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.     

Spatial scale and sampling resolution affect measures of gap disturbance in a lowland tropical forest: implications for understanding forest regeneration and carbon storage

Treefall gaps play an important role in tropical forest dynamics and in determining above-ground biomass (AGB). However, our understanding of gap disturbance regimes is largely based either on surveys of forest plots that are small relative to spatial variation in gap disturbance, or on satellite imagery, which cannot accurately detect small gaps. We used high-resolution light detection and ranging data from a 1500 ha forest in Panama to: (i) determine how gap disturbance parameters are influenced by study area size, and the criteria used to define gaps; and (ii) to evaluate how accurately previous ground-based canopy height sampling can determine the size and location of gaps. We found that plot-scale disturbance parameters frequently differed significantly from those measured at the landscape-level, and that canopy height thresholds used to define gaps strongly influenced the gap-size distribution, an important metric influencing AGB. Furthermore, simulated ground surveys of canopy height frequently misrepresented the true location of gaps, which may affect conclusions about how relatively small canopy gaps affect successional processes and contribute to the maintenance of diversity. Across site comparisons need to consider how gap definition, scale and spatial resolution affect characterizations of gap disturbance, and its inferred importance for carbon storage and community composition.     

Predators of bird nests in the Neotropics: a review

Predation is the main cause of nest failure among birds and, therefore, a strong selective agent. To fully understand patterns of nest predation, determining the identities of nest predators is crucial. Information about nest predators in the Neotropics, however, is largely anecdotal and not easily accessible in the literature. Our objective was to search the literature and compile a list of the known predators of nests in the Neotropics. We identified 256 species belonging to 67 families of birds, reptiles, mammals, and arthropods as nest predators. Families with at least 10 species of identified nest predators included Colubridae, Accipitridae, Corvidae, Ramphastidae, Falconidae, Furnariidae, Icteridae, and Didelphidae. Species in the first five of these families, plus the family Cebidae, predated nests of at least 30 species of birds. Many species not included on our list are also likely nest predators, e.g., 79 species identified as nest predators in the Nearctic that also occur in the Neotropics, but have not yet been confirmed as predators there. Increased use of video technology in the future should lead to an increase in the numbers of nest predators identified, particularly those that are nocturnal. By determining which species on our list occur in a given study area, researchers can now consider the likely nest predators in their study areas when designing hypotheses and conservation plans.     

An artificial bird nest experiment in urban environments: Lessons from a school‐based citizen science programme

Maintaining suitable vegetation within urban environments is crucial for wildlife conservation in the face of anthropogenic habitat change. Here, we report on a citizen science project, involving students from seven schools across south‐eastern Australia, that investigated the effectiveness of urban vegetation as habitat for bird nests. The ‘nest concealment hypothesis’ posits that vegetation should obscure the nest from predator detection, thus reducing the likelihood of predation. To test this, participating school‐aged citizen scientists constructed artificial nests, which were placed in garden trees within school grounds and monitored for signs of predation. We found no evidence to support the nest concealment hypothesis, with no relationship between the density of vegetation immediately surrounding a nest and its likelihood of predation (binomial model:  = 1.714, P = 0.190). It was observed that 80% of the nests experienced predation. This aligns with mounting evidence suggesting that other factors, such as olfaction and adult defence, may be more important factors in the protection of bird nests. It is important to note that artificial nests are unreliable, and therefore, the veracity of the overall conclusions is limited. However, in conducting this experiment, we demonstrate the suitability of this method as a school‐based citizen science activity. This study exemplifies that field‐based experiments can used to engage future generations with conservation science.     

Diversity and evolutionary origins of fungi associated with seeds of a neotropical pioneer tree: a case study for analysing fungal environmental samples

