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 habitat selection by the Austral parakeet in north‐western Patagonia

Identifying habitat or nesting microhabitat variables associated with high levels of nest success is important to understand nest site preferences and bird–habitat relationships. Little is known about cavity availability and nest site requirements of cavity nesters in southern hemisphere temperate forests, although nest site limitation is suggested. Here we ask which characteristics are selected by the Austral parakeet (Enicognathus ferrugineus) for nesting in Araucaria araucana–Nothofagus pumilio forest in Argentine Patagonia. We compared nest plot and tree characteristics with unused plots and trees among areas of different A. araucana–N. pumilio density. We also examine whether nest plot and tree use and selection, and the associated consequences for fitness of Austral parakeets are spatially related to forest composition. Austral parakeets showed selectivity for nests at different spatial scales, consistently choosing isolated live and large trees with particular nest features in a non‐random way from available cavities. Mixed A. araucana–N. pumilio forests are ideal habitat for the Austral parakeets of northern Patagonia, offering numerous potential cavities, mainly in N. pumilio. We argue that Austral parakeet reproduction and fitness is currently very unlikely to be limited by cavity availability, although this situation may be rapidly changing. Natural and human disturbances are modifying south temperate forests with even‐aged mid‐successional stands replacing old growth forests. Cavity nesting species use and need old growth forests, due to the abundance of cavities in large trees and the abundance of larvae in old wood. Neither of the latter resources is sufficiently abundant in mid‐successional forests, increasing the vulnerability and threatening the survival of the Austral.     

The role of body size in nest-site selection by secondary cavity-nesting birds in a subtropical Chaco forest

Most bird species that nest in tree cavities globally occur in diverse assemblages in little-studied tropical and subtropical forests which have high rates of habitat loss. Conservation of these communities will require an understanding of how species traits, such as body size, influence nest-site selection. We examined patterns of nest-site selection of secondary cavity-nesting birds at the nest patch, tree and cavity scale, and investigated how these patterns are influenced by body size. Using conditional logistic regression, we compared characteristics of 155 nest tree cavities paired with 155 unused tree cavities in quebracho Schinopsis balansae forests in Chaco National Park, Argentina (2016–2018). The odds of a cavity being used for nesting increased with its depth and height above ground, decreased with entrance size, and were greater for dead trees than live. Small-bodied (13–90 g) species used floor diameters in proportion to availability, but medium- (150–200 g) and large-bodied (400–700 g) species selected cavities with larger floors. Model selection indicated that characteristics at the nest patch scale (canopy cover, tree density) had little effect on nest-site selection when cavity-scale variables were included. Cavity floor diameter, entrance size, cavity height and tree diameter (but not cavity depth) increased with body mass, and larger bird species more often used live trees. Two tree species proved to be key for the community: large and medium-sized birds used almost exclusively large live Schinopsis balansae, whereas small birds used live and dead Prosopis spp. in a proportion greater than its availability. Small birds could be differentiated according to species-specific cavity characteristics, but medium and large species overlapped considerably with one another. Although body mass explained much of the overall variation in tree and cavity characteristics between small and medium/large species, several small-bodied species consistently used cavities outside of the expected characteristics for their body size, suggesting that other natural history traits may play important roles in nest-site selection by small-bodied birds. To retain the full suite of secondary cavity-nesters in species-rich tropical and subtropical forests, it is necessary to conserve a diversity of trees and cavities that meet the full range of nesting requirements of these trait-diverse communities.     

