Vertical stratification of an avian community in New Guinean tropical rainforest

Vertical stratification of avian communities has been studied in both temperate and tropical forests; however, the majority of studies used ground-based methods. In this study we used ground-to-canopy mist nets to collect detailed data on vertical bird distribution in primary rain forest in Wanang Conservation Area in Papua New Guinea (Madang Province). In total 850 birds from 86 species were caught. Bird abundance was highest in the canopy followed by the understory and lowest in the midstory. Overall bird diversity increased towards the canopy zone. Insectivorous birds represented the most abundant and species-rich trophic guild and their abundances decreased from the ground to canopy. The highest diversity of frugivorous and omnivorous birds was confined to higher vertical strata. Insectivorous birds did not show any pattern of diversity along the vertical gradient. Further, insectivores preferred strata with thick vegetation, while abundance and diversity of frugivores increased with decreasing foliage density. Our ground-to-canopy (0–27 m) mist netting, when compared to standard ground mist netting (0–3 m), greatly improved bird diversity assessment and revealed interesting patterns of avian community stratification along vertical forest strata.     

Bird Assemblage and Visitation Pattern at Fruiting Elmerrillia tsiampaca (Magnoliaceae) Trees in Papua New Guinea

Most tropical trees produce fleshy fruits that attract frugivores that disperse their seeds. Early demography and distribution for these tree species depend on the effects of frugivores and their behavior. Anthropogenic changes that affect frugivore communities could ultimately result in changes in tree distribution and population demography. We studied the frugivore assemblage at 38 fruiting Elmerrillia tsiampaca, a rain forest canopy tree species in Papua New Guinea. Elmerrillia tsiampaca is an important resource for frugivorous birds at our study site because it produces abundant lipid-rich fruits at a time of low fruit availability. We classified avian frugivores into functional disperser groups and quantified visitation rates and behavior at trees during 56 canopy and 35 ground observation periods. We tested predictions derived from other studies of plant–frugivore interactions with this little-studied frugivore assemblage in an undisturbed rain forest. Elmerrillia tsiampaca fruits were consumed by 26 bird species, but most seeds were removed by eight species. The most important visitors (Columbidae, Paradisaeidae and Rhyticeros plicatus) were of a larger size than predicted based on diaspore size. Columbidae efficiently exploited the structurally protected fruit, which was inconsistent with other studies in New Guinea where structurally protected fruits were predominantly consumed by Paradisaeidae. Birds vulnerable to predation foraged for short time periods, consistent with the hypothesis that predator avoidance enhances seed dispersal. We identified seven functional disperser groups, indicating there is little redundancy in disperser groups among the regular and frequent visitors to this tropical rain forest tree species.     

Patterns of resource use in an assemblage of birds in the canopy of a temperate alluvial forest

In our study, we assessed patterns of resource use in an assemblage of birds by observing their foraging behaviour from a crane in the canopy of a temperate alluvial forest. We selected 12 bird species and addressed seasonal changes in feeding activity during a 2-month period in spring focussing on average staying time and utilisation of crown strata in two tree species, the common oak (Quercus robur) and the sycamore maple (Acer pseudoplatanus). We further examined ecological characteristics of the trees (i. e., crown density) that are likely to influence resource use in birds. The selected birds differed in their preference for the tree species. Most birds preferred common oaks. This preference was probably associated with higher food abundance related to substrate characteristics (i.e., roughness of bark) which offer more microhabitats for arthropods and thus permit higher densities of potential prey. Some bird species switched feeding preferences within the study period from sycamore maples to common oaks in association with tree phenology. We found two main foraging techniques. All birds searched for prey at short distance (≤50 cm) and gleaned food from substrate except the Pied Flycatcher that foraged by hovering and searched over longer distances (>50 cm). Overall, we demonstrate in our study that canopy access with mobile crane systems provides excellent opportunities to observe canopy birds and enables detailed analysis of their foraging behaviour. The main result of our study reveals fine-grained resource partitioning of birds within the canopy as an important factor structuring assemblages, with species-specific and in part also seasonal differences in stratification and substrate use.     

