Vegetation structure influences foraging decisions in a declining grassland bird: the importance of fine-scale habitat and grazing regime

Capsule: Whinchat Saxicola rubetra foraging behaviour was significantly influenced by habitat structure and grazing.

Aims: To assess how foraging habitats selected by breeding Whinchats differed from wider territory attributes under contrasting grazing management in multiple upland areas in Scotland: principally sheep grazed, Red Deer grazed or ungrazed, and to identify how differing land use may limit suitable foraging areas.

Methods: We compared fine-scale vegetation structure in patches chosen for foraging by Whinchats in contrasting grazing management regimes.

Results: Whinchats were less likely to forage in patches with a greater cover of bracken and tall non-bracken vegetation, regardless of grazing regime. Grass cover influenced foraging behaviour in ungrazed habitats only, where Whinchats were less likely to forage in areas with high grass cover.

Conclusion: Whinchats appear to require a mosaic or range of sward structures within breeding territories, highlighting the importance of establishing how vegetation structure influences breeding birds at different spatial scales. Our results suggest that suitable foraging patches were plentiful within grazed habitats but potentially limited in ungrazed habitats. Further work is needed to identify management regimes and interventions to maintain conditions suitable for breeding Whinchats that are compatible with other land use and conservation objectives.     

Density and behaviour of Whinchats Saxicola rubetra on African farmland suggest that winter habitat conditions do not limit European breeding populations

The Whinchat Saxicola rubetra is an Afro‐Palaearctic migrant undergoing widespread population decline. Whinchats winter in West Africa but there are almost no data on their habitat use and behaviour there that may help to explain the cause of this decline. We measured the density of Whinchats, the habitat characteristics associated with their occurrence on farmland, and the relationships between behavioural and habitat variation on farmland around Jos, central Nigeria, over three winters. Whinchats occurred in many fields harvested in the dry season, the density at three sites varying from 0.03 to 0.43 birds/ha, but they were absent at a fourth site. Whinchats were less likely to be found in farmland without particular crops (e.g. structural stem crops such as maize and millet), with more trees, lower amounts of short vegetation (grass, weeds, crops and crop stubble less than 10 cm in height), and higher amounts of medium vegetation (coverage of vegetation 10–100 cm in height) and litter (dead, unburned, vegetation on the ground). Whinchat abundance in areas of farmland where they were present was independent of most variables considered, but density was higher where there was more short vegetation cover. Foraging behaviour did not vary significantly between farmland habitats. All predictors were consistent between season, years and across sites. The presence/absence model was very poor at predicting presence and there were no strong predictors of abundance or foraging variation. This is consistent with a species well below carrying capacity within its environment so that many suitable areas do not have birds present and there is little aggregation at better sites. Overall, Whinchats were abundant and appeared to have plentiful habitat; densities have probably increased alongside the intensification of agriculture (presence of fallow farmland, short vegetation and structural crops). The results suggest that West African farmland in the dry season can support large numbers of Whinchats and that recent population declines in Europe are unlikely to be caused primarily by lack of suitable wintering habitat.     

Using habitat selection theories to predict the spatiotemporal distribution of migratory birds during stopover – a case study of pink‐footed geese Anser brachyrhynchus

