Drivers of low breeding success in the Lesser Spotted Woodpecker Dendrocopos minor in England: testing hypotheses for the decline

Capsule The breeding success of Lesser Spotted Woodpeckers Dendrocopos minor is now lower in England than previously reported and also lower than found in studies elsewhere in Europe.

Aims To quantify the breeding success and identify the causes of nest failure. To test the hypotheses that breeding success is related to aspects of food limitation and parental care, and inclement weather during the nesting period, or to interactions with Great Spotted Woodpeckers.

Methods Nests were monitored in three regions of England, recording survival and causes of failure. We measured aspects of food limitation and parental care, rainfall and Great Spotted Woodpecker interactions at nests, to explore whether there was any evidence that these factors were related to breeding success. We compared results to other studies from the UK and continental Europe.

Results Nest survival was 52%. The average number of chicks produced from successful nests was 2.8. Chick-stage daily nest survival was positively related to provisioning rates, indicating that food supply may be limiting. The most common cause of nest failure was presumed starvation of chicks after the disappearance of an adult. Some females ceased visiting nests, leaving provisioning solely to the male. This behaviour has been reported elsewhere in Europe, but in the present study males were unable to compensate fully by increasing their provisioning rates, leading to poor nest survival. Provisioning rates and chick-stage daily nest survival were negatively associated with rainfall. Nest predation by Great Spotted Woodpeckers occurred but was a less frequent cause of failure. Aggressive interactions were recorded between the two woodpecker species but these were unrelated to breeding parameters.

Conclusions Low breeding success is most probably related to food shortages in the breeding period. Simple population modelling using parameters from the present study and from published work shows that if the low productivity that we have observed is replicated throughout Britain, it would be sufficient to account for the observed population decline. However, the possibility that survival rates are also low cannot be ruled out.     

Provisioning behaviour at the nest in single-parent versus biparental nests and male versus female parents in the common redstart (<Emphasis Type="Italic">Phoenicurus phoenicurus</Emphasis>)

We analysed video-sequences of undisturbed parental provisioning behaviour on 12 nests of common redstart (Phoenicurus phoenicurus). In 4 of the 12 nests, chicks were fed by a single parent only. We compared provisioning rate of chicks, time spent on the nest and food allocation rules between nests with uniparental and biparental care and between male and female parents in biparental nests. In nests with a single parent, the frequency of feeding visits per parent was higher than in biparental nests. As a result, the rate of food provisioning of chicks was similar in uniparental and biparental nests. The food allocation rules did not differ between uniparental and biparental nests. In biparental nests, male and female provisioning behaviour was similar though with two exceptions: males had a strong preference for feeding chicks in front positions in the nest and females spent a longer time on the nest after feeding. We conclude that single common redstart parents are able to compensate fully for the absence of the other parent through increased provisioning efforts, and that in biparental nests, males and females contribute equally to the provisioning of the young.     

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.     

Immediate and delayed effects of a gale in late spring on the breeding of the Shag Phalacrocorax aristotelis

At the beginning of May 1982, a westerly gale destroyed or damaged 49% of 761 Shag Phalacrocorax aristotelis nests on the exposed west side of the Isle of May, southeast Scotland; 518 nests on the sheltered east side acted as controls against which to assess subsequent effects of the gale on Shag reproduction and recruitment.
On sites affected by the gale, 90% of pairs rebuilt their nest and laid a second clutch, on average 18 days after the loss of the first one. First-time breeders nested closer to the sea, suffered greater damage to their nests, and deserted their damaged nests more readily after the gale than did experienced ones; a quarter of potential recruits to the west side were deterred from breeding in that year.
By the end of the season, the number of chicks fledged per pair was 31% lower on the west side than on the east. The difference was caused partly by nest desertion, partly by greater hatching failure of full clutches in apparently undamaged nests and especially by lower fledging success of pairs that renested. Irrespective of breeding experience, delayed early breeders produced more chicks than late breeders nesting at the same time, showing that late breeders were of poorer intrinsic quality than early breeders. This was one reason for a seasonal decline in Shag productivity, although environmental factors also played a role.
During April-July, gales like that of May 1982 occur on average once every 5 years on the Isle of May. Despite the disruption that they may cause to nesting Shags, the impact on the long-term growth of the colony is small.     

Golden Plover Pluvialis apricaria breeding success on a moor managed for shooting Red Grouse Lagopus lagopus

Capsule The best estimate of breeding success was a mean of 0.57 fledglings per pair, which when combined with adult survival rates, successfully explained the observed population trend.

