Manipulation of laying effort reveals habitat-specific variation in egg production constraints in Great Tits (Parus major)

It has been suggested that a bird's clutch size is not limited by the amount of resources available at the time of laying but that differences in the availability of food for nestlings is the ultimate underlying factor determining spatio-temporal variations in clutch size. However, habitat-related variations in egg production ability has yet to be investigated explicitly. We studied the breeding of Great Tits Parus major in deciduous and coniferous forests in the same area. The sizes of both the clutches and the eggs were, on average, larger in the former habitat than in the latter. A number of females were induced to lay more eggs than usual by removing four eggs from designated experimental clutches early in the laying period. These manipulated females laid approximately one egg more than control females, with the number of additional eggs laid not differing between the habitats. However, in both study years the relative size of the extra eggs – relative to the mean size of earlier laid eggs of the same clutch – was smaller in the coniferous habitat than in the deciduous habitat, while there was no habitat-related difference in the relative size of the last-laid eggs of control clutches. This result indicates that some form of proximate limitation during egg-laying period can contribute to the relatively small clutches and eggs in the coniferous habitat. Our results emphasize the need to take egg production costs into account when attempting to account for spatial variation in the reproductive behaviour of birds.     

Geographic variation in breeding parameters of the Pied Flycatcher Ficedula hypoleuca

Variation in laying date, clutch-size and number of fledglings of the Pied Flycatcher Ficedula hypoleuca was studied in relation to altitude, habitat and latitude, using data from 99 breeding areas from 35̀ N to 70̀ N. Laying date significantly increased with altitude and showed a significant quadratic relationship with latitude, with the earliest values at about 50̀ N. This quadratic relationship shows the same pattern as the relationship between arrival time of adults on the breeding areas and latitude. Clutch-size variation was significantly related to altitude and habitat, small clutches being laid at high altitude and in coniferous forests. Number of fledglings was significantly affected by habitat type, with lower values in the coniferous than in the deciduous forest. Mean clutch-size and mean number of fledglings showed the same quadratic relationship with latitude. In contrast to some earlier studies, an increased clutch-size with latitude was found, mainly because of new data from areas in the southern part of the species' distribution.     

Providing nest boxes for hole-nesting birds – Does habitat matter?

Nest boxes are a popular management tool to increase nest site availability for hole-nesting birds, but biological consequences of this technique in different habitats are poorly studied. In our study area in southwestern Estonia, nest boxes for small passerines were set up in deciduous and coniferous woods. Great tits Parus major preferred the food-rich deciduous habitat for breeding, as judged by higher nest-box occupation, earlier egg-laying and larger clutches and eggs. However, in coniferous habitat more and heavier young fledged per nest, and the return rate of both fledglings and adults was higher. We propose two mutually non-exclusive explanations, both related to the maladaptive outcome of the provision of nest boxes: (i) in the preferred habitat, nest boxes caused a supra-optimal breeding density leading to an ecological trap; (ii) boxes drastically improved the non-preferred habitat, but birds were unable to exploit the breeding habitat fully. One should be careful in providing large numbers of artificial nest sites in preferred habitats. Sometimes it would be more preferable to improve less favourable habitats by removing critical constraints.     

Experimental evidence for a novel mechanism driving variation in habitat quality in a food-caching bird

Variation in habitat quality can have important consequences for fitness and population dynamics. For food-caching species, a critical determinant of habitat quality is normally the density of storable food, but it is also possible that quality is driven by the ability of habitats to preserve food items. The food-caching gray jay (Perisoreus canadensis) occupies year-round territories in the coniferous boreal and subalpine forests of North America, but does not use conifer seed crops as a source of food. Over the last 33 years, we found that the occupancy rate of territories in Algonquin Park (ON, Canada) has declined at a higher rate in territories with a lower proportion of conifers compared to those with a higher proportion. Individuals occupying territories with a low proportion of conifers were also less likely to successfully fledge young. Using chambers to simulate food caches, we conducted an experiment to examine the hypothesis that coniferous trees are better able to preserve the perishable food items stored in summer and fall than deciduous trees due to their antibacterial and antifungal properties. Over a 1–4 month exposure period, we found that mealworms, blueberries, and raisins all lost less weight when stored on spruce and pine trees compared to deciduous and other coniferous trees. Our results indicate a novel mechanism to explain how habitat quality may influence the fitness and population dynamics of food-caching animals, and has important implications for understanding range limits for boreal breeding animals.     