Understanding how animals select for habitat and foraging resources therein is a crucial component of basic and applied ecology. The selection process is typically influenced by a variety of environmental conditions including the spatial and temporal variation in the quantity and quality of food resources, predation or disturbance risks, and inter‐ and intraspecific competition. Indeed, some of the most commonly employed ecological theories used to describe how animals choose foraging sites are: nutrient intake maximisation, density‐dependent habitat selection, central‐place foraging, and predation risk effects. Even though these theories are not mutually exclusive, rarely are multiple theoretical models considered concomitantly to assess which theory, or combination thereof, best predicts observed changes in habitat selection over space and time. Here, we tested which of the above theories best‐predicted habitat selection of Svalbard‐breeding pink‐footed geese at their main spring migration stopover site in mid‐Norway by computing a series of resource selection functions (RSFs) and their predictive ability (k‐fold cross validation scores). At this stopover site geese fuel intensively as a preparation for breeding and further migration. We found that the predation risk model and a combination of the density‐dependent and central‐place foraging models best‐predicted habitat selection during stopover as geese selected for larger fields where predation risk is typically lower and selection for foraging sites changed as a function of both distance to the roost site (i.e. central‐place) and changes in local density. In contrast to many other studies, the nutritional value of the available food resources did not appear to be a major limiting factor as geese used different food resources proportional to their availability. Our study shows that in an agricultural landscape where nutritional value of food resources is homogeneously high and resource availability changes rapidly; foraging behaviour of geese is largely a tradeoff between fast refuelling and disturbance/predator avoidance.     

No Habitat Selection during Spring Migration at a Meso-Scale Range across Mosaic Landscapes: A Case Study with the Woodcock (Scolopax rusticola)

Success of migration in birds in part depends on habitat selection. Overall, it is still poorly known whether there is habitat selection amongst landbird migrants moving across landscapes. Europe is chiefly covered by agro-forestry mosaic landscapes, so migratory species associated to either agricultural landscapes or woodland habitats should theoretically find suitable stopover sites along migration. During migration from wintering to breeding quarters, woodcocks (Scolopax rusticola) tagged with PTT satellite-tracking transmitters were used to test for the hypothesis that migrants associated to agro-forest habitats have no habitat selection during migration, at a meso-scale level. Using a GIS platform we extracted at a meso-scale range habitat cover at stopover localities. Results obtained from comparisons of soil covers between points randomly selected and true stopover localities sites revealed, as expected, the species may not select for particular habitats at a meso-scale range, because the habitat (or habitats) required by the species can be found virtually everywhere on their migration route. However, those birds stopping over in places richer in cropland or mosaic habitats including both cropland and forest and with proportionally less closed forest stayed for longer than in areas with lower surfaces of cropland and mosaic and more closed forest. This suggests that areas rich in cropland or mosaic habitat were optimal.     

Fine‐scale habitat use during the non‐breeding season suggests that winter habitat does not limit breeding populations of a declining long‐distance Palearctic migrant

For migrant birds, which habitats are suitable during the non‐breeding season influences habitat availability, population resilience to habitat loss, and ultimately survival. Consequently, habitat preferences during winter and whether habitat segregation according to age and sex occurs directly influences migration ecology, survival and breeding success. We tested the fine‐scale habitat preferences of a declining Palearctic migrant, the whinchat Saxicola rubetra, on its wintering grounds in west Africa. We explored the influence of habitat at the territory‐scale and whether dominance‐based habitat occupancy occurs by describing the variation in habitat characteristics across wintering territories, the degree of habitat change within territories held throughout winter, and whether habitat characteristics influenced territory size and space‐use within territories or differed with age and sex. Habitat characteristics varied substantially across territories and birds maintained the same territories even though habitat changed significantly throughout winter. We found no evidence of dominance‐based habitat occupancy; instead, territories were smaller if they contained more perching shrubs or maize crops, and areas with more perching shrubs were used more often within territories, likely because perches are important for foraging and territory defence. Our findings suggest that whinchats have non‐specialised habitat requirements within their wintering habitat of open savannah and farmland, and respond to habitat variation by adjusting territory size and space‐use within their territories instead of competing with conspecifics. Whinchats show a tolerance for human‐modified habitats and results support previous findings that some crop types may provide high‐quality wintering habitat by increasing perch density and foraging opportunities. By having non‐specialised requirements within broad winter habitat types, migrants are likely to be flexible to changing wintering conditions in Africa, both within and across winters, so possibly engendering some resilience to the rapid anthropogenic habitat degradation occurring throughout their wintering range.     

