Impacts of Roads, Hunting, and Habitat Alteration on Nocturnal Mammals in African Rainforests

Abstract:  Nocturnal mammals are poorly studied in Central Africa, a region experiencing dramatic increases in logging, roads, and hunting activity. In the rainforests of southern Gabon, we used spotlighting surveys to estimate abundances of nocturnal mammal species and guilds at varying distances from forest roads and between hunted and unhunted treatments (comparing a 130-km² oil concession that was nearly free of hunting, with nearby areas outside the concession that had moderate hunting pressure). At each of 12 study sites that were evenly divided between hunted and unhunted areas, we established standardized 1-km transects along road verges and at 50, 300, and 600 m from the road. We then repeatedly surveyed mammals at each site during 2006. Hunting had few apparent effects on this assemblage. Nevertheless, the species richness and often the abundance of nocturnal primates, smaller ungulates, and carnivores were significantly depressed within approximately 30 m of roads. Scansorial rodents increased in abundance in hunted forests, possibly in response to habitat changes caused by logging or nearby swidden farming. In multiple-regression models many species and guilds were significantly influenced by forest-canopy and understory cover, both of which are altered by logging and by certain abiotic variables. In general, nocturnal species, many of which are arboreal or relatively small in size (<10 kg), were less strongly influenced by hunting and more strongly affected by human-induced changes in forest structure than were larger mammal species in our study area.     

Distribution of a Community of Mammals in Relation to Roads and Other Human Disturbances in Gabon,Central Africa

We present the first community‐level study of the associations of both roads and other human disturbances with the distribution of mammals in Gabon (central Africa). Our study site was in an oil concession within a littoral mosaic landscape. We conducted surveys along 199 line transects and installed camera traps on 99 of these transects to document mammal presence and abundance. We used generalized linear mixed‐effect models to document associations between variables related to the ecosystem (land cover, topography, and hydrology), roads (coating, width of rights of way, condition, type of vehicle used on the road, traffic level, affiliation of users, and general type of road), and other human disturbances (urbanization, agriculture, hunting, logging, gathering, and industrial activities) and the abundance or presence of 17 species or groups of mammals including elephant (Loxodonta cyclotis), buffalo (Syncerus caffer), sitatunga (Tragelaphus spekei), red river hog (Potamochoerus porcus), smaller ungulates, gorilla (Gorilla gorilla), chimpanzee (Pan troglodytes), side‐striped jackal (Canis adustus), carnivores, monkeys, and large rodents. Some types of roads and other human disturbances were negatively associated with the abundance or presence of elephants, buffalos, gorillas, sitatungas, some monkeys, and duikers. The pattern of associations of mammals with roads and other human disturbances was diverse and included positive associations with road presence (red river hog, some monkeys, and duikers), agriculture (sitatunga, small carnivores, and large rodents) and industrial activities (sitatunga, red river hog, red duikers, and side‐striped jackal). Our results suggest that the community of mammals we studied was mostly affected by hunting, agriculture, and urbanization, which are facilitated by road presence. We recommend increased regulation of agriculture, hunting, and road building in the area. Distribución de una Comunidad de Mamíferos en Relación a Carreteras y Otras Perturbaciones Humanas en Gabón, Africa Central     

Logging Concessions Can Extend the Conservation Estate for Central African Tropical Forests

Abstract:  The management of tropical forest in timber concessions has been proposed as a solution to prevent further biodiversity loss. The effectiveness of this strategy will likely depend on species-specific, population-level responses to logging. We conducted a survey (749 line transects over 3450 km) in logging concessions (1.2 million ha) in the northern Republic of Congo to examine the impact of logging on large mammal populations, including endangered species such as the elephant ( Loxodonta africana ), gorilla ( Gorilla gorilla ), chimpanzee ( Pan troglodytes ), and bongo ( Tragelaphus eurycerus ). When we estimated species abundance without consideration of transect characteristics, species abundances in logged and unlogged forests were not different for most species. When we modeled the data with a hurdle model approach, however, analyzing species presence and conditional abundance separately with generalized additive models and then combining them to calculate the mean species abundance, species abundance varied strongly depending on transect characteristics. The mean species abundance was often related to the distance to unlogged forest, which suggests that intact forest serves as source habitat for several species. The mean species abundance responded nonlinearly to logging history, changing over 30 years as the forest recovered from logging. Finally the distance away from roads, natural forest clearings, and villages also determined the abundance of mammals. Our results suggest that logged forest can extend the conservation estate for many of Central Africa's most threatened species if managed appropriately. In addition to limiting hunting, logging concessions must be large, contain patches of unlogged forest, and include forest with different logging histories.     

