpredictability promotes species diversity and coexistence in an avian scavenger guild: a field experiment

Chance per se plays a key role in ecology and evolution, e.g., genetic mutation, resource spatiotemporal unpredictability. In community ecology, chance is recognized as a key factor in community assemblage, but less is known about its role in intraguild processes leading to species coexistence. Here we study the relevance of resource unpredictability per se as a promoter of intraguild positive interspecific interactions and as a biodiversity enhancer in an Old World avian scavenger guild, which has evolved to feed upon spatially and temporally unpredictable resources, i.e., carcasses. We performed a large-scale field experiment in which 58 carcasses were disposed of and observed until complete consumption, either in continuously active supplementary feeding stations (predictable carcasses) or disposed of at random in the field (unpredictable carcasses). Richness of scavenger species was similar at unpredictable and predictable carcasses, but their relative abundances were highly uneven at predictable carcasses leading to higher scavenger diversity (Shannon index) at unpredictable carcasses. Facilitatory interspecific processes only occurred at unpredictable resources but were disrupted in predictable conditions because the dominant specialist species (in our case, the Griffon Vulture Gyps fulvus) arrived earlier and in larger numbers, monopolizing the resource. Small, endangered scavengers congregated at supplementary feeding stations but profited less compared to unpredictable carcasses, suggesting that they could constitute an ecological trap. Our findings offer new insights into the relevance of unpredictability of trophic resources in promoting both positive facilitatory interspecific interactions and species diversity and thus maintaining the function of guilds. Finally, the preservation of randomness in resource availability and the processes associated with its exploitation should be a major goal of conservation strategies aimed to preserve scavenger guilds evolved under naturally unpredictable trophic resources.

A heuristic model on the role of plasticity in adaptive evolution: Plasticity increases adaptation, population viability and genetic variation

An ongoing new synthesis in evolutionary theory is expanding our view of the sources of heritable variation beyond point mutations of fixed phenotypic effects to include environmentally sensitive changes in gene regulation. This expansion of the paradigm is necessary given ample evidence for a heritable ability to alter gene expression in response to environmental cues. In consequence, single genotypes are often capable of adaptively expressing different phenotypes in different environments, i.e. are adaptively plastic. We present an individual-based heuristic model to compare the adaptive dynamics of populations composed of plastic or non-plastic genotypes under a wide range of scenarios where we modify environmental variation, mutation rate and costs of plasticity. The model shows that adaptive plasticity contributes to the maintenance of genetic variation within populations, reduces bottlenecks when facing rapid environmental changes and confers an overall faster rate of adaptation. In fluctuating environments, plasticity is favoured by selection and maintained in the population. However, if the environment stabilizes and costs of plasticity are high, plasticity is reduced by selection, leading to genetic assimilation, which could result in species diversification. More broadly, our model shows that adaptive plasticity is a common consequence of selection under environmental heterogeneity, and hence a potentially common phenomenon in nature. Thus, taking adaptive plasticity into account substantially extends our view of adaptive evolution.

Adaptive allocation of stress-induced deformities on bird feathers

Physiological stress during ontogeny is known to cause abnormalities in keratin structures of vertebrates, but little is known about if and how organisms have evolved mechanisms to reduce the negative effects of these abnormalities. Stress experienced during avian feather growth is known to lead to the formation of fault bars, and thereby to the weakening of feathers because of shortage and slimming of barbules. Here we propose and test a new hypothesis (the ‘fault bar allocation hypothesis’) according to which birds should have evolved adaptive strategies to counteract this evolutionary pressure. In particular, we predicted and tested the idea that in flying birds, natural selection should have selected for mechanisms to reduce fault bar load on feathers with high strength requirements during flight. Data on the growth of feathers of nestling white storks (Ciconia ciconia) revealed a consistent allocation of more, and more intense, fault bars in innermost than in outermost wing feathers as predicted by our hypothesis. Moreover, the same pattern emerged from feathers of adult storks. We discuss the generality of our results, and suggest avenues for further investigations in this area.

