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.     

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.     

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.     

Impacts of Roads and Hunting on Central African Rainforest Mammals

Abstract:  Road expansion and associated increases in hunting pressure are a rapidly growing threat to African tropical wildlife. In the rainforests of southern Gabon, we compared abundances of larger (>1 kg) mammal species at varying distances from forest roads and between hunted and unhunted treatments (comparing a 130-km² oil concession that was almost entirely protected from 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 at five distances (50, 300, 600, 900, and 1200 m) from an unpaved road, and then repeatedly surveyed mammals during the 2004 dry and wet seasons. Hunting had the greatest impact on duikers ( Cephalophus spp. ), forest buffalo ( Syncerus caffer nanus ), and red river hogs ( Potamochoerus porcus ), which declined in abundance outside the oil concession, and lesser effects on lowland gorillas ( Gorilla gorilla gorilla ) and carnivores. Roads depressed abundances of duikers, sitatungas ( Tragelaphus spekei gratus ), and forest elephants ( Loxondonta africana cyclotis ), with avoidance of roads being stronger outside than inside the concession. Five monkey species showed little response to roads or hunting, whereas some rodents and pangolins increased in abundance outside the concession, possibly in response to greater forest disturbance. Our findings suggest that even moderate hunting pressure can markedly alter the structure of mammal communities in central Africa. Roads had the greatest impacts on large and small ungulates, with the magnitude of road avoidance increasing with local hunting pressure.     

Forests Too Deer: Edge Effects in Northern Wisconsin

Abstract: Browsing by white-tailed deer (Odocoileus virginianus) can profoundly affect the abundance and population structure of several woody and herbaceous plant species. Enclosure studies and population surveys reveal that past and current deer densities as low as 4 deer/km² may prevent regeneration of the once common woody species, Canada yew (Taxus canadensis), eastern hemlock (Tsuja canadensis), and white cedar Puja occidentalis), as well as several herbaceous species. Prior to European settlement, forests in northern Wisconsin contained relatively sparse deer populations (<4/km²), but extensive timber cutting in the late nineteenth century boosted deer populations. Continued habitat fragmentation resulting from scattered timber harvests and the creation of "wildlife openings" to improve deer forage maintain these high densities throughout much of the Northeast.
Because deer wander widely, the effects of high deer densities penetrate deeply into remaining stands of old and mature forest, greatly modifying their composition Thus, abundant early successional and "edge" habitat, and the high deer densities they engender, represent significant external threats to these plant communities. We hypothesize that establishing large (200–400 km²) continuous areas of maturing forest, especially in conjunction with increased hunting, could reduce local deer densities and so provide a simple and inexpensive method for retaining species sensitive to the deleterious effects of browsing.     

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.     

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.     

Long-Term Landscape Change and Bird Abundance in Amazonian Rainforest Fragments

Abstract:  The rainforests of the Amazon basin are being cut by humans at a rate >20,000 km²/year, leading to smaller and more isolated patches of forest, with remaining fragments often in the range of 1–100 ha. We analyzed samples of understory birds collected over 20 years from a standardized mist-netting program in 1– to 100-ha rainforest fragments in a dynamic Amazonian landscape near Manaus, Brazil. Across bird guilds, the condition of second growth immediately surrounding fragments was often as important as fragment size or local forest cover in explaining variation in abundance. Some fragments surrounded by 100 m of open pasture showed reductions in insectivorous bird abundance of over 95%, even in landscapes dominated by continuous forest and old second growth. These extreme reductions may be typical throughout Amazonia in small (≤10 ha), isolated fragments of rainforest. Abundance for some guilds returned to preisolation levels in 10- and 100-ha fragments connected to continuous forest by 20-year-old second growth. Our results show that the consequences of Amazonian forest loss cannot be accurately described without explicit consideration of vegetation dynamics in matrix habitat. Any dichotomous classification of the landscape into "forest" and "nonforest" misses essential information about the matrix.     

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.     

