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Dispersal costs can be classified into energetic, time, risk and opportunity costs and may be levied directly or deferred during departure, transfer and settlement. They may equally be incurred during life stages before the actual dispersal event through investments in special morphologies. Because costs will eventually determine the performance of dispersing individuals and the evolution of dispersal, we here provide an extensive review on the different cost types that occur during dispersal in a wide array of organisms, ranging from micro-organisms to plants, invertebrates and vertebrates. In general, costs of transfer have been more widely documented in actively dispersing organisms, in contrast to a greater focus on costs during departure and settlement in plants and animals with a passive transfer phase. Costs related to the development of specific dispersal attributes appear to be much more prominent than previously accepted. Because costs induce trade-offs, they give rise to covariation between dispersal and other life-history traits at different scales of organismal organisation. The consequences of (i) the presence and magnitude of different costs during different phases of the dispersal process, and (ii) their internal organisation through covariation with other life-history traits, are synthesised with respect to potential consequences for species conservation and the need for development of a new generation of spatial simulation models.

Questions To what extent does species-specific variation in gut passage time (GPT), habitat use and mobility of three key avian frugivores synergistically affect the distribution of Xymalos monospora seeds within and among isolated forest fragments? Location Three fragments of a severely fragmented cloud forest, Taita Hills, southeast Kenya. Methods We experimentally determined GPTs of X. monospora seeds and recorded movements and habitat use by Turdus helleri, Andropadus milanjensis and Tauraco hartlaubi through radiotelemetry, and combined these data to generate species-specific seed dispersal patterns. Results Differences in mobility and habitat use among the three frugivores caused significant complementarity in seed dispersal, despite the fact that gut transit times were highly comparable. While the most sedentary and forest-dependent species mainly led to short-distance dispersal away from parent trees, two more mobile species dispersed seeds further away from the source trees, both within indigenous forest patches and towards exotic plantations and isolated fruiting trees in the landscape matrix. A. milanjensis inhabiting a very small forest fragment spent significantly more time in the landscape matrix than conspecifics residing in the two larger fragments. Conclusions By varying distances over which seeds are carried away from parent trees and the habitat types in which they are ultimately deposited, avian frugivores affect the spatial distribution of seeds and early plant recruits in a distinct and complementary manner. Because landscape properties are expected to lead to different constraints on avian mobility for habitat specialists and for generalists, ecosystem processes such as avian seed dispersal are shaped by complex interactions between disperser behaviour and the environment.

Frugivorous species heavily depend on patchy food resources and are believed to track these in space and time, thereby providing an important seed dispersal function that might be critical toward the regeneration of fruiting plants. However, isolation of suitable food patches due to habitat fragmentation or changes in landscape connectivity may hamper food tracking behaviour and adversely affect populations of both frugivores (through starvation) and food plants (through interruption of seed dispersal). We here test whether density fluctuations in four frugivorous Afrotropical bird species were larger and/or matched fluctuations in ripe fruit densities better in study plots embedded in large tracts of indigenous forest than in equally-sized plots embedded in cultivated lands. We compared these results with those of four non-frugivorous species (out-group) which were not expected to track fruit resources. Whereas densities of both frugivores and fruit crops strongly fluctuated in space and time, these fluctuations were not synchronised, nor did the level of synchrony differ in relation to matrix type. For some but not all bird species, lower densities and smaller temporal fluctuations in forest plots surrounded by cultivation may reflect decreased mobility. The observed fluctuations in bird densities most likely reflect exchange with the surrounding landscape matrix, suggesting that small pockets of fruiting trees in farmland may comprise critical food resources for frugivores inhabiting highly fragmented landscapes, apart from increasing connectivity for both bird and seed dispersal.

