Decaying trees improve nesting opportunities for cavity‐nesting birds in temperate and boreal forests: A meta‐analysis and implications for retention forestry

Many studies have dealt with the habitat requirements of cavity‐nesting birds, but there is no meta‐analysis on the subject and individual study results remain vague or contradictory. We conducted a meta‐analysis to increase the available evidence for nest‐site selection of cavity‐nesting birds. Literature was searched in Web of Science and Google Scholar and included studies that provide data on the habitat requirements of cavity‐nesting birds in temperate and boreal forests of varying naturalness. To compare nest and non‐nest‐tree characteristics, the following data were collected from the literature: diameter at breast height (DBH) and its standard deviation (SD), sample size of trees with and without active nest, amount of nest and available trees described as dead or with a broken crown, and amount of nest and available trees that were lacking these characteristics. Further collected data included bird species nesting in the cavities and nest‐building type (nonexcavator/excavator), forest type (coniferous/deciduous/mixed), biome (temperate/boreal), and naturalness (managed/natural). From these data, three effect sizes were calculated that describe potential nest trees in terms of DBH, vital status (dead/alive), and crown status (broken/intact). These tree characteristics can be easily recognized by foresters. The results show that on average large‐diameter trees, dead trees, and trees with broken crowns were selected for nesting. The magnitude of this effect varied depending primarily on bird species and the explanatory variables forest type and naturalness. Biome had lowest influence (indicated by ΔAIC). We conclude that diameter at breast height, vitality, and crown status can be used as tree characteristics for the selection of trees that should be retained in selectively harvested forests.     

Nest sites as limiting resources for cavity‐nesting birds in mature forest ecosystems: a review of the evidence

ABSTRACT Assumptions that populations of cavity‐nesting birds are limited by access to nest sites have largely been based on anecdotal reports or correlative data. Nest‐box‐addition experiments or tree‐cavity‐blocking experiments are potentially rigorous ways to investigate how densities of breeding birds are affected by access to nest cavities. Experimental evidence indicates that natural tree holes are limited in human‐altered landscapes, but the possibility that cavity nests are limited in old growth (unmanaged) forests is less clear. I reviewed 31 nest‐cavity‐removal or addition experiments conducted with 20 species of cavity‐nesting birds in mature forests. Of these 31 experiments conducted with a variety of different species of birds, only 19% reported statistically significant changes in breeding densities. However, none of these studies included data about the reproductive history of individuals colonizing the boxes (i.e., whether birds using the boxes would have otherwise been floaters or that birds excluded from blocked cavities on the plots did not simply move elsewhere), so they provided no strong evidence that the number of breeding pairs was limited by availability of nest sites at the population scale. Although some studies indicate that nest sites are limited at local (plot) scales in old growth forests, there is still little empirical evidence for nest‐site limitation at the population‐ and landscape‐level in mature, unmanaged forests. I review the challenges in designing and interpreting box‐addition experiments and highlight the main gaps in knowledge that should be targeted in the future.     

A novel nest‐monitoring camera system using a Raspberry Pi micro‐computer

The utility, availability, cost‐effectiveness, and reliability of prefabricated video systems designed to monitor wildlife have lagged behind the unique and varied needs of many researchers. Many systems are limited by inflexible video settings, lack of adequate data storage, and cannot be programmed by the user. More sophisticated systems can be cost prohibitive, and the literature describing remote wildlife video monitoring has, for the most part, not incorporated advances in camera and computer technology. Here, we present details of a pilot study to design and construct a lower cost (US $340) nest camera system to record the behavior of Acorn Woodpeckers (Melanerpes formicivorus) in artificial tree cavity nests. This system incorporates a Raspberry Pi micro‐computer, Pi NoIR infrared camera, a wireless adapter to transmit video over the Internet, and Deka rechargeable gel batteries for power. We programmed the system to motion‐sense, to record exclusively during daylight hours, and to automatically upload videos to the cloud over wireless Internet. The Raspberry Pi micro‐computer does not require advanced programming or electrical engineering skills to build and configure and, because it is programmable, provides unprecedented flexibility for field researchers who wish to configure the system to the specific needs of their study.     

