Breeding ecology of Yellow Wagtails Motacilla flava in an arable landscape dominated by autumn-sown crops

Capsule Yellow Wagtails successfully raised two consecutive broods in landscapes dominated by autumn-sown crops and did not require spring crops or fallow plots for later nesting.

Aims To assess whether arable landscapes dominated by winter cropping provide habitats that allow Yellow Wagtails to raise two successful broods. To assess the utility of spring cultivated agri-environment fallow plots as a nesting and foraging habitat for Yellow Wagtails.

Methods Nesting success and foraging behaviour of Yellow Wagtails was monitored on lowland arable farmland dominated by winter cropping during two successive breeding seasons.

Results Yellow Wagtails successfully raised first and second broods mainly in winter cereals, with later nests being more successful. Some nests were initiated in bean fields where egg-stage failure rates were high, probably as a consequence of depredation. Fallow plots were not used for nesting but, along with other areas of sparse vegetation, were regularly used for foraging.

Conclusions Yellow Wagtails breeding at relatively low densities achieved high rates of reproductive success in a landscape dominated by winter cereals and with few spring-sown crops. This study suggests that spring-sown crops may not be necessary for Yellow Wagtails to rear two successful broods each summer within arable landscapes.     

Assessing breeding success in common woodland birds using a novel method

Capsule Repeated counts of fledged broods can provide a useful estimate of breeding success for most common woodland birds.

Aims To assess the efficacy of comparing fledged-brood survey data with territory mapping using simple mark–recapture analysis techniques to provide an estimate of breeding success for common woodland birds that does not involve finding nests.

Methods Three observers undertook territory mapping surveys of adults, followed by counts of fledged broods four times a week during May–July 2007 in two 15 ha woods each, both in southern England. Using known fledging to maturity periods, these counts were used to calculate daily detection probabilities for broods of ubiquitous species. These enabled fledged brood territory occupancy probabilities (i.e. brood to territory ratios) to be estimated that take account of the possibility that broods were present but missed by surveys.

Results Of the 19 species found in all six woods, mean daily detection probability estimates for fledged broods of 17 species ranged from 0.17 to 0.50 with significant variation between woods for 12 species, but within region/observer for four species. The mean probability of detecting a brood at least once was over 75% using four visits per week and over 50% using two visits. Only for Great Spotted Woodpeckers Dendrocopos major and Garden Warblers Sylvia borin was the fledging period too short and the daily detection probability too low to provide a reasonable estimate of the territory occupancy probability.

Conclusion Daily detection probabilities for fledged broods of most common woodland birds were sufficiently high to enable useable estimates of fledged-brood territory occupancy probabilities to be made based on a survey programme involving two or three visits per week between late May and the end June. The method used may have application as a means of providing a relatively easily derived productivity index for woodland bird monitoring programmes or for research studies.     

Timing of nest departure by broods of Dotterel Charadrius morinellus

Capsule Most broods left during early to mid-morning, which may allow the small, largely exothermic chicks to emerge into a favourable thermal environment for feeding.     

The pattern of Dunlin Calidris alpina distribution and abundance in relation to habitat variation in the Flow Country of northern Scotland

Capsule Folivorous caterpillars constituted the majority of nestlings’ food in a primeval forest. Blue Tit broods only partially matched the caterpillar peak, and the mismatch did not affect food composition or nesting success.

Aims To describe factors influencing the timing of reproduction in Blue Tits under primeval conditions (Bia?owie?a National Park, Poland) and to check whether they schedule breeding so as to synchronize broods with a seasonal caterpillar peak.

Methods We gathered information on phenology of leaf development, seasonal availability of folivorous caterpillars (frass collection), timing of Blue Tit breeding, composition of its nestling food, and nest fate over a three-year period.

Results Caterpillars constituted c. 74% of nestling diet, but only 17–65% of broods matched the caterpillar peak in any season. Neither total nest loss, nor frequency of brood reduction depended on the level of mismatch. Caterpillar availability was probably adequate every year, regardless of the amount of mismatch, and no selective advantage of precise matching was detectable. Phenological events at all trophic levels occurred earlier in warmer springs. Egg-laying coincided with tree bud burst and appearance of caterpillars, but was not critically dependent on their timing.

Conclusion The observations are consistent with the view that Blue Tits under primeval conditions in Bia?owie?a National Park, Poland, breed as early as possible, rather than synchronizing their breeding with the caterpillar peak later in the season.     

Car traffic along hedgerows affects breeding success of Great Tits Parus major

Capsule?In hedgerows near roads with fast and frequent traffic, the mortality of Great Tit Parus major broods was higher than in hedgerows with less traffic and hedgerows with no disturbance.     

