Demographic Limitations of the Ability of Habitat Restoration to Rescue Declining Populations

Abstract:  Habitat restoration is often recommended in conservation without first evaluating whether populations are in fact habitat limited and thus whether declining populations can be stabilized or recovered through habitat restoration. We used a spatially structured demographic model coupled with a dynamic neutral landscape model to evaluate whether habitat restoration could rescue populations of several generic migratory songbirds that differed in their sensitivity to habitat fragmentation (i.e., severity of edge effects on nesting success). Simulating a best-case scenario, landscapes were instantly restored to 100% habitat before, at, or after habitat loss exceeded the species' vulnerability threshold. The vulnerability threshold is a measure of extinction risk, in which the change in population growth rate ( δλ ) scaled to the rate of habitat loss ( δh ) falls below −1% ( δλ/δh ≤ −0.01). Habitat restoration was most effective for species with low-to-moderate edge sensitivities and in landscapes that had not previously experienced extensive fragmentation. To stabilize populations of species that were highly edge sensitive or any species in heavily fragmented landscapes, restoration needed to be initiated long before the vulnerability threshold was reached. In practice, habitat restoration is generally not initiated until a population is at risk of extinction, but our model results demonstrate that some populations cannot be recovered at this point through habitat restoration alone. At this stage, habitat loss and fragmentation have seriously eroded the species' demographic potential such that halting population declines is limited more by demographic factors than the amount of available habitat. Evidence that populations decline in response to habitat loss is thus not sufficient to conclude that habitat restoration will be sufficient to rescue declining populations.     

How Corridors Reduce Indigo Bunting Nest Success

Abstract:  Corridors are a popular strategy to conserve biodiversity and promote gene flow in fragmented landscapes. Corridor effectiveness has been bolstered by the fact that no empirical field studies have shown negative effects on populations or communities. I tested the hypothesis that corridors increase nest predation in connected habitat fragments relative to unconnected fragments. I evaluated this hypothesis in a large-scale experimental system of open-habitat fragments that varied in shape and connectivity. Corridors increased nest predation rates in connected fragments relative to unconnected fragments with lower edge:area ratios. Nest predation rates were similar between connected and unconnected fragments with higher edge:area ratios. These results suggest that the increase in predator activity is largely attributable to edge effects incurred through the addition of a corridor. This is the first field study to demonstrate that corridors can negatively impact animal populations occupying connected fragments.     

Effects of anthropogenic landscapes on population maintenance of waterbirds

Anthropogenic impacts have reduced natural areas but increased the area of anthropogenic landscapes. There is debate about whether anthropogenic landscapes (e.g., farmlands, orchards, and fish ponds) provide alternatives to natural habitat and under what circumstances. We considered whether anthropogenic landscapes can mitigate population declines for waterbirds. We collected data on population trends and biological traits of 1203 populations of 579 species across the planet. Using Bayesian generalized linear mixed models, we tested whether the ability of a species to use an anthropogenic landscape can predict population trends of waterbird globally and of species of conservation concern. Anthropogenic landscapes benefited population maintenance of common but not less-common species. Conversely, the use of anthropogenic landscapes was associated with population declines for threatened species. Our findings delineate some limitations to the ability of anthropogenic landscapes to mitigate population declines, suggesting that the maintenance of global waterbird populations depends on protecting remaining natural areas and improving the habitat quality in anthropogenic landscapes. Article impact statement: Protecting natural areas and improving the quality of anthropogenic landscapes as habitat are both needed to achieve effective conservation.     

Synergistic interactions between edge and area effects in a heavily fragmented landscape

Both area and edge effects have a strong influence on ecological processes in fragmented landscapes, but there is little understanding of how these two factors might interact to exacerbate local species declines. To test for synergistic interactions between area and edge effects, we sampled a diverse beetle community in a heavily fragmented landscape in New Zealand. More than 35,000 beetles of approximately 900 species were sampled over large gradients in habitat area (10(-2) 10(6) ha) and distance from patch edge (2(0)-2(10) m from the forest edge into both the forest and adjacent matrix). Using a new approach to partition variance following an ordination analysis, we found that a synergistic interaction between habitat area and distance to edge was a more important determinant of patterns in beetle community composition than direct edge or area effects alone. The strength of edge effects in beetle-species composition increased nonlinearly with increasing fragment area. One important consequence of the synergy is that the slopes of species area (SA) curves constructed from habitat islands depend sensitively on the distance from edge at which sampling is conducted. Surprisingly, we found negative SA curves for communities sampled at intermediate distances from habitat edges, caused by differential edge responses of matrix- vs. forest-specialist species in fragments of increasing area. Our data indicate that distance to habitat edge has a consistently greater impact on beetle community composition than habitat area and that variation in the strength of edge effects may underlie many patterns that are superficially related to habitat area.     