Flocking behaviour of foraging birds in a neotropical rain forest and the antipredator defence hypothesis

Randomly encountered foraging birds were recorded in a primary rain forest of French Guiana (13,550 records of 216 species), together with their size, diet and habitat use, to assess the relative frequencies of different types of flocking behaviour and some of their ecological correlates. Overall, 42% of birds foraged singly, primarily carnivores (raptors), nectarivores (hummingbirds) and lek-mating frugivores (manakins, some cotingas). For-aging in pairs (26.6%) was widespread, notably among insectivores in the 17–32-g size class. The remaining 31.4% of records were birds in groups of different composition and function, including, in order of decreasing frequency, (1) multispecies upper canopy flocks (83 member species identified)—the largest and most mobile associations of small insectivores, nectarivores and frugivores, mostly tanagers; (2) understorey mixed species flocks of small insectivores, at midlevels of closed forest interior, with 12 core, obligate members and 74 occasional species, mostly active foliage or bark gleaners and probers sharing a unique set of ecological characteristics; (3) monospecific groups (29 species), either gregarious foragers but solitary breeders (large frugivores in canopy) or also breeding colonially or several permanently group living cooperative breeders; (4) opportunistic gatherings of frugivores at fruiting trees (at least 40 species); (5) army ant followers near ground of closed understorey (29 species of mid- to large-sized insectivores); (6) followers of Red-throated Caracaras Daptrius americanus (23 species, usually canopy frugivores entering understorey with caracaras); (7) two raptors following monkeys. Attributes of vulnerability to predators defined by habitat structure (vegetation density or openness) and foraging behaviour (conspicuousness, speed, degree of vigilance) were important determinants of flocking propensity, at least in flocks that were not attracted by a particular food source. The results suggest that the permanent mixed-species flocks in the mature forest under-storey may be an antipredator defence to compensate for the conspicuousness and reduced vigilance resulting from active foraging behaviour in semi-open vegetation, where early detection of predators is difficult.     

Nectar as food for birds: the physiological consequences of drinking dilute sugar solutions

 Nectarivory has evolved many times in birds: although best known in hummingbirds, sunbirds and honeyeaters, it also occurs on an opportunistic basis in a varied assortment of birds. We present a phylogenetic analysis of the distribution of nectarivory in birds. Specialised avian nectarivores are generally small, with an energetic lifestyle and high metabolic rates. Their high degree of dependence on nectar as a food source has led to convergence in morphological, physiological and behavioural adaptations. We examine the constituents of nectar which are most important to bird consumers, and how the birds deal with them in terms of physiology and behaviour. There are still unanswered questions: for example, the dichotomy between sucrose-rich nectars in hummingbird-pollinated plants and predominantly hexose-rich nectars in sunbird-pollinated plants appears to have little to do with bird physiologies and may rather reflect patterns of nectar secretion. Received November 28, 2002; accepted January 26, 2003 Published online: June 2, 2003     

Avifauna response to hurricanes: regional changes in community similarity

Global climate models predict increases in the frequency and intensity of extreme climatic events such as hurricanes, which may abruptly alter ecological processes in forests and thus affect avian diversity. Developing appropriate conservation measures necessitates identifying patterns of avifauna response to hurricanes. We sought to answer two questions: (1) does avian diversity, measured as community similarity, abundance, and species richness, change in areas affected by hurricane compared with unaffected areas, and (2) what factors are associated with the change(s) in avian diversity? We used North American Breeding Bird Survey data, hurricane track information, and a time series of Landsat images in a repeated measures framework to answer these questions. Our results show a decrease in community similarity in the first posthurricane breeding season for all species as a group, and for species that nest in the midstory and canopy. We also found significant effects of hurricanes on abundance for species that breed in urban and woodland habitats, but not on the richness of any guild. In total, hurricanes produced regional changes in community similarity largely without significant loss of richness or overall avian abundance. We identified several potential mechanisms for these changes in avian diversity, including hurricane‐induced changes in forest habitat and the use of refugia by birds displaced from hurricane‐damaged forests. The prospect of increasing frequency and intensity of hurricanes is not likely to invoke a conservation crisis for birds provided we maintain sufficient forest habitat so that avifauna can respond to hurricanes by shifting to areas of suitable habitat.     