Seasonal habitat-dependent change in nest box occupation by Tengmalm’s owl associated with a corresponding change in nest predation

We investigated the nest-site settlement of the forest-dwelling Tengmalm’s owl (Aegolius funereus) in the temperate zone of Central Europe (Czech Republic) over a 14-year period. We analysed the effect of changes in nest-site suitability (expressed as the proportion of deciduous forest) within the immediate surroundings of the nest boxes (a buffer radius of 25 m) in relation to nest box occupancy, the mean standardized laying date (further the laying date), occurrence of nest predation, and breeding density. In line with the predictions of the site dependence model, we found a positive relationship between the laying date and the proportion of deciduous trees in the nest box surrounding. Further, we found the interactive effect of the laying date and the proportion of deciduous forest surrounding a nest box on the occurrence of breeding attempts predated by pine martens. A higher proportion of deciduous forest was found in the surroundings of predated than non-predated breeding attempts, and a steeper positive relationship between the laying date and the proportion of deciduous forest was found in the surroundings of non-predated than predated breeding attempts. The change in the owls’ habitat utilization can be explained by the process of tree leafing which makes the deciduous forest a less suitable habitat with high risk of nest predation before tree foliage and a more suitable habitat with low risk of nest predation after tree foliage. We conclude that our results are in agreement with the site dependence model, and that the degree of foliage of deciduous forest in the nest surrounding of Tengmalm’s owl is a key factor determining site suitability due to predation risk from pine martens in Central Europe.     

Provision of nestboxes raises the breeding density of Great Tits Parus major equally in coniferous and deciduous woodland

Nestbox provision is a technique used to increase nest-site availability for secondary cavity-nesting birds. However, little is known about the demographic consequences of nestbox provision in different habitat types. To assess how nestbox provision affects the density of hole-nesting birds simultaneously in two contrasting habitats, we compared the breeding density of Great Tits along transects without nestboxes with that in transects where nestboxes were provided. Although the initial density of breeders was considerably higher in the deciduous habitat than in the coniferous habitat, provision of nestboxes increased density by a similar number of additional pairs in each habitat type. Thus, the provision of nestboxes in managed coniferous forests may be as effective in increasing the breeding opportunities of cavity nesters as in deciduous stands. Moreover, previous research showed that pairs in deciduous habitat with nestboxes have consistently lower breeding success than those in coniferous habitat with nestboxes. It is possible that the addition of nestboxes in the preferred habitat increased density to such an extent that density-dependent effects became apparent.     

Nest site selection and predation driven despotic distribution of breeding long-eared owls Asio otus

Long-eared owls Asio otus do not build their own nests. They occupy old corvid nests in either coniferous or deciduous trees. In my study area owls favoured coniferous over deciduous trees for nesting and occupied nest sites in that order. Comparing overall breeding output, I found no difference between the two kinds of nest sites, but among early nests there was a higher proportion of unsuccessful nests in deciduous trees. This difference was due to differences in predation rate, indicating that the owls' pattern of nest site occupation followed predictions from the ideal despotic distribution model. I also show that, due to a seasonal change in habitat quality, predator driven despotism, although present, can be well masked and hence difficult to detect.     

