Breeding success of a Lesser Spotted Woodpecker Dendrocopos minor population

We made intensive observations of 50 pairs of Lesser Spotted Woodpecker Dendrocopos minor in southern Sweden from 1989 to 1992. Between 7% and 22% of the pairs each year defended a territory but laid no eggs. Overall, 76% of the pairs laying eggs raised fledglings, but this proportion was only 33% in 1991 when the weather was cold and wet. The mean clutch size (5.9) and brood size (4.5) in successful nests did not vary significantly between years. Incubation time was 1 day longer in 1991 (11–12 days) than in 1990 and 1992 (10–11 days). Number of young fledged per nesting attempt (3.5) and per pair (2.7) in the population also varied significantly between years. The initiation of egg laying varied between years but coincided each year remarkably well with the date when oak came into leaf. Breeding success is discussed, and comparisons are made with other hole-nesting species.     

Annual and seasonal reproductive trends in the Lesser Spotted Woodpecker Dendrocopos minor

Annual and seasonal variation in reproductive timing and performance were studied in a population of the Lesser Spotted Woodpecker Dendrocopos minor over 10 years in southern Sweden. The median laying date of the first egg varied by up to 17 days between years, being generally larger than the variation of laying dates within years. Neither clutch size, brood size in successful nests, fledging success in successful nests nor mean nestling weight differed significantly between years. There was no trend for mean clutch size to vary between early and late years. In spite of a more than threefold variation in population size, no reproductive variable demonstrated an apparent density-dependence. Within the season, clutch size declined steeply with increasing clutch initiation date, whereas fledging success and nesting success did not, leading to a trend in brood size almost identical to the trend in clutch size. The survival prospects of fledged young declined with increasing clutch initiation date, and it is argued that the clutch size laid is a strategic adjustment to laying date. Out of 124 breeding attempts, 34% did not produce fledged young. In 9% of the breeding attempts, pairs laid no eggs. At least 20% of the breeding attempts failed after egg-laying. The most common cause of breeding failure was loss of the breeding partner followed by nest abandonment (40% of the failures). Only 16–28% of the failures were due to predation on the nest. Most complete failures, and also partial losses from nests, occurred at the early breeding stages. It is argued that the early nestling phase may be a critical stage, which the woodpeckers adjust to coincide with the seasonal food peak, explaining the strikingly late breeding season compared with other non-migrant species.     

Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor

The sexes’ share in parental care and the social mating system in a marked population of the single‐brooded Lesser Spotted Woodpecker Dendrocopos minor were studied in 17 woodpecker territories in southern Sweden during 10 years. The birds showed a very strong mate fidelity between years; the divorce rate was 3.4%. In monogamous pairs, the male provided more parental care than the female. The male did most of the nest building and all incubation and brooding at night. Daytime incubation and brooding were shared equally by the sexes, and biparental care at these early breeding stages is probably necessary for successful breeding. In 42% of the nests, however, though still alive the female deserted the brood the last week of the nestling period, whereas the male invariably fed until fledging and fully compensated for the absent female. Post‐fledging care could not be quantified, but was likely shared by both parents. Females who ceased feeding at the late nestling stage resumed care after fledging. We argue that the high premium on breeding with the same mate for consecutive years and the overall lower survival of females have shaped this male‐biased organisation of parental care. In the six years with best data, most social matings were monogamous, but 8.5% of the females (N=59) exhibited simultaneous multi‐nest (classical) polyandry and 2.9% of the males (N=68) exhibited multi‐nest polygyny. Polyandrous females raised 39% more young than monogamous pairs. These females invested equal amounts of parental care at all their nests, but their investment at each nest was lower than that of monogamous females. The polyandrously mated males fully compensated for this lower female investment. Polygynous males invested mainly in their primary nest and appeared to be less successful than polyandrous females. Polyandry and polygyny occurred only when the population sex ratio was biased, and due to strong intra‐sexual competition this is likely a prerequisite for polygamous mating in Lesser Spotted Woodpeckers.     