Fungi associated with seeds of tropical trees pervasively affect seed survival and germination, and thus are an important, but understudied, component of forest ecology. Here, we examine the diversity and evolutionary origins of fungi isolated from seeds of an important pioneer tree (Cecropia insignis, Cecropiaceae) following burial in soil for five months in a tropical moist forest in Panama. Our approach, which relied on molecular sequence data because most isolates did not sporulate in culture, provides an opportunity to evaluate several methods currently used to analyse environmental samples of fungi. First, intra- and interspecific divergence were estimated for the nu-rITS and 5.8S gene for four genera of Ascomycota that are commonly recovered from seeds. Using these values we estimated species boundaries for 527 isolates, showing that seed-associated fungi are highly diverse, horizontally transmitted, and genotypically congruent with some foliar endophytes from the same site. We then examined methods for inferring the taxonomic placement and phylogenetic relationships of these fungi, evaluating the effects of manual versus automated alignment, model selection, and inference methods, as well as the quality of BLAST-based identification using GenBank. We found that common methods such as neighbor-joining and Bayesian inference differ in their sensitivity to alignment methods; analyses of particular fungal genera differ in their sensitivity to alignments; and numerous and sometimes intricate disparities exist between BLAST-based versus phylogeny-based identification methods. Lastly, we used our most robust methods to infer phylogenetic relationships of seed-associated fungi in four focal genera, and reconstructed ancestral states to generate preliminary hypotheses regarding the evolutionary origins of this guild. Our results illustrate the dynamic evolutionary relationships among endophytic fungi, pathogens, and seed-associated fungi, and the apparent evolutionary distinctiveness of saprotrophs. Our study also elucidates the diversity, taxonomy, and ecology of an important group of plant-associated fungi and highlights some of the advantages and challenges inherent in the use of ITS data for environmental sampling of fungi.     

An alternative strategy for maintenance of eusociality after nest destruction: new nest construction in a primitively eusocial wasp

Although nests are central to colonial life in social insects, nests are sometimes damaged by predators or natural disasters. After nest destruction, individuals usually construct new nests. In this case, a sophisticated mechanism like the scent trail pheromone used in large insect colonies that recruit individuals to new nest sites would be important for the maintenance of eusociality. In independent-founding Polistes wasps, it is well known that queens enforce workers physiologically on the natal nests even if evidence of trail pheromone use has not been exhibited. We investigated the effect of the queen on an alternative strategy for the maintenance of eusociality by first females after nest destruction in the primitively eusocial wasp Polistes chinensis. We predicted that the first females in queen-absent colonies have various behavioral options after nest destruction. Even if the females construct new nests cooperatively with other individuals, the new nest construction should be conducted more smoothly in queen-present colonies because the queens regulate the behavior of wasps. We made wasps construct new nests by removing the entire brood from existing nests. The presence of the queen did not cause variation in the alternative strategy of the first females, as the first females (workers) usually constructed new nests cooperatively irrespective of the queen-presence. Thus, the workers in the queenpresent colonies affiliated to the new nest construction more smoothly and constructed new nests more efficiently than workers in the queen-absent colonies. Our results suggest that the presence of the queen is important for maintaining eusociality in primitively eusocial wasps after nest destruction. Received 8 February 2005; revised 5 October 2005; accepted 17 October 2005.     

Symbiotic nesting of birds with formidable animals: a review with applications to biodiversity conservation

Increased predation and parasitism of bird nests has become a major problem in many biological communities altered by human activities, often causing declines in bird populations. To help solve this threat to biodiversity, I propose restoring the abundance of symbiotic nest-protecting animals in habitats where birds face an increased risk from predators and parasites, so that birds there can increase their chances of reproductive success by nesting close to these protectors. The re-establishment of such protective nesting associations to increase avian reproductive success differs from other proposed solutions to this problem in that it involves point defense of bird nests themselves. Rather than diminishing the number of nest predators and brood parasites in the whole habitat or community, as proposed with other approaches, the presence, activity and success of these enemies are reduced only within the microhabitat defended by the protector. The animal protecting the nest need not be larger in size than the predators or brood parasites, and is often many times smaller. In addition, it need not be from a higher trophic position, and in many cases comes from the same or a lower trophic level. Research suggests that an informed and careful use of nest protecting animals by wildlife managers can help reverse or prevent the decline of many bird populations, especially when used in combination with other approaches such as restoration of top predator populations and habitats. Although wildlife biologists have long recognized the important role that plants play in concealing and protecting bird nests from enemies, and regularly recommend manipulation of vegetation to enhance nest survival, they have generally ignored the important role that formidable animals play in protecting bird nests, and failed to incorporate animal protectors into management strategies. Because of this neglect, a host of new studies and experiments are urgently needed to provide managers with the critical information needed to use protective nesting associations effectively in integrated strategies to preserve avian biodiversity.     

Tri-trophic interactions affect density dependence of seed fate in a tropical forest palm

Natural enemies, especially host-specific enemies, are hypothesised to facilitate the coexistence of plant species by disproportionately inflicting more damage at increasing host abundance. However, few studies have assessed such Janzen-Connell mechanisms on a scale relevant for coexistence and no study has evaluated potential top-down influences on the specialized pests. We quantified seed predation by specialist invertebrates and generalist vertebrates, as well as larval predation on these invertebrates, for the Neotropical palm Attalea butyracea across ten 4-ha plots spanning 20-fold variation in palm density. As palm density increased, seed attack by bruchid beetles increased, whereas seed predation by rodents held constant. But because rodent predation on bruchid larvae increased disproportionately with increasing palm density, bruchid emergence rates and total seed predation by rodents and bruchids combined were both density-independent. Our results demonstrate that top-down effects can limit the potential of host-specific insects to induce negative-density dependence in plant populations.     