Aims To quantify Golden Plover breeding success on a moor managed for shooting Red Grouse Lagopus lagopus.

Methods An intensive study recorded the fate of individual Golden Plover nests and, using radiotelemetry, chicks. The factors associated with mortality were examined, allowing the construction of a model of breeding success. Adult survival was estimated from return rates of colour-ringed birds.

Results Estimated rates of daily nest survival during laying (0.8636) were significantly lower than during incubation (0.9913). The daily survival rate of chicks less than nine days (0.8868) was significantly lower than for older chicks (0.9792). A population model based on these parameters overestimated the rate of nest losses, but accurately described brood survival and fledging success. Although predation rates were low, poor survival of young chicks through starvation or exposure suggest other factors were limiting breeding success at the study site.

Conclusions Predation rates of Golden Plover nests and chicks can be low on moorlands managed for shooting Red Grouse. However, in the absence of predation, other factors may still reduce chick survival and limit breeding success.     

Proceedings of the New Zealand Society for Parasitology Inc.

Abstract

A study of the nesting habits and breeding biology of blue penguin Eudyptula minor was undertaken over the 1995–96 and 1996–97 breeding seasons on Matiu‐Somes Island, Wellington, New Zealand. Male and female blue penguins tended to be faithful to both mates and nest sites, although there was insufficient evidence to detect any association between a bird's breeding success in 1995 and a subsequent change of mate or nest in 1996. Over the 1995 and 1996 seasons the recorded hatching success (0.51 ±0.11 and 0.63 ± 0.10 respectively), fledging success (0.81 ±0.12 and 0.85 ±0.10 respectively) and reproductive success (0.41 + 0.11 and 0.54 ± 0.11 respectively) were similar each season. There was no significant difference between the proportion of eggs laid, or eggs hatched and chicks fledged, between the two seasons. The mean number of chicks raised over the two seasons was 0.94 ± 0.05 per nest. Replacement clutches were laid by 11 per cent of failed breeders in each season, but only in 1996 were they successful in fledging chicks.

No significant difference was found between the breeding success of the Matiu‐Somes Island blue penguin colony recorded during this study and a previous study undertaken on the island 40 years ago.     

Nest-site selection and characteristics in a mixed-species colony of Avocets Recurvirostra avosetta and Black-winged Stilts Himantopus himantopus

Capsule Causes of breeding failure determined optimal nest location, Black-winged Stilts being better adapted than Avocets to nest close to water.

Aims To compare nest size and location between the two species and to test whether there were inter-species differences that might suggest specific nesting adaptations, and to investigate possible correlates between hatching success and nest location.

Methods Nest-site selection, characteristics and hatching success were studied during 1989 in a large mixed-species colony located in southern Spain.

Results Black-winged Stilt nests differed in composition and size depending on distance to water (nests touching the water were larger and included mud), but Avocet nests did not. Avocets nests were more aggregated and central within the colony than those of Black-winged Stilts. Causes of breeding failure were flooding and rat predation. Successful or unsuccessful Black-winged Stilt nests did not differ significantly in characteristics or location. Successful Avocet nests were further from water than flooded nests and further from dry land than depredated ones. Distance to the centre of the colony did not affect hatching success.

Conclusion Black-winged Stilts were better adapted than Avocets to nest near water, due to their greater plasticity in nest-building behaviour. Causes of breeding failure determined optimal nest location.     

Effect of nest-site microclimatic conditions on nesting success in the Lesser Kestrel Falco naumanni

Capsule: Microclimatic conditions in the nests of the Lesser Kestrel Falco naumanni, particularly the percentage of time of extremely low humidity, affect breeding success.

Aim: To study the effect of within-nest temperature and humidity on nest productivity, and the correlation between nest productivity and the order of dates on which nests were occupied by the parents. To compare microclimatic conditions in the nest, breeding success and order of occupation between nests under tile roofs and in artificial nest boxes.

Methods: Three different Lesser Kestrel colonies were monitored in Israel – one rural, one urban and one in an open country habitat. Data loggers were placed in 39 nests for the entire breeding period to measure temperature and humidity. The number of fledglings was recorded for each nest, as well as the date of occupation.

Results: Full microclimatic data from 35 nests suggest that percentage of time of extremely low humidity is the major predictor of nest productivity – low humidity is negatively correlated with nest productivity. Sites of more successful nests were occupied earlier. Considering only successful nests, the urban colony had the lowest breeding success of the three colonies. There was no significant difference in mean productivity between nests in roofs and nest boxes, but nests in roofs were occupied earlier.