Secondhand homes: The multilayered influence of woodpeckers as ecosystem engineers

Ecosystem engineers alter, and can be influenced in turn by, the ecosystems they live in. Woodpeckers choose foraging and nesting sites based, in part, on food availability. Once abandoned, these cavities, particularly within areas of high forage, may be crucial to secondary cavity‐nesting birds otherwise limited by cavities formed through decay. Our study examined factors that influence the nesting success of primary cavity nesters and the subsequent impact on secondary cavity‐nesting birds. Using 5 years of point count data, we monitored the outcomes of cavity‐nesting birds in South Texas. We used logistic‐exposure models to predict daily survival rates based on cavity metrics and used woodpecker foraging trends and insect surveys to determine if nesting where woodpeckers actively forage benefits secondary cavity‐nesting birds. Both woodpeckers and secondary cavity nesters shared predictors of daily survival; nests were more successful in cavities with small openings in minimally decayed trees. All secondary cavity nesters had higher probabilities of success when nesting in an abandoned woodpecker cavity, opposed to ones formed by decay. Woodpeckers tended to forage in areas with higher‐than‐average levels of the insect orders Coleoptera, Hymenoptera, and Orthoptera, and secondary cavity nesters had higher rates of success when nesting in these areas. Our results suggest abandoned woodpecker cavities may be constructed in a way that directly benefit secondary cavity nesters. Additionally, we suggest an interplay between these ecosystem engineers, food availability, and secondary cavity nesters: Woodpeckers engineer superior nesting cavities in areas where food is more abundant, and the resultant cavities in areas of high forage may benefit local secondary cavity nesters. Our findings indicate that there is still much to be explored in the role of ecosystem engineers, and how they influence local communities on multiple trophic levels.     

Characteristics of nest cavities of Barrow's Goldeneyes in eastern Canada

ABSTRACT Barrow's Goldeneyes (Bucephala islandica) are secondary cavity nesters found in western North America and, to a lesser extent, in eastern North America. The eastern North American population is concentrated in the province of Québec and totals about 2000 pairs. Characteristics of nest cavities used by Barrow's Goldeneyes have been described in western North America, but no nest cavities have been found in eastern North America. From 2004 to 2008, we searched for nest cavities in the species’ core breeding area in the boreal forests north of the St. Lawrence River. We captured 12 adult females on their breeding grounds and fitted them with transmitters, but none apparently nested so we conducted ground searches in areas near lakes where paired birds were observed. We found 11 cavities, with 10 in dead, decaying trees and one in the dead part of a dying tree. Nine cavities were in white birch (Betula papyrifera) trees. Mean cavity height was 3.5 ± 1.6 (SD) m (range = 1.2–6.6 m) and mean diameter at breast height (DBH) of cavity trees was 37.8 ± 4.7 cm (range = 32.2–47.5 cm). In contrast to the population in western North America, Barrow's Goldeneyes in eastern North America appear to rely on the availability of natural cavities formed in large, decaying trees for nesting. Current forestry regulations in Québec do not promote the retention of either large trees or older forests, reducing the availability of potential nest cavities for Barrow's Goldeneyes and likely threatening their long‐term conservation. Therefore, we recommend that guidelines be developed to promote silvicultural practices aimed at preserving the long‐term availability of large (DBH ≥ 30 cm) decaying trees across the breeding range of Barrow's Goldeneyes.     

The conservation value of tree decay processes as a key driver structuring tree cavity nest webs in South American temperate rainforests

South American temperate rainforests, a global biodiversity hotspot, have been reduced to nearly 30% of their original extent and most remaining stands are being degraded. Cavity-nesting vertebrate communities are dependent on cavity-bearing trees and hierarchically structured within nest webs. Evaluating the actual degree of cavity dependence (obligate, non-obligate) and the preferred attributes of trees by cavity nesters is critical to design conservation strategies in areas undergoing habitat loss. During three breeding seasons (2010–2013), we studied the cavity-nesting bird community in temperate rainforests of Chile. We found the highest reported proportion of tree cavity nesters (n = 29 species; 57%) compared to non-cavity-using birds for any forest system. Four species were excavators and 25 were secondary cavity nesters (SCNs). Among SCNs, ten species were obligate and 15 were non-obligate cavity nesters. Seventy-five percent of nests of SCNs were located in cavities produced by tree decay processes and the remaining 25% were in cavities excavated mainly by Pygarrhichas albogularis and Campephilus magellanicus. Nest web structure had a low dominance and evenness, with most network interactions occurring between SCNs and large decaying trees. Tree diameter at breast height (DBH) was larger in nest-trees (57.3 cm) than in available trees (26.1 cm). Cavity nesters showed a strong preference for dead trees, both standing and fallen (58% of nests). Our results stress that retaining large decaying and standing dead trees (DBH > 57 cm), and large fallen trees, should be a priority for retention in forest management plans in this globally threatened ecosystem.     