Host-parasite kinship in a female-philopatric bird population: evidence from relatedness trend analysis

Conspecific brood parasitism (CBP), an alternative reproductive tactic where some females lay eggs in the nests of other females of the same species, occurs in many animals with egg care. It is particularly common in waterfowl, for reasons that are debated. Many waterfowl females nest near their birthplace, making it likely that some local females are relatives. We analyse brood parasitism in a Hudson Bay population of common eiders, testing predictions from two alternative hypotheses on the role of relatedness in CBP. Some models predict host-parasite relatedness, others predict that parasites avoid close relatives as hosts. To distinguish between the alternatives, we use a novel approach, where the relatedness of host-parasite pairs is tested against the spatial population trend in pairwise relatedness. We estimate parasitism, nest take-over and relatedness with protein fingerprinting and bandsharing analysis of egg albumen, nondestructively sampled from each new egg in the nest throughout the laying period. The results refute the hypothesis that parasites avoid laying eggs in the nests of related hosts, and corroborate the alternative of host-parasite relatedness. With an estimated r of 0.12-0.14, females laying eggs in the same nest are on average closer kin than nesting neighbour females. Absence of a population trend in female pairwise relatedness vs. distance implies that host-parasite relatedness is not only an effect of strong natal philopatry: some additional form of kin bias is also involved.     

Natural history and behaviour of Dunatothrips aneurae Mound (Thysanoptera: Phlaeothripidae), a phyllode‐gluing thrips with facultative pleometrosis

Thrips of the genus Dunatothrips (Thysanoptera: Phlaeothripidae) construct domiciles by tying phyllodes of Australian Acacia trees together with silk‐like glue. Females often co‐found domiciles (pleometrosis), an apparently cooperative behaviour that potentially provides insights into social evolution. However, little is known about their basic natural history, limiting the scope for testable predictions. Here, we address this crucial prerequisite step by investigating some key outstanding questions in the most common species, D. aneurae, on its host, Acacia aneura. We detail distribution in space and time, mating, dispersal, domicile building and defence. Dunatothrips aneurae was distributed in loosely reproductively synchronized patches, and tended to prefer east‐facing, terminal phyllodes on thin‐phyllode A. aneura varieties. Mature domiciles contained middens, concentrated areas of waste, suggesting active maintenance of domiciles and the potential for the division of labour. We observed inbreeding and outbreeding. Dunatothrips aneurae males engaged in short, truncated matings with sisters before dispersing locally, mating with females in nearby immature domiciles; longer distance dispersal, although it must happen, is still undocumented. Males and females mated multiply. Lone females required male presence to initiate domiciles, constructed them without male help and lost wings on nesting by abscission. Silk production occurred well before egg laying. Aggression or defence appeared to be entirely absent. Taken together, these observations suggest that research into co‐founding behaviour should focus on: (1) local crowding; (2) lack of aggression; and (3) potential division of labour with respect to egg production, silk production and domicile maintenance. These results should provide a springboard for questions on the potential evolution of cooperation in this species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 802‐816.     

A highly resolved food web for insect seed predators in a species‐rich tropical forest

The top‐down and indirect effects of insects on plant communities depend on patterns of host use, which are often poorly documented, particularly in species‐rich tropical forests. At Barro Colorado Island, Panama, we compiled the first food web quantifying trophic interactions between the majority of co‐occurring woody plant species and their internally feeding insect seed predators. Our study is based on more than 200 000 fruits representing 478 plant species, associated with 369 insect species. Insect host‐specificity was remarkably high: only 20% of seed predator species were associated with more than one plant species, while each tree species experienced seed predation from a median of two insect species. Phylogeny, but not plant traits, explained patterns of seed predator attack. These data suggest that seed predators are unlikely to mediate indirect interactions such as apparent competition between plant species, but are consistent with their proposed contribution to maintaining plant diversity via the Janzen–Connell mechanism.     

