Consequences of Fragmentation of Tropical Moist Forest for Birds and Their Role in Predation of Herbivorous Insects

The consequences of habitat alteration on the role of understory insectivorous birds as predators of herbivorous insects in tropical forests are poorly understood. To examine whether fragmentation may affect the top–down controls of herbivory, we compared the number of species, individuals, and the community structure of insectivorous birds between fragments and continuous tropical moist forest in Mexico. We also registered insect herbivore abundances and conducted a larvae predation experiment to evaluate the potential role of insectivorous birds as predators of herbivorous insects. We recorded 63 bird species from 22 families, 43 percent of which were insectivorous birds. Species richness, abundance, and diversity of the avian community were higher in continuous forest compared with forest fragments. For insectivorous birds in particular, there was low similarity in avian insectivore communities between forest types, and forest fragments had more heavily dominated communities of avian insectivores. During the dry season, forest fragments presented significantly higher predation rates on artificial caterpillars, and lower abundance of herbivorous Lepidoptera larvae, compared with continuous forest. Furthermore, there was a significant negative correlation between artificial caterpillar predation rate and larval Lepidoptera abundance, with higher rates of predation in sample sites of low Lepidoptera abundance. Hence, the potentially greater light in the dry season combined with a more dominated avian insectivore community in forest fragments may facilitate increased predation by avian insectivores, resulting in a decline in abundance of larval Lepidoptera, with implications for the process of insect‐driven herbivory in forest fragments.     

Predators do not spill over from forest fragments to maize fields in a landscape mosaic in central Argentina

South America is undergoing a rapid and large‐scale conversion of natural habitats to cultivated land. Ecosystem services still remain important but their level and sustainability are not known. We quantified predation intensity in an Argentinian agricultural landscape containing remnants of the original chaco serrano forest using artificial sentinel prey. We sought to identify the main predators and the effect of landscape configuration and maize phenology on predation pressure by invertebrate and vertebrate predators in this landscape. The most common predators were chewing insects (50.4% predation events), birds (22.7%), and ants (17.5%). Overall predation rates in forest fragments (41.6% per day) were significantly higher than in the surrounding maize fields (21.5% per day). Invertebrate predation was higher inside and at the edge of forest fragments than within fields, and did not change with increasing distance from a fragment edge, indicating a lack of spillover from the native habitat remnants to the cultivated matrix at the local scale. Distance from a continuous forest had a positive impact on predation by invertebrates and a negative impact on vertebrate predation.     

Forest fragmentation relaxes natural nest predation in an Afromontane forest

Nest predation is widely regarded as a major driver underlying the population dynamics of small forest birds. Following forest fragmentation and the subsequent invasion by species from non-forested landscape matrices, shifts in predator communities may increase nest predation near forest edges. However, effects of human-driven habitat change on nest predation have mainly been inferred from studies with artificial nests, despite being regarded as poor surrogates for natural ones. We studied variation in predation rates, and relationships with timing of breeding and characteristics of microhabitats and fragments, on natural white-starred robin Pogonocichla stellata nests during three consecutive breeding seasons (2004–2007) in a Kenyan fragmented cloud forest. More than 70% of all initiated nests were predated during each breeding season. Predation rates nearly quadrupled between the earliest and the latest nests within a single breeding season, increased with distance to the forest edge, and decreased with the edge-to-area ratio of forest fragments. These spatial relationships oppose the traditional perception of edge and fragmentation effects on nest predation, but are in line with results from artificial nest experiments in other East African forests. In case of inverse edge and fragmentation effects on nest predation, such as shown in this study, species that tolerate edges for breeding may be affected positively, rather than negatively, by forest fragmentation, while the opposite can be expected for species restricted to the forest interior. The possibility of inverse edge effects, and its conservation implications, should therefore be taken into account when drafting habitat restoration plans.     