Use of autonomous audio recordings for the rapid inventory of birds in the white‐sand forests of the Peruvian Amazon

White‐sand forests are patchily distributed ecosystems covering just 5% of Amazonia that host many specialist species of birds not found elsewhere, and these forests are threatened due to their small size and human exploitation of sand for construction projects. As a result, many species of birds that are white‐sand specialists are at risk of extinction, and immediate conservation action is paramount for their survival. Our objective was to evaluate current survey methods and determine the relative effect of the size of patches of these forests on the presence or absence of white‐sand specialists. Using point counts and autonomous recorders, we surveyed avian assemblages occupying patches of white‐sand forest in the Peruvian Amazon in April 2018. Overall, we detected 126 species, including 21 white‐sand forest specialists. We detected significantly more species of birds per survey point with autonomous recorders than point counts. We also found a negative relationship between avian species richness and distance from the edge of patches of white‐sand forest, but a significant, positive relationship when only counting white‐sand specialists. Although we detected more species with autonomous recorders, point counts were more effective for detecting canopy‐dwelling passerines. Therefore, we recommend that investigators conducting surveys for rare and patchily distributed species in the tropics use a mixed‐method approach that incorporates both autonomous recorders and visual observation. Finally, our results suggest that conserving large, continuous patches of white‐sand forest may increase the likelihood of survival of species of birds that are white‐sand specialists.     

The Influence of Habitat Variables on Bird Communities in Forest Remnants in Costa Rica

This study examined the effects of forest structure (tree species richness, canopy height, percent canopy cover, understory density, tree density and DBH) and avian species traits (nest type and indicator list status) on the diversity, abundance and dissimilarity of bird communities in forest remnants and reforestation areas adjacent to Costa Rican banana plantations. Bird species richness and abundance were significantly related to tree species richness, canopy height and canopy cover in multiple linear regressions, the latter two forest structure variables being the best statistical predictors. Stratification of analyses by bird species indicator categories improved fits of regressions, because correlations with environmental variables differed in sign for different guilds of birds, a result likely to hold for other avifaunas. Analysis of avifauna dissimilarities among sites demonstrated that the species composition of bird communities was highly correlated with forest structure and tree species composition. Logistic regressions indicated that birds making protected (cavity, burrow, pendant, sphere and covered) nests were 2–6 times more likely to be present in the study avifauna than birds making open (cup, saucer, platform and scrape) nests and indicators of disturbed habitats were 11 times more likely to be present than indicators of primary forest. The forest structure data used were simple and inexpensive to collect, and data on avian traits were drawn from the literature. Thus, these methods could easily be replicated at other locations and would be valuable management aids and biodiversity assessment tools for conservation planning.     

The influence of agricultural system,stand structural complexity and landscape context on foraging birds in oil palm landscapes

Functional diversity, an important element of avian biodiversity, can be examined by quantifying foraging guild composition. Understanding the ecological processes that underpin functional diversity of birds in oil palm Elaeis guineensis landscapes is important because different foraging guilds are likely to be influenced in different ways by land use practices. We surveyed birds at 55 sites within oil palm landscapes and at 20 sites within logged peat swamp forest, recording 208 species belonging to 19 foraging guilds. Oil palm landscapes supported a lower abundance of insectivorous, granivorous and omnivorous birds than did logged peat swamp forest despite the latter being severely degraded due to intensive timber extraction. However, abundances of other groups of foraging birds, such as raptors and wetland taxa, were higher in oil palm landscapes than logged peat swamp forest. Frugivorous species were more abundant in smallholdings than plantation estates, probably because of the presence of native trees. Foraging guild diversity was explained by stand‐level attributes such as stand age, vegetation cover, epiphyte persistence and canopy cover. However, each foraging guild exhibited unique responses to different oil palm management regimes and stand‐level attributes. Only arboreal omnivores and terrestrial frugivores were affected by the proximity of nearby natural forest. This diversity of responses implies that the occurrence of particular avian foraging guilds may not be a suitable ecological indicator of best‐practice palm oil production. Our study also suggests that multiple conservation measures will be needed in oil palm landscapes irrespective of management regimes, including: (1) the maintenance of ground layer vegetation cover; (2) the pruning of oil palm canopy to permit light penetration to the ground layer; (3) re‐vegetation of parts of oil palm landscapes with native trees; and (4) retention of natural and/or secondary forest patches within the boundaries of plantations.     

