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.     

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.     

Habitat preferences and habitat restoration options for small‐bodied and juvenile fish species in the northern Murray–Darling Basin

Degradation of instream habitats in the northern Murray–Darling Basin has occurred through numerous stressors, including siltation, clearing of bankside vegetation, intrusion of livestock and impacts of pest species. A better understanding of habitat preferences of native fish species could help guide future instream habitat restoration actions. The habitat choices of seven native fish species, juvenile Murray Cod (Maccullochella peelii), juvenile Golden Perch (Macquaria ambigua ambigua), juvenile Silver Perch (Bidyanus bidyanus), adult Murray–Darling Rainbowfish (Melanotaenia fluviatilis), adult Olive Perchlet (Ambassis agassizii), adult Un‐specked Hardyhead (Craterocephalus stercusmuscarum fulvus) and adult carp gudgeons (Hypseleotris spp.) were tested in preference troughs to help inform potential habitat restoration actions in the Condamine catchment. Each species was given a choice between pair combinations of open sandy habitat, submerged macrophytes, emergent plants and rocky rubble. Habitat preferences varied between species. Murray Cod, Golden Perch, carp gudgeons and Olive Perchlets preferred structure over open sandy habitat, whilst juvenile Silver Perch, Un‐specked Hardyhead and Murray–Darling Rainbowfish did not avoid open sandy habitats. Juvenile Murray Cod preferred rocky rubble habitat over all other habitat choices. Use of complex rock piles to provide nursery habitat for Murray Cod populations is a potential restoration option. Introduction of rock could also benefit Golden Perch and carp gudgeons. Use of emergent plants, submerged macrophytes and rocky rubble for habitat restoration all appear to have merit for one or more species of small‐bodied fishes or juvenile stages of larger sized fishes. Rocky rubble or floating attached macrophytes could be viable restoration options in areas too turbid to establish submerged macrophytes. These habitat interventions would complement existing actions such as re‐snagging and provision of fish passage to assist with sustainable management of native fish populations.     

Seasonal patterns in reproductive success of temperate‐breeding birds: Experimental tests of the date and quality hypotheses

For organisms in seasonal environments, individuals that breed earlier in the season regularly attain higher fitness than their late‐breeding counterparts. Two primary hypotheses have been proposed to explain these patterns: The quality hypothesis contends that early breeders are of better phenotypic quality or breed on higher quality territories, whereas the date hypothesis predicts that seasonally declining reproductive success is a response to a seasonal deterioration in environmental quality. In birds, food availability is thought to drive deteriorating environmental conditions, but few experimental studies have demonstrated its importance while also controlling for parental quality. We tested predictions of the date hypothesis in tree swallows (Tachycineta bicolor) over two breeding seasons and in two locations within their breeding range in Canada. Nests were paired by clutch initiation date to control for parental quality, and we delayed the hatching date of one nest within each pair. Subsequently, brood sizes were manipulated to mimic changes in per capita food abundance, and we examined the effects of manipulations, as well as indices of environmental and parental quality, on nestling quality, fledging success, and return rates. Reduced reproductive success of late‐breeding individuals was causally related to a seasonal decline in environmental quality. Declining insect biomass and enlarged brood sizes resulted in nestlings that were lighter, in poorer body condition, structurally smaller, had shorter and slower growing flight feathers and were less likely to survive to fledge. Our results provide evidence for the importance of food resources in mediating seasonal declines in offspring quality and survival.     

Distribution,movement, and microhabitat use of the introduced predatory snail Euglandina rosea in Hawaii: implications for management

The purposeful introduction of the land snail Euglandina rosea, which feeds exclusively on snails and slugs, has been implicated as a major factor in the decline of diverse Pacific island land snail faunas. We report on the distribution, movement patterns, and microhabitat preferences of E. rosea in a gulch in the Waianae Mountains, Oahu, Hawaii, because such data will help focus management actions at a local scale to protect native snail populations in areas where E. rosea is established. The Waianae Mountains harbor many endangered or threatened snails, most currently found in isolated habitat patches near the ridges. Conversely, most living individuals (28/29) and shells (46/56) of E. rosea were collected within the gulch, which supported higher densities of other native and non‐native snails, and was cooler and more moist than the ridges. Thirteen individuals of E. rosea were tracked (eight directly using a bobbin and thread method, and five indirectly by mark–recapture); most (10/13) moved on average <2.5 m per week (range 0.1–25.21 m), and all stayed within the gulch. Members of E. rosea preferred leaf litter over open, fern/shrub, or wood microhabitats. There were large differences in the population density of E. rosea over small spatial scales, indicating that there may be places where native snail populations could persist even in areas where populations of E. rosea are established. Identifying areas with differing population densities of E. rosea is critical for not only understanding why some native snail species may be more vulnerable to extinction, but also to locate areas where predation pressure is low and conservation efforts will be most likely to succeed.     