Conclusion: Nest microclimate, particularly low humidity, affects nesting success in addition to colony location.     

Reproductive measures and chick provisioning of Cory's Shearwater Calonectris diomedea borealis in the Azores

Capsule Timing of breeding influenced wing-length at fledging, and egg size may be an indicator of fledging weight and the amount of food received by chicks.

Aims To investigate chick growth, temporal patterns of chick food provisioning and the importance of indices of parental condition or quality, egg size and hatching date, to predict nestling body mass and wing-length at fledging, and compare breeding and chick feeding characteristics between colonies in the northeast Atlantic.

Methods A survey of Cory's Shearwater nests was carried out at Vila islet. A sample of 52 chicks, ringed and weighed at hatching, was selected to study chick growth and food provisioning.

Results Hatching success (51%) was much lower than fledging success (87%). Both hatching date and egg size contributed to explain wing-length at fledging, but hatching date, which was negatively correlated with wing-length at fledging, had the most important contribution (22%). There was some indication that egg size may explain variation in fledging weight and the amount of food received by chicks. Food delivery and feeding frequency of chicks varied throughout the chick development stage and three phases were distinguished: (1) 0–29 days, the highest feeding frequency values and a linear increase in food delivery; (2) 30–69 days, an oscillation in food delivery and medium feeding frequencies; (3) 70–90+ days, a sharp decrease in both food delivery and feeding frequency.

Conclusion Variation in food availability did not seem sufficient to override the overall importance of indices of parental quality in determining reproductive measures and chick provisioning. Breeding and feeding characteristics were similar between colonies in the northeast Atlantic, with variability in chick provisioning higher further south.     

Nestbox provisioning in a rural population of Eurasian Kestrels: breeding performance,nest predation and parasitism

The breeding biology of the Eurasian Kestrel Falco tinnunculusin nestboxes in farmland was studied to test for differences between artificial and natural sites. We report on the direct effect of nestbox provisioning on some life-history traits and how nestbox use affects nest predation and parasitism. Five types of nest-sites were available: nestboxes on poles and trees (artificial sites), stick nests on trees, stick nests on pylons and holes in buildings (‘natural’ sites). The Kestrel population increased from 23 pairs in 1993 (prior to nestbox installation) to 55 in 1998 as nestboxes were provided. In general, pairs breeding in trees started to lay later than those nesting in nestboxes on poles or in building holes, but this difference was probably associated with habitat quality rather than nest type. Differences in clutch size were found between nest-sites in some years, and were associated with laying date and, probably, with variation in territory quality. Using only data from successful nests, pairs breeding in nestboxes produce more fledglings than those in building holes or pylons. The frequency of nest predation was higher in natural sites than in nestboxes. The number of fledglings from pairs breeding in nestboxes was higher than from those breeding in old stick nests in trees when all nests were considered. Nestbox provisioning had no effect on the occurrence of the ectoparasite Carnus hemapterus, but chicks from nestboxes showed higher intensity of infection. Our results suggest that nestbox provisioning increases reproductive success and the frequency of nest predation or intensity of parasite infestation in Kestrels.     

Parental behaviour of Spanish Imperial Eagles Aquila adalberti: sexual differences in a moderately dimorphic raptor

Capsule The sexes make significantly different contributions in the tasks of nest-building, incubation and food provisioning to chicks.

Aims To determine the division of parental activities during breeding (nest-building, incubation, brooding, food provisioning and feeding).

Methods Between 1991 and 1998 focal observations were made at nests using 20–60× telescopes. During the incubation period, 11 pairs were studied involving a total of 2812 observation hours over 269 days. During the chick-rearing period, seven pairs were studied involving a total of 5499 observation hours spread over 503 days. The birds were sexed and identified individually on the basis of size, position during copulation, vocalization and plumage differences.

Results Significant differences were found between the sexes in parental behaviour. Males scarcely participated in incubation and contributed significantly more than the females only in food provisioning, both during incubation (food transfers to feed the female) and during chick-rearing (to feed the chicks). Females were responsible for most of the incubation, nest-building (during incubation and chick-rearing), brooding, shading and feeding of chicks. Nest attendance by females decreased over time. In both sexes, food provisioning increased with brood size.