Experimental pyrethroid treatment underestimates the effects of ectoparasites in cavity‐nesting birds due to toxicity

Nest‐dwelling ectoparasites may result in costs for nestlings of cavity nesters in terms of compromised growth and condition before fledging. The reduction or elimination of nest ectoparasites to study their effects on avian hosts can be conducted through physical methods such as heat‐treatment or through chemical methods using insecticides. Pyrethroids are the most frequently used of the latter, although some studies have shown that they may compromise the development and future survival of birds. In this study conducted in central Spain we analysed the differences between a group of fumigated Pied Flycatcher Ficedula hypoleuca nests and a heat‐treated group, both rendered ectoparasite‐free by these treatments. We also compared these ectoparasite‐free nests with a control group with natural ectoparasite loads. Our aim was to test the possible effects of a pyrethroid‐based insecticide on reproductive success, parental care behaviours and body condition of adult females and nestlings. We also determined the effects of treatment on a biochemical biomarker, the total glutathione (tGSH) level, involved in detoxification of xenobiotics and considered the most important intracellular antioxidant. Although behavioural variables were not affected by treatment, results showed lighter 3‐day‐old chicks and shorter tarsi and wings in nestlings shortly before fledging in fumigated nests, together with depletion of tGSH levels in both females and nestlings. Fumigation with pyrethroids in ectoparasite load reduction experiments may introduce undesired systematic variability associated with toxicity, leading to underestimation of the effects of ectoparasites on avian hosts.     

Observations on the breeding behaviour of the Stripe‐breasted tit (Parus fasciiventer) in Bwindi Impenetrable National Park,Uganda

The motivation of this study was to investigate some hitherto unknown information on the breeding ecology of the Stripe‐breasted Tit (Parus fasciiventer) in Bwindi Impenetrable National Park, south‐western Uganda. Parus fasciiventer is one of the least studied and endemic bird species restricted to the montane forests of the Albertine Rift. Regionally, it is classified as near‐threatened. The study was carried out around the Institute of Tropical Forest Conservation Ruhija camp and the period of study was from January to June 2003. Data were generated through direct observation at the nest box sites of three active nests. Each of the nest boxes was monitored from the time of nest building to the time the chicks fledged. Results and comparative assessments from this study demonstrate that P. fasciiventer, compared with its temperate congeners like Great Tits (Parus major), Marsh Tits (Parus palustris), Crested Tits (Parus cristatus), Coal Tits (Parus ater) and Blue Tits (Parus caeruleus), raised small broods and had longer nestling period. The findings further revealed that the species is capable of raising more than one brood in a single breeding season and provide further evidence that it is a cooperative breeder. Parents participated equally in raising the young, an indication of pure parenting in the species.     

Egg concealment is an antipredatory strategy in a cavity‐nesting bird

Birds have developed different behavioural strategies to reduce the risk of predation during the breeding period. Bird species that nest in the open often cover their eggs to decrease the risk of predators detecting the clutches. However, in cavity nesters, the potential functions of egg covering have not been explored despite some bird species that nest in cavities also covering their eggs as open nesters do. We analysed whether egg covering is an antipredatory behaviour in the blue tit (Cyanistes caeruleus). We simulated an increase in the perceived risk of predation at experimental nests by adding predator scent inside the nest boxes during the egg‐laying period, whilst adding lemon essence or water to control nest boxes. Birds exposed to predator chemical cues in the nest of experimental pairs more frequently covered their eggs than birds exposed to an odorous control. These results suggest that egg covering may have evolved as an antipredatory behaviour also in cavity nesters to reduce the risk of egg predation and thus increase reproductive success in birds.     

Sex-specific provisioning behaviour in a monomorphic seabird with a bimodal foraging strategy

Sexual differences in foraging and provisioning behaviour have been observed in several size-dimorphic seabird species. These differences are usually thought to be driven by size-related mechanisms such as the ability to compete for food or defend the nest. However, recent studies on monomorphic species suggest that sexual differences in foraging may arise independently of size. Selective forces driving sex-specific patterns are poorly known but essential to understand parental strategies. In this study, we examine sex differences in the provisioning behaviour of a monomorphic species, the Little Auk Alle alle . Using automated recording systems during two consecutive seasons at two colonies, we found that both sexes used a bimodal foraging strategy in which they regularly alternated single foraging trips of long duration with a cycle of several short-trips. The duration of long-trips was substantially longer in females than in males, and the sexes differed in the number of short-trips they performed in between long-trips, resulting in male-biased provisioning rates in both years. In species with a bimodal foraging strategy, long-trips have been interpreted as self-feeding trips to replenish body reserves. Our results therefore suggest that female Little Auks allocate more time to self-maintenance at the cost of chick provisioning, possibly due to different energetic constraints of the sexes prior and/or subsequent to chick-rearing. Our findings contribute to accumulating evidence that sex-specific foraging patterns may be widespread in sexually size-monomorphic seabird species.     