Cavity construction by reintroduced populations of Red‐cockaded Woodpeckers (Dryobates borealis)

Although species reintroduction attempts are now common, monitoring of reintroduction attempts rarely extends beyond initial population establishment. This short timespan likely fails to document long‐term population stability, subtle changes in behavior, and the potentially larger effects that some reintroduced species may have on other species. The Red‐cockaded Woodpecker (RCW; Dryobates borealis) is an important habitat specialist and ecosystem engineer that excavates cavities in living trees. Excavation of natural RCW cavities can take years to complete, but they also persist for many years and are used by many other species. We quantified characteristics of cavity trees excavated by RCWs (n = 44) in two populations that were reintroduced to unoccupied areas more than 10 years earlier. We measured features associated with heartwood rot and used generalized linear mixed effects regression to determine whether these features differed for trees selected for cavity excavation compared with random neighboring trees. We also assessed population trends for cavity‐nesting species that commonly used RCW cavities on one of the sites. Height of first live limb was the only factor distinguishing natural RCW cavity trees from control trees. Four of six cavity‐nesting species monitored increased significantly following RCW establishment. The increases may relate to the many natural and artificial cavities created during the reintroduction effort. Future reintroductions of the RCW should lead to successful natural cavity excavation if sufficiently large trees with smaller live crowns are present. Future efforts may also benefit the broader community of cavity‐nesting birds.     

Nest-site fidelity and cavity reoccupation by Blue-fronted Parrots Amazona aestiva in the dry Chaco of Argentina

The frequency of cavity reoccupation in secondary cavity nesters depends on several factors including quality of cavities, degree of nest-site fidelity, competition with other cavity nesters and availability of new cavities. Blue-fronted Parrots Amazona aestiva are secondary cavity nesters that live in subtropical forests and savannas of central South America. We examined the characteristics of the trees and cavities used by this species in a protected area of the dry Chaco of Argentina and estimated nest-site fidelity and cavity reoccupation. We also assessed whether the probability of cavity reoccupation was associated with cavity characteristics and nesting success during the previous year. Nest-site fidelity of banded females was 68% and cavity reoccupation by banded and unbanded individuals 62%. Probability of reoccupation was associated with wall thickness and depth of the cavity, and was lower if the nest failed the previous year than if it was successful. The high rate of cavity reoccupation in Blue-fronted Parrots is largely attributable to strong nest-site fidelity and may reflect preferences for cavities whose characteristics are associated with higher nesting success.     

Dark tree cavities – a challenge for hole nesting birds?

Holes provide the safest nest sites for birds, but they are an underutilized resource; in natural forests there are usually more holes than birds that could use them. Some bird species could be prevented from nesting in holes because of their inability to operate in the low light conditions which occur in cavities. As no visual system can operate in complete darkness some nest cavities could be too dark to be useable even by hole‐nesters. Thus, the light conditions within tree cavities could constrain both the evolution of the hole nesting habit, and the nest site choice of the hole‐nesting birds. These ideas cannot be tested because little is known about the light conditions in cavities. We took an opportunity provided by ongoing studies of marsh tits Poecile palustris and great tits Parus major breeding in a primeval forest (Bia?owie?a National Park, Poland) to measure illumination inside their nest cavities. We measured illuminance in cavities at daybreak, which is just after the parents commenced feeding nestlings. Only ca 1% of incoming light reached the level of the nest. Illuminance at nests of both species (median = 0.1–0.2 lx) fell within mesopic‐scotopic range, where colour vision is impaired. Measurements in model cavities showed strong declines in illumination with distance from the entrance, with light levels typically as low as 0.01 lx at 40 cm from the cavity entrance. Thus cavities can be very dark, often too dark for the use of colour vision, and we suggest that ‘lighting’ requirements can affect the adoption of specific nest sites by hole nesting birds. We discuss implications of the findings for understanding the adaptations for hole‐breeding in birds.     