Conclusion Intersexual differences are discussed in the context of the reversed sexual size dimorphism and parental investment strategies. We suggest that in Spanish Imperial Eagles reversed sexual size dimorphism is best explained by the prey capture difficulty hypothesis, than by parental role division during reproduction.     

Chick survival in relation to nest site: is the Antarctic petrel hiding from its predator?

In birds, the characteristics of the nest site may affect reproductive success. We found that shelter is an important characteristic of the Antarctic petrel (Thalassoica antarctica) nests because shelter prevents chick predation. However, the benefit of shelter was countervailed by melt water which mainly entered well-sheltered nests. Chick survival was monitored until the chick was left unattended for the first time. Late-hatched chicks had a higher survival probability than early-hatched chicks, possibly because late hatchers swamp the predator, the south polar skua (Catharacta maccormicki). Poorly sheltered nests tended to be occupied by parents with low body mass and late-hatched eggs. The results suggest that both shelter per se and parental characteristics may explain the relationship between predation risk and shelter. We need experiments to study the influence of nest site on reproductive success, and we need to map the frequency of melt water as a cause of reproductive failure.     

DO NEST SITE CHARACTERISTICS AFFECT THE BREEDING SUCCESS OF RED BISHOPS EUPLECTES ORIX?

Summary

Ferguson, J.W.H. 1994. Do nest site characteristics affect the breeding success of Red Bishops Euplectes orix? Ostrich 65:274-280.

A colony of Red Bishop birds Euplectes orix in a reed bed near Pretoria was studied during two breeding seasons. Breeding activity was greater during the second year, following increased rainfall. Predation was the most important source of mortality with 25–30% of the nests yielding fledglings. Between 7 and 11% of the nests were parasitised by Diederik Cuckoos. Reed height and distance from the edge of the reed bed were strongly correlated with nest density. From a temporal perspective breeding success was highest in late January following the peak in breeding activity. These observations suggest that overall breeding success is higher when nest densities are low. The mean fledging rate per nest was lower in areas of high nest density. This was, however, a statistical artifact. Multivariate analyses could not identify any environmental factors as predictors of breeding success within a colony.     

Alternative Reproductive Strategies of a Gregarious Ground-Nesting Bee, Dieunomia triangulifera (Hymenoptera: Halictidae)

I studied reproductive behavior of females of the gregarious ground-nesting bee Dieunomia triangulifera (Halictidae). During peak bloom of the host plant, Helianthus annuus, some females consistently brought pollen to one nest (provisioning bees), whereas others visited many nests without taking pollen to any (searching bees). Searching bees were more likely to have two or more developed oocytes and crops full of pollen. The ingested pollen probably provided protein for egg production. The differing behavior of provisioning and searching bees combined with contrasting internal morphology, indicated that these two sets of behaviors were alternative reproductive strategies. Three possible explanations for the searching bees' behavior included: usurpation, floating, or intraspecific cleptoparasitism. Each of these is evaluated as a possible function of searching behavior. Given the evidence I conclude that intraspecific cleptoparasitism is the most likely explanation.     

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.     

New is not always better: low breeding success and different occupancy patterns in newly built nests of a long-lived species,the white stork Ciconia ciconia

Capsule An increase in new nest building in a white stork population revealed that they were built further from human settlement and on non-typical structures; such nests had lower breeding success resulting from later breeding.

Aim To determine why some birds build new nests rather than occupy older ones, and how new nests affect breeding performance compared to old nests, in a long-lived bird, the white stork.

Methods We compared new nest construction in 2010 with a long-term data set on white stork in Western Poland from 1974 to 2009. For data from 2010, we analysed nest location and breeding biology in detail.

Results Since 1974, the proportion of new build nests was ca. 1.6%; in 2010 this was 13.2%. Pairs in new nests bred later than pairs in old, and had smaller clutches and lower breeding success. New nests were located further from settlements and tended to be built on different structures. A significantly lower proportion of new nests were re-occupied in subsequent years.

Conclusions Pairs may build new nests to gain experience in nest building, cooperation and foraging for subsequent seasons or because of competitive pressure when the environment is close to carrying capacity. Breeding success can be initially very low.     