Discriminating the Drivers of Edge Effects on Nest Predation: Forest Edges Reduce Capture Rates of Ship Rats (Rattus rattus), a Globally Invasive Nest Predator,by Altering Vegetation Structure

Forest edges can strongly affect avian nest success by altering nest predation rates, but this relationship is inconsistent and context dependent. There is a need for researchers to improve the predictability of edge effects on nest predation rates by examining the mechanisms driving their occurrence and variability. In this study, we examined how the capture rates of ship rats, an invasive nest predator responsible for avian declines globally, varied with distance from the forest edge within forest fragments in a pastoral landscape in New Zealand. We hypothesised that forest edges would affect capture rates by altering vegetation structure within fragments, and that the strength of edge effects would depend on whether fragments were grazed by livestock. We measured vegetation structure and rat capture rates at 488 locations ranging from 0–212 m from the forest edge in 15 forest fragments, seven of which were grazed. Contrary to the vast majority of previous studies of edge effects on nest predation, ship rat capture rates increased with increasing distance from the forest edge. For grazed fragments, capture rates were estimated to be 78% lower at the forest edge than 118 m into the forest interior (the farthest distance for grazed fragments). This relationship was similar for ungrazed fragments, with capture rates estimated to be 51% lower at the forest edge than 118 m into the forest interior. A subsequent path analysis suggested that these ‘reverse’ edge effects were largely or entirely mediated by changes in vegetation structure, implying that edge effects on ship rats can be predicted from the response of vegetation structure to forest edges. We suggest the occurrence, strength, and direction of edge effects on nest predation rates may depend on edge-driven changes in local habitat when the dominant predator is primarily restricted to forest patches.     

Does land use change influence predation of bird nests?

Worldwide, many areas of agricultural land which were once covered with native vegetation have been converted to tree plantations. Such landscape transformation can influence the dynamics of wildlife populations through, for example, altering rates of predation (e.g. predation of nests of birds). Nest predation can influence reproductive success and, in turn, may alter populations by affecting juvenile recruitment. We quantified predation of bird nests in woodland remnants surrounded by two types of land use, grazing farmland and exotic Radiata pine (Pinus radiata) plantation. We also examined differences in predation rates between artificial and natural nests. We found both artificial and natural nests were more susceptible to nest predation in woodland remnants surrounded by a pine plantation than in woodland remnants located within farmland. Our study suggests that higher levels of nest predation may reduce occupancy of woodland remnants by small‐bodied birds over time, including species of conservation concern. This may have been occurred as a result of the conversion of semi‐cleared grazing land to exotic pine plantation.     

Seed predation as a barrier to alien conifer invasions

Interactions between exotic plants and animals can play a major role in determining success or failure of plant introductions. Seed predation has been seen as important in explaining biotic resistance to plant invasion, but this hypothesis has rarely been tested. We studied seed predation on exotic forest plants on an island in Patagonia, Argentina where 43 pine species, including 60% of the world’s known invasive Pinaceae, were introduced ca. 80 years ago, but where exotics attain relatively high densities only near the original plantings. To test if seed predation limits exotic conifer establishment in this area, we compared seed predation in areas close to plantations (colonized by exotics) and far from them (not invaded). Seeds of exotics were preferred over seeds of native species, possibly because exotic seeds are bigger. Predation was more intense in areas far from plantations than in areas close to them, substantially reducing the chances of exotic seed establishment. Using automatic cameras, we found that both rodents and birds preyed on exotic seeds. This study suggests that native seed predators can be an important component of biological resistance to plant invasion.     

Predation of birds in mist nets by callitrichids (primates): how to prevent similar events

We report predation of four birds while caught in mist nets and recommend some means of prevention. Two birds were attacked by Callitrhix jacchus and one by Saguinus midas. The predator in the fourth case was unidentified. These cases were relatively rare, affecting 0.4–4.4% of the captured birds. Two of the predated birds were caught more than 1 m above the ground and may have been accessed from branches. The other two were caught close to the ground. Reducing time intervals between net checks and cutting off branches close to nets may reduce bird predation in mist nets.     

