Heritability estimates and maternal effects on tarsus length in pied flycatchers,Ficedula hypoleuca

Tarsus length has previously been shown to have an additive genetic component in some pied flycatcher populations. In addition to estimation of additive genetic variation by means of repeatability analyses at various ontogenetic stages and degrees of genetic resemblance, we explore in this paper variation among parent-offspring regressions between sexes and years, as well as the influence of hatching date, ectoparasite abundance, egg volume, and male and female condition on the tarsus length of their offspring. Mother-offspring regressions gave heritability estimates consistently higher than father-offspring regressions, although variation among years was large and both types of estimates yielded lower heritability values than those estimated by means of full-sib resemblance. This indicates that common environmental effects were inflating heritability estimates. There existed maternal effects via egg size, larger eggs fledging chicks with larger tarsi. Mean tarsus length of broods decreased with hatching date and, independently, with high loads of ectoparasitic, blood-feeding mites (Acari). The maternal effect via egg size persisted into the adulthood, and confounded the interpretation of differences between heritability slopes. We address the method of examining differences in parent-offspring regressions as a shorthand for estimating extra-pair copulation (EPC) rates. In our population, this method would give an EPC rate of 0–59%, depending on whether the analysis is performed with fledglings recruited to the breeding population or with offspring at the nest.     

Development of stress response in nestling pied flycatchers

Birds respond to unpredictable events by secreting corticosterone, which induces various responses to cope with stressful situations. However, the evidence is still elusive whether altricial nestlings perceive and respond to external stressors. We investigated the development of adrenocortical stress response to handling-related stressor in nestlings of a small passerine bird, the pied flycatcher (Ficedula hypoleuca). Nestlings were held in isolation from their parents during the experiment to ensure that they indeed respond to handling, not to parental alarm calls. We found that both 9- and 13-day-old nestlings were able to elicit hormonal stress response. Although baseline as well as stress-induced corticosterone levels rose slightly with age, the magnitude of difference between the control and stress-induced levels remained similar in both age groups. However, comparison with adults showed that the stress response of nestlings prior to fledging was still incomplete and significantly lower than in adults. Overall, our results indicate that altricial nestlings do respond to acute stressors, but on the contrary to previous predictions the development of corticosterone stress response during growth period is not gradual and varies remarkably between different passerine species.     

Ectoparasites,nest site choice and breeding success in the pied flycatcher

It has recently been suggested that nest box studies might bias the measurement of behavioural and life-history traits, because the removal of old nests may reduce the load of ectoparasites. This experimental artefact may have notable effects on nest site choice and breeding success in cavity-breeding birds. We tested (i) if pied flycatchers Ficedula hypoleuca prefer clean nest boxes and (ii) if old nest material affects the number of parasites and the breeding success of pied flycatchers. In the first experiment we offered birds one cleaned nest box and one nest box with old nest material from the previous year. The two nest boxes were placed in very similar sites near each other. In this experiment all pied flycatchers clearly preferred dirty nest boxes. In the second part of the study we distributed clean nest boxes and dirty ones on the same study area. After breeding was over we counted the number of fleas Ceratophyllus gallinae in the nest material. This flea species was the most abundant and probably the most hazardous parasite in the nests. Surprisingly, we found that there were significantly more fleas in the nest boxes with nests of the current year only than in the boxes with nests of both current and previous year. This might explain the preference for the dirty boxes. However, our results do not indicate that the number of fleas affects breeding success in the pied flycatcher.     

Experimentally induced clutch size enlargements affect reproductive success in the Pied Flycatcher

We performed a food provisioning experiment in a population of Pied Flycatchers Ficedula hypoleuca breeding at high altitude in central Spain to test if food availability before and during laying determines clutch size. Food was provided to one of two pairs with the same date of initiation of nest-building (15 dyads of subsequently reproducing pairs were thus created). Food provisioning began on the day of initiation of nest-building and ended on the day after the last egg was laid. Although laying date was unaffected by the experiment, clutch size in the experimental treatment was significantly larger. This result could indicate that food availability at laying (1) proximately constrained clutch size or (2) that females evaluated future conditions for incubating eggs and feeding nestlings based on food availability at laying. Reproductive success (proportion of eggs that resulted in fledged young) was significantly reduced in the experimental treatment. This effect suggest that supplemented females were tricked by the experiment into laying more eggs than the number of eggs they were able to incubate with success and the number of nestlings they were able to feed, a source of error in clutch size adjustment which could be common in non-experimental situations.     