Validation of a landscape‐scale planning tool for cavity‐dependent wildlife

Tree cavities provide important habitat for wildlife. Effective landscape‐scale management of cavity‐dependent wildlife requires an understanding of where cavities occur, but tree cavities can be cryptic and difficult to survey. We assessed whether a landscape‐scale map of mature forest habitat availability, derived from aerial photographs, reflected the relative availability of mature trees and tree cavities. We assessed cavities for their suitability for use by wildlife, and whether the map reflected the availability of such cavities. There were significant differences between map categories in several characteristics of mature trees that can be used to predict cavity abundance (i.e. tree form and diameter at breast height). There were significant differences between map categories in the number of potential cavity bearing trees and potential cavities per tree. However, the index of cavity abundance based on observations made from the ground provided an overestimate of true cavity availability. By climbing a sample of mature trees we showed that only 5.1% of potential tree cavities detected from the ground were suitable for wildlife, and these were found in only 12.5% of the trees sampled. We conclude that management tools developed from remotely sensed data can be useful to guide decision‐making in the conservation management of tree cavities but stress that the errors inherent in these data limit the scale at which such tools can be applied. The rarity of tree cavities suitable for wildlife in our study highlights the need to conserve the tree cavity resource across the landscape, but also the importance of increasing the accuracy of management tools for decision‐making at different scales. Mapping mature forest habitat availability at the landscape scale is a useful first step in managing habitat for cavity‐dependent wildlife, but the potential for overestimating actual cavity abundance in a particular area highlights the need for complementary on‐ground surveys.     

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.     

Co-occurrence patterns of cavity-nesting birds in cottonwood-willow communities

Summary We studied the covariation of six species of cavity-nesting birds to determine the extent to which species' co-occurrence patterns were attributable to interspecific relations. We tested two null hypotheses: (1) inverse relations did not exist among potential competitors for nest trees or nest cavities, and (2) there were no positive relations among potential commensals for nest cavities. Statistical power analyses, and simple, partial, and multiple partial correlation and regression analyses provided little evidence of species-species or higher-order inverse or positive relations. Most of the variation in co-occurrence patterns was associated with stand characteristics. Collectively, an abundance of nest trees and nest cavities, differences among species in acceptable nest-tree and nest-cavity dimensions, and different periods of peak nesting activity appear to be responsible for the lack of negative relations among potential competitors. Positive relations among potential commensals were weak presumably because (1) secondary cavity nesters (SCNs) frequently used cavities that originated from wind-, lightning-, and/or disease-damaged limbs and were thus not dependent completely on primary cavity nesters for cavities, and (2) the lag time for cavity use by SCNs is often two or more seasons. Interspecific relations were apparently not major determinants of community structure during our investigation; assemblages of cavity nesters appeared instead to be influenced more by independent species-specific habitat responses.     

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.     

Nest site selection in middle and great spotted woodpeckers <Emphasis Type="Italic">Dendrocopos medius</Emphasis> &; <Emphasis Type="Italic">D. major</Emphasis>: implications for forest management and conservation