Habitat Split as a Cause of Local Population Declines of Amphibians with Aquatic Larvae

Abstract:  Most amphibian species have biphasic life histories and undergo an ontogenetic shift from aquatic to terrestrial habitats. In deforested landscapes, streams and forest fragments are frequently disjunct, jeopardizing the life cycle of forest-associated amphibians with aquatic larvae. We tested the impact of habitat split—defined as human-induced disconnection between habitats used by different life-history stages of a species—on four forest-associated amphibian species in a severely fragmented landscape of the Brazilian Atlantic Forest. We surveyed amphibians in forest fragments with and without streams (referred to as wet and dry fragments, respectively), including the adjacent grass-field matrix. Our comparison of capture rates in dry fragments and nearby streams in the matrix allowed us to evaluate the number of individuals that engaged in high-risk migrations through nonforested habitats. Adult amphibians moved from dry fragments to matrix streams at the beginning of the rainy season, reproduced, and returned at the end of the breeding period. Juveniles of the year moved to dry fragments along with adults. These risky reproductive migrations through nonforested habitats that expose individuals to dehydration, predation, and other hazards may cause population declines in dry fragments. Indeed, capture rates were significantly lower in dry fragments compared with wet fragments. Declining amphibians would strongly benefit from investments in the conservation and restoration of riparian vegetation and corridors linking breeding and nonbreeding areas.     

Changes in Wildlife Communities Near Edges

Abstract: Wildlife managers and land managers have traditionally considered edges as beneficial to wildlife because species diversity generally increases near habitat edges. Explanations for this edge effect include greater vegetative complexity at edges or the simultaneous availability of more than one landscape element. However, edges can have negative consequences for wildlife by modifying distribution and dispersal and by increasing incidence of nest predation and parasitism Edges also may be detrimental to species requiring large undisturbed areas because increases in edge generally result in concommitant reductions in size and possible isolation of patches and corridots. Thus, both wildlife and land managers should be cautious when describing the benefits of edges to wildlife: particularly when dealing with species that require forest interiors.
Changes in wildlife communities associated with habitat edges are not easily assessed because defining edge species and measuring edge dimensions can be difficult in field studies Also, there is no general consensus as to how edge effect is best measured. Well-designed long-term studies of edges in various landscapes are needed (1) to better understand the positive and negative impacts of edges on wildlife communities, guilds, or key species, and (2) to effectively quantify edge effect and thereby develop management recommendations to improve the quality of edges for wildlife. Additional studies of edge effect are timely because greater amounts of edge will continue to be created in future landscapes due to extensive agriculture and other land-use-practices, and because developing knowledge in conservation biology and landscape ecology will facilitate multidisciplinary approaches to edge and landscape management for the benefit of wildlife.     

The Early Development of Forest Fragmentation Effects on Birds

The early development of forest fragmentation effects on forest organisms is poorly understood partly because most fragmentation studies have been done in agricultural or suburban landscapes, long after the onset of fragmentation. We develop a temporal model of forest fragmentation effects on densities of forest-breeding birds and provide data from an active industrial forest landscape to test the model. The model and our empirical data indicate that densities of several forest-dwelling bird species can increase within a forest stand soon after the onset of fragmentation as a result of displaced individuals packing into remaining habitat. Along with higher densities in the newly formed fragments, pairing success in one species, the Ovenbird ( Seiurus aurocapillus ), was lower in fragments than nonfragments, possibly due to behavioral dysfunction resulting from high densities. Thus, density was inversely related to productivity. The duration and extent of increased densities following onset of fragmentation depends on many factors, including the sensitivity of a species to edge and area effects, the duration and rate of habitat loss and fragmentation, and the proximity of a forest stand to the disturbance. Incipient forest fragmentation may affect populations differently from later stages of fragmentation when the geometry of the landscape has reached a more stable configuration. Our model and data indicate, for reasons unrelated to traditional edge effects, that large tracts of forest can be important because they are relatively free from the variety of plant and animal population dynamics that might take place near new edges, including the encroachment of individuals displaced by habitat loss.     