Characteristics of woods used recently and historically by Lesser Spotted Woodpeckers Dendrocopos minor in England

Lesser Spotted Woodpecker Dendrocopos minor numbers have declined greatly in England since the early 1980s for reasons that are not yet fully understood. It has been suggested that the species’ decline may be linked to the increase in Great Spotted Woodpeckers Dendrocopos major, changes in woodland habitat quality (such as deadwood abundance) and landscape‐scale changes in tree abundance. We tested some of these hypotheses by comparing the characteristics of woods in southern England where the species is still relatively numerous with those of woods used in the 1980s before the major decline. In each time period, habitat, predator and landscape information from woods known to be occupied by Lesser Spotted Woodpeckers was compared with those found to be unoccupied during surveys. Before the main period of decline, Lesser Spotted Woodpeckers used oak‐dominated, mature, open woods with a large amount of standing deadwood. Habitat use assessed from recent data was very similar, the species being present in mature, open, oak‐dominated woodlands. There was a strong relationship between wood use probability and the extent of woodland within a 3‐km radius, suggesting selection for more heavily wooded landscapes. In recent surveys, there was no difference in deadwood abundance or potential predator densities between occupied and unoccupied woods. Habitat management should focus on creating and maintaining networks of connected woodlands in areas of mature, open woods. Finer‐scale habitat selection by Lesser Spotted Woodpecker within woodlands should be assessed to aid development of beneficial management actions.     

Two cooperative breeding cases in Lesser Spotted Woodpecker Dendrocopos minor

During a study of a colour-marked population of Lesser Spotted Woodpecker (LSW) in NE Spain we found two cases of cooperative breeding, in which two females and a male were rearing the brood. One of the females was involved in both cooperative breeding cases, in two consecutive years. We analysed the feeding contribution of each individual to nestling care as well as feeding rates to nestlings. In both cases the male showed a (compensatory) reduction in nestling feeding due to the involvement of the second female. Total feeding rate increased slightly compared to regular (noncooperative) monogamous pairs. The high breeding success found in the second case (six eggs/six nestlings) is thought to be due to the cooperation of the three adults in nestling care. The two cooperative breeding cases were found in territories that suffered from significant selective logging of dead trees. This logging is thought to be a possible anthropogenic example of the ecological constraints hypothesis and a significant factor that could force atypical cooperative breeding attempts. A locally biased sex-ratio might result in the cooperation of two females and a male in both cases. No evidence of kinship was found among members of the cooperative breeding teams. We could not determine whether the cooperative breeding cases were monogamous or polygamous. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Long-term trends in the nest survival and productivity of the Lesser Spotted Woodpecker Dryobates minor in Britain

ABSTRACT

Capsule: There has been no trend in nest survival of Lesser Spotted Woodpecker Dryobates minor in the 70 years since the 1940s whereas the numbers of young per nesting attempt has declined. Late nests in any year are now significantly less productive than early ones.

Aims: To test whether there has been a long-term decline in the nest survival and productivity of the Lesser Spotted Woodpecker in Britain.

Methods: Breeding data from 331 nests over the period 1949–2019 have been analysed. There were three sources – nest records submitted to the British Trust for Ornithology, a study by the Royal Society for the Protection of Birds from 2006 to 2009 and records submitted to the Woodpecker Network from 2015 to 2019. Generalized linear models were used to analyse the records for first egg date, clutch size, numbers of young fledged, and numbers of nest days for which the nest was under observation. Data were grouped into three periods reflecting population trends of the bird (pre-1980, 1980–1999, 2000–2019) and we also analysed trends with year and spring temperature.