Effect of Small‐Scale Heterogeneity of Prey and Hunter Distributions on the Sustainability of Bushmeat Hunting

Abstract: Bushmeat is the main source of protein and the most important source of income for rural people in the Congo Basin, but intensive hunting of bushmeat species is also a major concern for conservationists. Although spatial heterogeneity in hunting effort and in prey populations at the landscape level plays a key role in the sustainability of hunted populations, the role of small‐scale heterogeneity within a village hunting territory in the sustainability of hunting has remained understudied. We built a spatially explicit multiagent model to capture the dynamics of a system in which hunters and preys interact within a village hunting territory. We examined the case of hunting of bay duikers (Cephalophus dorsalis) in the village of Ntsiété, northeastern Gabon. The impact of hunting on prey populations depended on the spatial heterogeneity of hunting and prey distribution at small scales within a hunting area. Within a village territory, the existence of areas hunted throughout the year, areas hunted only during certain seasons, and unhunted areas contributed to the sustainability of the system. Prey abundance and offtake per hunter were particularly sensitive to the frequency and length of hunting sessions and to the number of hunters sharing an area. Some biological parameters of the prey species, such as dispersal rate and territory size, determined their spatial distribution in a hunting area, which in turn influenced the sustainability of hunting. Detailed knowledge of species ecology and behavior, and of hunting practices are crucial to understanding the distribution of potential sinks and sources in space and time. Given the recognized failure of simple biological models to assess maximum sustainable yields, multiagent models provide an innovative path toward new approaches for the assessment of hunting sustainability, provided further research is conducted to increase knowledge of prey species’ and hunter behavior.     

Forest Roads as Partial Barriers to Terrestrial Salamander Movement

Abstract:  Roads can fragment animal populations by disrupting movement among formerly continuous habitats. Although models have demonstrated that disrupted movement can contribute to long-term extinction, there are few empirical data on the effects of roads on animal movement. We used displacement and homing experiments to determine whether forest roads are barriers to the movement of terrestrial salamanders. We displaced 1471 red-backed salamanders ( Plethodon cinereus ) across five forest roads and compared return rates to those of salamanders displaced equal distances toward the forest interior. Roads significantly reduced the return rate of salamanders, with a mean reduction of 51%. Steep roadside verges further reduced return rates, particularly for salamanders moving downhill across verges. The permeability of roads to salamander movement did not appear to be related to road surface type. Gravel roads had both the highest and lowest observed permeability with the two paved roads intermediate between these. We conclude that narrow forest roads are partial barriers to salamander movement and that steep roadside verges may exacerbate these effects.     

Hunting for Sustainability in Tropical Secondary Forests

Abstract:  The interaction between land-use change and the sustainability of hunting is poorly understood but is critical for sustaining hunted vertebrate populations and a protein supply for the rural poor. We investigated sustainability of hunting in an Amazonian landscape mosaic, where a small human population had access to large areas of both primary and secondary forest. Harvestable production of mammals and birds was calculated from density estimates. We compared production with offtake from three villages and used catch-per-unit-effort as an independent measure of prey abundance. Most species were hunted unsustainably in primary forest, leading to local depletion of the largest primates and birds. The estimated sustainable supply of wild meat was higher for primary (39 kg · km⁻²· yr⁻¹) than secondary forest (22 kg · km⁻²· yr⁻¹) because four species were absent and three species at low abundance in secondary forests. Production of three disturbance-tolerant mammal species was 3 times higher in secondary than in primary forest, but hunting led to overexploitation of one species. Our data suggest that an average Amazonian smallholder would require ≥3.1 km² of secondary regrowth to ensure a sustainable harvest of forest vertebrates. We conclude that secondary forests can sustainably provide only 2% of the required protein intake of Amazonian smallholders and are unlikely to be sufficient for sustainable hunting in other tropical forest regions.     

