Pollen dispersal of tropical trees (Dinizia excelsa: Fabaceae) by native insects and African honeybees in pristine and fragmented Amazonian rainforest

Tropical rainforest trees typically occur in low population densities and rely on animals for cross-pollination. It is of conservation interest therefore to understand how rainforest fragmentation may alter the pollination and breeding structure of remnant trees. Previous studies of the Amazonian tree Dinizia excelsa (Fabaceae) found African honeybees (Apis mellifera scutellata) as the predominant pollinators of trees in highly disturbed habitats, transporting pollen up to 3.2 km between pasture trees. Here, using microsatellite genotypes of seed arrays, we compare outcrossing rates and pollen dispersal distances of (i) remnant D. excelsa in three large ranches, and (ii) a population in undisturbed forest in which African honeybees were absent. Self-fertilization was more frequent in the disturbed habitats (14%, n = 277 seeds from 12 mothers) than in undisturbed forest (10%, n = 295 seeds from 13 mothers). Pollen dispersal was extensive in all three ranches compared to undisturbed forest, however. Using a twogener analysis, we estimated a mean pollen dispersal distance of 1509 m in Colosso ranch, assuming an exponential dispersal function, and 212 m in undisturbed forest. The low effective density of D. excelsa in undisturbed forest (approximately 0.1 trees/ha) indicates that large areas of rainforest must be preserved to maintain minimum viable populations. Our results also suggest, however, that in highly disturbed habitats Apis mellifera may expand genetic neighbourhood areas, thereby linking fragmented and continuous forest populations.     

Photosynthetic Responses to Light for Rainforest Seedlings Planted in Abandoned Pasture, Costa Rica

Efforts to reforest tropical pasture with native tree species have increased in recent years, yet little is known about the physiology of most tropical trees. The goal of this study was to assess the effect of habitat on photosynthetic responses to light for seedlings of four native rainforest species (Calophyllum brasiliense, Ocotea glaucosericea, Ocotea whitei, and Sideroxylon portoricense) planted to facilitate tropical rainforest recovery in southern Costa Rica. Seedlings were planted in primary forest, in open abandoned pasture, and in the shade of remnant trees within the pasture. Growth, morphology, photosynthetic gas exchange responses to light, and chlorophyll fluorescence (an indication of the integrity of photosynthetic processes) were measured in the three habitats. Height and leaf area were generally greater for seedlings in tree shade compared to those in the forest and open pasture. Photosynthetic rates were higher for plants in open pasture and tree shade compared to those in the forest for two of the four species. Chlorophyll fluorescence results indicated flexibility in the photosynthetic processing of light energy that may help plants tolerate the bright light of the pasture. This study demonstrates that, for certain species, seedlings under remnant pasture trees do not exhibit the level of photosynthetic stress experienced in open abandoned pasture. Seedling responses to light, in combination with other factors such as increased nutrient input through litterfall, help explain the enhanced growth of seedlings under remnant pasture trees. Planting seedlings under remnant trees may increase the success of future efforts to restore tropical forest in abandoned agricultural land.     

Pollen dispersal and genetic structure of the tropical tree Dipteryx panamensis in a fragmented Costa Rican landscape

In the face of widespread deforestation, the conservation of rainforest trees relies increasingly on their ability to maintain reproductive processes in fragmented landscapes. Here, we analysed nine microsatellite loci for 218 adults and 325 progeny of the tree Dipteryx panamensis in Costa Rica. Pollen dispersal distances, genetic diversity, genetic structure and spatial autocorrelation were determined for populations in four habitats: continuous forest, forest fragments, pastures adjacent to fragments and isolated pastures. We predicted longer but less frequent pollen movements among increasingly isolated trees. This pattern would lead to lower outcrossing rates for pasture trees, as well as lower genetic diversity and increased structure and spatial autocorrelation among their progeny. Results generally followed these expectations, with the shortest pollen dispersal among continuous forest trees (240 m), moderate distances for fragment (343 m) and adjacent pasture (317 m) populations, and distances of up to 2.3 km in isolated pastures (mean: 557 m). Variance around pollen dispersal estimates also increased with fragmentation, suggesting altered pollination conditions. Outcrossing rates were lower for pasture trees and we found greater spatial autocorrelation and genetic structure among their progeny, as well as a trend towards lower heterozygosity. Paternal reproductive dominance, the pollen contributions from individual fathers, did not vary among habitats, but we did document asymmetric pollen flow between pasture and adjacent fragment populations. We conclude that long-distance pollen dispersal helps maintain gene flow for D. panamensis in this fragmented landscape, but pasture and isolated pasture populations are still at risk of long-term genetic erosion.     