EVALUATION OF GENOTOXIC EFFECTS OF HEAVY METALS AND ARSENIC IN WILD NESTLING WHITE STORKS (CICONIA CICONIA) AND BLACK KITES (MILVUS MIGRANS) FROM SOUTHWESTERN SPAIN AFTER A MINING ACCIDENT

Studies of birds from Doñana (southwestern Spain) after the Aznalcóllar mining accident (April 1998) have reported high levels of genetic damage when compared to conspecifics from reference areas. However, potential relationships between DNA damage and metal pollution have not yet been reported. The aim of the present study was to investigate the current levels of Zn, Pb, As, Cu, and Cd and to determine if they were associated with the genetic damage observed in free-living, nestling white storks (Ciconia ciconia) and black kites (Milvus migrans) born in the Doñana area after the mining spill. Blood concentrations of heavy metals and of As were quantified and DNA damage (comet assay) was determined in 258 storks and 132 kites monitored during a four-year period (1999-2002). Correlations between these elements and genetic damage varied between species and throughout years within species. Some elements did not show any relationship with DNA damage (e.g., Pb), whereas others had a significant correlation (e.g., As in storks, and Cu and Cd in kites) or only marginal statistical effects (e.g., Zn and Cd in storks, and As in kites) in some years but not in others. These results suggest that nestling white storks and black kites were affected, in part, by the elements studied, but they alone do not satisfactorily explain the observed DNA damage. Moreover, our results show that species-specific differences should be carefully considered when planning schemes for pollution monitoring, and highlight the need for including the temporal scale into the study of the pollutants effects in the wild.

Fractal bird nest distribution produces scale-free colony sizes

The spatial distribution of organisms often differs across scales. For instance, colonial bird populations could be described, from large to small scale, as scattered clumps of otherwise regularly distributed breeding pairs. We analysed the distribution of nests of a large colonial population of white storks (Ciconia ciconia) and found a fractal pattern in each of the 4 study years. Moreover, we found that the often-observed, long-tailed frequency distribution of colony sizes was well described by a power law, regardless of the cut-off used to define colonies (from 16 to 1024 m). Thus, although storks were locally highly clumped even with tens of nests in a single tree, the population was not structured in colonies (a simple clustered distribution) as previously thought. Rather, they were distributed in a continuous hierarchical set of clusters within clusters across scales, clusters lacking the commonly assumed characteristic mean size. These quantitative solutions to previously perceived scaling problems will potentially improve our understanding of the ecology and evolution of bird coloniality and animal spacing patterns and group living in general.

Breeding synchrony in colonial birds: from local stress to global harmony

Why and how birds in colonies often breed in striking synchrony is an unsolved question. In colonies, conspecific birds often destroy eggs and kill chicks, either intentionally or not. We propose that social tranquillity at the time of laying can be achieved if a birds stress level is partly determined by the agitation of its neighbours. Moreover, we propose that this local process, together with environmental cues, can synchronize breeding between neighbours and through a whole colony. We tested our hypotheses using a generic individual-based model where the breeding predisposition of females was updated daily depending on an increase in the photoperiod (positively) and the stress level of neighbours: negatively if they were agitated, and positively otherwise. A female laid her eggs when her stress level fell to a critical value. Even giving only a low relevance to the neighbours stress level was enough to synchronize the laying date of neighbours and also of a huge colony. Moreover, females bred in a safer environment, which is known from field studies to increase fitness. Our study highlights the power of local adaptive (individual) behaviour to create global (colony) patterns. We argue that collective patterns such as breeding synchrony in colonial birds could have simple adaptive individual-level explanations.

Apparent Absence of Blood Parasites in the Patagonian Seabird Community: Is it Related to the Marine Environment?