Nesting success of understory forest birds in central Panama

Greater nest predation in tropical than temperate birds has been hypothesized to be a primary selective force generating latitudinal differences in avian life history traits. Few extensive data sets, however, have been available from tropical forests to compare with data from temperate forests. To increase the amount of empirical information available for addressing issues related to the evolution of life history traits of tropical birds, we measured the nesting success of understory birds in lowland forest of central Panama. We found and monitored the fates of 696 nests of 71 species over two breeding seasons. Daily nest predation rates for the ten species for which we obtained the largest samples ranged from 1.6 to 8.3%, equivalent to a loss of 43 to 92% of nests. These values overlapped extensively the range of daily predation rates experienced by ecologically similar species in North America. Proportion of nests fledging young, estimated with the Mayfield method, was significantly lower in tropical (range: 8 to 57%) than temperate (27 to 60%) species. Nesting success in Panama varied among years, however, being greater in 1996 than 1997. In 1996, nesting success was similar to that of species breeding in forest fragments of midwestern North America. When compared with success of nests in large, contiguous forest tracts of North America, however, tropical avian nesting success was consistently lower by approximately 23%. We conclude that nesting success in central Panama may be poor in most breeding seasons, but also may be punctuated by occasional years of relatively exceptional success, a possibility heretofore unappreciated because of a general paucity of data from the tropics. Furthermore, our results indicate substantial variation in levels of nesting success among species, and almost no variation in clutch size. Such large interspecific variation, as well as potentially large annual variation, in nesting success does not support the hypothesis that uniformly low levels of nesting success select for small tropical clutch sizes.     

Indicators of habitat quality and the reproductive output of a forest songbird in small and large fragments

The distribution of individuals among habitats that vary in quality (i.e. resource availability) may affect reproductive output at a population level. I compared indicators of habitat quality including the breeding experience ratios (inexperienced:experienced birds), turnover rates, pairing success, and densities of a forest songbird, the Eastern Yellow Robin Eopsaltria australis , in two small, food-poor forest fragments, with those in two large, food-rich fragments. I then evaluated the likelihood that the breeding experience ratio affected the reproductive output of populations. Inexperienced males occurred in small fragments eight times as often as in large fragments. Male turnover rates were 1.5 times higher in the small than large fragments, and 20% (10/50) of the male population were unpaired in the small fragments compared to 0% (0/25) in the large. None of these measures differed significantly for females. Experienced birds of both sexes produced almost all of the offspring compared with inexperienced birds. Despite these findings, reproductive output did not vary with fragment size for robins. Thus, while breeding experience clearly influenced the reproductive success of individuals, there were no obvious population consequences of having disproportionately more inexperienced males in the small fragments. I conclude that while male traits may be good indicators of resource levels within fragments (specifically, food availability), they may not be adequate predictors of population performance. Thus, the reproductive output of populations must be measured directly before conclusions concerning population performance can be made. Interestingly, breeding densities were not accurate indicators of either resource levels or population performance. Densities were two times higher in the smaller fragments, and I suggest that this result reflects problems with male dispersal among fragments caused by isolation.     

Forest fragmentation and rhinocryptid nest predation in central Chile

Fragmentation of forest landscapes can raise the intensity of nest predation by increasing the abundance and richness of generalist or introduced predators. Understory foraging birds, such as rhinocryptids, can be highly vulnerable to nest predation in fragmented landscapes because they often place their nests on the ground. Temperate deciduous forests in Chile have been intensively fragmented in the last centuries, causing changes in nest predator densities. We tested if predation of artificial nests, mimicking those of rhinocryptids, placed on and above ground was higher in the remnant fragments of central Chile due to an increase in predator abundance. The rate of nest predation in forest remnants was larger than in native continuous forest. Small mammals were the main nest predators. Despite high predation rates, the abundance of rhinocryptids is higher in forest remnants, suggesting that fragments might constitute ecological traps.     