Weather‐mediated decline in prey delivery rates causes food‐limitation in a top avian predator

Inclement weather can negatively affect breeding birds directly by exposure to factors such as severe temperature and rainfall, or indirectly by reducing food supply. During a three‐year study of Arctic peregrine falcons Falco peregrinus tundrius breeding in Nunavut, Canada, we estimated annual prey density at a biologically relevant scale (i.e. the home range of breeding pairs), and examined the manner in which prey density and within‐season weather conditions influenced occupancy of breeding sites, egg‐laying, hatch rate, prey delivery rates and growth and survivability of nestlings. The first two summers of our study (2010–2011) were warm and dry, while the third summer (2012) was cool and wet, and was preceded by a severe spring rain event. We found that occupancy of breeding sites was consistently high. As a proportion of the number of eggs laid, hatch rate did not change among years, but the number of eggs laid per occupied site declined in the third year of the study. In the first two years of the study, the number of nestlings per occupied sited was high, but declined in the third year. Total prey density at the home range scale was similar in 2010 and 2012, while the highest prey density was recorded in 2011. Total prey delivery rates to nestlings and nestling growth rates were significantly lower in 2012, which received more precipitation than 2010 and 2011. Nestling growth rates were similar in 2010 and 2011, but were markedly different in 2012; for both sexes the period of rapid growth was of shorter duration in 2012 and asymptotic weights were lower. This research contributes to the growing body of evidence that indicates severe rain events and ongoing periods of wet weather can reduce reproductive output of Arctic‐breeding raptors regardless of whether it occurs during laying, incubation or brood rearing.     

Isolation by time and habitat and coexistence of distinct host races of the common cuckoo

Isolation by time occurs when different populations of a single species reproduce at different times and thereby reduce the probability of interbreeding, potentially causing divergent adaptation to timing of reproduction, eventually resulting in ecological species separated by timing of reproduction. We analysed extensive data on timing of reproduction by different host races of the common cuckoo Cuculus canorus that is an obligate brood parasite laying eggs in the nests of many different species of passerine birds. Because different hosts breed at different times, specific host races of cuckoos have adapted to specific hosts by laying eggs when nests of these hosts are available, and such divergence may be further exaggerated by differences in timing of breeding among host races with similar habitat requirements. Host species accounted for a quarter of the variance in timing of breeding by the cuckoo. Common cuckoos reproduced at a similar, but narrower subset of dates as did possible hosts, showing that only a fraction of hosts with specific breeding dates were parasitized. Common cuckoo eggs laid in the 'right' kind of nests, phenotypically matching the eggs of the host, were laid later during the season than cuckoo eggs laid in the 'wrong' kind of nests where the eggs did not mimic those of the host. Pairs of sympatric cuckoo host races differed more in timing of breeding than pairs of allopatric host races, and pairs of cuckoo host races with similar breeding habitat differed more in breeding date than pairs of cuckoo host races with dissimilar habitat, as expected from reproductive character displacement. These findings are consistent with cuckoo host races being isolated by timing of breeding and habitat.     

Habitat quality as a predictor of spatial variation in blue tit reproductive performance: a multi-plot analysis in a heterogeneous landscape

Vertebrate studies have rarely investigated the influence of spatial variation in habitat richness on both short-term (breeding) and long-term (offspring recruitment) reproductive performance using simultaneously multi-patch, multi-habitat type and multi-year approaches at landscape level. Here we present results of such an approach using the influence of two oak tree (Quercus ilex, Q. humilis) species on reproductive performance in Corsican blue tits (Parus caeruleus ogliastrae) as a model system. We found that blue tits breeding in rich broad-leaved deciduous patches consistently laid eggs earlier in the season, and produced larger clutches and more fledglings of higher quality, than those breeding in poor evergreen patches. Also, parents, especially males, were in better physical condition in the broad-leaved deciduous than in the evergreen patches. Surprisingly, estimates of long-term effects of reproduction, such as recruitment rates of locally born offspring, did not differ between the two habitat types. Our results suggest that short-term breeding performance and phenotypic quality of both chicks and parents do not necessarily provide reliable information about contributions to following generations at a scale larger than that of the local study plot. Differences in reproductive performance between the two oak habitat types could not be attributed to density-dependent effects, differences in levels of nest predation, or differences in age structure of the birds. We suggest that habitats that are optimal for breeding are not necessarily optimal for survival after the breeding season.     

