Paolo Galeotti

Effects of elevated egg corticosterone levels on behavior, growth, and immunity of yellow-legged gull (Larus michahellis) chicks

Eggs of vertebrates contain steroid hormones of maternal origin that may influence offspring performance. Recently, it has been shown that glucocorticoids, which are the main hormones mediating the stress response in vertebrates, are transmitted from the mother to the egg in birds. In addition, mothers with experimentally elevated corticosterone levels lay eggs with larger concentrations of the hormone, which produce slow growing offspring with high activity of the hypothalamo-adrenal axis under acute stress. However, the effects and function of transfer of maternal corticosterone to the eggs are largely unknown. In the present study, we injected corticosterone in freshly laid eggs of yellow-legged gulls (Larus michahellis), thus increasing the concentration of the hormone within its natural range of variation, and analyzed the effect of manipulation on behavioral, morphological, and immune traits of the offspring in the wild. Eggs injected with corticosterone had similar hatching success to controls, but hatched later. Mass loss during incubation was greater for corticosterone-treated eggs, except for the last laid ones. Corticosterone injection reduced rate and loudness of late embryonic vocalizations and the intensity of chick begging display. Tonic immobility response, reflecting innate fearfulness, was unaffected by hormone treatment. Elevated egg corticosterone concentrations depressed T-cell-mediated immunity but had no detectable effects on humoral immune response to a novel antigen, viability at day 10, or growth. Present results suggest that egg corticosterone can affect the behavior and immunity of offspring in birds and disclose a mechanism mediating early maternal effects whereby stress experienced by females may negatively translate to offspring phenotypic quality.

Colour polymorphism in birds: causes and functions.

We studied polymorphism in all species of birds that are presently known to show intraspecific variation in plumage colour. At least three main mechanisms have been put forward to explain the maintenance of polymorphism: apostatic, disruptive and sexual selection. All of them make partly different predictions. Our aims were to investigate evolutionary causes and adaptive functions of colour polymorphism by taking into account a number of ecological and morphological features of polymorphic species. Overall, we found 334 species showing colour polymorphism, which is 3.5% of all bird species. The occurrence of colour polymorphism was very high in Strigiformes, Ciconiiformes, Cuculiformes and Galliformes. Phylogenetically corrected analysis using independent contrasts revealed that colour polymorphism was maximally expressed in species showing a daily activity rhythm extended to day/night, living in both open and closed habitats. All these findings support the hypothesis that colour polymorphism probably evolved under selective pressures linked to bird detectability as affected by variable light conditions during activity period. Thus, we conclude that selective agents may be prey, predators and competitors, and that colour polymorphism in birds may be maintained by disruptive selection.

Paternity and multiple signaling: effects of a secondary sexual character and song on paternity in the barn swallow.

Multiple signals may evolve because they provide independent information on the condition of a signaler. Females should pay attention to male characters relative to their reliability as signals of male attractiveness or quality. Since behavioral traits are flexible and, therefore, subject to strong environmental influences, females should weigh stable morphological signals higher in their choice of mates for genetic benefits than flexible behavioral traits, for example, by paying particular attention to phenotypically plastic traits when produced in combination with an exaggerated morphological signal. Consistent with this prediction, female barn swallows Hirundo rustica, which are known to prefer males with the longest tail feathers (a secondary sexual character), also preferred males with extreme expressions of a behavioral trait (song rate), as determined from patterns of paternity assessed by microsatellites. However, a statistical interaction between tail length and song rate implied that song rate was relatively unimportant for males with a short tail but more important for longtailed males. Since song rate is a flexible behavioral trait, females appear to have responded to this flexibility by devaluing the importance of song rate in assessment of unattractive sires.

Male-male combats in a polymorphic lizard: residency and size, but not color, affect fighting rules and contest outcome.

Theoretical models predict that the outcome of dyadic agonistic encounters between males is influenced by resource-holding potential, resource value, and intrinsic aggressiveness of contestants. Moreover, in territorial disputes residents enjoy a further obvious competitive advantage from the residency itself, owing to the intimate familiarity with their territory. Costs of physical combats are, however, dramatically high in many instances. Thus, signals reliably reflecting fighting ability of the opponents could easily evolve in order to reduce these costs. For example, variation in color morph in polymorphic species has been associated with dominance in several case studies. In this study, we staged asymmetric resident-intruder encounters in males of the common wall lizard Podarcis muralis, a species showing three discrete morphs (white, yellow, and red) to investigate the effects of asymmetries in color morph, residency, and size between contestants on the outcome of territorial contests. We collected aggression data by presenting each resident male with three intruders of different color morph, in three consecutive tests conducted in different days, and videotaping their interactions. The results showed that simple rules such as residency and body size differences could determine the outcome of agonistic interactions: residents were more aggressive than intruders, and larger males were competitively superior to smaller males. However, we did not find any effect of color on male aggression or fighting success, suggesting that color polymorphism in this species is not a signal of status or fighting ability in intermale conflicts.