Overlooked biodiversity loss in tropical smallholder agriculture

Smallholder agriculture is the main driver of deforestation in the western Amazon, where terrestrial biodiversity reaches its global maximum. Understanding the biodiversity value of the resulting mosaics of cultivated and secondary forest is therefore crucial for conservation planning. However, Amazonian communities are organized across multiple forest types that support distinct species assemblages, and little is known about smallholder impacts across the range of forest types that are essential for sustaining biodiversity. We addressed this issue with a large-scale field inventory of birds (point counts) and trees (transects) in primary forest and smallholder agriculture in northern Peru across 3 forest types that are key for Amazonian biodiversity. For birds smallholder agriculture supported species richness comparable to primary forest within each forest type, but biotic homogenization across forest types resulted in substantial losses of biodiversity overall. These overall losses are invisible to studies that focus solely on upland (terra firma) forest. For trees biodiversity losses in upland forests dominated the signal across all habitats combined and homogenization across habitats did not exacerbate biodiversity loss. Proximity to forest strongly predicted the persistence of forest-associated bird and tree species in the smallholder mosaic, and because intact forest is ubiquitous in our study area, our results probably represent a best-case scenario for biodiversity in Amazonian agriculture. Land-use planning inside and outside protected areas should recognize that tropical smallholder agriculture has pervasive biodiversity impacts that are not apparent in typical studies that cover a single forest type. The full range of forest types must be surveyed to accurately assess biodiversity losses, and primary forests must be protected to prevent landscape-scale biodiversity loss.     

The Sustainability of Subsistence Hunting by Matsigenka Native Communities in Manu National Park, Peru

Abstract:  The presence of indigenous people in tropical parks has fueled a debate over whether people in parks are conservation allies or direct threats to biodiversity. A well-known example is the Matsigenka (or Machiguenga) population residing in Manu National Park in Peruvian Amazonia. Because the exploitation of wild meat (or bushmeat), especially large vertebrates, represents the most significant internal threat to biodiversity in Manu, we analyzed 1 year of participatory monitoring of game offtake in two Matsigenka native communities within Manu Park (102,397 consumer days and 2,089 prey items). We used the Robinson and Redford (1991) index to identify five prey species hunted at or above maximum sustainable yield within the ∼150-km² core hunting zones of the two communities: woolly monkey (Lagothrix lagotricha), spider monkey (Ateles chamek), white-lipped peccary (Tayassu pecari), Razor-billed Currasow (Mitu tuberosa), and Spix's Guan (Penelope jacquacu). There was little or no evidence that any of these five species has become depleted, other than locally, despite a near doubling of the human population since 1988. Hunter–prey profiles have not changed since 1988, and there has been little change in per capita consumption rates or mean prey weights. The current offtake by the Matsigenka appears to be sustainable, apparently due to source–sink dynamics. Source–sink dynamics imply that even with continued human population growth within a settlement, offtake for each hunted species will eventually reach an asymptote. Thus, stabilizing the Matsigenka population around existing settlements should be a primary policy goal for Manu Park.     

Meta‐Analysis of the Effects of Human Disturbance on Seed Dispersal by Animals

Animal‐mediated seed dispersal is important for sustaining biological diversity in forest ecosystems, particularly in the tropics. Forest fragmentation, hunting, and selective logging modify forests in myriad ways and their effects on animal‐mediated seed dispersal have been examined in many case studies. However, the overall effects of different types of human disturbance on animal‐mediated seed dispersal are still unknown. We identified 35 articles that provided 83 comparisons of animal‐mediated seed dispersal between disturbed and undisturbed forests; all comparisons except one were conducted in tropical or subtropical ecosystems. We assessed the effects of forest fragmentation, hunting, and selective logging on seed dispersal of fleshy‐fruited tree species. We carried out a meta‐analysis to test whether forest fragmentation, hunting, and selective logging affected 3 components of animal‐mediated seed dispersal: frugivore visitation rate, number of seeds removed, and distance of seed dispersal. Forest fragmentation, hunting, and selective logging did not affect visitation rate and were marginally associated with a reduction in seed‐dispersal distance. Hunting and selective logging, but not fragmentation, were associated with a large reduction in the number of seeds removed. Fewer seeds of large‐seeded than of small‐seeded tree species were removed in hunted or selectively logged forests. A plausible explanation for the consistently negative effects of hunting and selective logging on large‐seeded plant species is that large frugivores, as the predominant seed dispersers for large‐seeded plant species, are the first animals to be extirpated from hunted or logged forests. The reduction in forest area after fragmentation appeared to have weaker effects on frugivore communities and animal‐mediated seed dispersal than hunting and selective logging. The differential effects of hunting and selective logging on large‐ and small‐seeded tree species underpinned case studies that showed disrupted plant‐frugivore interactions could trigger a homogenization of seed traits in tree communities in hunted or logged tropical forests. Meta Análisis de los Efectos de la Perturbación Humana sobre la Dispersión de Semillas por Animales     