Local and Landscape Correlates of Primate Distribution and Persistence in the Remnant Lowland Rainforests of the Upper Brahmaputra Valley,Northeastern India

Habitat fragmentation affects species distribution and abundance, and drives extinctions. Escalated tropical deforestation and fragmentation have confined many species populations to habitat remnants. How worthwhile is it to invest scarce resources in conserving habitat remnants within densely settled production landscapes? Are these fragments fated to lose species anyway? If not, do other ecological, anthropogenic, and species‐related factors mitigate the effect of fragmentation and offer conservation opportunities? We evaluated, using generalized linear models in an information‐theoretic framework, the effect of local‐ and landscape‐scale factors on the richness, abundance, distribution, and local extinction of 6 primate species in 42 lowland tropical rainforest fragments of the Upper Brahmaputra Valley, northeastern India. On average, the forest fragments lost at least one species in the last 30 years but retained half their original species complement. Species richness declined as proportion of habitat lost increased but was not significantly affected by fragment size and isolation. The occurrence of western hoolock gibbon (Hoolock hoolock) and capped langur (Trachypithecus pileatus) in fragments was inversely related to their isolation and loss of habitat, respectively. Fragment area determined stump‐tailed (Macaca arctoides) and northern pig‐tailed macaque occurrence (Macaca leonina). Assamese macaque (Macaca assamensis) distribution was affected negatively by illegal tree felling, and rhesus macaque (Macaca mulatta) abundance increased as habitat heterogeneity increased. Primate extinction in a fragment was primarily governed by the extent of divergence in its food tree species richness from that in contiguous forests. We suggest the conservation value of these fragments is high because collectively they retained the entire original species pool and individually retained half of it, even a century after fragmentation. Given the extensive habitat and species loss, however, these fragments urgently require protection and active ecological restoration to sustain this rich primate assemblage. Correlaciones Locales y de Paisaje de la Distribución y Persistencia de Primates en los Bosques Lluviosos Remanentes en el Valle del Alto Brahmaputra, Noreste de India     

Edge Effects in the Great Tit: Analyses of Long-term Data with GIS Techniques

Abstract:  In contemporary fragmented landscapes, edges are commonplace, and understanding the effects of edge environments is thus essential for the conservation of forest communities. The reproductive output of forest passerines is often reduced close to forest edges. Possible explanations include overcrowding by conspecifics, elevated rates of predation, and the occurrence of lower-quality habitat and/or individuals at forest edges. We attempted to separate these processes by examining edge effects in the absence of nest predation and by effectively controlling for differences in breeding density and the quality of habitats and individuals. We used an edge distance index (EDI), which accounts for the number and distribution of edges in close proximity to a breeding location, to help explain variation in breeding density, nesting success, and reproductive traits of 8308 pairs of Great Tits ( Parus major ) breeding between 1965 and 2005, in Wytham, near Oxford, United Kingdom. Results from linear mixed modeling confirmed higher breeding density and a higher proportion of immigrant individuals at forest edges. Nevertheless, independently of these effects, we also found that birds laying later, with smaller clutches but larger eggs, were typical of edge environments. The number of offspring recruited to the breeding offspring per breeding attempt was also reduced at edges, both directly and mediated through changes in clutch size and laying date. Edge effects on life histories were detectable within individual females and up to 500 m from the woodland edge. Woodland edges are increasingly common in contemporary fragmented landscapes. Therefore these results, which suggest a pervasive effect of edges on reproduction, are of considerable importance to the management and conservation of forest communities.     

Countryside Biogeography of Moths in a Fragmented Landscape: Biodiversity in Native and Agricultural Habitats

Abstract: Studies of fragmented landscapes, especially in the tropics, have traditionally focused on the native fragments themselves, ignoring species distributions in surrounding agricultural or other human-dominated areas. We sampled moth species richness within a 227-ha forest fragment and in four surrounding agricultural habitats (coffee, shade coffee, pasture, and mixed farms) in southern Costa Rica. We found no significant difference in moth species richness or abundance among agricultural habitats, but agricultural sites within 1 km of the forest fragment had significantly higher richness and abundance than sites farther than 3.5 km from the fragment. In addition, species composition differed significantly between distance classes ( but not among agricultural habitats), with near sites more similar to forest than far sites. These results suggest that (1) different agricultural production regimes in this region may offer similar habitat elements and thus may not differ substantially in their capacities to support native moth populations and (2) that the majority of moths may utilize both native and agricultural habitats and move frequently between them, forming "halos" of relatively high species richness and abundance around forest fragments. Correlations between species richness and the amount of nearby forest cover, measured over circles of various radii around the sites, suggest that halos extend approximately 1.0–1.4 km from the forest edge. The extent of these halos likely differs among taxa and may influence their ability to survive in fragmented landscapes.     