Effect of Hunting in Source-Sink Systems in the Neotropics

Abstract: Previous studies of the sustainability of wildlife hunting in the Neotropics have not considered the potential dispersal of animals into hunted areas. A literature review of studies of subsistence hunting in the Neotropics suggests that hunting is often conducted in areas adjacent to relatively undisturbed habitat that may act as sources of animals for the hunted sites. We compared studies of tapir (   Tapirus terrestris ) hunting at different sites to illustrate the potential bias of sustainability evaluations based on local productivity. The limited information available suggests that dispersal could have a key role in rebuilding animal populations depleted by hunting. Thus, factors that strongly affect dispersal—such as spatial distribution and size of areas with and without hunting, population size in source areas, and social behavior—should be considered when the sustainability of hunting is evaluated in areas with heterogeneous hunting pressure. We suggest the application of two models that use spatial controls (recognizing the potential source-sink nature of some hunted systems and protecting unhunted refugia) to avoid wildlife overexploitation when biological data and enforcement capabilities to regulate harvests are limited. This approach may produce more reliable evaluations of sustainability, provide information on the dynamics of hunting systems, and help local communities and policymakers conserve key areas (including protected areas) that may act as game sources.     

Effects of Subsistence Hunting on Vertebrate Community Structure in Amazonian Forests

Abstract: Subsistence hunting affects vast tracts of tropical wilderness that otherwise remain structurally unaltered, yet distinguishing hunted from nonhunted tropical forests presents a difficult problem because this diffuse form of resource extraction leaves few visible signs of its occurrence. I used a standardized series of line-transect censuses conducted over a 10-year period to examine the effects of subsistence game harvest on the structure of vertebrate communities in 25 Amazonian forest sites subjected to varying levels of hunting pressure. Crude vertebrate biomass, which was highly correlated with hunting pressure, gradually declined from nearly 1200 kg km⁻² at nonhunted sites to less than 200 kg km⁻² at heavily hunted sites. Hunting had a negative effect on the total biomass and relative abundance of vertebrate species in different size classes at these forest sites, but it did not affect their overall density. In particular, persistent hunting markedly reduced the density of large-bodied game species (>5 kg), which contributed a large proportion of the overall community biomass at nonhunted sites (65–78%) and lightly hunted sites (55–71%). Nutrient-rich floodplain forests contained a consistently greater game biomass than nutrient-poor unflooded forests, once I controlled for the effects of hunting pressure. Conservative estimates of game yields indicate that as many as 23.5 million game vertebrates, equivalent to 89,224 tons of bushmeat with a market value of US$190.7 million, are consumed each year by the rural population of Brazilian Amazonia, which illustrates the enormous socioeconomic value of game resources in the region. My cross-site comparison documents the staggering effect of subsistence hunters on tropical forest vertebrate communities and highlights the importance of considering forest types and forest productivity in game management programs.     

Effects of Forest Roads on Macroinvertebrate Soil Fauna of the Southern Appalachian Mountains

Abstract: Many forested landscapes are fragmented by roads, but our understanding of the effects of these roads on the function and diversity of the surrounding forest is in its infancy. I investigated the effect of roads in otherwise continuous forests on the macroinvertebrate fauna of the soil. I took soil samples along transects leading away from the edges of unpaved roads in the Cherokee National Forest in the Southern Appalachian mountains of the United States. Roads significantly depressed both the abundance and the richness of the macroinvertebrate soil fauna. Roads also significantly reduced the depth of the leaf-litter layer. These effects persisted up to 100 m into the forest. Wider roads and roads with more open canopies tended to produce steeper declines in abundance, richness, and leaf-litter depth, but these effects were significant only for canopy cover and litter depth. The macroinvertebrate fauna of the leaf litter plays a pivotal role in the ability of the soil to process energy and nutrients. These macroinvertebrates also provide prey for vertebrate species such as salamanders and ground-foraging birds. The effect of roads on the surrounding forest is compounded by the sprawling nature of the road system in this and many other forests. My data suggest that even relatively narrow roads through forests can produce marked edge effects that may have negative consequences for the function and diversity of the forest ecosystem.     