Nest quality limits the number of hole-nesting passerines in their natural cavity-rich habitat

We experimentally tested whether the lack of high-quality nest holes may limit the number of secondary cavity-nesters in cavity-rich habitats. We mapped and measured natural tree-holes in 10 mature aspen forests in Estonia. In five experimental plots, we provided nest-boxes on trees having suitable cavities for hole-nesting passerines. This improved the quality of available cavities but retained their total number (due to territoriality, sites in the same tree could not be occupied simultaneously) and location on the landscape. In the next breeding season, the density of hole-nesting passerines increased in experimental plots but remained at the same level in control plots. The increase was due to the species that used boxes: Parus major that preferred these over natural cavities, and Ficedula hypoleuca that used both nest types more opportunistically. We concluded that the quality of cavities can influence the numbers of hole-nesting birds in old cavity-rich forests, also; probably, adaptive habitat selection is the proximate mechanism in this case.     

Population dynamics of japanese serow in relation to social organization and habitat conditions. I. Stability of Japanese serow density in stable habitat conditions

The density of a Japanese serow (Capricornis crispus) population, which has been in stable habitat conditions and free from hunting and predation, was investigated by direct observation of serows at Kusoudomari (336 ha), Wakinosawa village, Aomori Prefecture, Japan, during 1977–89. The vegetation, composed of mature mixed forest ofThujopsis with deciduous trees (43.2%), secondary forest of deciduous trees (23.4%) and a plantation of coniferous trees (30.0%) showed little change during the study period. The serows were counted 17 times each winter for 14 consecutive years. The serow population maintained a stable density with a mean of 12.5 individuals km⁻² (SD=1.4) ranging from 8.6 to 14.6 km⁻². The ratio of the proportion of kids (<1 year old) to serows more than 1 year old decreased, but density did not decrease during the study period. Though extremely severe winters with heavy snowfall in 1984–86 caused a decline in serow density, the density recovered within a few years. It was considered that the stability of the serow density resulted mainly from the stable food supply and the lack of human disturbance, such as forest cutting in the habitat. Stable population density for a long period is probably related to the social organization of the serow.     

Seasonal use by birds of stream-side riparian habitat in coniferous forest of northcentral British Columbia

To determine use of riparian habitats by birds in the northern coniferous forest of British Columbia, we censused birds and vegetation along 500 m transects placed parallel and perpendicular to three second-order streams. Censuses were conducted during spring, summer, autumn, and winter to investigate how use of riparian habitat changed seasonally. Stream-side riparian zones were characterized by a dense understorey of deciduous vegetation not found in the upslope forest. Nine bird species preferred the riparian understorey for breeding, six preferred it only during migration. Neotropical migrants (16 of 46 species) were more closely associated with stream-sides than year-round residents (11 species). Some breeding birds (five species) were significantly negatively associated with riparian habitats. The density of riparian birds declined with distance upstream but did not decline up to 250 m away from the stream. The more extensive riparian areas downstream supported a greater density of birds in all seasons compared to upstream areas, but more species only in spring and autumn. Species that nested in non-riparian areas in summer used riparian habitat in autumn, making riparian corridors in the northern coniferous forest important during migration. Maintaining both riparian and upslope habitats is necessary to preserve species diversity al the landscape level.     