Chick‐provisioning behaviour of two shearwaters breeding in south‐eastern Australia

The chick‐provisioning behaviour of the short‐tailed shearwater Puffinus tenuirostris and the wedge‐tailed shearwater Puffinus pacificus was investigated in a mixed colony on Montague Island, New South Wales, Australia, over two breeding seasons. This colony is located at opposite edges of the breeding distribution of the two species. Frequent weighing techniques were used to determine chick feeding frequency, feed timing, meal size, chick weight loss and indices of food conversion efficiency of the chicks. Short‐tailed shearwater parents fed their chicks larger more infrequent meals than wedge‐tailed shearwater parents. Short‐tailed shearwater chicks demonstrated higher food conversion efficiencies and lower weight loss than wedge‐tailed shearwater chicks, indicating either differences in diet or metabolic rates. The feeding frequency in wedge‐tailed shearwaters also fluctuated more widely than for short‐tailed shearwaters over the two breeding seasons. Despite the fact that the timing of the breeding cycle on Montague Island is almost identical for the two species, these differences in chick provisioning are probably a result of differences in prey type and location, so they may help explain variations in annual breeding success and limits to the distribution of the two species.     

Nonmotorized recreation and motorized recreation in shrub‐steppe habitats affects behavior and reproduction of golden eagles (Aquila chrysaetos)

Different forms of outdoor recreation have different spatiotemporal activity patterns that may have interactive or cumulative effects on wildlife through human disturbance, physical habitat change, or both. In western North America, shrub‐steppe habitats near urban areas are popular sites for motorized recreation and nonmotorized recreation and can provide important habitat for protected species, including golden eagles. Our objective was to determine whether recreation use (i.e., number of recreationists) or recreation features (e.g., trails or campsites) predicted golden eagle territory occupancy, egg‐laying, or the probability a breeding attempt resulted in ≥1 offspring (nest survival). We monitored egg‐laying, hatching and fledging success, eagle behavior, and recreation activity within 23 eagle territories near Boise, Idaho, USA. Territories with more off‐road vehicle (ORV) use were less likely to be occupied than territories with less ORV use (β = ?1.6, 85% CI: ?2.8 to ?0.8). At occupied territories, early season pedestrian use (β = ?1.6, 85% CI: ?3.8 to ?0.2) and other nonmotorized use (β = ?3.6, 85% CI: ?10.7 to ?0.3) reduced the probability of egg‐laying. At territories where eagles laid eggs, short, interval‐specific peaks in ORV use were associated with decreased nest survival (β = ?0.5, 85% CI: ?0.8 to ?0.2). Pedestrians, who often arrived near eagle nests via motorized vehicles, were associated with reduced nest attendance (β = ?11.9, 85% CI: ?19.2 to ?4.5), an important predictor of nest survival. Multiple forms of recreation may have cumulative effects on local populations by reducing occupancy at otherwise suitable territories, decreasing breeding attempts, and causing nesting failure. Seasonal no‐stopping zones for motorized vehicles may be an alternative to trail closures for managing disturbance. This study demonstrates the importance of considering human disturbance across different parts of the annual cycle, particularly where multiple forms of recreation have varying spatiotemporal use patterns that create human–wildlife interactions.     