Results: There was no trend in first egg date up to 1980 but subsequently it has advanced by 13 days. The mean clutch was 5.3 with no trend with period or year. There were no trends in nest survival during the egg stage or chick rearing. The mean number of young fledged was 2.6, with nests since 1980 fledging lower numbers of young than those pre-1980 and a decline with year. Loss of chicks, probably to starvation, was the main cause of low productivity. In the last period (2000–2019) nests started early in the season fledged more young than those started later, a trend not apparent in the earlier periods.

Conclusions: Low productivity is a widespread problem for the Lesser Spotted Woodpecker which has probably been exacerbated by the trend towards warm springs.     

Sexual dimorphism and foraging niche partitioning in the Middle Spotted Woodpecker Dendrocopos medius

I investigated sexual differences in morphology and foraging behaviour in the nearly monomorphic Middle Spotted Woodpecker Dendrocopos medius. In the northeastern Swiss lowlands, radiotracked birds were observed from 1994 to 1996 between January and June. The sexes overlapped in all morphological characters, with males ( n = 25) being on average larger than females ( n = 17). Bill-length and length of the crimson-red crown patch were most useful to discriminate the sexes. Sex-specific differences in foraging behaviour were found both within and between the prebreeding and breeding periods concerning the use of tree species, foraging techniques, relative height zones and living and dead substrates within a tree. Overall, both sexes had broader niches in the prebreeding than in the breeding period and niche overlap was larger during the latter period. Hence, sex-specific differences were more pronounced in the prebreeding than in the breeding period, indicating different niche use of males and females in the two periods. These varying patterns of sex-specific differences imply that niche segregation results from behavioural plasticity, probably based on dominance relationships between the sexes, rather than from sexual dimorphism.     

A male destroying an egg in a cooperative breeding attempt in Lesser Spotted Woodpecker Dendrocopos minor

A cooperative breeding case was found in Lesser Spotted Woodpecker (LSW) in which a male and three different females were involved. The contributions of each member of the cooperative breeding team were quantified during nest excavation and incubation. This is the first such report on LSW. During daytime, the nest was mainly occupied by the male during the final excavation phase, nest guarding, and egg laying. Aggressive interactions were recorded between two of the females during nest excavation, egg laying and incubation. The male was recorded destroying an egg presumably laid by the dominant female. The highest contribution during daytime incubation was made by the dominant female (39%), followed by the male (34%) and then by a second female (27%). The third female contributed very little at any stage. A picture of the male removing the egg was analyzed to estimate the egg maximum width, ruling out a possible runt egg. Motivations behind the behaviour of the male destroying the egg remain unclear.     

Reproductive success and nestling diet in the Lesser Spotted Woodpecker ( Picoides minor ): the early bird gets the caterpillar

Populations of Lesser Spotted Woodpecker (Picoides minor) are decreasing in size, necessitating management strategies. However, data on the reproductive biology of this species are scarce. The aim of this study was to investigate the influence of timing of breeding and nestling diet on the reproductive success of the Lesser Spotted Woodpecker and thereby contribute to an understanding of the determinants influencing its reproductive success. During 6 study years between 1996–2003, we investigated various variables of reproductive success in a Lesser Spotted Woodpecker population in the Taunus low mountain range, Germany. We observed nestling feeding at breeding holes to assess the composition of nestling diet and how it changes during the season. Clutch size, number of fledglings as well as body mass of the nestlings declined with the start of egg laying. Pairs composed of individuals that had breed together in previous years started egg laying earlier than newly established pairs. Nestling diet consisted mainly of aphids, caterpillars, craneflies and wood-living larvae. The composition changed considerably within the breeding season in both early and late broods. However, early broods were provided with caterpillars more frequently, whereas late broods received wood-dwelling larvae more often. Our results suggest that Lesser Spotted Woodpeckers time their breeding so as to coincide with the early nestling stage with the highest availability of caterpillars, since this is the time of highest energy demand. Moreover, food availability declines during the breeding season, and the decline in reproductive performance seems to be an effect of this development.     