Rapid Evolution in the Wild: Changes in Body Size, Life-History Traits, and Behavior in Hunted Populations of the Japanese Mamushi Snake

Abstract:  Rapid evolution caused by human exploitation of wildlife is not usually addressed in studies of the impacts of such exploitation despite its direct relevance to population persistence. Japanese mamushi ( Gloydius blomhoffii ), an endemic venomous snake of the Japanese archipelago, has been heavily hunted by humans, and many populations appear to be declining or are already extirpated. We compared local populations that have been hunted regularly with populations that have not been hunted. Mamushi in hunted populations were smaller, had fewer vertebrae, produced more and smaller offspring, had increased reproductive effort among smaller females, and in nature fled at greater distances from an approaching human and were less defensive than mamushi in unhunted populations, as predicted from life-history theory. Heritability estimates for body size, number of vertebrae, and antipredator behavior were statistically significant, and neonates from hunted sites showed the same distribution of altered characters (compared with those from unhunted sites) as adults. Thus, distribution of the divergent trait between hunted and unhunted sites appeared in part to be genetically based, which suggests rapid evolution to human predation pressures. Trait distributions in hunted populations probably deviate from naturally (as opposed to anthropogenically) selected optima and, therefore, may have long-term negative repercussions on population persistence. Because rapid evolution affects a suite of parameters that characterize exploited populations, accurate understanding of the impacts of exploitation and effective resource management and conservation can only be achieved if evolutionary consequences are considered explicitly.     

Human Hunting of Nongame Birds at Zuni, New Mexico, U.S.A.

Abstract: Nongame bird hunting is a critical activity of the Zuni people of the southwestern United States. To help determine whether their current hunting practices may be negatively affecting bird populations on the Zuni Reservation, we interviewed 98 Zuni hunters. Nongame bird hunting was practiced by a large portion of Zuni males (45%); the most active age group was 20- to 49-year-olds. The rate a species was hunted was not a function of its abundance at Zuni but seemed instead to be related to its cultural demand. Five "species"—bluebirds ( Sialia currucoides , S. mexicana ), Northern Flicker ( Colaptes auratus ), woodpeckers ( Picoides villosus , P. pubescens , Melanerpes lewis ), Steller's Jay ( Cyanocitta stelleri ), and American Kestrel ( Falco sparverius )—made up 77% of all birds taken. The two most heavily hunted species were each taken in numbers> 10,000 individuals per year. Although the greatest number of hunters were active in autumn, 31% hunted in spring, despite discouragement by the tribal government. Habitats favored by hunters were coniferous forests and riparian areas. Rates of hunting of nongame birds at Zuni equaled or exceeded those reported for game birds hunted by indigenous hunters in the Neotropics and New Mexico. The Zuni believe that some heavily hunted species are decreasing in number because of hunting. Although we cannot prove this at present, we suggest measures to mitigate possible overhunting, including the creation of refugia and a reduction of spring hunting.     

Salamander Abundance along Road Edges and within Abandoned Logging Roads in Appalachian Forests

Abstract:  Roads may be one of the most common disturbances in otherwise continuous forested habitat in the southern Appalachian Mountains. Despite their obvious presence on the landscape, there is limited data on the ecological effects along a road edge or the size of the "road-effect zone." We sampled salamanders at current and abandoned road sites within the Nantahala National Forest, North Carolina (U.S.A.) to determine the road-effect zone for an assemblage of woodland salamanders. Salamander abundance near the road was reduced significantly, and salamanders along the edges were predominantly large individuals. These results indicate that the road-effect zone for these salamanders extended 35 m on either side of the relatively narrow, low-use forest roads along which we sampled. Furthermore, salamander abundance was significantly lower on old, abandoned logging roads compared with the adjacent upslope sites. These results indicate that forest roads and abandoned logging roads have negative effects on forest-dependent species such as plethodontid salamanders. Our results may apply to other protected forests in the southern Appalachians and may exemplify a problem created by current and past land use activities in all forested regions, especially those related to road building for natural-resource extraction. Our results show that the effect of roads reached well beyond their boundary and that abandonment or the decommissioning of roads did not reverse detrimental ecological effects; rather, our results indicate that management decisions have significant repercussions for generations to come. Furthermore, the quantity of suitable forested habitat in the protected areas we studied was significantly reduced: between 28.6% and 36.9% of the area was affected by roads. Management and policy decisions must use current and historical data on land use to understand cumulative impacts on forest-dependent species and to fully protect biodiversity on national lands     

Effects of Landscape Features on the Distribution and Sustainability of Ungulate Hunting in Northern Congo