Howling on the edge: Mantled howler monkey (Alouatta palliata) howling behaviour and anthropogenic edge effects in a fragmented tropical rainforest in Costa Rica

The function of long calling is a subject of interest across animal behaviour study, particularly within primatology. Many primate species have male‐specific long‐distance calls, including platyrrhines like the folivorous howler monkey (Alouatta spp.). Howler monkeys may howl to defend resources such as feeding trees or areas of rich vegetation from other monkey groups. This study tests the ecological resource defence hypothesis for howling behaviour in the mantled howler monkey (Alouatta palliata) and investigates how anthropogenic forest fragmentation may influence howling behaviour. More specifically, this study examines how howling bout rate, duration, precursors and tree species richness, DBH, and canopy cover vary in 100 m anthropogenic edge and interior forest zones at La Suerte Biological Research Station (LSBRS), a fragmented tropical rainforest in Costa Rica. Results show that tree species richness and canopy cover are higher in forest interior at this site, suggesting that monkeys should howl at greater rates in the interior to defend access to these higher‐quality vegetation resources. Overall, our results supported the ecological resource defence hypothesis. The main howl precursor was howling from neighbouring groups. Although howling rate did not differ between forest zones, howling bouts from forest interior were longer, had a greater number of howls per bout and were preceded by different precursors than howls from anthropogenic edge zones, including more howls from neighbouring groups. Our findings provide some of the first evidence for behavioural edge effects in primate vocal communication behaviour.     

Habitat selection by a despotic passerine,the Bell Miner (Manorina melanophrys): When restoring habitat through Lantana (Lantana camara) removal is not enough

The Bell Miner (Manorina melanophrys) occurs in logged eucalypt forest in northern NSW with a dense understorey of the invasive Neotropical shrub Lantana (Lantana camara) that is used for nesting. The link between Bell Miners and Lantana is important as the birds aggressively exclude all smaller and similar‐sized birds from their colonies, reducing avian diversity in forest occupied by the species. We monitored the impact of Lantana removal on Bell Miner persistence in several plots in two logged forest sites, along with untreated control plots at one of the sites. Lantana control was successful over 7 years at both sites, with regeneration of native understorey, midstorey and canopy species compensating for the loss of live Lantana cover in the understorey. Bell Miner individuals vacated the treated plots in one site (Creek's Bend) but persisted in the control and treated plots at the second site (Toonumbar National Park). Bell Miner response was correlated with forest structure: birds vacated forest with a sparse understorey (<5 m) but dense midstorey (5–15 m) and canopy (>15 m) at Creek's Bend, but remained at the site with a dense understorey but sparse midstorey and canopy at Toonumbar. We therefore predict that forest restoration that simultaneously reduces Lantana understorey and increases midstorey density will be most successful in reducing the abundance of the despotic Bell Miner and increasing avian diversity in rehabilitated sites.     