Breeding biology and breeding success of the Lesser Grey Shrike Lanius minor in a stable and dense population

The Lesser Grey Shrike Lanius minor is highly endangered throughout Europe, having declined markedly in abundance and range. Long-term changes in climate and agricultural practices have been identified as the main reasons for its decline. To determine which factors influence short-term changes in breeding success, we examined several aspects of its breeding biology. Our investigation revealed that our study area bears a large and stable population of this species. In 1996 and 1997, we recorded 84 and 77 breeding pairs in an area of 20 km², with an average of 4.20 and 3.85 pairs/km² respectively. Data on breeding density, clutch size and fledging success from 1989 to 1997 (excluding 1992) indicate a stable breeding population with a constant high breeding success. Reproductive success declined through the season, mainly through seasonal variation in clutch size rather than chick mortality. However, breeding success was generally high (69% and 79% of the nests produced chicks], with low hatching failure and few nest losses. The main cause of breeding failure was nest predation (at least 50% of nest losses), mainly by magpies (at least 66% of depredated nests). Although in this population the Lesser Grey Shrike tends to aggregate in clusters, breeding density had no obvious effect on breeding success and nest predation.     

The carotenoid‐based red cap of the Middle Spotted Woodpecker Dendrocopos medius reflects individual quality and territory size

Carotenoid‐based plumage ornaments have the potential to signal individual condition and health in many species of birds. However, very little is known about the function of red plumage in woodpeckers. We assessed whether the red cap displayed by both male and female Middle Spotted Woodpeckers reflects individual quality, finding that the size of the cap is sex‐dependent, whereas the brightness of the cap correlates with the body condition of an individual. Furthermore, birds with brighter caps had larger clutches, suggesting that cap coloration may be an honest signal of parental quality in woodpeckers. Interestingly, more colourful individuals also occupied smaller territories, suggesting that territory size and territory quality may be inversely related in the Middle Spotted Woodpecker.     

Relationships between patterns of habitat cover and the historical distribution of the Marsh Tit,Willow Tit and Lesser Spotted Woodpecker in Britain

Spatial analysis of remotely-sensed land cover data in conjunction with species distribution atlases can reveal large-scale relationships between animal taxa and their habitats. We investigated the historical distribution patterns of three declining woodland birds, the Marsh Tit (Poecile palustris), Willow Tit (Poecile montana) and Lesser Spotted Woodpecker (Dendrocopos minor), in relation to a parsimonious landscape metric for describing habitat availability in Britain. Bird distributions were derived from two field-based atlas surveys, conducted in 1968–1972 and 1988–1991, and used to classify areas of the landscape for each species as retained, lost or gained between atlas periods, or unoccupied in both. We used remotely-sensed land cover data from 1990 to compare percentage habitat cover between landscape areas classified by bird occupation, and regional summary data from national woodland inventories to investigate changes in habitat cover and bird distributions. Percentage habitat cover was a sufficient landscape metric to explain the distribution pattern of all three bird species; habitat cover was greatest in areas where each species was retained between atlas surveys, significantly less in areas from which species were lost, and least in areas that remained unoccupied. Reductions in Marsh Tit distribution were less in regions that showed greater increases in habitat cover, but there was no such relationship for other species. Results indicated that spatial studies could be used to infer aspects of the spatial ecology of species where field data is lacking: by comparing distribution patterns with the relatively well-studied Marsh Tit, we found support for the assumption that the Lesser Spotted Woodpecker occupies very large territories in Britain, and provided evidence that the spatial habitat requirements of the Marsh Tit could be used as a proxy for the data-poor Willow Tit. The results showed that the habitat cover required to retain each species in the landscape had increased over time, illustrating how spatial studies can be used to identify priorities for further research and suggest conservation measures for declining species, and these are discussed.     

Habitat preferences of the Syrian Woodpecker Dendrocopos syriacus in urban environments: an ambiguous effect of pollution

Capsule The occurrence of the Syrian Woodpecker Dendrocopos syriacus in an urbanized habitat is determined by the number of trees and pollutant emissions. Air pollution may weaken trees and increase the number of insects inhabiting them.