Assemblage structure and dynamics of terrestrial birds in the southwest Amazon: a camera‐trap case study

Habitat spatial distribution, seasonal variation, and activity patterns influence changes in vertebrate assemblages over time. Terrestrial birds play major roles in the dynamics of tropical forests, but there are few effective methods to study these species due to their cryptic coloration and elusive behavior. We used camera‐trap data collected during 16 mo (February 2017–June 2018) to describe the terrestrial avifauna in southeastern Peru, assess to what extent the composition of terrestrial avifauna changes among seasons and across two major habitats (terra firme and floodplain forests), and determine daily activity patterns of terrestrial birds. We used overlap analyses to examine temporal co‐occurrence between ecologically similar and sympatric species. Camera traps recorded 16 species, including eight species in the family Tinamidae. Capture rates were highest for Pale‐winged Trumpeters (Psophia leucoptera; Psophiidae) and Gray‐fronted Doves (Leptolila rufaxilla; Columbidae). Species composition did not differ between habitats or seasons, and capture rates between habitats only differed for White‐throated Tinamous (Tinamus guttatus). Overlaps of activity patterns were high between ecologically similar species and species found in terra firme habitats (White‐throated Tinamous and Cinereous Tinamous, Crypturellus cinereus) and in both habitat types (Pale‐winged Trumpeters and Gray‐fronted Doves). Low numbers of captures of possibly locally rare or less abundant species hindered a complete analysis of spatial and seasonal patterns of terrestrial bird assemblages. We suggest a greater sampling effort and greater spatial replication to better understand the spatial and seasonal dynamics of the terrestrial avifauna. Further studies that assess the mechanisms that allow the coexistence of sympatric tinamous would be valuable, both in our study area and elsewhere. The use of camera traps in long‐term monitoring projects proved to be an effective tool for monitoring terrestrial birds, identifying cryptic and often rare animals to species level, and providing valuable ecological information at species and community levels.     

Pockmarks enhance deep‐sea benthic biodiversity: a case study in the western Mediterranean Sea

Aim Pockmarks are craters on the sea floor formed by sub‐sea‐floor fluid expulsions, which occur world‐wide at all ocean depths. These habitats potentially host a highly specialized fauna that can exploit the hydrocarbons released. Pockmarks at relatively shallow depths can be easily destroyed by human activities, such as bottom trawling. In the present study, we investigated the combined effects of sea‐floor heterogeneity, rate of fluid emission and trophic conditions of different pockmarks on the biodiversity of the deep‐sea assemblages. Location Continental slope of the Gulf of Lions, western Mediterranean Sea, at water depths from 265 to 434 m. Methods We investigated the biodiversity associated with sea‐floor pockmarks that are both inactive and that have active gas emissions. Control sites were selected on the sea floor outside the influence of the gas seepage, both within and outside the pockmark fields. We examined the combined effects of: (i) sea‐floor heterogeneity; (ii) variable levels of fluid (gas) emissions; and (iii) trophic characteristics of the meiofaunal assemblage structure and nematode diversity. Results Sediments within the pockmark fields had lower meiofaunal abundance and biomass when compared with the surrounding sediments that were not influenced by the gas seepage. Although several higher taxa were absent in the pockmarks (e.g. Turbellaria, Tardigrada, Cumacea, Isopoda, Tanaidacea, Nemertina and Priapulida, which were present in the control areas), the richness of the nematode species within all of these pockmarks was very high. About 25% of the total species encountered in the deep‐sea sediments of the investigated areas was exclusively associated with these pockmarks. Main conclusions We conclude that both active and inactive pockmarks provide significant contributions to the regional (gamma) diversity of the continental slope in the western Mediterranean Sea, and thus the protection of these special and fragile habitats is highly relevant to the conservation of deep‐sea biodiversity.     