Experimental evidence for the interplay of exogenous and endogenous factors on the movement ecology of a migrating songbird

Movement patterns during songbird migration remain poorly understood despite their expected fitness consequences in terms of survival, energetic condition and timing of migration that will carry over to subsequent phases of the annual cycle. We took an experimental approach to test hypotheses regarding the influence of habitat, energetic condition, time of season and sex on the hour-by-hour, local movement decisions of a songbird during spring stopover. To simulate arrival of nocturnal migrants at unfamiliar stopover sites, we translocated and continuously tracked migratory red-eyed vireos (Vireo olivaceus) throughout spring stopover with and without energetic reserves that were released in two replicates of three forested habitat types. Migrants moved the most upon release, during which time they selected habitat characterized by greater food abundance and higher foraging attack rates. Presumably under pressure to replenish fuel stores necessary to continue migration in a timely fashion, migrants released in poorer energetic condition moved faster and further than migrants in better condition and the same pattern was true for migrants released late in spring relative to those released earlier. However, a migrant's energetic condition had less influence on their behavior when they were in poor quality habitat. Movement did not differ between sexes. Our study illustrates the importance of quickly finding suitable habitat at each stopover site, especially for energetically constrained migrants later in the season. If an initial period prior to foraging were necessary at each stop along a migrant's journey, non-foraging periods would cumulatively result in a significant energetic and time cost to migration. However, we suggest behavior during stopover is not solely a function of underlying resource distributions but is a complex response to a combination of endogenous and exogenous factors.     

Foraging,roosting, and nesting habitats of the avian fauna of the Agmon wetland,northern Israel

The foraging, nesting and roosting habitats of the avian fauna of a newly created Agmon wetland and surrounding cultivated peat land (5 km2) in the Hula Valley, northern Israel, were evaluated (January 1996–February 1997) to assess the value as a habitat and for wildlife tourism. We recorded 180 bird species (herons, dabbling ducks, kingfishers, waders, wagtails and raptors) in different habitats (the lake, shores, cattail and reed-bed stands, trees, temporary inundated areas). The most heavily used habitat for foraging, breeding, and roosting was a large cattail stand in the southern third of the lake. The foraging habitat and diet data of 97 avian species were determined. The most intensively used foraging habitats were cultivated fields, lake shore reed-beds, shallow canals and the cattail stand. Forty six species nested in the wetland during March-October, with 2,040 colonial and solitary nests mostly in the cattail stand, near-shore reed-beds and young trees. Roosts, also mostly in the cattail and reed-bed stands and in trees, were used by 55 species. Based on this survey of available habitats, we identified several key habitats that are either missing or require further development (e.g., temporarily inundated mud flats, reed-beds and bare islands). We also provide suggestions for improving the delicate balance between requirements of this developing wetland ecosystem and of the adjacent agricultural areas.     

Foraging behaviour and fuel accumulation of capital breeders during spring migration as derived from a combination of satellite‐ and ground‐based observations

The migration strategy of many capital breeders is to garner body stores along the flyway at distinct stopover sites. The rate at which they can fuel is likely to be strongly influenced by a range of factors, such as physiology, food availability, time available for foraging and perceived predation. We analysed the foraging behaviour and fuel accumulation of pink‐footed geese, an Arctic capital breeder, at their mid‐flyway spring stopover site and evaluated to what extent their behaviour and fuelling were related to physiological and external factors and how it differed from other stopovers along the flyway. We found that fuel accumulation rates of geese at the mid‐flyway site were limited by habitat availability rather than by digestive constraints. However, as the time available for foraging increased over the stopover season, geese were able to keep constant fuelling rate. Putting this in perspective, geese increased their daily net energy intake along the flyway corresponding to the increase in time available for foraging. The net energy intake per hour of foraging remained the same. Geese showed differences in their reaction to predators/disturbance between the sites, taking higher risks particularly at the final stopover site. Hence, perceived predation along the flyway may force birds to postpone the final fuel accumulation to the last stopover along the flyway. Flexibility in behaviour appears to be an important trait to ensure fitness in this capital breeder. Our findings are based on a new, improved method for estimating fuel accumulation of animals foraging in heterogeneous landscapes based on data obtained from satellite telemetry and habitat specific intake rates.     