A review of predation as a limiting factor for bird populations in mesopredator‐rich landscapes: a case study of the UK

The impact of increasing vertebrate predator numbers on bird populations is widely debated among the general public, game managers and conservationists across Europe. However, there are few systematic reviews of whether predation limits the population sizes of European bird species. Views on the impacts of predation are particularly polarised in the UK, probably because the UK has a globally exceptional culture of intensive, high‐yield gamebird management where predator removal is the norm. In addition, most apex predators have been exterminated or much depleted in numbers, contributing to a widely held perception that the UK has high numbers of mesopredators. This has resulted in many high‐quality studies of mesopredator impacts over several decades. Here we present results from a systematic review of predator trends and abundance, and assess whether predation limits the population sizes of 90 bird species in the UK. Our results confirm that the generalist predators Red Fox (Vulpes vulpes) and Crows (Corvus corone and C. cornix) occur at high densities in the UK compared with other European countries. In addition, some avian and mammalian predators have increased numerically in the UK during recent decades. Despite these high and increasing densities of predators, we found little evidence that predation limits populations of pigeons, woodpeckers and passerines, whereas evidence suggests that ground‐nesting seabirds, waders and gamebirds can be limited by predation. Using life‐history characteristics of prey species, we found that mainly long‐lived species with high adult survival and late onset of breeding were limited by predation. Single‐brooded species were also more likely to be limited by predation than multi‐brooded species. Predators that depredate prey species during all life stages (i.e. from nest to adult stages) limited prey numbers more than predators that depredated only specific life stages (e.g. solely during the nest phase). The Red Fox and non‐native mammals (e.g. the American Mink Neovison vison) were frequently identified as numerically limiting their prey species. Our review has identified predator–prey interactions that are particularly likely to result in population declines of prey species. In the short term, traditional predator‐management techniques (e.g. lethal control or fencing to reduce predation by a small number of predator species) could be used to protect these vulnerable species. However, as these techniques are costly and time‐consuming, we advocate that future research should identify land‐use practices and landscape configurations that would reduce predator numbers and predation rates.     

Predation risk alters interactions among species: competition and facilitation between ants and nesting birds in a boreal forest

Although interactions between species are often assumed to be fixed, theory and empirical evidence suggest that they may be quite variable, changing in the presence of other species or environmental conditions. The interaction between ants and nesting birds exhibits such variability, ants sometimes being predators of bird nests and other times protectors of them. Hypothesizing that predation risk might be a critical factor in altering the interaction of ants with birds, I investigated the interaction of wood ants Formica aquilonia with nesting birds under different levels of predation risk. In a controlled field experiment, I allowed tits ( Parus major, P. caeruleus, P. ater ) and pied flycatchers ( Ficedula hypoleuca ) to select nest boxes in trees with ants (ant trees) or trees without ants. I found that birds usually nested in trees without ants, apparently to avoid the danger of injury from encounters with ants. Nesting in ant trees occurred mainly in the habitat where risk of predation was highest (along the forest edge), and with the bird taxa that lost nests most frequently in trees without ants (tits). Tits nesting on the forest edge achieved significantly greater nesting success, and fledged significantly more young, in ant trees compared with trees without ants. As the season progressed, ant traffic increased in trees without nesting birds, but decreased in trees with nesting birds, indicating that the outcome of interference competition between ants and nesting birds was reversed under increased predation risk. These results support the idea that predation risk can shift species interactions from predominately competitive processes to predominately facilitative processes.     

Food, vigilance, and sperm: the role of male direct benefits in the evolution of female preference in a polygamous bird

The adaptive significance of female selection of copulationpartners remains unresolved, particularly in polygamous specieswhere males do not provide paternal care. In these species thepossibility that direct benefits other than paternal care mayplay an important role in the evolution of female choice hasreceived little attention. I tested whether direct benefitsare associated with female choice in the polygamous feral fowl,Gallus g. domesticus, where females prefer socially dominantcopulation partners and males do not care for the young butdo provide females with three commodities: food, vigilance,and sperm. I used a combination of empirical and experimentaldata to show that male propensity to offer food and vigilance,but not sperm, was positively associated with male social status,suggesting that the provision of these resources may be costlyand condition dependent in males. Copulation success was correlatedwith male status but not with the number of feedings a femalereceived from a male, indicating that a female preferred dominantpartners that in general provided any female with more food,rather than partners that provided only her with more food,consistent with the idea that females may use male resourceprovisioning as a proximate mechanism to assess male condition.Together, these results indicate that male resources provisioningis (1) tightly linked to male social status, (2) a potentialindicator of male condition and possibly genetic quality, and(3) a potential criterion for females to select dominant partners,thus playing an important role in the evolution of partner choiceeven in polygamous species lacking paternal care.     

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.