Parental and genetic factors rather than nesting forest patch size affect physiological traits of offspring in an area‐sensitive passerine: an experimental study

Our understanding of the effects of habitat loss on individual performance is limited due to a lack of experimental studies that take the potential genetic and parental effects producing phenotypic variation into consideration. To assess the relative role of habitat loss on offspring phenotype while controlling for the confounding effects of genetic and parental variation we performed a partial cross‐fostering experiment using the Eurasian Treecreeper Certhia familiaris. The experiment was carried out by swapping half the nestlings in a brood between small and large nesting forest patches to determine the effect of nesting forest patch size on five nestling traits reflecting morphological size, body condition, physiological stress and inflammation status. There was no effect of nesting forest patch size on the offspring traits examined. Instead, we found evidence of genetic and early parental effects on all traits except inflammation status, as well as parental effects after cross‐fostering for all of the measured offspring traits. Our results suggest that genetic and parental effects should be taken into account when making inferences about species’ responses to habitat loss.     

Diversification in tropics and subtropics following the mid‐Miocene climate change: A case study of the spider genus Nesticella

Caves may offer suitable refugia for troglophilic invertebrates during periods of unfavourable climatic conditions because of their stable microclimates. As a consequence, allopatric divergence from their epigean counterparts may occur, leading to formation of truly hypogean communities (the Climatic Relict Hypothesis). Unlike the well‐studied effects of Pleistocene glaciations, we know little about how ancient climate changes drove the development of cave‐dwelling organisms living at both middle and lower latitudes. We investigate the evolutionary history of the troglophilic spider genus Nesticella (Araneae, Nesticidae) in relation to Asian Neogene (23–2.6 Ma) climatic changes. Our analyses discern clear differences in the evolution of the two main clades of Nesticella, which occur in temperate/subtropical and tropical latitudes. Eastern Asian Nesticella gradually evolved greater sedentariness and a strict subterranean lifestyle starting from the middle Miocene Epoch (~15–14 Ma) in conjunction with the progressive deterioration of the climate and vegetational shifts. Caves appear to have acted as refugia because of their internally uniform temperature and humidity, which allowed these spiders to survive increasing external seasonality and habitat loss. In contrast, a uniform accumulation of lineages, long‐lasting times for dispersals and the lack of a comparable habitat shifting characterized the tropical lineage. This difference in pattern likely owes to the mild effects of climate change at low latitudes and the consequent lack of strong climatic drivers in tropical environments. Thus, the mid‐Miocene climatic shift appears to be the major evolutionary force shaping the ecological differences between Asian troglophilic invertebrates and the driver of the permanent hypogean communities in middle latitudes.     

Determinants of fine‐scale homogenization and differentiation of native freshwater fish faunas in a Mediterranean Basin: implications for conservation

Aim Increasing threats to freshwater biodiversity are rapidly changing the distinctiveness of regional species pools and local assemblages. Biotic homogenization/differentiation processes are threatening the integrity and persistence of native biodiversity patterns at a range of spatial scales and pose a challenge for effective conservation planning. Here, we evaluate the extent and determinants of fine‐scale alteration in native freshwater fish assemblages among stream reaches throughout a large river basin and consider the implications of these changes for the long‐term conservation of native fishes. Location Guadiana River basin (South‐Western Iberian Peninsula). Methods We quantified the magnitude of change in compositional similarity between observed and reference assemblages and its potential effect on natural patterns of compositional distinctiveness. Reference assemblages were defined as the native species expected to occur naturally (in absence of anthropogenic alterations) and were reconstructed using a multivariate adaptive regression splines predictive model. We also evaluated the role of habitat degradation and introduced species as determinants of biotic homogenization/differentiation. Results We found a significant trend towards homogenization for native fish assemblages. Changes in native fish distributions led to the loss of distinctiveness patterns along natural environmental gradients. Introduced species were the most important factor explaining the homogenization process. Homogenization of native assemblages was stronger in areas close to reservoirs and in lowland reaches where introduced species were more abundant. Main conclusions The implementation of efficient conservation for the maintenance of native fish diversity is seriously threatened by the homogenization processes. The identification of priority areas for conservation is hindered by the fact that the most diverse communities are vanishing, which would require the selection of broader areas to adequately protect all the species. Given the principal role that introduced species play in the homogenization process and their relation with reservoirs, special attention must be paid to mitigating or preventing these threats.     