Forest bird diversity in Mediterranean areas affected by wildfires: a multi-scale approach

Fire is a key mechanism creating and maintaining habitat heterogeneity in Mediterranean landscapes by turning continuous woody landscapes into mosaics of forests and shrublands. Due to the long historical role of fires in the Mediterranean, we hypothesised a moderate negative effect of this type of perturbation on forest bird distribution at a landscape level. We conducted point bird censuses in Aleppo pine forest patches surrounded by burnt shrublands and studied the relationships between three ecological groups of bird species (forest canopy species, forest understorey species, and ubiquitous species) and the features of local habitat, whole patch and surrounding landscape. We used a multi-scale approach to assess the effects of landscape variables at increasing spatial scales on point bird richness. Regarding local habitat components, canopy species were positively associated with tall pines while understorey species with the cover of shrubs and plants from holm-oak forests. Forest birds were positively related to patch size and irregular forest shapes, that is, with high perimeter/size ratios. Thus, these species did not seem to perceive edges as low quality but rather favourable microhabitats. We did not detect any negative effect of isolation or cover of woodlands in the landscape on the presence of forest species after local habitat factors had been accounted for. Finally, only local habitat factors entered the model for ubiquitous species. We suggest that mosaic-like landscapes shaped by fires in the Mediterranean basin are not strongly associated with negative effects fragmentation on forest birds other than those related with habitat loss.     

Behavioural adaptations of birds to environments where evaporation is high and water is in short supply

1. Behaviour that reduces the heat load or evaporation experienced by birds living in arid areas is reviewed. Many species have evolved hunting behaviour that enables them to remain inactive during the hottest parts of the day and thus greatly reduce the amount of metabolic heat that they need to dissipate. Flights to water are made at low ambient temperatures, either early in the morning or late in the evening. Fighting is rare in many species of desert birds, avoiding the excess generation of heat by this activity. Many arid zone birds maintain long-lasting pair bonds, avoiding the necessity for active, elaborate display before breeding and again reducing activity. 2. The observations on nomadism are discussed. No unifying principles that might control the behaviour of birds seeking widely separated areas of abundance of food have yet emerged. 3. Some species have evolved mechanisms, embodied in behavioural characteristics, that ensure that the eggs and chicks are sheltered from high temperatures and are provided with adequate moisture. 4. Birds have evolved many different kinds of behavioural adaptation to arid zones and representatives from many avian families live there, apparently successfully.     

Primate origins and the evolution of angiosperms

Traditionally, the morphological traits of primates were assumed to be adaptations to an arboreal way of life. However, Cartmill [1972] pointed out that a number of morphological traits characteristic of primates are not found in many other arboreal mammals. He contends that orbital convergence and grasping extremities indicate that the initial divergence of primates involved visual predation on insects in the lower canopy and undergrowth of the tropical forest. However, recent research on nocturnal primates does not support the visually-oriented predation theory. Although insects were most likely important components of the diets of the earliest euprimates, it is argued here that visual predation was not the major impetus for the evolution of the adaptive traits of primates. Recent paleobotanical research has yielded evidence that a major evolutionary event occurred during the Eocene, involving the angiosperms and their dispersal agents. As a result of long-term diffuse coevolutionary interactions with flowering plants, modern primates, bats, and plant-feeding birds all first arose around the Paleocene-Eocene boundary and became the major seed dispersers of modern tropical flora during the Eocene. Thus, it is suggested here that the multitude of resources available on the terminal branches of the newly evolved angiosperm, rain forest trees led to the morphological adaptations of primates of modern aspect.     

Vertical gradients in leaf trait diversity in a New Zealand forest

Leaves come in a remarkable diversity of sizes and shapes. However, spatial patterns in leaf trait diversity are rarely investigated and poorly resolved. We used a hierarchical approach to evaluate vertical variability in leaf morphology (i.e., leaf trait diversity) in 16 common tree and shrub species inhabiting a New Zealand forest. Height-related heterogeneity in leaf area, specific leaf area, circularity and length to width ratio was analyzed at three scales: (1) among leaves within plants, (2) among plants within species and (3) among species within functional groups (i.e., trees vs. shrubs). Results were scale dependent. Among-leaf morphological diversity was unrelated to plant height. Among-individual morphological diversity increased with the average height of each species, indicating that taller plant species express a greater range of leaf traits than shorter species. Among-species morphological diversity was higher in shrubs than in trees. We hypothesize that scale-dependent patterns in leaf trait diversity result from scale-dependent adaptations to forest environmental conditions. As trees grow from the forest floor into the canopy, they are exposed to a range of environmental conditions, which may select for a range of leaf traits through ontogeny. Conversely, shrubs never reach the forest canopy and may instead be differentially adapted to suites of environmental conditions associated with different stages of forest recovery from tree-fall disturbances. Overall results indicate that vertical patterns in leaf trait diversity exist. However, their strength and directionality are strongly scale-dependent, suggesting that different processes govern leaf shape diversity at different levels of ecological organization.     