Success of species conservation depends to a large extent on comprehensive management that considers all critical aspects of a species’ niche. Many studies have examined habitat factors in relation to occurrence, abundance or foraging behaviour of European woodpecker species, while relatively little is known about nest site selection. I compared habitat structures used for nesting by middle and great spotted woodpeckers Dendrocopos medius and D. major with available structures in an oak forest in the Swiss lowlands. I first tested if nest trees were randomly selected among available trees by focusing on species, condition and diameter of nest trees, and on the presence of the fruiting body (hereafter sporophore) of polypores (wood-decomposing fungi). Second, I examined if the nesting niches of the two species were differentiated. Both species showed strong preferences for oaks, large trees, dead trees and for trees with sporophores. Nest sites of the two species differed most strongly with respect to the presence of sporophores, cavity age and tree condition, pointing towards interspecific competition for nest sites. Old living or dead trees with sporophores are central components of the nesting niche of middle and great spotted woodpeckers. Conservation plans for the threatened middle spotted woodpecker have so far mostly focused on the needs in terms of distribution and foraging; future conservation strategies and forest management must take into account the preference for dead and decaying trees with sporophores as another vital resource. This will also provide benefits for other woodpecker species as well as for the community of secondary cavity nesters.     

The relationship between introduced European Starlings and the reproductive activities of Mountain Bluebirds and Tree Swallows in British Columbia,Canada

The European Starling Sturnus vulgaris is an introduced species in North America and is an aggressive competitor for tree cavity nest‐sites. Starlings are commonly considered to influence nest‐site selection and reproductive success of native cavity‐nesting species negatively. We examined the relationship between Starling nest density and the fecundity of two native secondary cavity‐using passerines, Mountain Bluebird Sialia currucoides and Tree Swallow Tachycineta bicolor. We monitored a total of 622 nests (approximately equal numbers for each of the three species) in woodpecker‐excavated and naturally occurring cavities in 29 small forest groves in central British Columbia, Canada, between 2000 and 2009. The dimensions of cavities used and the timing of nest initiation overlapped for all species, although Starlings initiated clutches earliest. Mixed‐effects models were used to assess whether nest abundance, clutch size or nest success were affected directly by Starling nest abundance, or indirectly via a shift in cavity selection or timing of breeding. Starlings and Mountain Bluebirds showed inverse trends in nest abundance. Mountain Bluebird clutch sizes were smaller if they were initiated later in the breeding season. There was weak evidence that Tree Swallow clutch size decreased with cavity depth when Starling nests were abundant, and increased with cavity depth where there were few Starling nests. We conclude that despite the aggressive nature of this exotic cavity‐nester, the influence of Starlings on native secondary cavity‐nesting passerines is modest where cavities are abundant.     

Cavity-nesting birds are limited by nesting habitat in Neotropical agricultural landscapes

Most studies comparing biodiversity between natural and human-modified landscapes focus on patterns in species occurrence or abundance, but do not consider how different habitat types meet species' breeding requirements. Organisms that use or nest in tree cavities may be especially threatened by habitat conversion due to the loss of their nesting sites. Although cavity-nesting bird diversity is highest in the tropics, little is known about how tropical birds use cavities, how agriculture affects their reproductive biology, and how effective nest boxes could be as a conservation strategy in tropical agriculture. Here, we explored how habitat conversion from tropical forests to pasture affects the abundance, nesting habitat availability, and nest success of cavity-nesting birds in Northwest Ecuador. We conducted bird surveys and measured natural cavity availability and use in forest and agriculture. We also added artificial nest boxes to forest and agriculture to see whether cavity limitation in agriculture would elicit higher use of artificial nest boxes. We found evidence of cavity limitation in agriculture—there were many more natural cavities in forest than in agriculture, as well as more avian use of nest boxes placed in agriculture as compared to forest. Our results suggest that it is important to retain remnant trees in tropical agriculture to provide critical nesting habitat for birds. In addition, adding nest boxes to tropical agricultural systems could be a good conservation strategy for certain species, including insectivores that could provide pest-control services to farmers. Abstract in Spanish is available with online material.     