Photographic identification in reptiles: a matter of scales

Photographic identification is a promising marking technique alternative to the toe-clipping, since it is completely harmless, cheap, and it allows long time identification of individuals. Its application to ecological studies is mainly limited by the time consuming to compare pictures within large datasets and the huge variation of ornamentation patterns among different species, which prevent the possibility that a single algorithm can effectively work for more than few species. Scales of Reptiles offer an effective alternative to ornamentations for computer aided identification procedures, since both shape and size of scales are unique to each individual, thus acting as a fingerprint like ornamentation patterns do. We used the Interactive Individual Identification System (I 3 S) software to assess whether different individuals of two species of European lizards ( Podarcis muralis and Lacerta bilineata ) can be reliably photographically identified using the pattern of the intersections among pectoral scales as fingerprints. We found that I 3 S was able to identify different individuals among two samples of 21 individuals for each species independently from the error associated to the ability of the operators in collecting pictures and in digitizing the pattern of intersections among pectoral scales. In a database of 1043 images of P. muralis collected between 2007 and 2008, the software recognized 98% of recaptures within each year, and 99% of the recaptures between years. In addition, 99% and 96% of matches were ranked among the top five, and no more than 5 minutes were needed for digitizing and processing each image. The lepidosis of reptiles is a reliable alternative to ornamentation patterns in photographic identification of reptiles, which can be effectively analysed using the I 3 S software. This result represents a significant improvement in photographic identification of reptiles since (i) this procedure can be easily extended to most other species of reptiles, (ii) all kind of individuals within a species can be marked (i.e., young, subadults and adults) despite the differences in ornamentation patterns, and (iii) it is the only technique for species, like the western green lizard, which lack a clear ornamentation pattern.

Female freshwater crayfish adjust egg and clutch size in relation to multiple male traits

Females may invest more in reproduction if they acquire mates of high phenotypic quality, because offspring sired by preferred partners may be fitter than offspring sired by non-preferred ones. In this study, we tested the differential maternal allocation hypothesis in the freshwater crayfish, Austropotamobius italicus, by means of a pairing experiment aimed at evaluating the effects of specific male traits (body size, chelae size and chelae asymmetry) on female primary reproductive effort. Our results showed that females laid larger but fewer eggs for relatively small-sized, large-clawed males, and smaller but more numerous eggs for relatively large-sized, small-clawed males. Chelae asymmetry had no effects on female reproductive investment. While the ultimate consequences of this pattern of female allocation remain unclear, females were nevertheless able to adjust their primary reproductive effort in relation to mate characteristics in a species where inter-male competition and sexual coercion may mask or obscure their sexual preferences. In addition, our results suggest that female allocation may differentially affect male characters, thus promoting a trade-off between the expression of different male traits.

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.

Differential parasitaemia in the tawny owl ( Strix aluco ): effects of colour morph and habitat

The relationships between haematozoan parasites, cell-mediated immune response, territory features and colour morph were investigated in tawny owls Strix aluco, in the Park of Monza in Milan, northern Italy. Rufous and grey birds were found to differ significantly in their blood parasite loads, particularly during the breeding season, when rufous birds hosted more parasites than grey birds. The increase in parasitaemia of rufous owl was not, however, owing to the breeding status of sampled individuals. Although body condition was similar between the two colour morphs, immune response was higher in grey than in rufous owls for the same level of parasites. Moreover, parasites seemed to influence breeding of rufous birds negatively, because only individuals with no or few parasites reproduced. Both these results suggested a higher susceptibility to parasites of rufous owls. Nevertheless, a habitat effect existed because parasite loads increased significantly with woodland extent and tree-density within owl territories, and rufous birds defended more wooded territories than grey ones. Thus, the differential parasitaemia between colour morphs in tawny owls may be the result of both a differential exposure to flying vectors depending on habitat selection and a differential colour-based susceptibility of individuals.

Rufous and Grey Colour Morphs in the Italian Tawny Owl: Geographical and Environmental Influences

We examined Tawny Owls Strix aluco from different regions of Europe, especially Italy, in order to determine whether a predominance of either rufous or grey colour morphs depended on geographical, weather or habitat features. Proportions of rufous and grey owls differed throughout Europe showing a mosaic pattern of variation. Although grey owls dominated strongly in the Italian sample, no effect of latitude, altitude, weather or habitat could be found. However, the annual frequencies of the two colour morphs differed significantly over a 101-year period, depending on the weather of the years of collection, with rufous birds suffering a higher mortality in cool-dry years and grey birds in warm-wet years. This provides potential for disruptive selection.

Individually distinct hooting in male Pygmy Owls Glaucidium passerinum: a multivariate approach

During a study of territorial behaviour of the Pygmy Owl in NE Italy we used spectrographic analyses of recorded territorial calls to distinguish individual males. Two temporal parameters and the mean fundamental frequency of the song were selected and measured. All parameters of calls varied significantly within the population and discriminant analysis correctly classified 84.6% of the individual Pygmy Owls. Most individuals differed from each other with respect to the three selected parameters.