Condition of large brood in Bonelli's Eagle Hieraaetus fasciatus

Capsule Young body condition is affected by the interaction of environment (rainfall) and brood size.

Aims To investigate factors affecting offspring condition using levels of urea in plasma.

Methods We used generalized linear mixed models (GLMMs) with the levels of urea in plasma as the dependent variable and laying date, brood size, sex and year as the explanatory ones.

Results Brood size had a significant effect on offspring condition only during a year of adverse weather (heavy rainfall). During this period, young from broods of two were in poor condition compared with single broods. Conversely, brood size had no effect in the other two years analysed. Neither sex nor laying date had a significant effect on young condition. Offspring condition was not related to first-year survival.

Conclusion There is a trade-off between reproduction (brood size) and offspring condition only in years of adverse environmental conditions.     

Parental differences in brood provisioning by Hen Harriers Circus cyaneus

Capsule Females varied their provisioning patterns according to brood age and brood size, whereas males did not.

Aims To quantify how parents balance the needs of their offspring for food and protection.

Methods We studied 13 nests from hides and spent on average 101 hours per nest monitoring prey types, provisioning rate and the time spent at the nest by both sexes in relation to brood size and brood age.

Results Males always provided more food than females. Males brought similar amounts of prey items irrespective of brood size and nestling age, whereas females brought more prey and bigger items to larger and older broods. Females spent less time brooding larger broods, particularly early on.

Conclusions Hen Harrier parents share the provisioning burden, with each parent delivering prey as a function of brood care requirements, hunting capability and the behaviour of the other parent.     

Habitat selection and diet of the Capercaillie Tetrao urogallus in Abernethy Forest,Strathspey, Scotland

Capsule Woodland characteristics are important for fully grown birds and for broods.

Aims To describe habitat selection and diet of the Capercaillie in ancient native pinewoods and Scots Pine Pinus sylvestris plantations, in order to provide a guide for forest management.

Methods Tree selection was studied in winter by locating droppings under trees along transects, and comparing those trees used by Capercaillie with those not used. Habitat selection by broods was studied by following radiotagged chicks or females with broods. Diet was described from the analysis of droppings.

Results In winter, Capercaillie preferred Scots Pines with wide crowns and large branches, and little surrounding Heather Calluna vulgaris, but avoided forest tracks. Female Capercaillie occurred in stands with a higher density of pines compared to males. The diet of fully grown birds comprised primarily Scots Pine needles between October and April. Scots Pine pollen cones were eaten during May, June and July. Bilberry Vaccinium myrtillus leaves and fruit were eaten in summer and autumn. Habitats used by broods had a higher percentage of Vaccinium sp. and smaller Scots Pines compared with random sites. Chicks hatch at the beginning of June, and in their first two weeks, their diet comprised Formicidae, Lepidoptera larvae and Coleoptera. In July and August, Lepidoptera larvae were rarely eaten but a higher percentage of chicks took vegetable material (particularly Bilberry fruit). In June, smaller numbers of invertebrates were generally trapped in mires of wooded bogs compared with areas of Heather and Vaccinium sp. in old woodland. Areas of Vaccinium sp. had the highest numbers of Formicidae, Coleoptera and Diptera, and especially Lepidoptera larvae. Numbers of Lepidoptera larvae declined during June and July as larvae pupated, explaining the reduction of Lepidoptera larvae in the diet of older chicks.

Conclusions Habitat management for Capercaillie should aim to provide old Scots Pine forest with a Vaccinium-rich field layer. Within Scotland, this may entail increasing the tree density in ancient native pinewoods but decreasing it in pine plantations. The removal or closure of tracks might increase the amount of woodland available to Capercaillie, by reducing human disturbance.     

Burrow depth,carbon dioxide and reproductive success in Sand Martins Riparia riparia

Capsule: Carbon dioxide (CO₂) concentrations in the burrows of Sand Martins Riparia riparia increase with depth but have no detectable impact on fledging success.

Aims: To investigate whether burrow depth and CO₂ concentrations influence reproductive success in Sand Martins.

Methods: We monitored two Sand Martin colonies along the River Lune, Lancashire, UK, to investigate the effect of burrow depth on reproductive success. We also measured CO₂ levels in a sample of burrows to test whether burrow depth predicts CO₂ concentration, and to test for a relationship between CO₂ concentration and breeding success.