Exotic shrubs as ephemeral ecological traps for nesting birds

Certain exotic plants may increase risk of nest predation, and, in this way, may act as ecological traps. We hypothesized that the greater vulnerability to predation was a consequence of either (1) reduced nest height due to architectural differences among plant species or (2) seasonal changes in the distribution of nests among forest strata. To test this, we examined temporal variation in nest survival of 888 nests of Northern Cardinal (Cardinalis cardinalis) in native substrates and two exotic shrubs (Lonicera maackii and Rosa multiflora) in Ohio, USA, 2001–2006. We evaluated evidence for an ecological trap by monitoring the annual reproductive productivity of 245 breeding pairs of cardinals. Only nests in Rosa experienced relatively constant survival rates across the season, whereas probability of survival increased over the season for nests in other substrates. Interestingly, the relative vulnerability of nests in different substrates varied across the season. Most strikingly, nests in Lonicera in early spring showed the lowest survival rates but exceeded survival rates of nests in native substrates late in the season. Nest height failed to explain seasonal changes in nest survival, as only nests in native plants significantly increased in height as the season progressed. Rather, predation risk seemed to be a function of the proportion of nests within each substrate, as illustrated by the decreased predation in Lonicera as the relative proportion of nests in native substrates increased. The patterns of temporal variation in predation risk that we detected show that impacts of Lonicera are not a function of plant architecture alone and may be related to leaf phenology, changes in nest density, nest site location, and/or nest synchrony. Examination of the reproductive productivity of cardinals showed that pairs that made their first nest attempt in Lonicera fledged 20% fewer cardinal young than birds that began the season using other substrates. Thus, we suggest that exotic plants may represent an ephemeral ecological trap for certain nesting birds, where negative effects persist only during certain periods.     

Nesting Success of Costa Rican Lowland Rain Forest Birds in Response to Edge and Isolation Effects

Although open-cup nesting birds generally face increased risk of nest depredation from forest edge predators and brood parasites in fragmented temperate landscapes, little information exists to assess such risks in tropical birds. We compared nesting success of real birds' nests in large and small forest fragments to a control site in Caribbean lowland wet forest of Costa Rica. Pooling across species, nesting success was significantly greater in unfragmented forest than in either small, isolated fragments or the La Selva Biological Reserve, which is at the tip of a forest 'peninsula' embedded in a largely deforested landscape. Nesting success in isolated fragments did not vary according to distance from edge, suggesting that predators in fragments act throughout these forest patches. The case for increased nest predation as a plausible mechanism to explain the documented decline of forest interior bird populations in this fragmented tropical landscape is enhanced by a simple demographic model that suggests nesting success is likely too low to maintain populations at La Selva and in the fragments. The fact that the large (> 1000 ha) La Selva forest reserve is experiencing nest predation rates similar to those in much smaller fragments is cause for concern. Our results make a strong case for additional studies to document the identities of nest predators in both fragmented and unfragmented forests in such tropical forest landscapes.     

Nest predation patterns in ground-nesting passerines on the Iberian Peninsula

The idea that ground-nesting birds have a high risk of predation is widely accepted This paper analyzes the nest predation patterns of ground-nesting passerines on the Iberian Peninsula The nest predation rates are higher m open land species x= 71%) than m countryside farmland (10%) or forest birds (29%) The latter species have no differences in daily survival between ground and above-ground nests The higher nest predation in ground-nesting open land species was similar m peninsular Spain and North America We conclude that it is incorrect to generalize that all ground nesters have high nest predation rates, and we discuss the possible link of the high nest predation rates of open land birds to the decline observed in their population trends in Spain     

Decreased frugivory and seed germination rate do not reduce seedling recruitment rates of <Emphasis Type="Italic">Aristotelia chilensis</Emphasis> in a fragmented forest

Habitat fragmentation reduces frugivorous bird abundance. Such a reduction may lead to a reduction in seed dispersal, thereby compromising seedling recruitment rate with far reaching consequences for plant population persistence. We assessed frugivory, seed germination, and seedling recruitment rates in a fragmented forest of central Chile by comparing a continuous forest with four forest fragments surrounded by pine plantations. Frugivory was 2.4 times higher in continuous forest than in forest fragments. Seeds eaten by birds germinated 1.7 and 3.7 times higher than non-eaten seeds from continuous forest and fragments respectively. Non-eaten seeds from continuous forest germinated 2.2 times higher than those from forest fragments, suggesting inbreeding depression. However, seedling recruitment rates at forest fragments were far higher than in continuous forest where no seedling recruited in the five years analysed. Therefore, despite forest fragmentation negatively affected frugivory, it did not translate into a decreased fitness of plants, thus highlighting the importance of considering the overall processes leading the reproductive success of plants following anthropogenic disturbances.     

