Competition, predation and nest niche shifts among tropical cavity nesters: phylogeny and natural history evolution of parrots (Psittaciformes) and trogons (Trogoniformes)

Nest site selection by birds is a critically important life history trait as competition for suitable sites can be intense, and because birds are at their most vulnerable to predators during nesting. Previous studies show that the clutch size and nestling period evolve in response to competition for nest sites and nest predation, respectively. This provides the opportunity to study the relative contribution of competition and predation to the evolution of nesting niche. Using previously published phylogenies for parrots and trogons, I found evidence for at least 13 independent evolutionary transitions from tree cavities to alternative nesting niches (including termitaria, cliffs, and burrows). I analyzed variations in clutch size, incubation period and nestling period for 16 phylogenetically controlled pairs of species to test the relative roles of competition for tree cavities and nest predation, in favoring evolutionary switches to alternative nest sites. Tree cavity nesting species did not have larger clutch sizes as predicted if competition for tree cavities leads birds to invest heavily in nesting once they obtain a nest site (the limited breeding opportunities hypothesis). Instead I found that shifts to alternative nesting niches were accompanied by an increase in nestling period. As nestling period is a surrogate measure for long-term nest predation rates, this finding suggests that nest predation has been more important than competition in niche diversification among cavity nesting parrots and trogons. The timing of events in South America suggests that the explosive radiation of mammalian nest predators during the Upper-Oligocene, Lower-Miocene (20–30 million years ago) corresponded with the radiation of parrot and trogon taxa that exploit novel nesting niches.     

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.     

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.     

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.     

Sources of variation in the nesting success of understory tropical birds

Survival of offspring is a key fitness component and, for birds, the threat of predation on nests is especially influential. Data on rates of nest success from tropical regions are comparatively few, conservation‐relevant, and essential for assessing the validity of models comparing the life histories and behavior or birds across latitudinal gradients. We monitored over 2 000 nests in the lowland forests of central Panama and, using the logistic exposure to model the fate of nests, explored the importance of variation in rate of nest success according to type of nest, height of nests, among years, in early versus late nests, and at different stages of the nest cycle. Analyses of over 1 400 nests for 18 species revealed considerable variation among species in the daily survival rate of nests (range among 18 species=0.91 to 0.98), but nest type and stage of the nesting cycle were generally influential on the probability of nest success. Cavity or enclosed nesters experienced greater nest success than open cup nesters and rates of nest loss were generally greatest in the nestling stage. We found limited evidence that height of nests affected probability of success, but no indication that timing of nesting effort was influential. Despite the occurrence of a severe ENSO event during our sampling, annual variation in nest success was not consistent among species. Interspecific variation in the rates and patterns of nest predation in our study, coupled with reports of high rates of nest loss at temperate latitudes, lead us to question long standing assumptions about latitudinal trends in rates of nest loss. We urge further work to understand the implications of nest predation on the evolutionary ecology of tropical birds.     

Characteristics of and competition for nest sites by the Rüppell's Parrot,Poicephalus ruepelli

A knowledge of the nesting requirements of Rüppell's Parrot can aid its conservation. Nests were found during 17 months of fieldwork in Namibia and characteristics of the sites are reported here. Nests were found in woodpecker cavities. 72% of the nests were in three tree species: Faidherbia albida, Acacia erioloba and Combretum imberbe, and 72% were in live branches. Relatively large trees were used and the entrance holes were generally tight-fitting for the birds. Both sexes worked to further excavate a cavity. Comments are included regarding territoriality, relationships with other cavity-nesting species and nest site limitation.     

Do bright colors at nests incur a cost due to predation?

Summary Birds show much interspecific variation in the coloration and brightness of their plumage. I examine the hypothesis that selection due to predation on incubating birds and their nest contents can explain part of this diversity. First, I argue that rather than using absolute rates of nest predation to make predictions about the costs of conspicuous colours, we should measure experimentally whether increases in plumage conspicuousness elevate rates of nest predation. Second, I present experimental data investigating the cost of red and brown colour at ground and tree nests. These data provide the first evidence that bright colours do attract predators to nests and that, in addition, this cost varies according to the nesting site. Natural selection seems to most strongly oppose the evolution of conspicuous colours in ground-nesting birds.     

