Matteo Griggio

Males prefer ornamented females: a field experiment of male choice in the rock sparrow

Recent evidence suggests that female ornaments in birds not only are genetic correlates of selection on males but may also have evolved by selection acting directly on females. However, most evidence comes from studies in captivity and it is therefore still debated whether male choice is important under natural conditions. The rock sparrow, Petronia petronia, is a monomorphic passerine in which both sexes show a yellow breast patch, although females have a smaller patch than males. Breast patch in females is positively correlated with body mass and fecundity (number of broods per year). We tested whether males preferentially mate with more ornamented females in this species. We examined male mate choice by reducing the size of the yellow breast patch of free-ranging females during the pair formation period in a colony in southeast Spain. We found that females with an experimentally reduced breast patch were courted less intensely and chased less frequently, than control females were. Furthermore, females with an experimentally reduced ornament were less likely to mate and, when mated, did so at a significantly later date than controls. These results are supported by correlational observations from another rock sparrow population, for which a larger data set was available, showing that pairs mated assortatively with respect to breast patch size. Altogether, these results therefore suggest that female ornamentation may be sexually selected in this species.

Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.

Moult speed affects structural feather ornaments in the blue tit

Growing evidence suggests that structural feather colours honestly reflect individual quality or body condition but, contrary to pigment‐based colours, it is not clear what mechanism links condition to reflectance in structural feather colours. We experimentally accelerated the moult speed of a group of blue tits (Cyanistes caeruleus) by exposing them to a rapidly decreasing photoperiod and compared the spectral characteristics of their structural feather colours with those of control birds. Blue tits were sexually dimorphic on the UV/blue crown and on the white cheek feathers. Moult speed, however, dramatically reduced brightness and the saturation only on the UV/blue crown feathers, whereas structural white on the cheek feathers was basically unaffected by moult speed. Given that the time available for moulting is usually confined to the period between the end of the breeding season and migration or wintering, UV/blue colours, but not structural white, may convey long‐term information about an individual’s performance during the previous breeding season. The trade‐off between fast moulting and structural colour expression may represent a previously unrecognized selective advantage for early‐breeding birds.

Male rock sparrows adjust their breeding strategy according to female ornamentation: parental or mating investment?

We investigated the relations between female quality and ornamentation and between male breeding investment and female ornamentation in the rock sparrow, Petronia petronia, a passerine in which both sexes have a yellow breast patch. Breast patch size in females was positively correlated with body mass and breeding status; double-brooding and primary females of polygynous males had a larger patch, and patch size could therefore be an indicator of female phenotypic quality. We conducted a field experiment to test whether males allocate their parental effort in relation to female quality, as predicted by the differential allocation hypothesis. We increased and reduced the ornament sizes of paired females and compared the behaviour of their males before and after manipulation. Frequency of brood feeding by the male was not affected by female ornament manipulation; there was a nonsignificant trend for females with enlarged ornaments, contrary to predictions, to increase their feeding rate. Reducing female ornaments resulted in a decrease in male nest attendance, a measure of passive brood defence, whereas enlarging the ornament had no effect. Males concurrently reduced their territorial (song output) and sexual activity (courtship and copulation). The reduction in sexual activity suggests that males may have changed their nest attendance in response to their mate’s renesting probability. Whatever the interpretation, these results provide some of the first evidence that not only female, but also male, birds change breeding strategy according to their mate’s phenotype in the wild.  2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.

Moult speed constrains the expression of a carotenoid-based sexual ornament

We investigated the effect of moult speed on the expression of a sexually selected, carotenoid‐based feather ornament in the rock sparrow (Petronia petronia). We experimentally accelerated the moult speed of a group of birds by exposing them to a rapidly decreasing photoperiod and compared the area and the spectral characteristics of their ornaments with those of control birds. Birds with accelerated moulting rate showed a smaller yellow patch with lower yellow reflectance compared to their slow‐moulting counterparts. Considering that the time available for moulting is usually constrained between the end of the breeding season and migration or wintering, carotenoid feather ornaments, whose expression is mediated by moult speed, may convey long term information about an individuals condition, potentially encompassing the previous breeding season. Furthermore, the observed trade‐off between moult speed and ornament expression may represent a previously unrecognized selective advantage for early breeding birds.

Extrapair paternity as a cost of polygyny in the rock sparrow: behavioural and genetic evidence of the ‘trade-off’ hypothesis

We studied the association between extrapair paternity (EPP) rate and male mating status in the rock sparrow, Petronia petronia, a facultative polygynous species. Overall, 32.0% (58/181) of the chicks were not sired by the social father and 57.1% (24/42) of the broods contained at least one extrapair young. Polygynous males allocated less time to guarding their mate during her fertile period than monogamous males but did not differ in the time spent guarding their nest. Polygynous males were cuckolded more frequently than monogamous males (50.5 and 6.6% of the young, respectively) and their paternity loss was positively correlated with the degree of overlap between the fertile periods of their primary and secondary females. Paternity loss did not differ between primary and secondary broods of polygynous males and acquiring a second mate was possible only at the expense of paternity in both broods. Late broods contained fewer extrapair young, despite no significant seasonal trend in the time allocated by the male to guarding his mate. Male yellow badge size was not associated with paternity. Old males were cuckolded less frequently than first-year males, but male age had a minor effect on paternity compared with male mating status. Reproductive success (number of young fledged/year) did not differ between monogamous and polygynous males once paternity was accounted for. Together, these results suggest that mate guarding can be efficient in preventing cuckoldry, and that there is a trade-off between attracting an additional mate and protecting paternity in the rock sparrow, whereas male age and phenotype were, at best, fair predictors of paternity.  2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour

Female ornamentation, parental quality, and competitive ability in the rock sparrow

The evolution of female ornaments is poorly understood. Recent evidence suggests not only that female ornaments may be genetic correlates of selection on males but may also have evolved through male mate choice and/or through female–female aggressive interactions. In the rock sparrow, Petronia petronia, both sexes have a carotenoid-based yellow patch that is sexually selected by both sexes. The benefits that male may gain from choosing an attractive female remain unidentified. Both parents participate in caring for the young, so there should be mutual mate choice because males and females should both benefit from choosing a good parent (good parent hypothesis; GPH). Moreover, it has already been demonstrated that the yellow patch in males is also a badge of status (armament). Therefore, the yellow patch could also serve as both ornament and armament in females (dual utility hypothesis; DUH). We investigated the hypothesis that male and female yellow patch size signals parental quality in the field. We tested by an experiment in captivity the signal function of the yellow patch in female–female aggressive interactions for access to food. Yellow patch size correlated with paternal, but not maternal, feeding rates. Thus, this study supports the hypothesis that yellow patch dimension signals male parental quality, but there is no evidence for the GPH to explain female ornamentation. In the experiment females with relatively large yellow patches had earlier access to food than those with small patches. These results seem to suggest that a sexually selected carotenoid-feather signal may be used in female–female competition, in agreement with the DUH. Males may benefit from choosing well ornamented females because these may be superior competitors.

Sexual conflict over parental care in a species with female and male brood desertion

Each parent has limited resources to invest in current reproduction, so each parent would benefit if its mate did more of the work, which generates a sexual conflict between parents. Parental care behaviour is an adaptive decision, involving trade-offs between remating (and consequently desertion of the brood) and continuing parental effort. The rock sparrow, Petronia petronia, is an unusual species in which brood desertion can occur in both sexes. Consequently, the rock sparrow is a good species to investigate the behavioural response of parents to the desertion by their mates and whether sexual difference in this response exists. We compared parental effort of pair members divided into three groups: pairs that cooperated, females that deserted the brood and males that deserted. During the period of biparental care, females fed offspring more often than males, but there were no differences among the three groups. Following desertion the total amount of care in biparental, female-only and male-only broods differed, because both sexes adjusted their care to the absence of their mates, but females overcompensated the mates absence while males only partially compensated. Nestling survival rate was positively correlated with feeding rate and the mean survival rate was lower in the male-only broods. We suggest that both parents, in a negotiation process, would benefit from withholding parental investment, as proposed in recent theoretical models on sexual conflict over parental care. Altogether our results show an unexpected plastic response of parents to care during the desertion process.

UV plumage color is an honest signal of quality in male budgerigars

Elaborate and colorful feathers are important traits in female mate choice in birds. Plumage coloration can result from pigments deposited in feathers such as carotenoids and melanins, or can be caused by nano-scale reflective tissues (structurally based coloration), usually producing ultraviolet (UV) coloration. Structural colorations remain the least studied of the three most important feather colorations. Previous studies have found a female preference for UV color in the budgerigar, Melopsittacus undulatus, but it is not clear what information this ornament conveys, nor what is the possible cost associated with its production. We investigated possible correlations between immune response and plumage color of wild-type (green) male budgerigars. In particular we measured the wing web swelling resulting from injection of phytohaemagglutinin (PHA). We did not detect any correlation between the sedimentation rate and morphological and color variables. We found that UV chroma is the best predictor for the cutaneous immune activity. Indeed, male budgerigars with high UV reflectance in the breast feathers showed stronger immune responses. These results are consistent with the idea that UV colors are special signals conveying information about a bird’s condition.

Female attractiveness affects paternal investment: experimental evidence for male differential allocation in blue tits

IntroductionThe differential allocation hypothesis (DAH) predicts that individuals should adjust their parental investment to their current mate’s quality. Although in principle the DAH holds for both sexes, male adjustment of parental investment has only been tested in a few experimental studies, revealing contradictory results. We conducted a field experiment to test whether male blue tits (Cyanistes caeruleus) allocate their parental effort in relation to female ornamentation (ultraviolet colouration of the crown), as predicted by the DAH.ResultsWe reduced the UV reflectance in a sample of females and compared parental care by their mates with that of males paired to sham-manipulated control females. As predicted by the DAH our results demonstrate that males paired with UV-reduced females invested less in feeding effort but did not defend the chicks less than males paired with control females.ConclusionsTo our knowledge, this is one of the first studies providing support for male differential allocation in response to female ornamentation.