Egg shell quality,clutch size and hatching success of the great tit (Parus major) and the pied flycatcher (Ficedula hypoleuca) in an air pollution gradient

Egg shell thickness, egg volume, clutch size and hatching success of Parus major and Ficedula hypoleuca were studied at 14 study sites around a copper smelter complex in Harjavalta, south-west Finland, in 1991–1993. In 1991–1992 unhatched eggs were collected to measure shell quality. F. hypoleuca was more susceptible to pollutants than P. major, the response of which was weaker in all aspects studied. Egg shells of F. hypoleuca were about 17% thinner and eggs were about 8% smaller in volume near the factory than at a distance of 10 km. The clutch size of F. hypoleuca was significantly smaller and hatching success markedly lower at a study site next to the factory complex than at all other sites. In P. major, variation in shell thickness and egg volume was not significantly related to the distance from the pollution source. Clutch size and hatching success of P. major did not significantly differ among study sites, although the trend in hatching success was in the same direction as in F. hypoleuca. Clutches of both species contained less shell material and both species had more nests without eggs near the factory than further away. The surface structure of the eggshells was studied by scanning electron microscope. Especially in F. hypoleuca, the egg shell surface was more rough and porous near the factory. The roles of Ca and heavy metals in shell thinning are discussed.     

Growth and mortality of nestling great tits (Parus major) and pied flycatchers (Ficedula hypoleuca) in a heavy metal pollution gradient

We studied nestling growth, growth abnormalities, mortality and breeding success of two hole-nesting passerines, the great tit (Parus major) and the pied fly-catcher (Ficedula hypoleuca), at 14 study sites around a copper smelter complex in Harjavalta, south-west Finland in 1991–1993. The main pollutants in the area are heavy metals and sulphuric oxides. Exposure of birds to heavy metals was shown by measuring their faecal concentrations. Copper, nickel and lead contents of nestling faeces were high near the factory and decreased with distance away from the pollution source. F. hypoleuca nestlings suffered high mortality very close to the factory complex, but did relatively well at all other sites. Breeding success of P. major was below background levels up to 3–4 km from the pollution source and nestlings grew poorly close to the factory. Growth abnormalities of legs and wings in F. hypoleuca nestlings were significantly more common near the factory than farther away. In contrast, F. hypoleuca nestlings grew equally well at all distances. The poor breeding success of F. hypoleuca close to the factory complex is probably related to the high amount of heavy metals in its diet, and low availability of calcium-rich food items may enhance this effect. We suggest that the poor breeding success of P. major is related to habitat changes that have taken place around the factory. The different responses of these two bird species are probably due to their different diet. Our results show convincingly that species-specific differences in response should be carefully considered when planning schemes for air pollution monitoring.     

Properties of food loads delivered to nestlings in the pied flycatcher: effects of clutch size manipulation, year, and sex

Experimental manipulation of the number of altricial offspring is supposed to modify parental expenditure in birds. In addition to the observed increase in parental feeding rate, it is also possible that the choice of prey or the size of load may change with the changing demand for food. Sexual differences in the provisioning response are also expected, on the basis of earlier studies. We examined the effect of brood size manipulation on choice of prey brought to nestlings and load size in the pied flycatcher. The composition and size of loads differed between years, possibly depending on varying availability of different prey types. Males responded to brood size enlargement by gathering heavier loads, whereas females showed no response. The alteration of load size in males was not explained by a larger number of prey items or mean prey size, but was a combination of these components. It is likely that males also increased their work rate in response to increased food demand at the nest. The absence of response in females might be because they are unable to increase work rate any further, or because food delivery rate in females can not be optimized by changing load properties. Received: 18 December 1997 / Accepted: 1 March 1998     

Breeding site fidelity in willow ptarmigan: the influence of previous reproductive success and familiarity with partner and territory

Summary Breeding site fidelity is high in willow ptarmigan: only 9% of males and 31% of females switched territories between years. Unpaired males were more likely to switch territories than paired males. For paired males, survival of their previous partner and reproductive success in year x did not influence probability of switching in year x+1. A female was more likely to switch territories if her previous partner disappeared. If her partner returned, she had a higher probability of switching if she did not produce chicks the previous year. Most hens moved to the territories of older males, although hens paired with unfamiliar older males did not have higher reproductive success than those paired with yearlings. Individuals that paired with their previous partner laid earlier and produced heavier chicks than those paired with unfamiliar partners. Excluding birds paired with familiar partners, survival and reproductive success in year x+1 was similar for males and females that did or did not switch territories. Males had a higher probability of producing chicks after switching than before, but females were more likely to lose their clutch after switching. For both sexes, birds that switched territories were as successful as the birds that replaced them on their former territories. We conclude that high site fidelity in willow ptarmigan is maintained because of the benefits of pairing with a familiar partner.     