Landscape genetics of a tropical rescue pollinator

Pollination services are increasingly threatened by the loss and modification of natural habitats, posing a risk to the maintenance of both native plant biodiversity and agricultural production. In order to safeguard pollination services, it is essential to examine the impacts of habitat degradation on the population dynamics of key pollinators and identify potential “rescue pollinators” capable of persisting in these human-altered landscapes. Using a landscape genetic approach, we assessed the impact of landscape structure on genetic differentiation in the widely-distributed tropical stingless bee Trigona spinipes (Apidae: Meliponini) across agricultural landscape mosaics composed of coffee plantations and Atlantic forest fragments in southeastern Brazil. We genotyped 115 bees at 16 specific and highly polymorphic microsatellite loci, developed using next-generation sequencing. Our results reveal that T. spinipes is capable of dispersing across remarkably long distances, as we did not find genetic differentiation across a 200 km range, nor fine-scale spatial genetic structure. Furthermore, gene flow was not affected by forest cover, land cover, or elevation, indicating that reproductive individuals are able to disperse well through agricultural landscapes and across altitudinal gradients. We also found evidence of a recent population expansion, suggesting that this opportunistic stingless bee is capable of colonizing degraded habitats. Our results thus suggest that T. spinipes can persist in heavily-altered landscapes and can be regarded as a rescue pollinator, potentially compensating for the decline of other native pollinators in degraded tropical landscapes.     

Effects of habitat disruption on the activity of nectarivorous bats (Chiroptera: Phyllostomidae) in a dry tropical forest: implications for the reproductive success of the neotropical tree Ceiba grandiflora

In the tropical dry forest of the central Pacific coast of Mexico the pollination and reproductive success of the bombacaceous tree Ceiba grandiflora was negatively affected by habitat disruption. Two of the three bat species that function as effective pollinators for this species ( Glossophaga soricina and Musonycteris harrisoni) visited flowers found in trees in disturbed habitats significantly less than trees found in undisturbed habitats. A similar pattern was observed for the effective bat pollinator, Leptonycteris curasoae; however the difference was not significant. The three nectarivorous bats that functioned as effective pollinators of C. grandiflora also visited flowers to exclusively feed on pollen by biting or pulling off an anther (see Fig. S1 of Electronic Supplementary Material). The number of pollen grains deposited on stigmas from flowers in undisturbed areas was significantly greater than from flowers in disturbed habitats. The greater visitation rate and the greater number of pollen grains deposited on flowers from trees in undisturbed forest resulted in a significantly greater fruit set for trees in these areas. Our study demonstrates the negative effect that habitat disruption has on bat pollinators in tropical dry forest ecosystems and documents the negative consequences for the plants they pollinate.     

Effects of habitat transitions on rainforest bird communities across an anthropogenic landscape mosaic

We compared bird community responses to the habitat transitions of rainforest‐to‐pasture conversion, consequent habitat fragmentation, and post‐agricultural regeneration, across a landscape mosaic of about 600 km² in the eastern Australian subtropics. Birds were surveyed in seven habitats: continuous mature rainforest; two size classes of mature rainforest fragment (4–21 ha and 1–3 ha); regrowth forest patches dominated by a non‐native tree (2–20 ha, 30–50 years old); two types of isolated mature trees in pasture; and treeless pasture, with six sites per habitat. We compared the avifauna among habitats and among sites, at the levels of species, functional guilds, and community‐wide. Community‐wide species richness and abundance of birds in pasture sites were about one‐fifth and one‐third, respectively, of their values in mature rainforest (irrespective of patch size). Many measured attributes changed progressively across a gradient of increased habitat simplification. Rainforest specialists became less common and less diverse with decreased habitat patch size and vegetation maturity. However, even rainforest fragments of 1–3 ha supported about half of these species. Forest generalist species were largely insensitive to patch size and successional stage. Few species reached their greatest abundance in either small rainforest fragments or regrowth. All pastures were dominated by bird species whose typical native habitats were grassland, wetland, and open eucalypt forest, while pasture trees modestly enhanced local bird communities. Overall, even small scattered patches of mature and regrowth forest contributed substantial bird diversity to local landscapes. Therefore, maximizing the aggregate rainforest area is a useful regional conservation strategy.     