Demographic Stability Metrics for Conservation Prioritization of Isolated Populations

Abstract:  Systems of geographically isolated habitat patches house species that occur naturally as small, disjunct populations. Many of these species are of conservation concern, particularly under the interacting influences of isolation and rapid global change. One potential conservation strategy is to prioritize the populations most likely to persist through change and act as sources for future recolonization of less stable localities. We propose an approach to classify long-term population stability (and, presumably, future persistence potential) with composite demographic metrics derived from standard population-genetic data. Stability metrics can be related to simple habitat measures for a straightforward method of classifying localities to inform conservation management. We tested these ideas in a system of isolated desert headwater streams with mitochondrial sequence data from 16 populations of a flightless aquatic insect. Populations exhibited a wide range of stability scores, which were significantly predicted by dry-season aquatic habitat size. This preliminary test suggests strong potential for our proposed method of classifying isolated populations according to persistence potential. The approach is complementary to existing methods for prioritizing local habitats according to diversity patterns and should be tested further in other systems and with additional loci to inform composite demographic stability scores.     

Distribution of Carabid Beetles (Coleoptera, Carabidae) across a Boreal Forest–Clearcut Ecotone

Abstract: We studied the occurrence of carabid beetles (Coleoptera, Carabidae) in boreal forest fragments, their edges, and adjacent clearcuts in central Finland. Beetles were collected with pitfall traps along transects extending 60 m from the edge into clearcuts and 60 m into forest interior. Our main findings were that (1) species richness was significantly higher in the clearcut than in the forest fragments, (2) clearcuts hosted many open-habitat species, which increased overall species richness in these sites, (3) carabid assemblages in the edges were more similar to forest assemblages than to those found in the clearcuts, (4) no edge specialists were found, and (5) open-habitat species did not penetrate into the forest fragments from the clearcut. Because forest specialists occurred all the way to the edge on the forest side, it seems that edge effects per se do not adversely affect these species, at least in the short term. In the long term, however, habitat conditions in the edges may deteriorate for interior species because of trees falling over in strong winds, thereby reducing the size of the fragments and widening the edge zone.     

Nesting Success of Neotropical Migrant Songbirds in a Highly Fragmented Landscape

Abstract: Small forests in agricultural landscapes are generally thought to be population sinks where the nesting success of Neotropical migrant songbirds is too low to sustain populations. In 1996 and 1997, we assessed the nesting success and productivity of Wood Thrushes (  Hylocichla mustelina ) and Rose-breasted Grosbeaks (  Pheucticus ludovicianus ) in the Region of Waterloo, a highly fragmented agricultural landscape with 14% forest cover in southwestern Ontario. We located 154 Wood Thrush nests and 63 Rose-breasted Grosbeak nests in 26 small forest fragments (3–14 ha) and 15 large fragments (26–140 ha). We used the Mayfield method to determine nesting success. Across all sites nesting success was 51% for Wood Thrushes and 46% for Rose-breasted Grosbeaks. Nesting success and productivity rates were used in a simple population growth model that suggested that the Wood Thrushes but not the Rose-breasted Grosbeaks were a self-sustaining population. Forest size and distance of nests to the nearest forest edge (measured in five distance classes ranging from 0 to over 100 m) did not significantly affect the nesting success or productivity of either species. Brown-headed Cowbirds (   Molothrus ater ) parasitized 47% of all Wood Thrush nests and 10% of all Rose-breasted Grosbeak nests; parasitism significantly reduced Wood Thrush productivity but not that of Rose-breasted Grosbeaks. Our findings challenge the prevailing notion that small woodlots in a farmed landscape are invariably habitat sinks and offer additional incentive for habitat protection in settled landscapes where small forest fragments are often all that remain for conservation purposes.     