Synergistic Effects of Subsistence Hunting and Habitat Fragmentation on Amazonian Forest Vertebrates

Abstract: Subsistence game hunting has profound negative effects on the species diversity, standing biomass, and size structure of vertebrate assemblages in Amazonian forests that otherwise remain largely undisturbed. These effects are likely to be considerably aggravated by forest fragmentation because fragments are more accessible to hunters, allow no (or very low rates of  ) recolonization from nonharvested source populations, and may provide a lower-quality resource base for the frugivore-granivore vertebrate fauna. I examined the likelihood of midsized to large-bodied bird and mammal populations persisting in Amazonian forest fragments of variable sizes whenever they continue to be harvested by subsistence hunters in the aftermath of isolation. I used data from a comprehensive compilation of game-harvest studies throughout Neotropical forests to estimate the degree to which different species and populations have been overharvested and then calculated the range of minimum forest areas required to maintain a sustainable harvest. The size distribution of 5564 Amazonian forest fragments—estimated from Landsat images of six regions of southern and eastern Brazilian Amazonia—clearly shows that these are predominantly small and rarely exceed 10 ha, suggesting that persistent overhunting is likely to drive most midsized to large vertebrate populations to local extinction in fragmented forest landscapes. Although experimental studies on this negative synergism remain largely unavailable, the prospect that increasingly fragmented Neotropical forest regions can retain their full assemblages of avian and mammalian species is unlikely.     

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.     

The importance of agricultural lands for Himalayan birds in winter

The impacts of land‐use change on biodiversity in the Himalayas are poorly known, notwithstanding widespread deforestation and agricultural intensification in this highly biodiverse region. Although intact primary forests harbor many Himalayan birds during breeding, a large number of bird species use agricultural lands during winter. We assessed how Himalayan bird species richness, abundance, and composition during winter are affected by forest loss stemming from agriculture and grazing. Bird surveys along 12 elevational transects within primary forest, low‐intensity agriculture, mixed subsistence agriculture, and intensively grazed pastures in winter revealed that bird species richness and abundance were greatest in low‐intensity and mixed agriculture, intermediate in grazed pastures, and lowest in primary forest at both local and landscape scales; over twice as many species and individuals were recorded in low‐intensity agriculture than in primary forest. Bird communities in primary forests were distinct from those in all other land‐use classes, but only 4 species were unique to primary forests. Low‐, medium‐, and high‐intensity agriculture harbored 32 unique species. Of the species observed in primary forest, 80% had equal or greater abundance in low‐intensity agricultural lands, underscoring the value of these lands in retaining diverse community assemblages at high densities in winter. Among disturbed landscapes, bird species richness and abundance declined as land‐use intensity increased, especially in high‐intensity pastures. Our results suggest that agricultural landscapes are important for most Himalayan bird species in winter. But agricultural intensification—especially increased grazing—will likely result in biodiversity losses. Given that forest reserves alone may inadequately conserve Himalayan birds in winter, comprehensive conservation strategies in the region must go beyond protecting intact primary forests and ensure that low‐intensity agricultural lands are not extensively converted to high‐intensity pastures.     

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.     