Abstract: Understanding the spatial dimensions of hunting and prey population dynamics is important in order to estimate the sustainability of hunting in tropical forests. We investigated how hunting offtake of vertebrates differed in mixed forest and monodominant forest (composed of Gilbertiodendron dewevrei) and over different spatial extents within the hunting catchment around the logging town of Kabo, Congo. In 9 months of recall surveys with hunters, we gathered information on over 1500 hunting trips in which ungulates were 65% of the species killed and 82% of harvested biomass. Hunters supplied information on animals killed and the hunting trip, including the area visited (i.e., hunting zone; 11 separate zones within a 506 km² catchment or commonly hunted area). Over 65% of all animals were killed in monodominant forest, which made up 28% of the hunting catchment, and zones with small amounts of monodominant forest were used most frequently by hunters. Given the large offtakes from monodominant forests, we suggest that animal dispersal may be maintaining high, localized harvests in these areas. We believe hunters preferred to hunt in monodominant forest because the understory was accessible and that areas with small amounts of monodominant forest and large amounts of mixed forest were more productive. The variation in hunting pressure we found between and within hunting zones differs from past examinations of spatial variation in hunting offtake, where entire hunting catchments were considered population sinks and areas with low to no hunting (no‐take zones) were outside hunting catchments. Future use of no‐take zones to manage hunting should incorporate variability in offtake within hunting catchments.     

Disturbance-Mediated Mammal Persistence and Abundance-Area Relationships in Amazonian Forest Fragments

Abstract:  Few researchers have investigated the synergistic effects of tropical forest fragmentation and disturbance on species persistence and abundance. We examined effects of both forest-patch metrics and forest disturbance in determining richness and abundance of midsized to large-bodied mammal species in a highly fragmented Amazonian forest landscape. Twenty-one forest fragments, ranging from 2 to 14,480 ha, and two continuous forest sites were sampled based on sightings, tracks, line-transect censuses, armadillo burrow censuses, and camera trapping. Patch occupancy of 37 species recorded ranged from 4% to all forest sites surveyed. Forest fragment size was the strongest predictor of species persistence, explaining 90% of the variation in species richness. Information-theoretic analysis confirmed that fragment area was the most important explanatory variable for the overall species richness and abundance of mammal species, followed by surface fires, which affected the abundance of seven species. Large mammal species were typically absent from fragments <100 ha, whereas some ubiquitous species were favored by fragmentation, exhibiting hyperabundance in small patches. Our findings highlight the importance of large (>10,000 ha), relatively undisturbed forest patches to maximize persistence and maintain baseline abundances of Neotropical forest mammal species.     

Bushmeat Supply and Consumption in a Tropical Logging Concession in Northern Congo

Abstract: Unsustainable hunting of wildlife for food empties tropical forests of many species critical to forest maintenance and livelihoods of forest people. Extractive industries, including logging, can accelerate exploitation of wildlife by opening forests to hunters and creating markets for bushmeat. We monitored human demographics, bushmeat supply in markets, and household bushmeat consumption in five logging towns in the northern Republic of Congo. Over 6 years we recorded 29,570 animals in town markets and collected 48,920 household meal records. Development of industrial logging operations led to a 69% increase in the population of logging towns and a 64% increase in bushmeat supply. The immigration of workers, jobseekers, and their families altered hunting patterns and was associated with increased use of wire snares and increased diversity in the species hunted and consumed. Immigrants hunted 72% of all bushmeat, which suggests the short‐term benefits of hunting accrue disproportionately to “outsiders” to the detriment of indigenous peoples who have prior, legitimate claims to wildlife resources. Our results suggest that the greatest threat of logging to biodiversity may be the permanent urbanization of frontier forests. Although enforcement of hunting laws and promotion of alternative sources of protein may help curb the pressure on wildlife, the best strategy for biodiversity conservation may be to keep saw mills and the towns that develop around them out of forests.     

Hunting and the Likelihood of Extinction of Amazonian Mammals

Species inhabiting tropical forests are thought to be on the verge of mass extinction. Much work has focused on extinction rates caused by deforestation; however, many of the recorded extinctions that have occurred since 1600 were a result of overhunting. We collected data on the relative abundance of large-bodied mammals in the northeastern Peruvian Amazon in areas with persistent hunting pressure and in areas with infrequent hunting pressure. We quantified the effects of hunting by calculating the change in abundance of species between the infrequently and persistently hunted sites. We report that in Amazonian mammals weighing more than 1 kg the degree of population declines caused by hunting is correlated with the species' intrinsic rate of natural increase (r_max  ), longevity, and generation time. Our results show that species with long-lived individuals, low rates of increase, and long generation times are more vulnerable to extinction than species with short-lived individuals, high rates of increase, and shorter generations.     