Selection of Nest Trees by Cavity‐nesting Birds in the Neotropical Atlantic Forest

One of the five most important global biodiversity hotspots, the Neotropical Atlantic forest supports a diverse community of birds that nest in tree cavities. Cavity‐nesting birds may be particularly sensitive to forestry and agricultural practices that remove potential nest trees; however, there have been few efforts to determine what constitutes a potential nest tree in Neotropical forests. We aimed to determine the characteristics of trees and cavities used in nesting by excavators (species that excavate their own nest cavity) and secondary cavity‐nesters (species that rely on existing cavities), and to identify the characteristics of trees most likely to contain suitable cavities in the Atlantic forest of Argentina. We used univariate analyses and conditional logistic regression models to compare characteristics of nest trees paired with unused trees found over three breeding seasons (2006–2008). Excavators selected dead or unhealthy trees. Secondary cavity‐nesters primarily selected cavities that were deep and high on the tree, using live and dead cavity‐bearing trees in proportion to their availability. Nonexcavated cavities suitable for birds occurred primarily in live trees. They were most likely to develop in large‐diameter trees, especially grapia Apuleia leiocarpa and trees in co‐dominant or suppressed crown classes. To conserve cavity‐nesting birds of the Atlantic forest, we recommend a combination of policies, economic assistance, environmental education, and technical support for forest managers and small‐scale farmers, to maintain large healthy and unhealthy trees in commercial logging operations and on farms.     

Nest‐site selection by Slender‐billed Parakeets in a Chilean agricultural‐forest mosaic

ABSTRACT Species in the family Psittacidae may be particularly vulnerable to anthropogenic habitat transformations that reduce availability of suitable breeding sites at different spatial scales. In southern Chile, loss of native forest cover due to agricultural conversion may impact populations of Slender‐billed Parakeets (Enicognathus leptorhynchus), endemic secondary cavity‐nesting psittacids. Our objective was to assess nest‐site selection by Slender‐billed Parakeets in an agricultural‐forest mosaic of southern Chile at two spatial scales: nest trees and the habitat surrounding those trees. During the 2008–2009 breeding seasons, we identified nest sites (N= 31) by observing parakeet behavior and using information provided by local residents. Most (29/31) nests were in mature Nothofagus obliqua trees. By comparing trees used for nesting with randomly selected, unused trees, we found that the probability of a tree being selected as a nest site was positively related to the number of cavity entrances, less dead crown, and more basal injuries (e.g., fire scars). At the nesting‐habitat scale, nest site selection was positively associated with the extent of basal injuries and number of cavity entrances in trees within 50 m of nest trees. These variables are likely important because they allow nesting parakeets to minimize cavity search times in potential nesting areas, thereby reducing energetic demands and potential exposure to predators. Slender‐billed Parakeets may thus use a hierarchical process to select nest sites; after a habitat patch is chosen, parakeets may then inspect individual trees in search of a suitable nest site. Effective strategies to ensure persistence of Slender‐billed Parakeets in agricultural‐forest mosaics should include preservation of both individual and groups of scattered mature trees.     

Strong influences of a dominant,ground‐nesting ant on recruitment,and establishment of ant colonies and communities

Many factors drive the organization of communities including environmental factors, dispersal abilities, and competition. In particular, ant communities have high levels of interspecific competition and dominance that may affect community assembly processes. We used a combination of surveys and nest supplementation experiments to examine effects of a dominant ground‐nesting ant (Pheidole synanthropica) on (1) arboreal twig‐nesting, (2) ground‐foraging, and (3) coffee‐foraging ant communities in coffee agroecosystems. We surveyed these communities in high‐ and low‐density areas of P. synanthropica over 2 years. To test for effects on twig ant recruitment, we placed artificial nesting resources on coffee plants in areas with and without P. synanthropica. The first sampling period revealed differences in ant species composition on the ground, in coffee plants, and artificial nests between high‐ and low‐density sites of P. synanthropica. High‐density sites also had significantly lower recruitment of twig ants and had species‐specific effects on twig ant species. Prior to the second survey period, abundance of P. synanthropica declined in the high‐density sites, such that P. synanthropica densities no longer differed. Subsequent sampling revealed no difference in total recruitment of twig ants to artificial nests between treatments. Likewise, surveys of ground and coffee ants no longer showed significant differences in community composition. The results from the first experimental period, followed by survey results after the decline in P. synanthropica densities suggest that dominant ants can drive community assembly via both recruitment and establishment of colonies within the community.     

