Comparing aerodynamic efficiency in birds and bats suggests better flight performance in birds

Flight is one of the energetically most costly activities in the animal kingdom, suggesting that natural selection should work to optimize flight performance. The similar size and flight speed of birds and bats may therefore suggest convergent aerodynamic performance; alternatively, flight performance could be restricted by phylogenetic constraints. We test which of these scenarios fit to two measures of aerodynamic flight efficiency in two passerine bird species and two New World leaf-nosed bat species. Using time-resolved particle image velocimetry measurements of the wake of the animals flying in a wind tunnel, we derived the span efficiency, a metric for the efficiency of generating lift, and the lift-to-drag ratio, a metric for mechanical energetic flight efficiency. We show that the birds significantly outperform the bats in both metrics, which we ascribe to variation in aerodynamic function of body and wing upstroke: Bird bodies generated relatively more lift than bat bodies, resulting in a more uniform spanwise lift distribution and higher span efficiency. A likely explanation would be that the bat ears and nose leaf, associated with echolocation, disturb the flow over the body. During the upstroke, the birds retract their wings to make them aerodynamically inactive, while the membranous bat wings generate thrust and negative lift. Despite the differences in performance, the wake morphology of both birds and bats resemble the optimal wake for their respective lift-to-drag ratio regimes. This suggests that evolution has optimized performance relative to the respective conditions of birds and bats, but that maximum performance is possibly limited by phylogenetic constraints. Although ecological differences between birds and bats are subjected to many conspiring variables, the different aerodynamic flight efficiency for the bird and bat species studied here may help explain why birds typically fly faster, migrate more frequently and migrate longer distances than bats.     

Conservation of tropical forest birds in countryside habitats

The pressing need to increase agricultural production often seems at odds with conserving biodiversity. We find that if managed properly, the tropical countryside may provide a substantial opportunity for tropical bird conservation. We detected 144 bird species from 29 families in agricultural areas outside of extensive native forest in southern Costa Rica. The majority of the species detected were observed foraging, often kilometres from extensive native forest. We estimate that 46% of those native to this region (excluding nocturnal species and waterfowl) are utilizing the countryside in some manner. Forecasts of biodiversity change under various land-use scenarios indicate that policies that affect habitat composition could greatly impact the persistence of these species in the countryside. In particular, if tall trees and edge habitats were removed from this landscape, we predict that bird richness in the countryside would decline by approximately 40%.     

Analysis of the aerodynamic performance of birds during bounding flight

A new analysis of bounding flight is presented. The rise and fall in this style of flight are explicitly taken into account, and the aerodynamic performance required to achieve curvature of the flight path during the powered phase is included. It is shown that this flight style may be viewed as a strategy for increasing flight speed without incurring the penalty of increased energy expenditure associated with increased flight speed under conditions of steady horizontal flight.     

Conservation value of degraded habitats for forest birds in southern Peninsular Malaysia

Clearance of tropical forest for agricultural purposes is generally assumed to seriously threaten the survival of forest species. In this study, we quantified the conservation value, for forest bird species, of three degraded habitat types in Peninsular Malaysia, namely rubber tree plantations, oil palm plantations, and open areas. We surveyed these degraded habitats using point counts to estimate their forest bird species richness and abundance. We assessed whether richness, abundance, and activities of different avian dietary groups (i.e. insectivores and frugivores) varied among the habitats. We identified the critical habitat elements that accounted for the distribution of forest avifauna in these degraded habitats. Our results showed that these habitats harboured a moderate fraction of forest avifauna (approximately 46–76 species) and their functions were complementary (i.e. rubber tree plantations for moving; open habitats for perching; shrubs in oil palm plantations for foraging). In terms of species richness and abundance, rubber tree plantations were more important than oil palm plantations and open habitats. The relatively high species richness of this agricultural landscape was partly due to the contiguity of our study areas with extensive forest areas. Forecasts of forest-species presence under various canopy cover scenarios suggest that leaving isolated trees among non-arboreal crops could greatly attract relatively tolerant species that require tree canopy. The conservation value of degraded habitats in agricultural landscapes seems to depend on factors such as the type of crops planted and distance to primary forest remnants.     