Artificial nest predation in hedgerows and scrub forest in a human-dominated landscape of central Mexico

Hedgerows as well as other narrow corridors could be valuable habitats for birds in regions of intensive agriculture, however, it is still not clear how successful breeding birds are in different types of hedgerows as compared to birds nesting in their natural habitats. We used artificial nests to examine whether hedgerows were sinks (ecological traps) for birds by comparing rates of predation in two types of hedgerows with different vegetation structure (simple and complex), and in a tract of scrub forest in an agricultural landscape of central Mexico. We determined also the types of predators responsible for egg predation. Ground and elevated nests were baited with one Japanese quail Coturnix japonica egg and one plasticine egg and placed alternately along transects. Significantly, greater predation rates were found in scrub forest and complex hedgerows than in simple hedgerows. Higher predation rates in complex habitats seemed to reflect the higher number of predator types found there. The most important predator types were carnivores followed by rodents, birds, and humans. Carnivores and rodents mainly predated ground nests, whereas birds and humans predated elevated nests. Simple hedgerows in this landscape appeared to offer relatively safe nest sites in terms of predation pressure when compared to more complex habitats (complex hedgerows and scrub forest).     

Forest fragmentation and avian nest predation in forested landscapes

Summary The size of forest fragments, the use of land bordering fragments, and the distance of nests from an edge all affect the frequency of predation upon bird nests in Maine (USA), an area where the forest has been fragmented by roads, but not significantly reduced in area. We placed artificial nests containing quail eggs in forests of different sizes and at various distances from the edge to test which of these factors was most important in describing predation. Predation was greatest in small tracts surrounded completely by land. Large areas and those bordered on at least one side by a large water body had lower predation rates. This suggests that influx of predators from nearby habitats may be responsible for much of the nest predation in forest fragments.     

Introduced mammalian predators induce behavioural changes in parental care in an endemic New Zealand bird

The introduction of predatory mammals to oceanic islands has led to the extinction of many endemic birds. Although introduced predators should favour changes that reduce predation risk in surviving bird species, the ability of island birds to respond to such novel changes remains unstudied. We tested whether novel predation risk imposed by introduced mammalian predators has altered the parental behaviour of the endemic New Zealand bellbird (Anthornis melanura). We examined parental behaviour of bellbirds at three woodland sites in New Zealand that differed in predation risk: 1) a mainland site with exotic predators present (high predation risk), 2) a mainland site with exotic predators experimentally removed (low risk recently) and, 3) an off-shore island where exotic predators were never introduced (low risk always). We also compared parental behaviour of bellbirds with two closely related Tasmanian honeyeaters (Phylidonyris spp.) that evolved with native nest predators (high risk always). Increased nest predation risk has been postulated to favour reduced parental activity, and we tested whether island bellbirds responded to variation in predation risk. We found that females spent more time on the nest per incubating bout with increased risk of predation, a strategy that minimised activity at the nest during incubation. Parental activity during the nestling period, measured as number of feeding visits/hr, also decreased with increasing nest predation risk across sites, and was lowest among the honeyeaters in Tasmania that evolved with native predators. These results demonstrate that some island birds are able to respond to increased risk of predation by novel predators in ways that appear adaptive. We suggest that conservation efforts may be more effective if they take advantage of the ability of island birds to respond to novel predators, especially when the elimination of exotic predators is not possible.     