Contrasting trophic‐cascade effects driven by variation in morphology of the perches used by a larval damselfly

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Foraging in a landscape mosaic: selection for energy and minerals in free-ranging cattle

Several studies have indicated the potential importance of nutrients, other than energy, in determining foraging decisions. A model was developed to test this idea, on the assumption of an intake maximization for different nutrients (energy, sodium and phosphorus). The model predictions were tested using field data from cattle grazing in a landscape mosaic of Pleistocene cover-sand and riverine grassland. Observations on foraging behaviour, food intake and diet composition were collected in thirteen 4-day-periods over 2 years. Habitat selection was determined by comparing the proportion of grazing time in different vegetation units with the available area proportion of the units. Two levels of habitat selection were examined: a micro-level (fine-scale, where vegetation units were considered separately) and a macro-level (coarse-scale, where vegetation units were combined to give selection at the landscape level). At the micro-level of habitat selection, no selection was apparent between the vegetation units of the riverine landscape, but the Deschampsia flexuosa unit was significantly selected for in the cover-sand landscape. At the landscape (macro-) level, the animals preferred the riverine landscape. The model revealed poor predictions of habitat occupancy on a micro-level. A much better prediction was obtained when vegetation units were combined at a macro-level. The D. flexuosa unit provided a higher energy intake, whereas the intake of sodium was higher in riverine grassland. Phosphorus proved relatively significant in determining habitat occupancy. Based on energy maximization alone, the model was a very poor predictor of habitat occupancy. It is argued that selection occurred mainly at the macro-level. The incorporation of different nutrient constraints in foraging models can then prove fruitful when seeking explanations of habitat occupancy. At the micro-level, difficulties for the animal in assessing nutrient availability may result in a less selective foraging pattern. However, the costs of increased selectivity may be greater than the benefits.     

Songbirds are resilient to hurricane disturbed habitats during spring migration

The Gulf of Mexico is a conspicuous feature of the Neotropical–Nearctic bird migration system. Traveling long distances across ecological barriers comes with considerable risks, and mortality associated with intercontinental migration may be substantial, including that caused by storms or other adverse weather events. However, little, if anything, is known about how migratory birds respond to disturbance‐induced changes in stopover habitat. Isolated, forested cheniere habitat along the northern coast of the Gulf of Mexico often concentrate migrants, during weather conditions unfavorable for northward movement or when birds are energetically stressed. We expected hurricane induced degradation of this habitat to negatively affect the abundance, propensity to stopover, and fueling trends of songbirds that stopover in coastal habitat. We used spring banding data collected in coastal Louisiana to compare migrant abundance and fueling trends before (1993–1996 and 1998–2005) and after hurricanes Rita (2006) and Ike (2009). We also characterized changes in vegetative structure before (1995) and after (2010) the hurricanes. The hurricanes caused dramatic changes to the vegetative structure, which likely decreased resources. Surprisingly, abundance, propensity to stopover, and fueling trends of most migrant species were not influenced by hurricane disturbance. Our results suggest that: 1) the function of chenieres as a refuge for migrants after completing a trans‐Gulf flight may not have changed despite significant changes to habitat and decreases in resource availability, and 2) that most migrants may be able to cope with habitat disturbance during stopover. The fact that migrants use disturbed habitat points to their conservation value along the northern coast of the Gulf of Mexico.     