Tolerance to habitat fragmentation influences the colonization of new habitat by forest birds

We examined the relationship between the ability of bird species to persist in fragmented forests and their ability to colonize new forest habitat. Using a long-term data set on the colonization of a forest plantation, we tested the hypothesis that bird species tolerant to habitat fragmentation would detect and colonize the new habitat faster than intolerant species. The forest plantation under study is situated on an area of land reclaimed from the sea (a polder) in the central part of The Netherlands. We constructed an index of tolerance to habitat fragmentation and included it as a predictor variable in a set of three logistic regression models that compared the probability of colonization over four consecutive time periods. After controlling statistically for the effects of regional incidence, preferred habitat and life-history characteristics, there was a significant effect of tolerance to fragmentation on the ability of species to colonize the plantation, and a marginal effect on the timing of colonization. We then examined the effect of the same index of tolerance to fragmentation on colonization patterns over a larger spatial scale. Multivariate regression models showed that the proportion of three polders of different ages occupied by forest bird species was dependent upon the regional incidence of a species, its preferred habitat and its tolerance to fragmentation. The results support the hypothesis that species tolerant to habitat fragmentation detect and colonize new habitat faster than those intolerant to habitat fragmentation.     

Rapid colonization and turnover of birds in a tropical forest treefall gap

Ecological disturbance is an important factor that influences the abundance and distribution of species. Treefalls are a prominent source of disturbance in tropical forests, but robust characterization of community change after treefalls requires baseline data that are often not available. We capitalized on 25 yr of avian mark–recapture data from a lowland moist forest in central Panama to investigate the timescale of colonization and persistence of birds in a newly formed treefall gap. We compared bird species assemblages pre- and post-treefall to explore how the disturbance affected specific foraging guilds and overall assemblage structure (abundance and alpha diversity). We documented rapid colonization (i.e., within five months post-treefall) of the treefall gap by birds. Abundance and alpha diversity increased following the treefall, but both remained relatively constant in a nearby control plot. At the guild level, frugivores spiked in abundance and nectarivores (i.e., hummingbirds) increased in alpha diversity following the treefall. These results are in agreement with those of previous spatial studies of gap dynamics and suggest that certain tropical frugivores and nectarivores have a remarkable ability to rapidly find and exploit preferred resources and microhabitats embedded in a landscape matrix. Assemblage abundance and alpha diversity decreased back to pre-treefall levels within 1 and 4 yr of the treefall, respectively. Thus, even large gaps may provide only ephemeral benefits, highlighting the importance of periodic disturbance for landscape-level persistence of species that use gaps.     

Conservation value of secondary forest habitats for endemic birds,a perspective from two widely separated tropical ecosystems

Tropical secondary forests are increasingly widespread, but their potential for conserving endemic birds remains unclear. Previous studies report different results; however all have been restricted to geographically discreet locations. This is important as different ecosystems are influenced by different external factors, possibly influencing conservation potential. Here we use consistent survey methods to examine how endemic bird richness varies between primary and secondary forest habitats in two widely separated tropical ecosystems, providing a more global context for evaluating the conservation value of secondary forests. Research was completed in Lambusango Forest Reserve (LFR) on Buton Island, Sulawesi, and Cusuco National Park (CNP), a Honduran cloud forest reserve. Bird communities in both forests were surveyed using 50 m radius point counts. Vulnerability assessments based on ecological theory on avifaunal assemblages were then conducted, which suggested endemics in LFR to be more susceptible to disturbance than those in CNP. Contrary to the results from our vulnerability assessments, endemics in CNP were less tolerant of moderate habitat modification than those in LFR. Richness of Mesoamerican endemics per study site declined significantly between core zone forest (6.34 ± 0.81) and more degraded forest in the boundary zone (3.86 ± 0.69). Richness of Wallacean endemics was similar in primary (4.89 ± 1.68) and disturbed secondary forest (4.52 ± 1.62). We recommend considering local and regional biogeographical and ecological factors when determining the conservation value of secondary forests, and suggest examples of potential importance, including differential community richness, influence of figs and human settlement patterns.     