Are habitat islands islands? Woodliving beetles (Coleoptera) in deciduous forest fragments in boreal forest

Modern forestry has changed the fire dynamics in the boreal forest and as a result the size and number of deciduous forest patches have been reduced as well as the number of deciduous trees within the coniferous forest This has exaggerated the insularity of deciduous forest patches within the boreal coniferous forest zone In this paper I examine whether the diversity of beetles living in dead stems of deciduous trees follows the relationship with patch area to be predicted from island biogeographic theory, and to what extent the species assemblages differ between large and small patches and single deciduous trees within managed coniferous forests Three larger patches of deciduous forest (>120 ha) arisen as successions after forest fires in the late 1880's are compared with 6 small (<20 ha) patches of similar origin No difference in diversity could be detected between large and small patches A statistically significant difference between assembly composition was detected using matrix regression between a matrix of observed assembly similarities and a hypothetical similarity matrix based on the type of area in which the plot was situated (large- small or matrix) Finally I examined the distribution of the 56 most common beetle species over the habitat types investigated Thirty four species did not show any Significant habitat preference, 12 were found more often in clear-cut areas than expected 5 were over-represented in small patches, and 4 in large areas My explanation to this apparent lack of insularity effects is the relative recent commencement of intensive forestry in these areas and the fact that the matrix is of rather high quality that is inhabitable for numerous beetle species Thus the patches may be viewed as incipient islands still exchanging biota with their surroundings     

自然环境下影响扬子鳄繁殖的环境因子

生态因子对生物的生长、发育、生殖和分布有着直接和间接的影响.在1998～2005年对扬子鳄繁殖研究中心内自然环境下鳄卵及其繁殖地环境因子进行调查,包括窝卵数、产卵时间、出壳时间、孵化期内温度、湿度的变化情况、雏鳄孵出后的存活数量以及巢区的植被盖度.调查发现天气状况直接影响母鳄产卵时间、卵的孵化质量与雏鳄的出壳数量.盖度相对大于0.5的窝巢中多数卵基本能正常发育,盖度小于0.5的窝巢中可能只有少数几枚卵发育正常.     

Assessing the age of field female wheat bulb flies (Leptohylemyia coarctata)

From 1967 to 1971, wheat bulb fly females caught by sweeping from a number of localities contained mature eggs from the third week of July to the end of August. In 1971 when female wheat bulb flies were caught by sweeping, by white water traps, by suction traps as well as by light traps, the first flies containing mature eggs were caught 29 days after the first detected emergence while the first batch was laid 29–58 days after first emergence. The peak period of egg laying occurred in the fourth week of July, 24–38 days after the peak emergence. The second batch of eggs was laid when the flies were 8 weeks old, 29 days after the first; the peak for this batch was 35 days after the first. A few females laid a third batch of eggs 28 days after the first record of the laying of the second batch. A female that lives sufficiently long to lay the third batch of eggs is at least 12 weeks old. In 1970, when emergence started earlier in June, ripening and laying of eggs of the three batches was also earlier. After the harvest of cereal crops more females were caught by white water traps placed on the fallow than by other methods. The most important pathogen affecting adult wheat bulb flies was a fungus, probably Entomophthora muscae. When infection occurred early in July, fewer eggs from the first batch were laid than when infection developed later. Fewer flies were infected in 1971 than in 1970. To be effective against wheat bulb fly in the early spring, control measures-should be applied against adults before their eggs mature in early July of the preceding year.     

BREEDING BIOLOGY OF THE DARTFORD WARBLER SYLVIA UNDATA IN ENGLAND

The breeding biology of Dartford Warblers was studied during 1975–1976 in 1010 ha of heathland on the Purbeck peninsula, Dorset; comparisons are drawn from other areas of the species' range. Clutches were of 3,4 or 5 eggs, with the mean clutch size increasing progressively from early May to mid-June, declining thereafter. Incubation and brooding of the young were shared unequally between the sexes, with the female taking the greater part. In the number of feeding visits to nestlings there was no difference between the sexes until the last four days of nest-life, when the male greatly increased his share. Nesting was comparatively successful (about 80% of nests producing some young), though the incidence of single infertile eggs was high. There were large variations between years and sites in the timing of the starting of laying. Late-laying pairs reared only one brood, while early starters could rear two. Timing of starting, influencing the number of broods reared, appeared to be the most important factor causing variation in breeding productivity. No vegetational characteristics were entirely adequate to explain variation in starting dates. Past data, however, showed earlier starting in Sussex and Jersey where gorse was a more abundant component of the vegetation of Dartford Warbler habitat than in Dorset. It is suggested that the abundance of gorse is directly related to the quality of the habitat for Dartford Warblers, and hence to breeding productivity.     