Trans‐generational effects on ageing in a wild bird population

Ageing, long thought to be too infrequent to study effectively in natural populations, has recently been shown to be ubiquitous, even in the wild. A major challenge now is to explain variation in the rates of ageing within populations. Here, using 49 years of data from a population of great tits (Parus major), we show that offspring life‐history trajectories vary with maternal age. Offspring hatched from older mothers perform better early in life, but suffer from an earlier onset, and stronger rate, of reproductive senescence later in life. Offspring reproductive lifespan is, however, unaffected by maternal age, and the different life‐history trajectories result in a similar fitness payoff, measured as lifetime reproductive success. This study therefore identifies maternal age as a new factor underlying variation in rates of ageing, and, given the delayed trans‐generational nature of this effect, poses the question as to proximate mechanisms linking age‐effects across generations.     

Evaluating seasonal patterns of female aggression: Case study in a cavity‐nesting bird with intense female–female competition

Seasonal plasticity in aggression is likely to be shaped by the contexts in which aggression is beneficial, as well as the constraints inherent in its underlying mechanisms. In males, seasonal plasticity in testosterone (T) secretion is thought to underlie seasonal plasticity in conspecific aggression, but it is less clear how and why female aggression may vary across different breeding stages. Here, we integrate functional and mechanistic perspectives to begin to explore seasonal patterns of conspecific aggression in female tree swallows (Tachycineta bicolor), a songbird with intense female–female competition and T‐mediated aggression. Female tree swallows elevate T levels during early breeding stages, coinciding with competition for nest boxes, after which time T levels are roughly halved. However, females need to defend ownership of their nesting territory throughout the breeding season, suggesting it may be adaptive to maintain aggressive capabilities, despite low T levels. We performed simulated territorial intrusions using 3D‐printed decoys of female tree swallows to determine how their aggressive response to a simulated intrusion changes across the breeding season. First, we found that 3D‐printed decoys produce data comparable to stage‐matched studies using live decoys, providing researchers with a new, more economical method of decoy construction. Further, female aggressiveness remained relatively high through incubation, a period of time when T levels are quite low, suggesting that other mechanisms may regulate conspecific female aggression during parental periods. By showing that seasonal patterns of female aggression do not mirror the established patterns of T levels in this highly competitive bird, our findings provide a unique glimpse into how behavioural mechanisms and functions may interact across breeding stages to regulate plasticity.     

A Two‐Resonance Tapping Cavity for an Optimal Light Trapping in Thin‐Film Solar Cells

An optimal photon absorption in thin film photovoltaic technologies can only be reached by effectively trapping the light in the absorber layer provided a considerable portion of the photons is rejected or scattered out of such layer. Here, a new optical cavity is proposed that can be made to have a resonant character at two different nonharmonic frequencies when adjusting the materials or geometry configurations of the partially transmitting cavity layers. Specific configurations are found where a reminiscence of such two fundamental resonances coexists leading to a broadband light trapping. When a PTB7‐Th:PC₇₁BM organic cell is integrated within such cavity, a power conversion efficiency of 11.1% is measured. This study also demonstrates that when materials alternative to organics are used in the photoactive cell layer, a similar cavity can be implemented to also obtain the largest light absorption possible. Indeed, when it is applied to perovskite cells, an external quantum efficiency is predicted that closely matches its corresponding internal one for a broad wavelength range.     

Dispersal in Kin Coalition Throughout the Non‐Breeding Season to Facilitate Fine‐Scale Genetic Structure in the Breeding Season: Evidence From a Small Passerine

Evidence shows that social cooperation among kin may evolve even in birds with extensive dispersal. In such cases, maintaining kinship during dispersal is essential to the subsequent expression of kin cooperation. This hypothesis has not been examined for most bird species. We addressed it in the ground tit (Parus humilis), a passerine where kin frequently interact in terms of cooperative polygamy and extra‐pair mating despite fast annual turnover of the breeding population. Pedigree and genotype data showed that while groups varied in composition throughout the non‐breeding season due to continual individual emigration and immigration, they always contained kin coalitions consisting of either local or immigrant individuals of different age and sexes. The first‐order kin coalitions, according to the information from local individuals, stemmed from single‐family lineages (siblings and their parents), and the lower‐order ones from neighbouring, related family lineages that merged after fledging. It was probable that immigrants had formed kin coalitions in similar ways before dispersing. Groups broke up in the breeding season. Pairing between unrelated individuals from different coalitions within a group was more likely, whereas related individuals from the same coalition tended to nest near each other. The resulting fine‐scale population genetic structure is expected to facilitate breeding interactions among kin. Our findings give clues to understanding the evolution of social cooperation in relation to dispersal.     