Foraging strategies involved in habitat use of shorebirds at the intertidal area of Chongming Dongtan, China

Dense flocks of migratory shorebirds from diverse species often concentrate in the intertidal areas for stopover. Trophic structure, food partition, prey availability and selectivity, predation risk, and abiotic factors are often used to explain the differences in habitat use of coexisting shorebirds. We sampled the macrobenthos and surveyed the distribution of shorebird populations to study the effects of foraging strategies on the habitat use of shorebirds at Chongming Dongtan, an important stopover site for shorebirds on the East Asian–Australasian Flyway. Results show that the relative abundance of epifaunal macrobenthos in salt marshes was much higher than that in the bare flats, whereas the relative abundance of infaunal macrobenthos in salt marshes was much lower than that in bare flats. The relative abundance of two life forms of macrobenthos was similar in the transitional zones between the salt marshes and the bare flats. Shorebirds with different foraging strategies exhibited different habitat uses. Pause-travel shorebirds mainly utilized the salt-marsh fringes, while tactile continuous shorebirds relied heavily on the bare flats. There was no significant difference in habitat use for visual continuous shorebirds. The density of tactile continuous shorebirds was positively correlated with bivalve density, and that of visual continuous shorebirds positively with crustacean density. Meanwhile, the relative abundance of pause-travel foraging shorebirds was positively correlated with the relative abundance of epifaunal, but negatively with infaunal macrobenthos. In contrast, the relative abundance of tactile foraging shorebirds had a positive correlation with infaunal but a negative one with epifaunal life form. Therefore, foraging strategies may play important roles in shorebirds’ habitat use in intertidal areas.     

Perch selection in Sympetrum species (Odonata: Libellulidae): importance of vegetation structure and composition

1. Perching dragonflies are closely associated with the physical structure of vegetation because adults utilise plants when foraging, thermoregulating, and mate-seeking. However, little attention has been given to which structural attribute of vegetation is playing a key role within foraging habitat use. 2. This study focused on the influence of different features of perches on their selection by adult dragonflies. As a model group, a typical percher behavioural guild of Sympetrum was used and experimental plots with artificial perches and different structural properties were established. 3. It was found that adults preferred perch sticks with a wider diameter and larger spacing. It is assumed that these perching sites are advantageous because their base is more stable, they provide the best view for successful foraging, and there is no interference among individuals. 4. This study also revealed significant differences between male and female preferences. Females used less structured vegetation formed by thin perches in high densities. It is suggested that female discrimination is influenced by the higher competitiveness of males, which is related to their agonistic behaviour. 5. These results suggest that the availability of perches in the foraging habitat might be an essential requirement for adults. However, within the terrestrial surroundings of breeding sites, the structure of vegetation providing conditions for perching may be associated only with certain habitats. In intensive landscapes, physically structured vegetation can be limited or completely absent.     

Patterns in departure phenology and mass gain on African non‐breeding territories prior to the Sahara crossing in a long‐distance migrant

Afro‐Palaearctic migrants are declining to a greater degree than other European species, suggesting that processes occurring in Africa or on migration may be driving these trends. Constraints on food availability on the wintering grounds may contribute to these declines but little is known about when and where these resource constraints may occur. Sufficient resources are particularly important prior to spring migration, when migrants must cross the Sahara Desert. We examined mass gain and departure phenology in a long‐distance Palaearctic passerine migrant to determine the degree to which pre‐migratory fattening occurs in their long‐term non‐breeding territories in the Guinea Savannah region of Africa. We monitored 75 Whinchats Saxicola rubetra for departure from their non‐breeding territories in one spring, and analysed mass data of 377 Whinchats collected over three non‐breeding seasons plus 141 migrating Whinchats caught in April over 8 years, all within the same few square kilometres of human‐modified Guinea Savannah in central Nigeria. Whinchats left their winter territories throughout April, with males departing on average 8 days earlier than females. However, there was no evidence that time of departure from territory was linked to age, body size or mass at capture. Whinchats departed their territories with a predicted mass of 16.8 ± 0.3 g, considerably less than the c. 24 g required for the average Whinchat to cross the Sahara directly. Comparing departure dates with arrival dates in southern Europe showed a discrepancy of at least 2 weeks, suggesting that many Whinchats spend considerable time on pre‐migratory fuelling outside their winter territory prior to crossing the Sahara. Overwintering birds gained mass slowly during February and March (0.03 g/day), and non‐territorial or migrating birds at a much higher rate in April (at least 0.23 g/day), with up to 20% of migrating Whinchats in April potentially having sufficient fuel loads to cross the Sahara directly from central Nigeria. Our results suggest that most Whinchats leave their winter territories to fatten up locally or, possibly, by staging further north, closer to the southern limit of the Sahara. Resource constraints are therefore likely to be particularly focused in West Africa during mid‐April and possibly at staging areas before the crossing of the Sahara Desert.     