Conservation Alliances with Indigenous Peoples of the Amazon

Abstract:  Ongoing alliances between indigenous peoples and conservation organizations in the Brazilian Amazon have helped achieve the official recognition of ∼1 million km² of indigenous lands. The future of Amazonian indigenous reserves is of strategic importance for the fate of biodiversity in the region. We examined the legislation governing resource use on indigenous lands and summarize the history of the Kayapo people's consolidation of their >100,000 km² territory. Like many Amazonian indigenous peoples, the Kayapo have halted the expansion of the agricultural frontier on their lands but allow selective logging and gold mining. Prospects for long-term conservation and sustainability in these lands depend on indigenous peoples' understandings of their resource base and on available economic alternatives. Although forest conservation is not guaranteed by either tenure security or indigenous knowledge, indigenous societies' relatively egalitarian common-property resource management regimes—along with adequate incentives and long-term partnerships with conservation organizations—can achieve this result. Successful initiatives include Conservation International's long-term project with the A'ukre Kayapo village and incipient large-scale territorial monitoring and control in the Kayapo territory, and the Instituto SocioAmbiental (ISA) 15-year partnership with the peoples of the Xingu Indigenous Park, with projects centered on territorial monitoring and control, education, community organization, and economic alternatives. The recent agreement on ecological restoration of the Xingu River headwaters between ranchers and private companies, indigenous peoples, and environmentalists, brokered by ISA, marks the emergence of an indigenous and conservation alliance of sufficient cohesiveness and legitimacy to negotiate effectively at a regional scale.     

Modeling the impacts of hunting on the population dynamics of red howler monkeys (Alouatta seniculus)

Overexploitation of wildlife populations occurs across the humid tropics and is a significant threat to the long-term survival of large-bodied primates. To investigate the impacts of hunting on primates and ways to mitigate them, we developed a spatially explicit, individual-based model for a landscape that included hunted and un-hunted areas. We used the large-bodied neotropical red howler monkey (Alouatta seniculus) as our case study species because its life history characteristics make it vulnerable to hunting. We modeled the influence of different rates of harvest and proportions of landscape dedicated to un-hunted reserves on population persistence, population size, social dynamics, and hunting yields of red howler monkeys. In most scenarios, the un-hunted populations maintained a constant density regardless of hunting pressure elsewhere, and allowed the overall population to persist. Therefore, the overall population was quite resilient to extinction; only in scenarios without any un-hunted areas did the population go extinct. However, the total and hunted populations did experience large declines over 100 years under moderate and high hunting pressure. In addition, when reserve area decreased, population losses and losses per unit area increased disproportionately. Furthermore, hunting disrupted the social structure of troops. The number of male turnovers and infanticides increased in hunted populations, while birth rates decreased and exacerbated population losses due to hunting. Finally, our results indicated that when more than 55% of the landscape was harvested at high (30%) rates, hunting yields, as measured by kilograms of biomass, were less than those obtained from moderate harvest rates. Additionally, hunting yields, expressed as the number of individuals hunted/year/km², increased in proximity to un-hunted areas, and suggested that dispersal from un-hunted areas may have contributed to hunting sustainability. These results indicate that un-hunted areas serve to enhance hunting yields, population size, and population persistence in hunted landscapes. Therefore, spatial regulation of hunting via a reserve system may be an effective management strategy for sustainable hunting, and we recommend it because it may also be more feasible to implement than harvest quotas or restrictions on season length.     

The Significance of Protected Areas for the Land Birds of Southern Finland

Protected areas (nature reserves) cover 0.7% of the land area in southern Finland (162,000 km²), which belongs to the south- and mid-boreal coniferous forest zones. The value of these areas in preserving land birds was studied on the basis of quantitative censuses. The numbers of bird pairs were estimated in nature reserves and in the whole of southern Finland, and the significance of the reserves was evaluated based on how large a proportion of the total population in southern Finland was found in them. Forest habitat generalists and species of coniferous forests occurred in protected areas as expected by the proportion of these areas in southern Finland. Specimens of these species groups comprise two thirds of all the country's land birds. Species of old-growth forests and open peatlands clearly preferred protected areas, whereas species of bushes, and lush and deciduous forest were scarcer in protected areas than elsewhere in southern Finland. The latter fertile habitats are poorly represented in protected areas, even though due to their disappearance or alteration elsewhere several species confined to these habitats have declined and are even threatened. Drainage of open peatlands and clearcutting of old-growth forests has caused a decrease in the area of these habitats in southern Finland during the past decades. Therefore, protected areas have a high significance in preserving bird species preferring these habitats. Protected areas are particularly important for decreased species of old-growth forests, such as the Three-toed Woodpecker Picoides tridactylus and the Siberian Jay Perisoreus infaustus , as about 10% of their total population in southern Finland was estimated to breed in protected areas. The future protection of both old-growth and deciduous forests is important if we are to preserve bio-diversity of land birds in Finland.