Effects of patch size and basal area on avian taxonomic and functional diversity in pine forests: Implication for the influence of habitat quality on the species–area relationship

Relationships between avian diversity and habitat area are assumed to be positive; however, often little attention has given to how these relationships can be influenced by the habitat structure or quality. In addition, other components of biodiversity, such as functional diversity, are often overlooked in assessing habitat patch value. In the Sandhills Ecoregion of Georgia, USA, we investigated the relationship between avian species richness and functional diversity, forest basal area, and patch size in pine forests using basal area as a surrogate for overstory structure which in turn impacts vegetation structure and determines habitat quality within a patch. We conducted bird surveys in planted mature pine stands, during breeding season of 2011. We used three classes of stand basal area (BA): OS, overstocked (BA ≥ 23 m²/ha); FS, fully/densely stocked (13.8 m²/ha ≤ BA < 23 m²/ha); and MS, moderately stocked (2.3 m²/ha ≤ BA < 13.8 m²/ha). MS patches showed more structural diversity due to higher herbaceous vegetation cover than other two pine stocking classes of patches. Total species richness and functional richness increased with the size of MS patches, whereas functional divergence decreased with the size of OS patches (p < 0.05). Functional richness tended to be lower than expected as the size of OS patches increased. Greater richness of pine–grassland species was also found at MS patches. Percent cover of MS patches within a landscape influenced positively the richness of pine–grassland species (p < 0.05). Our results suggest that (a) avian species–habitat area relationship can be affected by habitat quality (structural diversity) and varies depending on diversity indices considered, and (b) it is important to maintain moderate or low levels of pine basal area and to preserve large‐sized patches of the level of basal area to enhance both taxonomic and functional diversity in managed pine forests.     

Biogeographical patterns of algal communities in the Mediterranean Sea: Cystoseira crinita‐dominated assemblages as a case study

Aim The aim of this study was to describe the composition, community structure and biogeographical variation of subtidal algal assemblages dominated by the brown alga Cystoseira crinita across the Mediterranean Sea. Location The Mediterranean coast, from Spain (1°25′ E) to Turkey (30°26′ E). Methods Data on the species composition and structure of assemblages dominated by the species C. crinita were collected from 101 sites in nine regions across the Mediterranean Sea. Multivariate and univariate statistical tools were used to investigate patterns of variation in the composition of the assemblages among sites and regions, and to compare these with previously defined biogeographical regions. Linear regressions of species richness versus longitude and versus latitude were also carried out to test previously formulated hypotheses of biodiversity gradients in the Mediterranean Sea. Results The main features characterizing C. crinita‐dominated assemblages across the Mediterranean included a similar total cover of species, a similar cover of C. crinita, and consistency in the presence of the epiphyte Haliptilon virgatum. Biogeographical variation was detected as shifts in relative abundances of species among regions, partly coinciding with previously described biogeographical sectors. A significant positive correlation was found between species richness and latitude, while no significant correlation was detected between species richness and longitude. Main conclusions The patterns of variation in community structure detected among the studied regions reflected their geographical positions quite well. However, latitude seemed to contribute more to the explanation of biological patterns of diversity than did geographical distances or boundaries, which classically have been used to delimit biogeographical sectors. Moreover, the positive correlation between species richness and latitude reinforced the idea that latitude, and possibly temperature as a related environmental factor, plays a primary role in structuring biogeographical patterns in the Mediterranean Sea. The lack of correlation between species richness and longitude contradicts the notion that there is a decrease in species richness from west to east in the Mediterranean, following the direction of species colonization from the Atlantic.     

Identifying spawning sites and other critical habitat in lotic systems using eDNA “snapshots”: A case study using the sea lamprey Petromyzon marinus L.