Fragmentation of Phragmites Habitats, Minimum Viable Population Size, Habitat Suitability, and Local Extinction of Moths, Midges, Flies, Aphids, and Birds

Results from populations of insects and birds inhabiting Phragmites habitats were used to analyze effects of fragmentation. Flush-crash cycles of the stem-boring moth Archanara geminipuncta (Lepidoptera, Noctuidae) showed regionally concurrent, local extinctions despite an originally enormous population size (more than 180,000 adults), emphasizing the importance of metapopulation dynamics. Further, A. geminipuncta could be considered a keystone species, since shoot damage facilitated more than twenty species of herbivores, saprovores (of the caterpillars' feces), and their parasitoids. The gall midge Lasioptera arundinis could survive only in side shoots induced by shoot damage of A. geminipuncta .
Small Phragmites stands had thinner shoots (due to a water or nutrient deficiency) and shoots with more leaves (due to a better light supply) than large stands, thereby influencing species-specific demands for habitat suitability and nutritiousness of reed tissue. In other words significance of habitat fragmentation could not be assessed by area alone. For example, two chloropid flies depending on thin, yellowish shoots survived only in small habitats or in the unmown edges of large habitats.
Local persistence of Phragmites herbivores depended on much larger population sizes than could be expected from a population size sufficient to maintain genetic variation. At least 11,000 adults of the gall midge Giraudiella inclusa (or more than 84,000 galls) were necessary to avoid local extinction.
With regard to conservation management of reed habitats, nature reserves should consist of old and unmown reeds, have fewer disturbed (particularly, fewer mown) habitat edges, measure more than two hectares (priority should go to the largest remaining fragments), and be surrounded by nearby reed habitats providing reservoir populations and diverse shoot types.     

Are shrubland birds edge specialists?

In studies of forest fragmentation, birds of scrubby, early-successional habitats are considered edge specialists. Because these birds are assumed to thrive in fragmented, edge-dominated areas, their landscape ecology has received little attention from ecologists. With populations of shrubland birds declining throughout the eastern United States, the question of whether or not these birds really prefer edge habitats has important conservation implications. We used a meta-analysis to test how edges affect the abundance of shrubland birds in early-successional habitats. We analyzed data for 17 species from seven studies that compared the abundances of birds in the interiors and edges of regenerating clearcuts surrounded by mature forest. The meta-analysis clearly showed that shrubland birds avoid edges. All 17 species tested had higher abundances in patch centers than along edges, and edge effects were significant for 8 of 17 species. The key implication of this result is that small or irregular patches, dominated by edge, are unlikely to provide suitable habitat for shrubland birds. Thus, management for these declining species should involve providing large patches and minimizing edges. These findings demonstrate the importance of testing widely accepted ecological classifications and the need to view landscape ecology from the perspective of non-forest wildlife.     

Edge Effects and Isolation: Red-Backed Voles Revisited

Abstract:  We examined demographic responses of California red-backed voles ( Clethrionomys californicus ) to forest fragmentation in southwestern Oregon at sites where this species has previously shown negative responses to fragmentation. Voles were captured in live traps and released. Voles were rarely caught in clearcuts surrounding 11 forest fragments, but relative vole density did not decrease from the forest-fragment interiors to edges. The first result agrees with previous findings at these sites 6 years earlier, but the latter result does not. There was no evidence that vole response to edge changes with fragment age. Two years of intensive mark-recapture efforts at two forest-fragment sites and two unfragmented (control) sites did not show negative effects of fragmentation on vole survival, an important demographic rate. Vole capture probabilities varied greatly across space and time on these four sites, which may explain the differences in vole responses to edge seen between this and the previous study. These results suggest that reliable appraisal of edge effects may be difficult for many species on small fragments because the data necessary to apply population estimators require great efforts to obtain and the use of indices leads to a confounding of detection probabilities with demographic change.     

The Botanist Effect Revisited: Plant Species Richness, County Area, and Human Population Size in the United States

Abstract:  The "botanist effect" is thought to be the reason for higher plant species richness in areas where botanists are disproportionately present as an artefactual consequence of a more thorough sampling. We examined whether this was the case for U.S. counties. We collated the number of species of vascular plants, human population size, and the area of U.S. counties. Controlling for spatial autocorrelation and county area, plant species richness increased with human population size and density in counties with and without universities and/or botanical gardens, with no significant differences in the relation between the two subsets. This is consistent with previous findings and further evidence of a broad-scale positive correlation between species richness and human population presence, which has important consequences for the experience of nature by inhabitants of densely populated regions. Combined with the many reports of a negative correlation between the two variables at a local scale, the positive relation between plant species richness in U.S. counties and human population presence stresses the need for the conservation of seminatural areas in urbanized ecosystems and for the containment of urban and suburban sprawl.     