Repopulation and colonization by birds in the Agmon Wetland,Israel

Lake Hula and its surrounding wetland in northern Israel were drained in the late 1950s and the dried wetlands were transformed into a diverse agricultural region with a 0.3 km2 nature reserve. A portion of the extinct Hula wetland was re-flooded in April 1994 by constructing a small lake, Agmon, and 90 km of canals. The purposes of this study were (a) to document the re-population and colonization of the new Agmon wetland by birds after its flooding, (b) to evaluate bird species richness in this new wetland in comparison to that in a nearby mature Hula Nature Reserve, and, (c) to investigate if the species originally present in the Hula Valley before the drainage had been re-established. The new lake has succeeded in attracting a large variety of water birds to the newly formed habitats, especially ducks, egrets and herons, plovers, waders and snipes. The distribution of bird species among the different habitats was not random. The northern section of the lake, which is shallow and has an open shore, had the largest number of species during 1995 and 1996 and was preferred by plovers, waders and snipes. A large roosting and breeding colony of thousands of egrets and herons was formed in the dense cattails ( Typha domingensis) in the southern section of the lake. Unlike the nearby Hula Nature Reserve, the new wetland lacks a large seasonally flooded area and is less attractive for feeding and breeding plovers, waders and stilts. The new wetland also has very little muddy area without vegetation and attracts few of the species that regularly feed in this habitat in the reserve. The new wetland enlarges the area of only a few of the habitats found in the nearby reserve and therefore attracts fewer species than it might otherwise. Recommendations for management of the new wetland include increased diversity of habitats, restriction of human activities and optimization of conflicts that have arisen between the needs of the wetland and those of nearby agriculture.     

Hybridization and the colonization of novel habitats by annual sunflowers

Although invasive plant species often have a hybrid ancestry, unambiguous evidence that hybridization has stimulated the evolution of invasive behaviors has been difficult to come by. Here, we briefly review how hybridization might contribute to the colonization of novel habitats, range expansions, and invasiveness and then describe work on hybrid sunflowers that forges a direct link between hybridization and ecological divergence. We first discuss the invasion of Texas by the common sunflower and show that the introgression of chromosomal segments from a locally adapted species may have facilitated range expansion. We then present evidence that the colonization of sand dune, desert floor, and salt marsh habitats by three hybrid sunflower species was made possible by selection on extreme or “transgressive” phenotypes generated by hybridization. This body of work corroborates earlier claims regarding the role of hybridization in adaptive evolution and provides an experimental and conceptual framework for ongoing studies in this area.     

Physical theory,origin of flight,and a synthesis proposed for birds

Neither flapping and running to take-off nor gliding from heights can be disproved as the assured evolutionary origin of self-powered flight observed in modern vertebrates. Gliding with set wings would utilize available potential energy from gravity but gain little from flapping. Bipedal running, important in avian phylogeny, possibly facilitated the evolution of flight. Based on physical principles, gliding is a better process for the origin of powered flight than the "ground-up" process, which physically is not feasible in space or time (considering air resistance, metabolic energy costs, and mechanical resistance to bipedal running). Proto-avian ancestors of Archaeopteryx and Microraptor probably flapped their sparsely feathered limbs synchronously while descending from leaps or heights, with such "flutter-gliding" presented as a synthesis of the two earlier theories of flight origin (making use of the available potential energy from gravity, involving wing thrusts and flapping, coping with air resistance that slows air speed, but effecting positive fitness value in providing lift and slowing dangerous falls).     

Diversity and abundance of birds in relation to forest fragmentation,habitat quality and heterogeneity

This study investigated the importance of habitat quality and habitat heterogeneity for the abundance and diversity of breeding birds in continuous forest and in forest fragments surrounded by farmland in central Sweden. Positive correlations were found between species number and area, volume of Aspen Populus tremula and habitat heterogeneity. Spatial segregation of habitats at a relatively fine-grained scale is suggested to allow for the co-occurrence of more species. The abundance of at least 18 of the species in this study was influenced by fragmentation, and nine of these species preferred fragments to forest sites. The total density of birds was higher in fragments than in forest sites, probably because several fragment species forage in farmland surrounding the sites and a few also forage at edges. Nine species were more common in forest sites than in fragments, but only one species was restricted to continuous forest. However, several fragments were relatively close to forests (150 m) and forest was common in larger scale contexts. The abundance of most species (25 of 33 species) in this study was correlated with habitat quality variables (i.e. variables measuring the size, volume and diversity of ‘tree species’). Among these habitat variables the most important was the occurrence of deciduous trees which seemed to be important for 14 species. The second most important habitat factor seemed to be the diameter of trees, which was positively correlated with the abundance of eight species of which five are hole-nesters. Among coniferous trees, six species were positively correlated with the volume of Norway Spruce Picea abies, whereas no species seemed to be correlated with the volume of Pine Pinus sylvestris.     