Direct and indirect effects of an insect outbreak increase the reproductive output for an avian insectivore and nest‐cavity excavator,the red‐breasted nuthatch Sitta canadensis

Community‐wide food pulses may ameliorate food constraints but may also result in increased competition for other resources and predation rates. In cavity‐nesting vertebrate communities, where the availability of tree cavities can limit reproduction and the reuse of cavities can increase nest predation by squirrels, excavators may maximize their fecundity by creating new cavities in competitor‐ and predator‐rich habitats that undergo food pulses. The reproductive cost associated with excavation (i.e. increased energy allocation early in the breeding season that often delays laying and thereby reduces clutch size), may be reduced if food pulses allow for a longer breeding season and larger clutches. A large‐scale mountain pine beetle Dendroctonus ponderosae outbreak that occurred during our long‐term study (1995–2009) provided a natural food supplementation experiment across 27 sites in British Columbia, Canada. We examined the effects of a reduction in food constraints accompanied with increases in excavation rates, conspecific density and nest predation risk on the fecundity of a facultative excavator, the red‐breasted nuthatch Sitta canadensis. We found a total of 420 nests in tree cavities. Nuthatch clutch sizes ranged from two to nine eggs, and broods from one to nine fledglings per nest. Later clutches were larger at sites and in years with high beetle abundance (mean clutch size of six eggs did not decline later in the season), second broods were produced in outbreak years (usually only one nesting attempt/normal year), and the number of fledglings per successful nest increased with increasing beetle abundance and nuthatch densities, but declined with increased squirrel densities. Since fecundity did not differ between new and reused cavities, the costs and benefits of excavation versus cavity reuse may be neutralized for nuthatches during strong resource pulses. Overall, the beetle outbreak reduced food constraints for nuthatches and provided alternate food for nest predators, allowing increased annual fecundity.     

Cavity use and breeding biology of endangered Bahama Swallows (Tachycineta cyaneoviridis): implications for conservation

Bird populations, especially on islands, have declined or gone extinct due to overhunting, habitat loss and fragmentation, and adverse effects of the introduction of non-native species. Bahama Swallows (Tachycineta cyaneoviridis), endangered secondary cavity nesters that breed on only three islands in the northern Bahamas, are an island species with a declining population, but the causes of this decline are unknown. During four breeding seasons on Great Abaco Island (2014–2017), we identified cavity-nesting resources used by breeding swallows in native pine forest and other habitats, and estimated phenology and breeding parameters from a subset of nests. Bahama Swallows nest in cavities in a variety of structures, but rely most on woodpecker-excavated cavities in pine snags and utility poles. Swallows nesting in cavities in pine snags had higher fledging success (92%) than those nesting in cavities in utility poles (50–62%), which were concentrated in non-pine habitat that may expose swallows to predation and increased competition for nest cavities from other species. The high reproductive success of Bahama Swallows nesting in the pine forest indicates that the decline in population cannot be attributed to poor productivity on southern Great Abaco. However, our results suggest that the dependence of Bahama Swallows on cavities excavated by Hairy Woodpeckers (Dryobates villosus) for nesting sites may be a factor in their decline, and highlight the potential importance of the protection and management of pine forests in future efforts to ensure the survival of Bahama Swallows.     

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.     

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.     

Breeding biology of Eared Quetzals in the Sierra Madre Occidental, Mexico

ABSTRACT.   Eared Quetzals ( Euptilotis neoxenus ), a threatened species, are one of the least studied trogons in Mexico. We monitored 29 Eared Quetzal nests in the Chihuahuan portion of the Sierra Madre Occidental from 1998 to 2003. All nests were in tree cavities, and the mean tree and nest cavity heights ( N = 14) were 16.9 ± 7.8 m and 11.4 ± 4.1 m, respectively. The mean clutch size was 2.8 ± 0.9 eggs ( N = 28), the incubation period lasted 22 d ( N = 1), and nestling periods ranged from 29 to 31 d ( N = 5). Both adults incubated eggs and fed nestlings. Of 80 eggs, 70 hatched (87.5%) and 67 of 70 young fledged (95.7%). Twenty-five of 29 nests (86.2%) produced at least one fledgling. One nest was predated, and two failed when nest trees fell. Higher rates of nest predation have been reported for other species of trogons. However, fewer potential predators, such as snakes and mammals, are present in the Sierra Madre than in tropical zones where most trogon species occur. In addition, antipredator behaviors, including nestlings with calls resembling a snake and nests with an unpleasant odor, may contribute to the high nesting success. The main limiting factors for Eared Quetzals in the northern Chihuahua may be competition for cavities with other secondary cavity-nesters, and the failure of nests when snags fall.     

Impact of brood parasitism and predation on nest survival of the fan-tailed gerygone in New Caledonia

Predation and brood parasitism are common reasons for nesting failure in passerine species and the additive impact by invasive species is a major conservation concern, particularly on tropical islands. Recognising the relative contribution of the different components of nesting failure rates is important to understand co-evolutionary interactions within brood parasite–host systems. In the remote archipelago of New Caledonia, the fan-tailed gerygone Gerygone flavolateralis is the exclusive host of the brood-parasitic shining bronze-cuckoo Chalcites lucidus. Additionally, invasive rodents also possibly have an impact on breeding success. To estimate the impact of potential nest predators, we 1) video monitored nests to identify predators, 2) estimated the probability of predation based on nest visibility and predator abundance and 3) tested the possibility that the location of experimental nests and lack of odour cues decrease the predation by rodents. In addition, we estimated nest survival rates using data collected in different habitats over the course of eight breeding seasons. Nesting success of fan-tailed gerygones was relatively low and predation was the main cause of nesting failure. We recorded mainly predation by native birds, including the shining bronze-cuckoo, whereas predation by rats was rare. In open habitats predation by cuckoos was much lower than predation by other avian predators. Neither predator activity around nests nor nest visibility influenced the probability of predation. Experimental nests in more accessible locations and containing an odorous bait were more exposed to rodent predation. Apparently, the fan-tailed gerygone has either never been specifically vulnerable to predation by rats or has developed anti-predator adaptations.     