Contribution of Roads to Forest Fragmentation in the Rocky Mountains

The contribution of roads to forest fragmentation has not been adequately analyzed. We quantified fragmentation due to roads in a 30,213-ha section of the Medicine Bow-Routt National Forest in sout heastern Wyoming with several indices of landscape structure using a geographic information system. The number of patches, mean patch area, mean interior area, mean area of edge influence, mean patch perimeter, total perimeter, and mean patch shape identified patch- and edge-related landscape changes. Shannon-Wiener diversity, dominance, contagion, contrast, and angular second moment indicated effects on landscape diversity and texture. Roads added to forest fragmentation more than clearcuts by dissecting large patches into smaller pieces and by converting forest interior habitat into edge habitat. Edge habitat created by roads was 1.54–1.98 times the edge habitat created by clearcuts. The total landscape area affected by clearcuts and roads was 2.5–3.5 times the actual area occupied by these disturbances. Fragmentation due to roads could be minimized if road construction is minimized or rerouted so that its fragmentation effects are reduced. Geographic information system technology can be used to quantify the potential fragmentation effects of individual roads and the cumulative effects of a road network on landscape structure.     

Plant-Mammal Interactions in Tropical Bolivian Forests with Different Hunting Pressures

Abstract: We studied plant-animal interactions and vegetation structure in two geographically close tropical Bolivian forests subjected to different hunting intensities. We hypothesized that reduction of mammals openface> 1 kg in an "intensively hunted forest," compared with an "occasionally hunted forest," should correlate with decreased seed predation and seedling trampling, increased seedling survival and density, and decreased tree-species diversity at the seedling stage in relation to the adult stage. The occasionally hunted forest held 1.7 times as many mammalian species as the intensively hunted forest. As predicted, predation of Astrocaryum murumuru seeds was 34.2% lower in the intensively hunted forest. Similarly, trampling of model seedlings was 5.4 times lower and seedling survival was 1.15 times greater in the intensively hunted forest than that in the occasionally hunted forest. But the intensively hunted forest displayed lower seedling densities and a higher ratio of seedling diversity to tree diversity than did the occasionally hunted forest. Reduction of peccaries from the intensively hunted forest may explain much of the between-site differences in seed predation, trampling, and seedling survival. Lack of consistent differences in seedling density and diversity could mean that reduced granivory and trampling may be counteracted by reduced seed dispersal. The reduction of mammalian populations may produce a complex mosaic of forest patches with distinct degrees of structural change, depending on the intensity of defaunation.     

Effects of Roads on Movements by Understory Birds in Mixed-Species Flocks in Central Amazonian Brazil

Abstract: Roads through tropical forest create linear disturbances that have unknown consequences for forest birds. We studied how a narrow, rarely used road through otherwise undisturbed Amazonian forest affected the movements and area requirements of understory birds that form mixed-species flocks. Differences in road maintenance led to two distinct treatments along the same road. Trees along the "closed" road formed a partial canopy connecting the two sides of the road, although the roadway itself was kept open. The "open" road was regularly maintained, making a complete opening 10–30 m wide.We followed 15 flocks, 5 each in interior forest, along the open road, and along the closed road. These flocks were led by Thamnomanes antshrikes, and each flock had a discreet, permanent territory. Flock territory size (mean = 8.5 ha) did not differ among the three locations. The open road formed the territorial boundary for all five flocks, although birds moved within a few meters of the edge of the road. The closed road was less of a barrier: 2 of 5 flocks used both sides of the closed road. Playback experiments showed that flocks readily crossed the closed road to approach agonistic vocalizations. Along the open road, even though birds responded to playback by becoming agitated and moving to the extreme edge of the roadside vegetation, they were less likely to cross the road and did so only after a longer duration of playback. Our results suggest that flocks respond to a road as they would to a long linear gap. They use the vegetation along the edges of the road, but because they are unwilling to cross the open area, it becomes a flock territory boundary. Similarly, as in forest gaps, successional change along the closed road produced suitable habitat for flocks. Although this suggests that roads are a trivial problem, we caution that this result applies only to narrow roads that are not accompanied by deforestation or other disturbance.