Blue penguin (Eudyptula minor) nest distribution and breeding success on Otago Peninsula, 1992 to 1998

Abstract

Annual counts of nests with eggs or chicks (known nests) were made at blue penguin (Eudyptula minor) breeding sites on the Otago Peninsula in each November from 1994 to 1997. Although the population has doubled to an estimated 600 known nests over this period, the number of breeding sites on the Otago Peninsula has reduced since the 1970s. Breeding success at three areas at Taiaroa Head were monitored by regular nest checks in the breeding season from 1992 to 1998. At Taiaroa Head reproductive success ranged from 41 to 78% at the three sites during the seven‐year study and was generally higher for pairs nesting in nest boxes than for those in burrows. The percentage of breeding pairs that laid a second clutch after fledging at least one chick from their first clutch (double brooded) varied between seasons (0–48%) and was correlated with the date of the onset of breeding. Egg loss, possibly through predation by Norway rats (Rattus norvegicus), influenced the significantly lower reproductive success at one area (Area A) at Taiaroa Head during the 1996 season.     

Effects of extra-pair paternity and maternity on the provisioning strategies of the Azure-winged Magpie Cyanopica cyanus

Altricial birds show enormous intraspecific diversity in their provisioning strategies, in terms of both the provisioning rate and the amount of food delivered per feeding bout. Extra-pair copulations (EPCs), which result in either extra-pair paternity (EPP) or maternity (EPM), provide an opportunity to demonstrate why provisioning strategies vary among individuals. Because EPP-cuckolded males and EPM-cuckolded females must raise unrelated young, whereas EPM-cuckolded males and EPP-cuckolded females need not, we hypothesized that the first two categories of breeders would reduce parental effort, whereas the latter two categories of breeders would increase parental effort. We tested this hypothesis in the Azure-winged Magpie Cyanopica cyanus by comparing the number and body mass of fledglings, provisioning rates and food amount delivered per feeding bout between EPP- and EPM-cuckolded breeders and faithful breeders. We found that (1) the number of fledglings did not differ significantly between cuckolded and faithful breeders, and (2) fledglings raised by EPM-cuckolded males and EPP-cuckolded females did not differ from faithful breeders, whereas fledglings raised by EPP-cuckolded males and EPM-cuckolded females were significantly smaller than those raised by faithful breeders. Compared with faithful breeders, cuckolded breeders increased parental efforts in nests that contained no unrelated young; hence, their loss in parentage may be compensated for by the enhanced quality of fledglings. In nests that contained unrelated young, cuckolded breeders did not reduce parental efforts; hence, their own offspring would not starve and could survive in competition with mixed brood-mates. Our findings suggest that the differences in parentage created by EPP and EPM scenarios and the potential fitness return of raising a brood for the cuckolded breeders can explain the intraspecific variation in provisioning strategies of altricial birds.     

Intraspecific kleptoparasitism improves chick growth and reproductive output in Common Terns Sterna hirundo

Kleptoparasitism is a well‐known foraging strategy used opportunistically by many seabirds. Here, we investigated the effect of intraspecific kleptoparasitism on chick growth and reproductive output in Common Terns Sterna hirundo. Effects were compared between two groups comprising (1) individuals using kleptoparasitism during the chick‐rearing period (kleptoparasitic group, n = 18), and (2) individuals in pairs that never performed kleptoparasitism throughout the season (‘honest’ group, n = 21). The null models best described variation in mass at day 3 and the pre‐fledging mass, indicating no significant effect of the explanatory variables. However, the best models describing the linear growth rate (days 3–13) and peak mass included the parents' foraging strategy (kleptoparasitic vs. honest parents) as an explanatory variable. These two growth parameters were higher in chicks of kleptoparasitic parents. Kleptoparasitic foraging strategy was also associated with higher pre‐fledging survival, as the reproductive performance (i.e. number of fledglings) was significantly higher in the kleptoparasitic than in the honest group. We suggest that by stealing food (and consequently feeding offspring more frequently with high‐quality prey), kleptoparasitic parents are able to produce higher quality chicks with enhanced survival.     

Effects of nest size and dispersion on brood production in a North American population of wood ant Formica fusca (Hymenoptera: Formicidae)

We examined several key parameters of the population ecology of a North American population of Formica fusca (L.), including nest dispersion, colony size and brood production. Physical nest size was significantly correlated with colony size, and colony size, in turn, was significantly correlated with brood production. Sex allocation was male biased, although larger nests were more likely to produce reproductive female brood (gynes). Neither nest temperature nor moisture level was significantly correlated with brood production. Formica fusca nests in this population had a comparatively low average nearest-neighbor distance with a significantly even pattern of dispersion, which suggests relatively high intraspecific competition. However, nearest-neighbor distance was not significantly associated with either colony size or relative brood production.