Nest niche dynamics of Meyer's Parrot Poicephalus meyeri in the Okavango Delta,Botswana

Meyer's Parrot Poicephalus meyeri is the only cavity-nesting bird species that breeds during winter in the Okavango Delta. This is facilitated by exclusive access to arthropod larvae incubating inside and feeding on fruits and pods in their diet. To minimise predation risk and overcome low overnight temperatures they have specialised, non-random nest cavity preferences that restrict them to 4.5% of the available nest cavities in the study area. Here we evaluated the nest niche of Meyer's Parrot by studying the nest tree preferences and ecological context of all nest cavities to determine factors that may restrict breeding success in disturbed or altered habitat. Although specific nest tree preferences were significantly different between host tree species, Meyer's Parrot preferred trees greater than 14 m in height that were in relatively poor condition (e.g. portion of the canopy dead). A comparison of nest tree characteristics (n = 75) and the availability of these tree specifications in a representative sample of the different habitat types (n = 1 129) within the sample area indicated that Meyer's Parrot are dependent on riverine forest, Acacia-Combretum marginal woodland and dry mopane woodland for nesting opportunities. Disturbance to hardwood trees by African elephants Loxodonta africana and fungal attack (e.g. Coriolus versicolor) are likely important dynamics in supporting healthy Meyer's Parrot populations and cavity-nesting bird communities.     

Nesting Success in Crimson Finches: Chance or Choice?

In avian systems, nest predation is one of the most significant influences on reproductive success. Selection for mechanisms and behaviours to minimise predation rates should be favoured. To avoid predation, breeding birds can often deter predators through active nest defence or by modifying behaviours around the nest (e.g. reducing feeding rates and vocalisations). Birds might also benefit from concealing nests or placing them in inaccessible locations. The relative importance of these strategies (behaviour vs. site selection) can be difficult to disentangle and may differ according to life history. Tropical birds are thought to experience higher rates of predation than temperate birds and invest less energy in nest defence. We monitored a population of crimson finches (Neochmia phaeton), in the Australian tropics, over two breeding seasons. We found no relationship between adult nest defence behaviour (towards a model reptile predator) and the likelihood of nest success. However, nest success was strongly related to the visibility of the nest and the structure of the vegetation. We found no evidence that adult nest building decisions were influenced by predation risk; individuals that re‐nested after a predation event did not build their nest in a more concealed location. Therefore, predator avoidance, and hence nest success, appears to be largely due to chance rather than due to the behaviour of the birds or their choice of nesting sites. To escape high predation pressures, multiple nesting attempts both within and between seasons may be necessary to increase reproductive success. Alternatively, birds may be limited in their nest‐site options; that is, high‐quality individuals dominate quality nest sites.     

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.     

Nesting success of understory forest birds in central Panama

Greater nest predation in tropical than temperate birds has been hypothesized to be a primary selective force generating latitudinal differences in avian life history traits. Few extensive data sets, however, have been available from tropical forests to compare with data from temperate forests. To increase the amount of empirical information available for addressing issues related to the evolution of life history traits of tropical birds, we measured the nesting success of understory birds in lowland forest of central Panama. We found and monitored the fates of 696 nests of 71 species over two breeding seasons. Daily nest predation rates for the ten species for which we obtained the largest samples ranged from 1.6 to 8.3%, equivalent to a loss of 43 to 92% of nests. These values overlapped extensively the range of daily predation rates experienced by ecologically similar species in North America. Proportion of nests fledging young, estimated with the Mayfield method, was significantly lower in tropical (range: 8 to 57%) than temperate (27 to 60%) species. Nesting success in Panama varied among years, however, being greater in 1996 than 1997. In 1996, nesting success was similar to that of species breeding in forest fragments of midwestern North America. When compared with success of nests in large, contiguous forest tracts of North America, however, tropical avian nesting success was consistently lower by approximately 23%. We conclude that nesting success in central Panama may be poor in most breeding seasons, but also may be punctuated by occasional years of relatively exceptional success, a possibility heretofore unappreciated because of a general paucity of data from the tropics. Furthermore, our results indicate substantial variation in levels of nesting success among species, and almost no variation in clutch size. Such large interspecific variation, as well as potentially large annual variation, in nesting success does not support the hypothesis that uniformly low levels of nesting success select for small tropical clutch sizes.     

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.