Many aquatic species of conservation concern exist at low densities and are inherently difficult to detect or monitor using conventional methods. However, the introduction of environmental (e)DNA has recently transformed our ability to detect these species and enables effective deployment of limited conservation resources. Identifying areas for breeding, as well as the ecological distribution of species, is vital to the survival or recovery of a conservation species (i.e., areas of critical habitat). In many species, spawning events are associated with a higher relative abundance of DNA released within an aquatic system (i.e., gametes, skin cells etc.), making this the ideal time to monitor these species using eDNA techniques. This study aims to examine whether a “snapshot” eDNA sampling approach (i.e., samples taken at fixed points in chronological time) could reveal areas of critical habitat including spawning sites for our target species Petromyzon marinus. We utilized a species‐specific qPCR assay to monitor spatial and temporal patterns in eDNA concentration within two river catchments in Ireland over three consecutive years. We found that eDNA concentration increased at the onset of observed spawning activity and patterns of concentration increased from downstream to upstream over time, suggesting dispersal into the higher reaches as the spawning season progressed. We found P. marinus to be present upstream of several potential barriers to migration, sometimes in significant numbers. Our results also show that the addition of a lamprey‐specific fish pass at an “impassable” weir, although assisting in ascent, did not have any significant impact on eDNA concentration upstream after the pass had been installed. eDNA concentration was also found to be significantly correlated with both the number of fish and the number of nests encountered. The application of snapshot sampling techniques for species monitoring therefore has substantial potential for the management of low‐density species in fast‐moving aquatic systems.     

Incorporating social preferences into the ecological limits of hydrologic alteration (ELOHA): a case study in the Yampa‐White River basin,Colorado

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Life history and population dynamics of a tree species in tropical semi‐arid climate: A case study with Cordia oncocalyx

In semi‐arid climates, plant population dynamics are strongly influenced by the amount and temporal distribution of rainfall. We monitored a population of the tree species Cordia oncocalyx (Boraginaceae) for 24 months in the dry thorny woodland of semi‐arid northeastern Brazil, to investigate which life‐history traits allow this tree to be locally dominant. We used horizontal life tables and a Lefkovitch matrix and tested for relationships among demographic parameters of seedling, infant, juvenile, immature, virginile and reproductive ontogenetic stages with rainfall and canopy openness. Germination and recruitment occurred in the rainy months, and dry‐season mortality occurred only in seedlings (76% and 100%, first and second years, respectively) and infants (3% and 6%). Juveniles showed greater height growth under more open canopies (Spearman correlation coefficient = 0.24), suggesting that light availability influences growth. The population growth rate was λ = 1.0336, and the highest sensitivity occurred in the infant‐juvenile transition. Our results show light as a restrictive growth factor for plants in the juvenile stage and confirm the strong influence of rainfall on the dynamics of trees in a seasonally dry environment. The formation of a persistent seed bank with germination concentrated at the rainfall onset but spreading over the rainy season are strategies that hedge bets before establishment. The formation of a bank of infants, which can resume growth as soon as there is water, hedges bets after establishment. We attribute the positive population growth rate of Cordia oncocalyx to survival strategies allowing bet‐hedging both before and after establishment.     

Nitrogen Depletion and Redistribution by Free‐Ranging Cattle in the Restoration Process of Mosaic Landscapes: The Role of Foraging Strategy and Habitat Proportion

In a mosaic landscape in N‐Belgium (W‐Europe), consisting of forest, grassland, and wooded pasture on former agricultural land, we assessed nitrogen redistribution by free‐ranging cattle (±0.2 animal units ha^?1 yr^?1). We examined if the spatial redistribution of nitrogen among habitats by cattle could restore nutrient‐poor conditions in preferred foraging habitats, and conversely whether such translocation could lead to extreme eutrophication in preferred resting habitats. We used nitrogen content of different diet classes, habitat use, foraging and defecation behavior, weight gain, and nitrogen losses in the actual situation to explore four different habitat proportion scenarios and two different foraging strategies to calculate a net nitrogen balance per habitat. An atmospheric deposition of 30 kg N ha^?1 yr^?1 with varying interception factors according to the habitat types was included in an integrated nitrogen balance. All scenarios showed a net nitrogen transport from grassland and wooded pasture to forest habitat. We found that nitrogen redistribution strongly depends on habitat proportion. Nitrogen losses from preferred grassland habitat can be high, given its proportion is small. Depletion is only to be expected at excretion‐free areas and probably is of minor importance to trigger the establishment of woody species. In general, nitrogen transported by cattle was much lower than input by atmospheric deposition, but grazing can compensate for high N inputs in excretion‐free areas and maintain grassland types that support critical loads of 20–25 kg N ha^?1 yr^?1. In none of the scenarios, N transport by cattle resulted in the exceeding of critical nitrogen loads to vulnerable forest ground vegetation.