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

Habitat fragmentation and disturbance are known to impact animals and plants in different ways, depending on species' characteristics and the type and scale of habitat modification involved. In contrast, direct or indirect ramifications on mutualistic relationships between plants and animals are less clear, possibly because general patterns are confounded by the diffuse nature of many of these interactions. Here, we examine how fragment size and/or severe disturbance of a Kenyan mountain cloud forest affects the frugivore community and seed removal of a large-seeded, bird-dispersed tree of the forest interior, for three consecutive years. Forest deterioration reduced avian visitation and seed removal rates independent of fragment size, consistently so despite strong temporal variation in fruit production over the three-year study. In disturbed forest fragments, seed removal rates were on average 3.5 times lower than in more intact ones. Strong differences in both visitation and seed removal rates were largely attributable to shifts in frugivore assemblages, characterized by loss or reduced abundance of the most effective seed dispersers, most of which were forest specialists. Although some disturbed fragments benefited from visits of non-forest dependent seed dispersers, such 'resilience' was not predictable or reliable in time or space. We conclude that disruption of seed disperser-seed interactions in highly fragmented and disturbed tropical forests may be persistent in time when resiliency is inadequate, possibly posing long-term effects on tree communities.

Tropical forest fragmentation affects animal and plant populations in different ways. For plants, early stages (seed to seedling) are more sensitive to habitat alteration than adults, and can shape their future spatial patterns. Therefore, studying how seed germination and seedling growth and survival vary at different spatiotemporal scales enhances our understanding about plant recruitment in fragmented ecosystems. In this study we examine if, and to what extent, recruitment at early life-stages of Xymalos monospora (Monomiaceae), a bird-dispersed Afrotropical tree, differs between and within forest fragments that vary in size, surrounding matrix and microhabitats. Three years of field experiments (2004–2006) in south-east Kenya, revealed that patterns of seed germination and seedling survival and growth were largely inconsistent, both in space and time. Recruitment was not consistently higher in larger or less disturbed fragments. At smaller spatial scales within forest fragments, recruitment was subject to high between-year variation too, with decreased germination in gaps only in the dry year of 2004. However, performance of seeds and seedlings was consistently better away from than under conspecific fruiting trees. Our results imply that fragmented tree populations of X. monospora may become age-structured, or ultimately go extinct, if recruitment fails in subsequent years. This may especially affect populations in small, disturbed forest fragments, where seed dispersal and buffering against stochastic processes are generally reduced. Exotic plantations bordering indigenous forest fragments may provide suitable conditions for native tree recruitment; hence, forest expansion through enrichment planting should be considered in future conservation plans.

We recorded the association between the planthopper Tettigometra laetus Herrich-Schaffer, 1835 (Hemiptera: Fulgoro- morpha: Tettigometridae) and three ant species belonging to the subfamilies Myrmicinae and Formicinae in a coastal dune area of Flanders (Belgium). Lasius psammophilus Seifert, Tetramorium caespitum L. and Formica cunicularia Latreille were observed attending and palpating the dorsal glandular area of this planthopper, taking honeydew directly from its anus, herding them and car- rying them into their nests when disturbed. The planthopper was rarely found in the absence of ants and probably develops within ant nests, which may provide protection against predation and adverse weather conditions. The natural history of temperate ant- hemipteran relationships is discussed.

Fragmentation of continuous habitat blocks into small, isolated patches may Cause gradual replacement of habitat specialists by generalists and edge-dwelling species, thereby altering trophic interactions within communities. When fragmentation progressively increases the edge-to-interior ratio of forest remnants, forest birds are typically exposed to higher levels of nest predation, especially so near forest edges. However, the strength and direction of this 'edge effect on nest predation' may vary with predator communities, which in turn call be affected by habitat type, landscape structure and geographical location. We studied avian nest predation in two isolated forest fragments in the Taita Hills of southeast Kenya, where small mammals, rather than birds, dominate predator communities. The use of artificial ground nests allowed us to reduce variation in laying date and microhabitat use, which are known to affect predation on natural nest in the study area. Predation rates did not increase near the forest boundary, but instead, were higher in the forest interior of one of the fragments. Such inverse edge effects may be caused by forest-dependent small mammals or snakes that can remove bird eggs. Although inverse edge effects may be more common in East African tropical forests than previously thought, and should therefore be taken into account when drafting conservation plans, the underlying mechanisms remain unclear. Further study of the population dynamics of small mammals and snakes, and their relative impact on nest predation in fragmented habitats, is therefore required.