Aims To determine the habitat preferences of the Syrian Woodpecker in the highly urbanized environment of the city of Kraków.

Methods A set of 13 habitat and environmental parameters of 50 breeding territories and 50 random points were evaluated. A logistic regression was used to determine the probability of species occurrence, and Akaike's information criterion was used for model selection.

Results The number of trees, coverage of woody vegetation, total vegetation cover and level of pollutant emissions were significantly higher in Syrian Woodpecker breeding territories than in the random points. The model best explaining the probability of species occurrence consisted of four parameters: the number of trees, coverage of the total built-up area, total vegetation cover and pollutant emissions. The parameters best explaining a high probability of species occurrence were high number of trees and high pollutant emissions.

Conclusion Air pollution may weaken trees and potentially increase the number of insects inhabiting them, an important component of the Syrian Woodpecker's diet. However, the negative impact of air pollution on birds may impact on their condition. Urban habitats could, therefore, act as an ecological trap for this species.     

Compensation for predator-induced reduction in nestling feeding rate in the Great Spotted Woodpecker

Parent birds make efforts to prevent the immediate costs of predation through plastic behavioural responses to the actual predation risk, but this may incur future costs for offspring due to reduced parental care. However, the temporary nature of predator encounters suggests that nestling feeding reduced during the risky periods may be later compensated for by an increased feeding effort (the predation risk allocation hypothesis). We tested this prediction in the Great Spotted Woodpecker (Dendrocopos major) confronted with its major nest predator/competitor, the European Starling (Sturnus vulgaris). A brief encounter with a live starling was followed by a reduction in the nestling feeding rate, but the lost feedings were subsequently compensated for by an increased feeding rate. This compensatory effect was higher in older nestlings that are highly demanding in terms of energy requirements and fitness value from the parents’ perspective. Thus, birds are potentially able to respond not only to the immediate risk of nest predation by nest defence but also to compensate for the potential costs of nest defence in terms of unfulfilled nestling demands. However, data on the amount of delivered food are necessary to distinguish whether parents truly compensate for lost feeding or whether the increased feeding frequency represents a nest guarding strategy.     

Forest Management Considerations for Conservation of Black Woodpecker Dryocopus martius and White-backed Woodpecker Dendrocopos leucotos Populations in Quinto Real (Spanish Western Pyrenees)

The woodlands of Quinto Real (Quinto Real, Erreguerena and Legua Acotada) are a 3,000 hectare beech (Fagus sylvatica) forest managed by the shelterwood system applied to even-aged (regular) stands. This study analyses how forest management determines the local distribution of the white-backed woodpecker (Dendrocopos leucotos) and black woodpecker (Dryocopus martius) and its relationship with the type, structure and size of the stands used for nesting by both species, as well as their dead wood requirements. The most suitable nesting habitat of both species is the mature forest (stands of regular large final crop trees), but the size of the mature fragments and a minimum quantity of dead wood is also important.     

Population decline is accompanied by loss of genetic diversity in the Lesser Grey Shrike Lanius minor

The Lesser Grey Shrike has suffered successive declines in population size and a marked contraction of its breeding range since the early 20th century, largely because of long‐term agricultural intensification. This has resulted in a severely fragmented distribution in Western Europe, with isolated breeding nuclei in Spain, France and Italy and a more continuous distribution in Eastern Europe and Asia. Using a combination of nuclear and mitochondrial markers, we assessed the genetic structure and diversity of Lesser Grey Shrike populations from Western Europe, Central Europe and Asia. There was significant genetic differentiation among three major regional groups, one European and two Asian. Genetic diversity measures were lowest in the smallest and most marginal Spanish population. Limited genetic diversity, combined with rapid population decline, suggests the Spanish population may face extinction in the near future.