Plant size,breeding system,and limits to reproductive success in two sister species of Ferocactus (Cactaceae)

Plant reproductive output can be limited by a variety of factors, bothintrinsic and extrinsic. I investigated the reproductive biologies of twospecies of unbranched short-columnar cacti, Ferocactuscylindraceus and F. wislizeni. I recordedfemalereproductive output (flowers produced, fruit set, seeds per fruit and seedmass), plant size and growth, and used hand-pollination experiments todeterminebreeding systems and pollen limitation. In both species, the ability to selfvaried among individuals, but self-pollination resulted in very few seeds,suggesting strong inbreeding depression. Neither species was pollen-limited.Numbers of flowers produced increases with plant size for both species, andseeds per fruit may also be related to plant size, although the relationship isunclear. Seed mass is not correlated with plant size. Flower production wassimilar in both species, but F. cylindraceus producedfewerseeds per fruit than F. wislizeni, and its seeds weighedless. Fruit set by F. cylindraceus was heavily impacted bya florivorous lepidopteran. Fruit set was very high (94 to 96%) inF. wislizeni, suggesting that architectural constraints(e.g., meristem limitation) are more limiting than resource levels or the levelof pollinator services. In F. cylindraceus, numbers ofseeds per fruit was positively correlated with seed mass, whereas inF. wislizeni, the relationship was negative (tradeoff).Thegrowth rates of F. wislizeni are affected by rainfall theprevious season, and growth rates increase as the plant ages.Ferocactus cylindraceus and F.wislizeni are thought to be sister species, meaning that observeddifferences between them are more likely to be the result of recentevolutionaryprocesses in their lineages rather than differing phylogenetic histories.     

Temperature during egg formation and the effect of climate warming on egg size in a small songbird

The predicted effects of recent climate warming on egg size in birds are controversial, as only two long-term studies have been reported, with contrasting results. Long-term data on egg size variation are analyzed in relation to ambient temperatures in a southern European population of pied flycatchers where breeding phenology has not matched the spring advancement in the last decades. Cross-sectional, population analyses indicated that egg breadth, but not egg length, has decreased significantly along the 16-year period, leading to marginally non-significant decreases in egg volume. Longitudinal, individual analyses revealed that despite females consistently laying larger eggs when they experienced warmer temperatures during the prelaying and laying periods, there was an overall negative response – i.e. decreasing egg volume and breadth with increasing spring (May) average temperatures – across individuals. This trend is hypothesised to be caused by the mismatched breeding phenology, in relation to climate warming, of this population. Except in the unlikely cases of populations capable of perfectly synchronising their phenology to changes in their environment, maladjustments are likely for traits such as egg size, which depend strongly on female condition. Slight changes or absence thereof in breeding dates may be followed by mismatched dates, in terms of food abundance, for optimal egg formation, which would be reflected in smaller average egg size, contrary to early predictions on the effects of climate warming on bird egg size.     

Biological control of an alien tree, Acacia cyclops, in South Africa: impact and dispersal of a seed-feeding weevil, Melanterius servulus

Several Australian Acacia species have become highly invasive in a number of ecologically sensitive areas of South Africa. Many have useful attributes that, to an extent, have hampered biocontrol efforts by restricting potential agents to those reducing seed production whilst not affecting vegetative growth. The outcome of the introduction of a seed-feeding weevil, Melanterius servulus, against Acacia cyclops in 1994, was assessed by investigating the plant’s reproductive phenology, as well as levels of damage caused by the beetles. The study provided essential information on host/agent interactions and, on the build up and spread of populations of the agent. Seed damage is commonly as high as 90% (exceptionally it reaches 100%), and dispersal rates of the beetles on average is almost 2 km per year. The consistently high levels of seed damage achieved hold promise for the future of the program, however, its ultimate success will be further enhanced through the integration of biological control with other manual clearing operations that are currently employed on a large scale in South Africa.     

Effects of superabundant food on breeding success and behavior of the red-winged blackbird

Summary The effects of food on breeding success and behavior of the red-winged blackbird (Icteridae: Agelaius phoeniceus) were investigated during 3 successive breeding seasons. In the second season, a 4-week pulse of abundant food in the form of a periodical cicada emergence (Homoptera: Cicadidae: Magicicada spp.) occurred in the forest adjacent to the marsh where the birds were breeding.During the cicada period, the bird population showed: 1) an increase in foraging trips to the forest and a decrease in trips per h, 2) increased biomass of nestlings, 3) increased nestling survival caused by decreased starvation, 4) increased fledging success, and 5) bimodal weight distributions of older nestlings (reflective of the sexual dimorphism in this species). These data suggest the temporary removal of food limitations on the breeding population when the pulse of food was available.     