Energy density and its variation in space limit species richness of boreal forest birds

Aim An area’s ability to support species may be dependent not only on the total amount of available energy it contains but also on energy density (i.e. available energy per unit area). Acknowledging these two aspects of energy availability may increase mechanistic understanding of how increased energy availability results in increased species richness. We studied the relationship between energy density, its variation in space and boreal forest bird species richness and investigated two possible mechanisms: (1) metabolic constraints of organisms, and (2) increased resource availability for specialists. Location Protected areas in Finland’s boreal forest. Methods We tested whether bird species richness was best determined by total energy availability in an area or by energy density and its variation within the area, before and after including bird abundance in the models. We evaluated two main explanatory variables: tree growth reflecting the rate of energy production and tree volume as a measure of biomass. In addition, we modelled individual species’ responses to energy density and its variation, and evaluated the prediction of the metabolic constraints hypothesis that small species are limited by energy density whereas large species are limited by total energy availability in the area. Results Energy density and its variation were good predictors of species richness: together with abundance they explained 84% of variation in species richness (compared with 74% for abundance alone). Pure metabolic constraints were unlikely to explain this relationship. Instead, the mechanism probably involved increased habitat heterogeneity benefiting specialist species. Total energy availability was also an important factor determining species richness but its effect was indirect via abundance. Main conclusions Our results corroborate the importance of energy availability as a driver of species richness in forest bird communities, and they indicate that energy density and its variation in the landscape strongly influence species richness even after accounting for abundance.     

Landscape size affects the relative importance of habitat amount,habitat fragmentation,and matrix quality on forest birds

It is important to understand the relative effects of landscape habitat loss, habitat fragmentation, and matrix quality on biodiversity, so that potential management options can be appropriately ranked. However, their effects and relative importance may change with the size of the landscape considered because the multiple (and potentially conflicting) ecological processes that are influenced by landscape structure occur at different spatial scales (e.g. dispersal, predation, foraging). We estimated the relative effects of habitat loss, habitat fragmentation, and matrix quality (measured as the amount of forest, the proportion of forest area contained in large core forests, and the density of roads respectively) on fragmentation‐sensitive forest birds in southern Ontario, Canada using a range of landscape sizes (0.8–310 km²). We used three complementary statistical approaches to estimate relative effects of these correlated landscape factors – 1) multiple regression, 2) information theoretic (AIC) estimates of the most parsimonious model, and 3) multi‐model inference to average effects across all supported models. We controlled for spatial autocorrelation, local habitat, roadside sampling bias, time of day, season, habitat heterogeneity, and the interaction between the effects of habitat amount and fragmentation. We found that relative effects of habitat amount and fragmentation were scale dependent; habitat amount had a consistently positive effect that was consistent over more than two orders of magnitude in landscape area (~1–300 km²). In contrast, the effects of habitat fragmentation depended on the size of the landscape considered. Indeed, for veery Catharus fuscescens, habitat fragmentation had positive effects at one scale and negative effects at another. The effects of matrix quality were generally weak and changed little with scale. For the number of fragmentation sensitive species and the presence of veery, habitat amount was most important in large landscapes and habitat fragmentation in small landscapes but for the presence of ovenbird Seiurus aurocapilla, habitat amount was most important at all scales.     

Nest predation in forest birds: influence of predator type and predator's habitat quality

We used the introduction of a generalist nest predator, the red squirrel Tamiasciurus hudsonicus, and of a large herbivore, the Sitka black-tailed deer Odocoileus hemionus sitkensis, to the islands of Haida Gwaii (Queen Charlotte Islands, British Columbia, Canada) to study how predator assemblage and habitat quality and structure influenced nest predation in forest birds. We compared losses of natural nests to predators on islands with and without squirrels. We selected nine islands with or without squirrel or deer and used 506 artificial nests put on the ground or in shrubs to further analyse variation of nest predation with predator assemblage and habitat quality for the predators. For both natural and artificial nests predation risk was higher in presence of squirrels. But predation risk varied within island categories. In presence of squirrels it was highest in stands with mature conifers where it fluctuated from year to year, in response to fluctuations in squirrel abundance. Vegetation cover around the nest had little effect on nest predation by squirrels. Where squirrels were absent, nest predation concentrated near predictable food sources for corvids, the main native predators, and increased with decreasing vegetation cover, suggesting that removal of the vegetation by deer increased the risk of predation by native avian nest predators that use visual cues. Predation risk in these forests therefore varies in space and time with predator composition and with quality of the habitat from the predators' perspective. This temporal and spatial variation in predation risk should promote trade-offs in the response of birds to nest predation, rather than fine-tuned adaptations to a given predation pattern.     