Results: Burrow depth was significantly correlated with fledging success, but the correlation was positive in first broods and negative in second broods. The highest CO₂ concentration recorded was 73?650?ppm and the mean concentration across burrows was 31?757?ppm. However, while CO₂ concentrations were positively correlated with burrow depth after controlling for the number and age of nestlings, they were not correlated with reproductive success.

Conclusion: There are reproductive costs associated with deeper burrows in second broods, but these could not be attributed to CO₂ concentrations despite the exceptionally high levels recorded. This study highlights the need for further investigation into gas exchange and the potential impacts of, or adaptations to, CO₂ accumulation in avian burrows.     

Sex‐specific differences in Sand Martin Riparia riparia nest attentiveness

Capsule Nest attentiveness during the night is significantly lower in males but varies with the activities of the breeding cycle. Male nest attentiveness shows minima during the burrowing and egg‐laying phases, and also during the brooding of chicks in second or re‐nest broods.     

Renesting by male Dotterel Charadrius morinellus after successfully rearing chicks

Although some waders (Charadrii) are known regularly to attempt to rear 2 broods in some summers (e.g. Ringed Plover Charadrius hiaticula, Little Ringed Plover Charadrius dubius ^l), this ability is not associated with Palearctic waders breeding at higher latitudes, except where one or more clutches are left in the sole care of previous mates. This note documents an attempt made by a male Dotterel to raise a second brood after its first brood of young became independent.     

Factors influencing the nesting success of Lapwings Vanellus vanellus and behaviour of Red Fox Vulpes vulpes in Lapwing nesting sites

Capsule Lapwing nest predation was negatively correlated to nest density, while Lapwing alarm duration in response to foxes was positively correlated with the number of Lapwing broods present.

Aims To identify factors affecting Lapwing nest predation and Red Fox search effort.

Methods Lapwing nest success was monitored at four sites in 1996, seven sites in 1997 and six sites in 1998. In 1997 we mapped the position of all Lapwing nests in order to determine distances between nests, and the proximity of linear features and potential avian predator perches to each nest. From April to June 1998 we carried out 199 hours of nocturnal observations at six Lapwing nesting sites using night vision equipment.

Results The risk of nest predation was significantly higher for more isolated nests. Nocturnal observations showed that of all the nocturnal predators, foxes were the most active at Lapwing nesting sites. However, fox search effort in Lapwing colonies was relatively low, averaging 57 s/ha per visit. Foxes spent significantly longer foraging near breeding Lapwings (measured as duration of alarm calls) when more broods were present. Fox search effort (s/ha per hour of observation) tended to be greater in areas of high waterbird density.

Conclusion The lack of positive density-dependent nest predation, the relatively low search effort of foxes near Lapwing nesting sites and the high nest success sometimes achieved in areas with foxes all suggest that Lapwing nest predation by foxes is ‘incidental’. Lapwing chicks are probably more vulnerable to predation by foxes than clutches.     

Intraseasonal effects of clutch manipulation on parental provisioning and residual reproductive value of eastern phoebes (Sayornis phoebe)

Summary First clutches of double-brooded eastern phoebes Sayornis phoebe were manipulated (up two eggs, down 2 eggs or no change) to test for intraseasonal reproductive tradeoffs and to test whether size of first brood influenced food delivery rates to nestlings and nestling quality in second broods.Considering all nests from both broods, rate of feeding nestlings increased linearly with brood size but nestling mass per nest decreased with increasing brood size. High nestling weights in small broods may have resulted from parents delivering better quality food, but we did not test this.Among treatment groups in first broods, nestlings from decreased broods weighed more than those in control or increased broods. Treatment did not influence the likelihood that second nests would be attempted after successful first nests nor did it alter the interval between nests. Nestlings of parents that renested weighed more than those of parents that did not, regardless of treatment, suggesting that post-fledging care may preclude renesting. Mass of individual females did not change between broods, regardless of brood size. Clutch sizes of second attempts were not affected by manipulations of first broods but increasing first broods reduced the number of nestlings parents were able to raise to day 11 in their second broods. However, manipulation of first broods did not affect mean nestling mass per nest of nestlings that survived to day 11.In phoebes, parents of small first broods are able to raise nestlings in better condition. We predict that in harsh years, parents of small first broods would be more likely to renest. Parents of enlarged first broods sacrificed quality of offspring in second broods, which seems a reasonable strategy if nestlings from second broods have lower reproductive value.     

Diversionary feeding and nestling diet of Hen Harriers Circus cyaneus

Capsule: Diversionary feeding reduced Hen Harrier Circus cyaneus nestlings’ natural food intake by half. Red Grouse Lagopus lagopus scotica chicks constituted 0–4% of all nestling food items. Annually, this reduced annual grouse chick production by 0–6%.