Weak migratory connectivity,loop migration and multiple non-breeding site use in British breeding Whinchats Saxicola rubetra

Determining the links between breeding populations and the pressures, threats and conditions they experience presents a challenge for the conservation of migratory birds which can use multiple sites separated by hundreds to thousands of kilometres. Furthermore, migratory connectivity – the connections made by migrating individuals between networks of breeding and non-breeding sites – has important implications for population dynamics. The Whinchat Saxicola rubetra is declining across its range, and tracking data from a single African non-breeding site implies high migratory spread. We used geolocators to describe the migration routes and non-breeding areas of 20 Whinchats from three British breeding populations. As expected, migratory spread was high, with birds from the three populations overlapping across a wide area of West Africa. On average, in non-breeding areas, British breeding Whinchats were located 652 km apart from one another, with some likely to share non-breeding areas with individuals from breeding populations as far east as Russia. Four males made a direct non-breeding season movement to a second, more westerly, non-breeding location in January. Autumn migration was through Iberia and around the western edge of the Sahara Desert, whereas spring migration was more direct, indicating an anticlockwise loop migration. Weak migratory connectivity implies that Whinchat populations are somewhat buffered against local changes in non-breeding conditions. If non-breeding season processes have played a role in the species’ decline, then large-scale drivers are likely to be the cause, although processes operating on migration, or interactions between breeding and non-breeding processes, cannot be ruled out.     

Extensive Use of Intertidal Habitat by Shorebirds Outside Protected Nesting Areas

Conservation of shorebirds throughout their breeding and migratory ranges has become a priority as shorebird populations decline globally. Along the North Atlantic Coast, management efforts have particularly focused on preserving nesting habitat for piping plovers (Charadrius melodus), which are protected under the Endangered Species Act. It is unclear whether these conservation measures suffice to protect foraging habitat for piping plovers and other shorebirds on stopover during migration along the Atlantic Flyway. To evaluate the extent to which conservation of piping plover nesting areas extends to all habitats used by plovers, and to determine whether these protections also benefited non-breeding migratory shorebirds in the region, we conducted weekly shorebird surveys, recording the number and locations of piping plovers and other species, during northward and southward migration on Fire Island and Westhampton Island, New York, USA, from 2014–2016. We used canonical correspondence analysis (CCA) to assess the degree of spatiotemporal overlap between breeding plovers, foraging plovers, and other migratory shorebirds that temporarily stage at the site. The spatiotemporal distribution of migratory shorebirds matched more closely with piping plovers seen during foraging than piping plovers observed tending nests and engaging in other breeding activities. Migratory shorebirds and foraging piping plovers were more abundant and frequent in wet intertidal zones outside of fenced-off nesting areas, which were not protected under current management regimes. Therefore, additional protection of piping plover foraging habitat could benefit plovers and migratory shorebirds that use similar feeding grounds during stopover on northward and southward migration. © 2020 The Wildlife Society.     