Conservation potential of semi-natural habitats for birds in intensively-used agricultural landscapes

Agricultural intensification resulted in substantial loss of farmland biodiversity. Semi-natural habitats may be viewed as potential buffers of these adverse impacts, but a rigorous assessment of their capacity for supporting farmland biodiversity is lacking. In this study, we explored conservation potential of two different types of semi-natural habitats for birds in intensively-used agricultural landscapes – farmland hedges (i.e., linear strips of shrubby and tree vegetation) and open scrubland (i.e., scattered shrubs and abandoned orchards). Specifically, we tested whether the abundance and species richness of birds differ between these habitats considering various species traits, such as habitat affinity (i.e., forest, farmland and urban species), diet specialization (i.e., animal eaters, plant eaters, and omnivores) and conservation status (Species of European Conservation Concern). We found that open scrubland hosted on average 37.9 bird species and 122.6 individuals per 1 km² of the transect, whereas farmland hedges hosted only 19 species and 61.8 individuals per 1 km² of the transect. However, results have substantially changed if we considered the area of suitable habitat into account. More specifically, open scrubland hosted more bird species and individuals when we considered open habitat species and the area of open habitats, whereas farmland hedges had higher species diversity and individuals of woodland bird species when we considered the area of woodland habitats. Similarly, analyses of habitat affiliations of individual species corresponded to the whole-community patterns; and revealed that several woodland bird species were mainly associated with farmland hedges (e.g., Chaffinch Fringilla coelebs, Common Nightingale Luscinia megarhynchos and Blackcap Sylvia atricapilla), whereas the open scrubland was preferred by open habitat bird species (e.g., Corn Bunting Emberiza calandra, Quail Coturnix coturnix and Skylark Alauda arvensis). These results demonstrate that semi-natural habitats, both open scrubland and farmland hedges, have large potential for promotion and conservation of bird communities within intensively used agricultural landscapes, as both may have represented suitable habitats for species with different ecological requirements. Therefore, management measures focused on the enlargement of the area of these habitats, in combination with suitable management (e.g., regulating the progress of natural succession in open scrubland; increasing structural diversity of existing farmland hedges), may substantially contribute to bird conservation within agricultural landscapes.     

The use of agricultural, open and forest habitats by juvenile Mauritius Kestrels Falco punctatus

Knowledge of tropical raptor habitat use is limited and yet a thorough understanding is vital when trying to conserve endangered species. We used a well studied, reintroduced population of the vulnerable Mauritius Kestrel Falco punctatus to investigate habitat preferences in a modified landscape. We constructed a high resolution digital habitat map and radiotracked 13 juvenile Kestrels to quantify habitat preferences. We distinguished seven habitat types in our study area and tracked Kestrels from 71 to 130 days old during which they dispersed from their natal territory and settled within a home‐range after reaching independence. Mean home‐range size was 0.95 km² characterized by a bimodal pattern of intensity around the natal site and post‐independence home‐range. Compositional analysis showed that home‐ranges were located non‐randomly with respect to habitat but there was no evidence to suggest differential use of habitats within home‐ranges. Native and semi‐invaded forest and grassland were consistently preferred, whereas agriculture was used significantly less than other habitats. No difference was found between the available length of edge dividing native forest and grassland within a home‐range when compared to that available within a 2.35‐km buffer around their nest‐site, based on the maximum distance a juvenile was found to disperse. Repeating the analysis in three dimensions gave very similar results. Our results suggest that Mauritius Kestrels are not obligate forest dwellers as was once thought but can also exploit open habitats such as grassland. Kestrels may be using isolated mature trees within grassland as vantage points for hunting in the same way as they use the natural stratified forest structure. We suggest that the avoidance of agriculture is partly due to a lack of such vantage points. The conservation importance of forest degradation and agricultural encroachment is highlighted and comparisons with the habitat preferences of other tropical falcons are discussed.     

Aerodynamic corrections for the flight of birds and bats in wind tunnels

Few wind tunnel studies of animal flight have controlled or corrected for distortions to behaviour, physiology or flight aerodynamics representing the difference between flight in the tunnel and flight in free air. Aerodynamic correction factors are derived based on lifting-line theory and the method of images for an animal flying freely within closed- and open-section wind tunnels; the method is very similar to that used to model flight in ground effect, and as in ground effect the corrections to induced drag may be substantial. These correction factors are used to estimate bound wing circulation, drag and mechanical power for comparison with free flight, and to derive testable predictions of optimum flight strategies for an animal in a tunnel. In an open-section tunnel, mechanical power is increased compared to free flight, and the animal should fly at the tunnel centre. In a closed tunnel mechanical power is usually reduced, and substantial savings are available, particularly at low speeds, if the animal flies close to the tunnel roof. Anecdotal observations confirm that birds and bats adopt this strategy. The mechanical power-speed curve in a closed tunnel is flatter than the curve for free flight, and this may explain the flat metabolic power-speed curves for birds and bats obtained in some measurements.