No effect of partner age and lifespan on female age‐specific reproductive performance in blue tits

Studies of age‐specific reproductive performance are fundamental to our understanding of population dynamics and the evolution of life‐history strategies. In species with bi‐parental care, reproductive ageing trajectories of either parent may be influenced by their partner's age, but this has rarely been investigated. We investigated within‐individual age‐specific performance (laying date and number of eggs laid) in wild female blue tits Cyanistes caeruleus and evaluated how the age and longevity of their male partner indirectly influenced the females’ reproductive performance. Females showed clear age‐dependence in both laying date and number of eggs laid. We found that female reproductive performance improved in early life, before showing a decline. Longer‐lived females had an earlier laying date throughout their lives than shorter‐lived females, but there was no difference in number of eggs laid between longer‐ and shorter‐lived females. Within breeding pairs, the female's (age‐specific) reproductive performance was not dependent on the age and longevity of the male partner. We conclude that the age and quality of the male partner may be of little importance for traits that are under direct female control.     

Parid foraging choices in urban habitat and their consequences for fitness

Urban environments are habitat mosaics, often with an abundance of exotic flora, and represent complex problems for foraging arboreal birds. In this study, we used compositional analysis to assess how Blue Tits Cyanistes caeruleus and Great Tits Parus major use heterogeneous urban habitat, with the aim of establishing whether breeding birds were selective in the habitat they used when foraging and how they responded to non‐native trees and shrubs. We also assessed whether they showed foraging preferences for certain plant taxa, such as oak Quercus, that are important to their breeding performance in native woodland. Additionally, we used mixed models to assess the impact of these different habitat types on breeding success (expressed as mean nestling mass). Blue Tits foraged significantly more in native than non‐native deciduous trees during incubation and when feeding fledglings, and significantly more in deciduous than evergreen plants throughout the breeding season. Great Tits used deciduous trees more than expected by chance when feeding nestlings, and a positive relationship was found between the availability of deciduous trees and mean nestling mass. Overall, the breeding performance of both species was poor and highly variable. Positive relationships were found between mean nestling mass and the abundance of Quercus for Great Tits, but not for Blue Tits. Our study shows the importance of native vegetation in the complex habitat matrix found in urban environments. The capacity of some, but not all, species to locate and benefit from isolated patches of native trees suggests that species vary in their response to urbanization and this has implications for urban ecosystem function.     

Woodland structure,rather than tree identity,determines the breeding habitat of Willow Warblers Phylloscopus trochilus in the northwest of England

Capsule Woodland structure, rather than tree species, is the most important determinant of breeding habitat selection by Willow Warblers in North West England.

Aims To examine how habitat characteristics predict the occurrence of male Willow Warbler territories.

Methods Woodland structure (trunk density, trunk diameter, canopy cover and understory cover), tree species and food abundance were compared between woodland areas within and outside of male territories at a site in the UK.

Results Territories contained higher trunk numbers, had a narrow range of trunk diameters, and intermediate canopy cover. Food abundance did not differ with occupancy. Willow and alder were the most common trees within territories, in contrast to birch which has been found in previous studies. The habitat structure matches young woodlands, where birches often grow. However, at the study site the birches were large and mature, and therefore unsuitable. Moreover, woodland structure variables were better predictors of occupancy than any particular tree genera.

Conclusion The results indicate that vegetation structure, but not tree species or food availability, influence breeding habitat selection by Willow Warblers. The preferred structure is similar to coppice woodlands; therefore, the Willow Warbler decline may be linked to the loss of this traditional management across south England.