Microsatellite markers reveal spatial genetic structure of Tetranychus urticae (Acari: Tetranychidae) populations along a latitudinal gradient in Europe

The genetic structure of populations of the two-spotted spider mite Tetranychus urticae was investigated along a south–north European transect spanning from southern France to The Netherlands. Mites were collected on Urtica dioica in 6 sampling zones. Microsatellite variation at 5 loci revealed considerable genetic variation with an average heterozygozity of 0.49. Significant heterozygote deficiency was found in 7 populations out of the 18 samples analyzed and one of them was completely monomorphic. Tetranychus urticae populations show some level of genetic structuring. First, genetic differentiation between localities (F _ST estimates) was significant for all comparisons. Second, the analysis of molecular variance, AMOVA, indicates that there is an effect, albeit low (9%), of the locality in accounting for allele frequency variance. Geographic distance emerges as a factor responsible for this genetic structure. The results are discussed in relation to the biological features of the species and the known patterns of migration. Related agronomical issues are addressed.     

The dispersal behaviour of the phoretic mite Poecilochirus carabi (Mesostigmata,Parasitidae): adaptation to the breeding biology of its carrier Necrophorus vespilloides (Coleoptera,Silphidae)

Summary When the phoretic mite Poecilochirus carabi reproduces in the brood chamber of its carrier Necrophorus vespilloides, a beetle with biparental brood care, the first deuteronymphs of the new mite generation aggregate on the male beetle. They do not use sex-specific traits to discriminate between male and female beetles in the brood chamber, but traits that are related to the beetles' behaviour and may be displayed by both parent beetles. When the male beetle departs, it carries virtually all deuteronymphs then present in the brood chamber. Deuteronymphs that develop later congregate on the female, which leaves the crypt some days after the male. Only those deuteronymphs that miss the female's departure disperse on the beetle larvae, meaning they have to wait in their pupal chambers until the beetles have completed their development. On average, 86% of the deuteronymphs leave the brood chamber on the parent beetles, thereby gaining the advantage of an early departure. As soon as their carrier arrives at one of the beetles' meeting places, the deuteronymphs can transfer between the beetles present. Choice experiments revealed that the deuteronymphs tend to even out density differences between congregating carriers, and prefer sexually mature to immature beetles. Therefore, transferring between beetles results in a dispersion of deuteronymphs on the sexually mature beetles of the population.     

Incorporating patterns of disperser behaviour into models of seed dispersal and its effects on estimated dispersal curves

The processes determining where seeds fall relative to their parent plant influence the spatial structure and dynamics of plant populations and communities. For animal dispersed species the factors influencing seed shadows are poorly understood. In this paper we test the hypothesis that the daily temporal distribution of disperser behaviours, for example, foraging and movement, influences dispersal outcomes, in particular the shape and scale of dispersal curves. To do this, we describe frugivory and the dispersal curves produced by the southern cassowary, Casuarius casuarius, the only large-bodied disperser in Australia’s rainforests. We found C. casuarius consumed fruits of 238 species and of all fleshy-fruit types. In feeding trials, seeds of 11 species were retained on average for 309 min (±256 SD). Sampling radio-telemetry data randomly, that is, assuming foraging occurs at random times during the day, gives an estimated average dispersal distance of 239 m (±207 SD) for seeds consumed by C. casuarius. Approximately 4% of seeds were dispersed further than 1,000 m. However, observation of wild birds indicated that foraging and movement occur more frequently early and late in the day. Seeds consumed early in the day were estimated to receive dispersal distances 1.4 times the ‘random’ average estimate, while afternoon consumed seeds received estimated mean dispersal distances of 0.46 times the ‘random’ estimate. Sampling movement data according to the daily distribution of C. casuarius foraging gives an estimated mean dispersal distance of 337 m (±194 SD). Most animals’ behaviour has a non-random temporal distribution. Consequently such effects should be common and need to be incorporated into seed shadow estimation. Our results point to dispersal curves being an emergent property of the plant–disperser interaction rather than being a property of a plant or species.     

A trematode infection with no effect on reproductive success of a sand-bubbler crab

The infection effects of the parasitic digenean trematode on the body weight and reproductive success of the sand-bubbler crab were examined. Gynaecotyla squatarolae (Trematoda: Microphallidae) infects the body cavity of Scopimera globosa (Decapoda: Scopimeridae) and uses the crab as its second intermediate host. The parasites infected all reproductive crabs examined to varying degrees. Larger crabs of both sexes had more parasites than smaller ones, probably because body size reflects age, and older crabs had a longer period of exposure to infection. Males had more parasites than females, probably because of sexual difference in acting time on the surface. Ovigerous females stay in closed burrows and do not act on the surface during incubation, and so have less chance of infection than males. The quantity of infecting parasites did not explain variations in either body weight or reproductive success of individual crabs in a field experiment. The life history of this parasite, relative body size of the crabs, and cost and the possible benefit of manipulation for the parasite may explain these results.