Mixed patterns of morphological adaptation to insularity in an aerial displaying bird,the Common Snipe Gallinago gallinago

On islands, colonizing birds may evolve behavioural and morphological adaptations to the new environment, often resulting in changes in body size and reduction or even total loss of flight. These island populations have therefore been used to test hypotheses related to adaptations for flight. However, in certain species in which flight is used not only in foraging and migration but also in mating displays, disentangling the effects of natural and social selection is difficult. Thus, sedentary populations of species that perform aerial displays (such as the Common Snipe Gallinago gallinago that breed in the Azores archipelago) may offer an opportunity to separate the effects of natural and social selection on morphology. If insular Common Snipe respond to the characteristic ecological context of oceanic islands, we expect them to differ from migratory conspecifics in body size and by having relatively smaller and more rounded wings. On the other hand, if social selection exerts a more powerful force over the morphology of this species, we expect that sedentary and migratory birds will not differ in flight‐related characters. We tested these hypotheses by comparing morphological characters measured on live Common Snipe captured in the Azores during the breeding season with those measured on migratory specimens hunted during autumn/winter in mainland Portugal. Sedentary Azorean birds were smaller and had relatively shorter tails but did not show the tendency for insular birds to possess more rounded wings as described in other taxa, including in the Azores. Bergman's rule might explain the difference in body size and shorter tails may be responsible for behavioural differences between populations. The lack of difference in wing shape might be explained by the need of the Common Snipe to perform aerial displays during courtship, suggesting an effect of social selection on the migratory strategy of this species.     

Comparison of wing morphology in three birds of prey: correlations with differences in flight behavior

Flight is the overriding characteristic of birds that has influenced most of their morphological, physiological, and behavioral features. Flight adaptations are essential for survival in the wide variety of environments that birds occupy. Therefore, locomotor structure, including skeletal and muscular characteristics, is adapted to reflect the flight style necessitated by different ecological niches. Red-tailed hawks (Buteo jamaicensis) soar to locate their prey, Cooper's hawks (Accipiter cooperii) actively chase down avian prey, and ospreys (Pandion haliaetus) soar and hover to locate fish. In this study, wing ratios, proportions of skeletal elements, and relative sizes of selected flight muscles were compared among these species. Oxidative and glycolytic enzyme activities of several muscles were also analyzed via assays for citrate synthase (CS) and for lactate dehydrogenase (LDH). It was found that structural characteristics of these three raptors differ in ways consistent with prevailing aerodynamic models. The similarity of enzymatic activities among different muscles of the three species shows low physiological differentiation and suggests that wing architecture may play a greater role in determining flight styles for these birds.     

西双版纳热带雨林与海南热带雨林的比较研究

西双版纳的热带雨林与海南低地热带雨林和热带季雨林有基本一致的植物区系组成,群落中优势科无论在种数百分比还是重要值排名上均较接近,显然属于同样性质的植物区系。在生态特征上,西双版纳热带雨林群落高大,分层不明显,B层为林冠层,散生巨树常见,在生活型谱上以高位芽植物占绝对优势,大、中高位芽植物相对较多,落叶树种比例小;以中叶、纸质,全缘和复叶比例较高为特征,具有最接近海南低地湿润雨林的群落垂直结构和生态外貌,其雨林特点虽不如湿润雨林浓厚,但明显强于海南的热带常绿季雨林和山地雨林,海南常绿季雨林群落高度明显较矮,小叶比例通常较高,革质叶比例亦较高,群落具遥明显的旱生特点,海南的山地雨林群落高度较矮,A层连续,成为林冠,无散生巨树,分层明显,在生活型谱上大高位芽植物比例减少,附生植物丰富,并具有相当比例的地面芽植物;叶级虽以中叶占优势,但通常革质,非全缘和单叶比例较高,明显由于热量不足的影响而带有亚热带森林特色,在物种多样性上,西双版纳热带雨林的乔木物种多样性指数似乎与海南的低地热带雨林相当,低于海南的山地雨林群落,海南的热带雨林群落种类丰富度不同人研究的结果差异较大,如果这些用于比较的数据可靠和具有可比性的话,西双版纳热带雨林的物种多样性要比海南的山地雨林低。     