Characteristics of snags and forest structure in southern mistbelt forests of the Amatole region,South Africa

Standing dead trees (or snags) are an important component of forest ecosystems, especially for tree cavity‐nesting vertebrate species, but their prevalence in South African forests remains under studied. Consequently, we investigated forest structure, and the presence and abundance of snags in six southern mistbelt forests in the Eastern Cape, South Africa. These forests have had varying levels of timber extraction over the past 150 years or more. We found snags were relatively rare in all six forests (<4.3% of trees sampled). Mean diameter at breast height (dbh) of snags ranged from 52 to 82 cm across the forests, with smaller snags in Kologha Forest and larger snags in Tyume Forest. A bimodal distribution of snag successional stages was found, with frequencies peaking at early and late stages, and few in the intermediate stages. Tree species diversity in the forests was relatively low (twelve–nineteen species across forests; only 28 species in total). There was no significant difference in dbh of trees between forests, with most occurring in the 20–29‐cm dbh size class. Future studies are required to identify trees that most likely support suitable cavities for tree cavity‐nesting bird species, and to determine cavity‐nester assemblage requirements in southern African forests.     

Nest cavity orientation in black‐capped chickadees Poecile atricapillus: do the acoustic properties of cavities influence sound reception in the nest and extra‐pair matings?

Birds that nest in cavities may regulate nest microclimate by orienting their nest entrance relative to the sun or prevailing winds. Alternatively, birds may orient their nest entrance relative to conspecific individuals around them, especially if the acoustic properties of cavities permit nesting birds to better hear individuals in front of their nest. We measured the cavity entrance orientation of 132 nests and 234 excavations in a colour‐banded population of black‐capped chickadees Poecile atricapillus for which the reproductive behaviour of nesting females was known. Most chickadees excavated cavities in rotten birch Betula papyrifera, aspen Populus tremuloides and maple Acer saccharum. Nest cavities showed random compass orientation around 360° demonstrating that chickadees do not orient their cavities relative to the sun or prevailing winds. We also presented chickadees with nest boxes arranged in groups of four, oriented at 90° intervals around the same tree. Nests constructed in these nest box quartets also showed random compass orientation. To test the acoustic properties of nest cavities, we conducted a sound transmission experiment using a microphone mounted inside a chickadee nest. Re‐recorded songs demonstrate that chickadee nest cavities have directional acoustic properties; songs recorded with the cavity entrance oriented towards the loudspeaker were louder than songs recorded with the cavity entrance oriented away from the loudspeaker. Thus, female chickadees, who roost inside their nest cavity in the early morning during their fertile period, should be better able to hear males singing the dawn chorus in front of their nest cavity. Using GIS analyses we tested for angular‐angular correlation between actual nest cavity orientation and the azimuth from the nest tree to the territories and nest cavities of nearby males. In general, nest cavity entrances showed no angular‐angular correlation with neighbourhood territory features. However, among birds who followed a mixed reproductive strategy and nested in the soft wood of birch and aspen trees, nest cavity entrances were oriented towards their extra‐pair partners. We conclude that nest cavity orientation in birds may be influenced by both ecological and social factors.     

Effects of nestboxes on the breeding biology of Southern House Wrens Troglodytes aedon bonariae in the southern temperate zone

Nestboxes are known to increase clutch size, enhance breeding success and affect the social mating system of several cavity nesters. Although in recent years various cavity nesters have been studied in nestboxes in South America, the effects of boxes on the biology of the study species are unknown. We evaluated the effects of nestboxes on the breeding biology and social mating system of Southern House Wrens Troglodytes aedon bonariae by comparing birds breeding in nestboxes and tree cavities in two cattle ranches in Buenos Aires Province, Argentina. Southern House Wrens nesting in boxes had higher breeding success but, contrary to studies on the temperate zone, we did not find differences in clutch size between Wrens breeding in nestboxes and tree cavities. The main causes of nest failure in tree cavities were nest predation and flooding of the cavity (70 and 23% of the failures, respectively) while in nestboxes predation and desertion were the most important causes of failure (38 and 34% of the failures, respectively). The social mating system of Southern House Wrens is monogamy with biparental care, and neither was affected by the boxes. Males did not attract secondary females to additional nestboxes; however, nestboxes are safer breeding sites than tree cavities, and females seemed to prefer males with nestboxes on their territory. These results suggest that nest quality alone might be not enough for secondary females to accept polygyny.