-The geographic and specific distribution of avian Haematozoa is poorly understood and much basic information is still needed. Studies of blood parasites in avian communities are scarce despite of their potential for disentangling the relative importance of host-specific and ecological factors shaping blood parasite distributions. Here we present the first study of blood parasites in a breeding community of seabirds by scanning blood smears obtained from 560 birds belonging to 13 species breeding in Patagonia, Argentina. Seven of these species have not been sampled previously for blood parasites. No blood parasites were detected. The scarcity of vectors due to the marine environment and the dry conditions around the colonies is the most plausible hypothesis for explaining the apparent absence of blood parasites in the Patagonian seabird community, although other hypotheses should be examined. Received 12 December 2000, accepted 8 May 2001.

Feather mites (Astigmata) avoid moulting wing feathers of passerine birds

The moult of birds could greatly perturb the life cycle of feather mites (Astigmata). These mites live on the surface of feathers and are unlikely to survive if they are on one when it is moulted. Therefore, we hypothesized that they would escape to other feathers to avoid this risk of mortality. We analysed the distribution of feather mites on the wings of 63 moulting individuals of 13 species of passerine birds. We used generalized linear mixed models to test whether the distribution of feather mites depends on the stage of moult of feathers. Feather mites were significantly less abundant on the next feather to fall out than on the other feathers. This ability to escape seems to be higher on primaries and secondaries than on tertials. At least on primaries, feather mites avoided not only the next feather to be moulted but also the second one. For primaries and secondaries the new full-grown feathers harboured more feather mites than those still growing. This study shows that feather mites have an escape behaviour from moulting wing feathers in passerine birds that is consistent with a model of adaptive behaviour.

Lognormal distribution of individual lifetime fecundity: insights from a 23-year study

Individual variance in lifetime fecundity within populations is a life-history parameter of crucial evolutionary and ecological significance. However, knowledge of its magnitude and underlying mechanisms in natural populations is biased toward short-lived taxa. This paper summarizes results of a 23-year study on a population of the Mediterranean shrub Lavandula latifolia. We document the within-population pattern of individual variation in instantaneous and lifetime fecundity (as estimated by inflorescence production) and explore the mechanisms producing the lognormal distribution of individual fecundities by means of an individual-based simulation model. Throughout the study period, a few individuals consistently produced most inflorescences while the majority of plants exhibited moderate-to-low fecundities. The shape of yearly distributions of annual fecundities varied little across years, and most annual fecundity distributions did not depart significantly from a lognormal. The distribution of individual lifetime fecundity did not depart from lognormality. Despite the simplicity of the premises of our simulation model, it was remarkably successful at predicting the shapes of fecundity distributions and the early establishment of a persistent fecundity hierarchy. The agreement between model results and empirical data supports the view that multiplicative interactions of randomly varying environmental effects can play a central role in determining individual variation in lifetime fecundity in L. latifolia, and suggests that environmental stochasticity can be decisive in the genesis of strong fecundity hierarchies in long-lived plants.

Multiple mating opportunities boost protandry in a pied flycatcher population

Protandry, the earlier arrival of males than females to breeding areas, is widespread in birds, but its underlying mechanisms are far from well understood. The two, not mutually exclusive most highly supported hypotheses to explain avian protandry postulate that it has evolved from intrasexual male competition to acquire the best territories (“rank advantage” hypothesis) and/or to maximize the number of mates (“mate opportunity” hypothesis). We studied for two consecutive years the relative importance of both hypotheses in a population of pied flycatchers (Ficedula hypoleuca), a territorial songbird with a mixed mating strategy. We measured territory quality using a long-term dataset on nest occupation and breeding output, and we used molecular techniques to assess male fitness across the range of social and genetic mating options. Territory quality was unrelated to breeding date and had no influence on extra-pair paternity or social polygynous events. However, males breeding early increased their chances of becoming socially polygynous and/or of attaining extra-pair paternity and, as a consequence, increased their total reproductive success. These results support the “mate opportunity” hypothesis, suggesting that sexual selection is the main mechanism driving protandry in this population.