Leaf surface specializations of birds and Arthropods in a Panamanian forest

Summary Most species of Panamanian lowland forest birds specialize on leaf undersurfaces when hunting foliage insects. The few species of leaf surface generalists and leaf upper surface specialists are omnivorous gleaners. We estimate that while over 90% of the avian understory insectivory is directed towards leaf undersurfaces, only 50% of canopy foliage insectivory is directed towards the undersides of leaves. In the low understory we found 70–80% of the arthropods on leaf undersides. The excess use of leaf-bottoms by understory birds may be a result of their greater visibility. It is hypothesized that less proficient insectivores are unable to take advantage of the greater effective density of underleaf insects because they can only efficiently attack the closest leaf surfaces; these closest surfaces will usually be the leaf tops from the branch on which the bird is perched. Alternatively, leaf-top specialists may have special foraging adaptations for overcoming the disadvantages of leaf-top foraging. These adaptations may involve attack behavior (Tachyphonus luctuosus) or searching behavior (Dacnis cayana). Dacnis often used leaf damage as a foraging cue; this may be the first report of a bird using leaf damage for searching for insects. The greater use of leaf upper surfaces by canopy birds may be influenced by four factors: greater seasonality of insects in the canopy favoring omnivores which may be less efficient insectivores; more insects on leaf tops; fewer planar leaf arrangements in canopy plants; or the greater visibility of leaf upper surfaces of the outer shell of foliage of massive trees. Based on the greater number of arthropods on leaf bottoms in the dry season, the higher abundance of smaller insects on leaf bottoms, as well as the greater proportion of insects on leaf tops at cooler higher elevations, we suggest that arthropods prefer leaf bottoms in tropical areas for physiological, not predator avoidance reasons.     

The effects of locomotion on the structural characteristics of avian limb bones

Despite the wide range of locomotor adaptations in birds, little detailed attention has been given to the relationships between the quantitative structural characteristics of avian limb bones and bird behaviour. Possible differences in forelimb relative to hindlimb strength across species have been especially neglected. We generated cross‐sectional, geometric data from peripheral quantitative computed tomography scans of the humerus and femur of 127 avian skeletons, representing 15 species of extant birds in 13 families. The sample includes terrestrial runners, arboreal perchers, hindlimb‐propelled divers, forelimb‐propelled divers and dynamic soarers. The hindlimb‐propelled diving class includes a recently flightless island form. Our results demonstrate that locomotor dynamics can be differentiated in most cases based on cross‐sectional properties, and that structural proportions are often more informative than bone length proportions for determining behaviour and locomotion. Recently flightless forms, for example, are more easily distinguished using structural ratios than using length ratios. A proper phylogenetic context is important for correctly interpreting structural characteristics, especially for recently flightless forms. Some of the most extreme adaptations to mechanical loading are seen in aquatic forms. Penguins have forelimbs adapted to very high loads. Aquatic species differ from non‐aquatic species on the basis of relative cortical thickness. The combination of bone structural strength and relative cortical area of the humerus successfully differentiates all of our locomotor groups. The methods used in this study are highly applicable to fossil taxa, for which morphology is known but behaviour is not. The use of bone structural characteristics is particularly useful in palaeontology not only because it generates strong signals for many locomotor guilds, but also because analysing such traits does not require knowledge of body mass, which can be difficult to estimate reliably for fossil taxa. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 601–624.