Expensive Traditions: Energy Expenditure of Aleutian Geese in Traditional and Recently Colonized Habitats

ABSTRACT In an effort to reduce goose depredation at a traditional spring migratory stopover site, private landowners implemented a coordinated hazing plan to scare Aleutian cackling geese (Branta hutchinsii leucopareia) from private lands to adjacent public pastures that were cultivated and set aside specifically for geese. Coincidentally, some Aleutian geese began using a new stopover site 150 km farther south in their spring migratory range; numbers at the new site continue to increase. We tested the idea that when their ability to acquire resources deteriorates geese are likely to seek improved foraging conditions, especially during spring migration when individuals strive to maximize nutrient stores and minimize energy expenditure. We quantified measures of goose foraging performance in traditional and new spring staging sites by calculating foraging opportunity, foraging effort, body condition, and daily energy expenditure. Geese staging at the site with higher levels of human disturbance had less foraging opportunity and, despite increased foraging effort and more nutritious food-plants at the site, birds there experienced an elevated energy expenditure and poorer body condition than birds at the new stopover site. Reduced foraging time and increased energy expenditure at the traditional spring staging site may have triggered the colonization process. Suitability assessment of habitat for migratory geese should include measures of foraging opportunity, disturbance risks, and daily energy expenditure in addition to quantity and quality of foods.     

Age-related changes in song repertoire size and song type sharing in the Whinchat Saxicola rubetra

In many oscine passerines males’ songs, the repertoire size increases with age. At the same time it often remains unknown when and where males learn new songs. To infer the Whinchat Saxicola rubetra song learning strategy, we described and catalogued song-type repertoire, revealed age differences and examined song sharing strategies among neighbouring and distant males. We recorded song vocalizations of 40 males in a limited (104 ha) study plot during four years. Whinchats produce short and discrete songs with clear intersong pauses. In total 45 song types were allocated, the individual repertoire size averaged 23.5 ± 7.6 song types (range 9–34 song types). The males’ age significantly influenced the song-type repertoire size. The second calendar year (first breeding) males had a lower repertoire size than the older males. The majority of song types were shared by less than half of males in our sample. The Jaccard similarity indexes varied from 0.5 to 0.7. We could not find a relationship between males’ song sharing and geographic distances between their nests. We assume that Whinchat males learned new songs in the local population before territory establishment.     

Flying Foxes Prefer to Forage in Farmland in a Tropical Dry Forest Landscape Mosaic in Fiji

To test flying fox adaptations to a habitat mosaic with extreme deforestation, the abundance, habitat choice and feeding behavior of the Pacific flying fox, Pteropus tonganus, were investigated across 16 islands of the Yasawa archipelago, Fiji. The habitat mosaic is formed by 4.3 percent tropical dry forest and 3.3 percent farmland, leaving exotic grasslands and stands of Leucaena leucocephala to overrun the vast majority of land. Pteropus tonganus abundance was high (5757 bats) despite deforestation and hunting. Roosting sites were restricted to native forest fragments. Grasslands and stands of L. leucocephala were completely void of bats at all times. The mean foraging density in farmland was four times higher than in forests and foraging competition was routinely observed in farmland but was extremely rare in forests. The author suggests that during the study, extensive foraging in farmland was supporting the high P. tonganus population. Additionally, the preferential foraging in farmland was responsible for the low foraging densities within forests and dramatically less intraspecies competition for forest resources. Further research is needed on seed dispersal within forests and to test for seasonal variations in bat abundance and feeding.     

Time-activity budgets and behaviour of the Amazilia hummingbird, Amazilia amazilia (Apodiformes: Trochilidae) in an urban environment

This study deals with the time-activity budgets of Amazilia amazilia, a territorial hummingbird, and its preferences for different flower species and perches in the gardens of Lima (Peru) in September 2001. A. amazilia spent an important part of its time resting on perches (ca. 80%) and only 15.5% for foraging, devoted essentially to flower visitation and only 0.3% for hunting and drinking water. Territorial defence accounted for 2% of total time, mostly against Coereba flaveola, an introduced nectarivorous species that seem to be an important competitor of A. amazilia. Flower use is not directly related to flower abundance (chi(2)9, = 1,546, p<0.0001), with Justicia brandegeana and red-flowered Salvia splendens being selected and Impatiens balsamina being rejected. The large amount of time spent on the perches makes them an important element of the habitat. The perches selected are typically on trees, close to the flowers visited, and in a low vertical and middle horizontal position, surrounded by low foliage density, probably to minimize heat loss.