Prevalence of nest predators in a sub-Arctic ecosystem

High nest loss is an important driver of gallinaceous bird population dynamics. Identifying factors determining the spatial distribution of potential nest predators and thereby indirectly risk of nest losses is therefore essential. The aim of this 1-year study was to estimate relative predation rates on artificial ground nests in willow ptarmigan (Lagopus lagopus) habitats, along replicate altitudinal gradients (transects, n?=?60) spanning from sub-Arctic birch forest to the low-alpine tundra in three locations in northern Norway. In each transect, one artificial nest (track board) was placed in three different habitats: (1) birch forest, (2) edge between birch forest and low-alpine tundra and (3) low-alpine tundra. Total predation rates over all habitats within locations ranged from 47.4% to 77.5% and did not vary systematically in space and time. The average predation rate by avian predators was consistently high (58%), and mammalian predation rate was consistently low (5.6%). The consistently high level of predation inflicted by birds was mainly due to omnipresent corvids, especially the hooded crow (Corvus cornix). Analysis of species-specific predation rates showed that habitat and location effects were insignificant for all species, except for raven (Corvus corax) that showed clearly higher predation in one of the locations. The results indicate that from the perspective of the spatial distribution of potential nest predators in sub-Arctic birch forest, ground nesting birds like willow ptarmigan should not be expected to be selective with respect to nesting habitat in the ecotone between birch forest and the low-alpine tundra.     

Effects of avian mobbing on roost use and diet of powerful owls,Ninox strenua

We observed the species and numbers of mobbing birds and their effects on a large, nocturnal, bird-eating predator, the powerful owl, together with the pattern of owl predation on mobbing and non-mobbing species. Owls were mobbed on 35 occasions by seven of 44 species of forest birds at a site composed of open forest (88% by area) and rainforest (12%). The majority of bouts involved individuals of a single species, although mixed groups were observed on nine occasions. Regular mobbers were between 4 and 26% of the owls’ body weight. Owls abandoned their daytime roosts during 20% of bouts and responded by calling or actively monitoring mobbers during 54% of bouts. Mobbing appeared to explain why owls roosted in rainforest significantly more often than expected by its availability, mobbing being significantly less frequent in rainforest than in open forest. Only one mobbing species regularly occupied rainforest and the canopy of roosts in rainforest was denser than that in open forest, thus reducing the chances of an owl being detected by potential mobbers. Twelve species of forest birds were within the range of prey size of the powerful owl (75–800 g): six were mobbers and six non-mobbers. The frequency of owl predation on non-mobbers was 8.75 times that on mobbers. The species in this study took a high risk by mobbing a very large predator, but benefited by greatly reducing their chances of predation.     

Predicting Summer Site Occupancy for an Invasive Species,the Common Brushtail Possum (Trichosurus vulpecula), in an Urban Environment

Invasive species are often favoured in fragmented, highly-modified, human-dominated landscapes such as urban areas. Because successful invasive urban adapters can occupy habitat that is quite different from that in their original range, effective management programmes for invasive species in urban areas require an understanding of distribution, habitat and resource requirements at a local scale that is tailored to the fine-scale heterogeneity typical of urban landscapes. The common brushtail possum (Trichosurus vulpecula) is one of New Zealand’s most destructive invasive pest species. As brushtail possums traditionally occupy forest habitat, control in New Zealand has focussed on rural and forest habitats, and forest fragments in cities. However, as successful urban adapters, possums may be occupying a wider range of habitats. Here we use site occupancy methods to determine the distribution of brushtail possums across five distinguishable urban habitat types during summer, which is when possums have the greatest impacts on breeding birds. We collected data on possum presence/absence and habitat characteristics, including possible sources of supplementary food (fruit trees, vegetable gardens, compost heaps), and the availability of forest fragments from 150 survey locations. Predictive distribution models constructed using the programme PRESENCE revealed that while occupancy rates were highest in forest fragments, possums were still present across a large proportion of residential habitat with occupancy decreasing as housing density increased and green cover decreased. The presence of supplementary food sources was important in predicting possum occupancy, which may reflect the high nutritional value of these food types. Additionally, occupancy decreased as the proportion of forest fragment decreased, indicating the importance of forest fragments in determining possum distribution. Control operations to protect native birds from possum predation in cities should include well-vegetated residential areas; these modified habitats not only support possums but provide a source for reinvasion of fragments.