Aim: To quantify proportions of diversionary and natural food (including grouse) delivered to Hen Harrier nestlings in relation to brood size, male status and natural prey abundance.

Methods: We recorded diversionary food provisioned to 25 Hen Harrier broods (2008–15) and studied the diet of 15 broods using observations from hides, nest cameras and regurgitated pellet analysis. Variation in nestling diet was analysed using compositional analysis.

Results: Hen Harriers took 76% of diversionary food provided. Depending on assessment method, average nestling diet was 44–53% diversionary food, 39–55% natural prey (including 24–45% passerines, 4–15% small mammals, 0–4% grouse chicks) and 0–9% unknown items. The amount of diversionary food consumed was not influenced by male status, brood size or natural prey abundance. The number of Red Grouse chicks delivered annually was 34–100% lower than expected under unfed conditions, however, the confidence intervals associated with these estimates were large.

Conclusion: Diversionary food influenced Hen Harrier nestling diet and reduced the number of Red Grouse chicks taken relative to modelled predictions. It may help reduce conflict between Hen Harrier conservation and Red Grouse shooting, but only if overall grouse productivity is thereby maintained or increased.     

Male death resulting from hybridization between subspecies of the gypsy moth,Lymantria dispar

We explored the origin of all-female broods resulting from male death in a Hokkaido population of Lymantria dispar through genetic crosses based on the earlier experiments done by Goldschmidt and by testing for the presence of endosymbionts that are known to cause male killing in some insect species. The mitochondrial DNA haplotypes of the all-female broods in Hokkaido were different from those of normal Hokkaido females and were the same as those widely distributed in Asia, including Tokyo (TK). Goldschmidt obtained all-female broods through backcrossing, that is, F1 females obtained by a cross between TK females (L. dispar japonica) and Hokkaido males (L. dispar praeterea) mated with Hokkaido males. He also obtained all-male broods by mating Hokkaido females with TK males. Goldschmidt inferred that female- and male-determining factors were weakest in the Hokkaido subspecies and stronger in the Honshu (TK) subspecies. According to his theory, the females of all-female broods mated with Honshu males should produce normal sex-ratio broods, whereas weaker Hokkaido sexes would be expected to disappear in F1 or F2 generations after crossing with the Honshu subspecies. We confirmed both of Goldschmidt''s results: in the case of all-female broods mated with Honshu males, normal sex-ratio broods were produced, but we obtained only all-female broods in the Goldschmidt backcross and obtained an all-male brood in the F1 generation of a Hokkaido female crossed with a TK male. We found no endosymbionts in all-female broods by 4,′6-diamidino-2-phenylindole (DAPI) staining. Therefore, the all-female broods observed in L. dispar are caused by some incompatibilities between Honshu and Hokkaido subspecies.     

Allocation of Male Parental Care in Relation to Paternity Within and Among Broods of the Common Yellowthroat (Geothlypis trichas)

The relationship between male parental care and paternity has been investigated in a number of avian species, but in many cases the influences of confounding factors, such as variation in male and territory quality, were not addressed. These sources of variation can be controlled for by making within-male comparisons between successive broods or within-brood comparisons between groups of fledglings in a divided brood. We studied the relationship between male parental care and paternity in the common yellowthroat ( Geothlypis trichas ) at three levels: between groups of fledglings in divided broods, between first and second broods of the same pair, and among all broods in the population. In this study we proposed three hypotheses: first, males in double-brooded pairs should provide relatively more parental care to broods in which they have higher paternity; secondly, after fledging and brood division, males should provide more care to related offspring; and finally, among all broods in the population, paternity should be related positively to male parental care. Brood division occurred in many of the broods studied; however, broods were not divided according to fledgling size or paternity. Furthermore, within divided broods, males fed within-pair and extra-pair fledglings at similar rates. For sequential broods of the same pair, male feeding rates were not associated with differences in paternity between broods. Among all broods in the population, males did not provide relatively less care to broods containing unrelated young. The lack of a relationship between male parental care and paternity suggests that either males cannot assess their paternity or the costs of reducing male parental care outweigh the benefits.     