Nesting Success of a Neotropical Migrant in a Multiple-Use, Forested Landscape

Abstract: We studied the nesting success of an individually marked population of Kentucky Warblers ( Oporornis formosus ), a species that nests in disturbed and undisturbed forests, in a heterogeneous, managed forest site in the Shawnee National Forest in southern Illinois from 1992 to 1995. We examined the effects of forest stand type (clearcuts of various ages, tree plantations, and older forest) and distance from habitat edges on rates of nest predation and brood parasitism by Brown-headed Cowbirds (   Molothrus ater ). Brood parasitism levels gradually decreased from 60% to 3% ( nests) over a distance of 2 km from an agricultural edge proximal to a known cowbird foraging site (a pig feedlot), but they did not vary with distance from any other kinds of edges or with forest stand type. Rates of nest predation ( nests) did not vary with distance from any edges, but they were significantly lower in older forest than within even-aged clearcuts, a tree plantation, and in successional vegetation adjacent to a residential facility. These results suggest that, even in fragmented landscapes with high overall levels of parasitism and nest predation, management practices within and immediately adjacent to forest tracts can affect the nesting success of some species, but not necessarily as a simple function of distance from edge. For the Kentucky Warbler, our results suggest that a management strategy that avoids even-age silviculture and leaves core stands of older forest far from cowbird feeding areas can increase nesting success to levels similar to those measured in more forested landscapes.     

Fencerows, Edges, and Implications of Changing Connectivity Illustrated by Two Contiguous Ohio Landscapes

We evaluated the amounts and implications of changes in habitat connectivity on rural landscapes by modeling the colonization success and subsequent habitat colonization of a model edge organism within real landscapes. We first inventoried the changes in the fencerow and forest-edge network of two contiguous Ohio (U.S.A.) landscapes, an agriculturally dominated till plain and a more diverse and dynamic moraine landscape, from 1940 to 1988. On the moraine the number of fencerows changed little from 1940 to 1971. The number increased by 86 during 1971–1988 as marginal farms were subdivided. The total length of fencerows on the moraine increased 2.5-fold over 48 years. On the till plain the number and total length of fencerows remained relatively constant through the study period. The sum of fencerows and forest edges was used as a measure of total ecotonal edge. On the moraine total edge increased through the study period, whereas on the till plain it decreased. We selected two levels of landscape connectivity, low and high, to model animal habitat colonization success. As connectivity increased the earliest successful colonists preempted an increasingly large proportion of the total suitable habitat, and the probability of successful colonization by later-arriving individuals decreased. The changes in connectivity that resulted from changes in both the fencerow network and the proportion of forested land have resulted in contiguous landscapes that present very different colonization potentials to organisms with long-distance dispersal capability. Given the current uncertainty of the effects of corridors on species-preservation efforts, we suggest that further modeling of this type prior to field testing will add useful insights, especially if conducted using specific species and landscape types.     

Conservation value of small reserves

The importance of large reserves has been long maintained in the scientific literature, often leading to dismissal of the conservation potential of small reserves. However, over half the global protected-area inventory is composed of protected areas that are <100 ha, and the median size of added protected area is decreasing. Studies of the conservation value of small reserves and fragments of natural area are relatively uncommon in the literature. We reviewed SCOPUS and WOK for studies on small reserve and fragment contributions to biodiversity conservation and ecosystem services, and fifty-eight taxon-specific studies were included in the review. Small reserves harbored substantial portions (upward of 50%) of regional species diversity for many taxa (birds, plants, amphibians, and small mammals) and even some endemic, specialist bird species. Unfortunately, small reserves and fragments almost always harbored more generalist and exotic species than large reserves. Community composition depended on habitat quality, surrounding land use (agricultural vs. urban), and reserve and fragment size, which presents opportunities for management and improvement. Small reserves also provided ecosystem services, such as pollination and biological pest control, and cultural services, such as recreation and improved human health. Limitations associated with small reserves, such as extinction debt and support of area-sensitive species, necessitate a complement of larger reserves. However, we argue that small reserves can make viable and significant contributions to conservation goals directly as habitat and indirectly by increasing landscape connectivity and quality to the benefit of large reserves. To effectively conserve biodiversity for future generations in landscapes fragmented by human development, small reserves and fragments must be included in conservation planning.