Restless roosts: Light pollution affects behavior,sleep, and physiology in a free‐living songbird

The natural nighttime environment is increasingly polluted by artificial light. Several studies have linked artificial light at night to negative impacts on human health. In free‐living animals, light pollution is associated with changes in circadian, reproductive, and social behavior, but whether these animals also suffer from physiologic costs remains unknown. To fill this gap, we made use of a unique network of field sites which are either completely unlit (control), or are artificially illuminated with white, green, or red light. We monitored nighttime activity of adult great tits, Parus major, and related this activity to within‐individual changes in physiologic indices. Because altered nighttime activity as a result of light pollution may affect health and well‐being, we measured oxalic acid concentrations as a biomarker for sleep restriction, acute phase protein concentrations and malaria infection as indices of immune function, and telomere lengths as an overall measure of metabolic costs. Compared to other treatments, individuals roosting in the white light were much more active at night. In these individuals, oxalic acid decreased over the course of the study. We also found that individuals roosting in the white light treatment had a higher probability of malaria infection. Our results indicate that white light at night increases nighttime activity levels and sleep debt and affects disease dynamics in a free‐living songbird. Our study offers the first evidence of detrimental effects of light pollution on the health of free‐ranging wild animals.     

Insect outbreaks increase populations and facilitate reproduction in a cavity‐dependent songbird,the Mountain Chickadee Poecile gambeli

Large‐scale insect outbreaks, typical of temperate ecosystems, may increase food availability and influence species interactions in insectivorous bird communities. We assessed how population densities of a secondary cavity nester, the Mountain Chickadee Poecile gambeli, varied with densities of avian cavity excavators, potential competitors and nest predators, during two large‐scale insect outbreaks of Western Spruce Budworm Choristoneura occidentalis and Mountain Pine Beetle Dendroctonus ponderosae. At the regional level, Mountain Chickadee densities doubled with increases in budworm and beetle availability, then declined with decreases in beetle availability, despite high budworm supply. At the site level, densities of excavators were the best predictor of site‐level variation in Chickadee densities in the following year. During and after the beetle outbreak, Mountain Chickadees used more cavities excavated by Red‐breasted Nuthatches Sitta canadensis and Downy Woodpeckers Picoides pubescens, both of which are bark insectivores that increased in densities concomitantly with the beetle outbreak and whose foraging activities may have facilitated the Mountain Chickadees' access to bark beetles. Thus, Mountain Chickadees showed a numerical response to the food pulse at the regional level, but a functional response to the pulse of nesting cavities at the site level. Plasticity in resource selection and heterospecific attraction may allow Mountain Chickadees to respond to resource pulses in highly variable environments.     

Diet affects female mating behaviour in a seed‐feeding beetle

In species where males provide nuptial gifts, females can improve their nutritional status and thus increase their fecundity by mating when in need of resources. However, mating can be costly, so females should only mate to acquire resources when the need for resources is large, such as when females are nutritionally‐deprived. Two populations of the seed‐feeding beetle Callosobruchus maculatus, a species that produces relatively large nuptial gifts, are used to test whether female nutritional status affects mating behaviour. Female access to water, sugar and yeast are manipulated and the fitness consequences of these manipulations are examined together with the effects of diet on the propensity of nonvirgin females to mate. Access to water has a small but significant effect on mass loss over time, lifespan and fecundity of females, relative to unfed controls. Access to sugar (dissolved in water) improves female fecundity and lifespan above that of hydrated females but access to yeast has no positive effects on female survival or reproduction. Diet has a large effect on both receptivity of nonvirgin females to a male and how quickly they accept that male. Unfed females are both more likely to mate, and accept a mate more quickly, than females provided access to water, which are more likely to mate and accept a mate more quickly than females provided with sugar. This rank order of behaviours matches the order predicted if females increase their mating rate when nutritionally deprived (i.e. it matches the effect of diet on female fitness). The results obtained also suggest that mate choice may be condition‐dependent: females from one population (Burkina Faso) show a preference for large males when well‐fed but not when unfed, although this result is not found in a second population (South India). It is concluded that nutritionally‐deprived females are more receptive to mates than are well‐fed females, consistent with the hypothesis that females ‘forage’ for nuptial gifts, or at least more willingly accept sperm in exchange for nuptial gifts, when they are nutritionally deprived.