Brood size and the economy of brood defence: examining Lack's hypothesis in a biparental cichlid fish

Synopsis We tested the explanatory value of two hypotheses reviewed by Lack (1954) in the maintenance of brood size in free-ranging convict cichlidsCichlasoma nigrofasciatum: (1) physiological constraints on egg production, and (2) behavioural constraints imposed by brood defence. Number of free-swimming young in 13 experimental (E) broods was augmented to the upper limit of the size distribution of natural broods (150 young); 18 control (C) broods were handled in the same way but brood size was not changed (mean ± SE = 69.5 ± 11.0). E and C brood sizes were measured at 5 day intervals. At day 20 (just before independence from parental care), 50.3 ± 9.4 (n = 9) young remained in E broods and 30.8 ± 7.8 (n = 8) young remained in C broods (p> 0.05). Offspring number did not differ significantly (p> 0.05) between C and E broods after day 10. Mean growth rate of offspring was significantly lower in E broods than in C broods, perhaps in response to increased density of young in the former. Both the convergence of offspring number in E and C broods and suppression of growth in E broods support a behavioural constraint; that during the first 10 days in which the young are free swimming, two parents are unable to defend large broods as successfully as small broods. A trade-off exists in parental investment between current and future reproduction. Extra-parental investment in current reproduction (eggs) does not result in an increased number of young at independence, therefore a behavioural constraint during brood defence should stabilize the evolution of clutch size.     

GEOGRAPHICAL PATTERNS AND POPULATION STRUCTURE IN PERIODICAL CICADAS BASED ON SPATIAL ANALYSIS OF ALLOZYME FREQUENCIES

There are twelve major broods, or year classes, of 17-year periodical cicadas and three major broods of 13-year cicadas. Previous allozyme studies of three 13-year broods and two 17-year broods suggested that broods are distinct evolutionary units, exhibiting homogeneity within broods and heterogeneity among broods. Results from numerical phylogenetic analysis of allozyme data from these broods supported Lloyd and Dybas's (1966a, 1966b) evolutionary hypothesis and were consistent with biogeographical patterns. In the present paper, we use spatial statistics to extend the analysis of biogeographical patterns for periodical cicadas by examining geographic variation in allozyme frequencies within and among ten broods for the three most variable loci. The results from our analysis show that each gene frequency surface exhibits marked spatial autocorrelation. Three major geographic areas of similarity are indicated by the data: the eastern seaboard, the Mississippi Valley, and the eastern edge of the Great Plains. When localities are adjusted for regional membership, the autocorrelation disappears. In biological terms, broods are homogeneous breeding units. Geographically distinct localities within the same brood show little differentiation. Differentiation among populations within broods and among broods within regions exists but is less pronounced than the differentiation among geographic regions. For the particular enzymes examined, allozyme frequency differences appear to reflect the phylogenetic history of the broods rather than their environmental surroundings.     

Habitat use,nest-sites and chick diet of Grey Partridge Perdix perdix on hill farms in north east England

Capsule: Breeding Grey Partridges Perdix perdix on hill farms preferred Soft Rush-infested rough grazing which provided nesting cover and invertebrates for foraging chicks.

Aims: To measure habitat use, nest sites, chick diet and breeding productivity of Grey Partridge on hill farms in north east England.

Methods: We measured actual habitat use of radio-tagged individuals in relation to its availability both within home ranges and over the wider study area. Nesting habitat and breeding productivity data were collected from radio-tagged pairs. Faecal samples from roost sites of broods were collected to assess chick diet.

Results: Grey Partridges preferred rough grazing habitats. Sixty-nine per cent of nests were found in rough grazing with females preferring to nest in tall rushes which provided cover. Chick diet differed between habitats and in relation to brood age. In rough grazing, sawfly larvae were the most numerous item eaten by young (46%) and older broods (27%).

Conclusion: On hill farms, breeding Grey Partridges preferred rough grazings, where broods preferred sawfly larvae. It is important that rush-infested rough grazings are not agriculturally improved and grazing regimes that provide abundant sawfly larvae are practised.     

Hatching asynchrony in the house wren, Troglodytes aedon: a test of the brood-reduction hypothesis

We tested the brood-reduction hypothesis by adding three nestlingsto naturally occurring synchronous and asynchronous broods ofthe house wren (Troglodytes aedon) in order to mimic food shortagesfor the broods. Two types of controls were established in whichbrood size remained unchanged: those in which nestlings wereexchanged among broods and those in which no nestlings wereexchanged. The critical test of the hypothesis was in 1988 whenthere was a food shortage for enlarged broods. Although broodreduction occurred, enlarged synchronous broods produced asmany fledglings as did enlarged asynchronous broods, and fledgingmass was similar. Juvenile recapture 2-8 weeks after fledgingand offspring recruitment to subsequent breeding populationswere not related to treatment. The results are not consistentwith the brood-reduction hypothesis as an explanation for theoccurrence of hatching asynchrony in the house wren.