Forests are critically important to global pollinator diversity and enhance pollination in adjacent crops

Although the importance of natural habitats to pollinator diversity is widely recognized, the value of forests to pollinating insects has been largely overlooked in many parts of the world. In this review, we (i) establish the importance of forests to global pollinator diversity, (ii) explore the relationship between forest cover and pollinator diversity in mixed-use landscapes, and (iii) highlight the contributions of forest-associated pollinators to pollination in adjacent crops. The literature shows unambiguously that native forests support a large number of forest-dependent species and are thus critically important to global pollinator diversity. Many pollinator taxa require or benefit greatly from resources that are restricted to forests, such as floral resources provided by forest plants (including wind-pollinated trees), dead wood for nesting, tree resins, and various non-floral sugar sources (e.g. honeydew). Although landscape-scale studies generally support the conclusion that forests enhance pollinator diversity, findings are often complicated by spatial scale, focal taxa, landscape context, temporal context, forest type, disturbance history, and external stressors. While some forest loss can be beneficial to pollinators by enhancing habitat complementarity, too much can result in the near-elimination of forest-associated species. There is strong evidence from studies of multiple crop types that forest cover can substantially increase yields in adjacent habitats, at least within the foraging ranges of the pollinators involved. The literature also suggests that forests may have enhanced importance to pollinators in the future given their role in mitigating the negative effects of pesticides and climate change. Many questions remain about the amount and configuration of forest cover required to promote the diversity of forest-associated pollinators and their services within forests and in neighbouring habitats. However, it is clear from the current body of knowledge that any effort to preserve native woody habitats, including the protection of individual trees, will benefit pollinating insects and help maintain the critical services they provide.     

The Effect of Spatial Structure of Pasture Tree Cover on Avian Frugivores in Eastern Amazonia

The movement of frugivores between remnant forests and successional areas is vital for tropical forest tree species to colonize successional habitats. The response of these species to the spatial structure of pasture tree cover is largely unknown. We studied avian frugivores that were found in primary forest edges and large pastures in eastern Amazonia, Brazil. We determined how the small‐scale spatial structure of pasture trees at forest edges affects five response variables: bird presence, visitation rate, duration of visit, species richness, and an index accounting for species’ level of frugivory and abundance in forests. We used hierarchical linear models to estimate the effect of four predictor variables on response variables: (1) clustering of pasture trees; (2) percent canopy cover of pasture trees; (3) distance of pasture tree to forest edge; and (4) tree crown area. The study species, many of which are widely distributed in the Neotropics, were generally insensitive to percent cover and clustering of trees. Frugivore visitation to individual trees remained constant as cover increased. Visitation was positively correlated with focal tree distance to forest edge and crown area. The positive relationship between distance and visitation rates may be due to the increased abundance of some resource further from forests. If pastures were abandoned the distance from forest edges would not likely limit frugivore visitation and seed deposition under large pasture trees in our study (i.e., up to 200 m distant).     

Rain forest provides pollinating beetles for atemoya crops

Small beetles, usually species of Nitidulidae, are the natural pollinators of atemoya (Annona squamosa L. x A. cherimola Mill. hybrids; custard apple) flowers but commercial atemoya growers often need to carry out labor-intensive hand pollination to produce enough high-quality fruit. Because Australian rain forest has plant species in the same family as atemoya (Annonaceae) and because many rain forest plants are beetle pollinated, we set out to discover whether tropical rain forest in far north Queensland harbors beetles that could provide this ecosystem service for atemoya crops. Orchards were chosen along a gradient of increasing distance from tropical rain forest (0.1-24 km). We sampled 100 flowers from each of nine atemoya orchards and determined the identity and abundance of insects within each flower. To assess the amount of pollination due to insects, we bagged six flowers per tree and left another six flowers per tree accessible to insects on 10 trees at an orchard near rain forest. Results indicated that atemoya orchards < or = 0.5 km from rain forest were predominantly visited by five previously unrecognized native beetle pollinators that are likely to originate in tropical rain forest. These native beetles occurred reliably enough in crops near rain forest to have a positive effect on the quantity of fruit produced but their contribution was not great enough to satisfy commercial production needs. Management changes, aimed at increasing native beetle abundance in crops, are required before these beetles could eliminate the need for growers to hand pollinate atemoya flowers. Appreciation of the value of this resource is necessary if we are to develop landscapes that both conserve native biodiversity and support agricultural production.     

Dung beetles in pasture landscapes of Central America: proliferation of synanthropogenic species and decline of forest specialists

This study examines the spread of synanthropogenic dung beetles (species favoured by human activities) in pasture landscapes in Central America, and evaluates the role of forest fragments and regenerating patches of native vegetation in maintaining beetle diversity. Pitfall trapping was carried out at nine locations in El Salvador and seven in Atlantic Nicaragua that included both pasture and remnant or regenerating native vegetation. More dung beetle species occurred in forest fragments than in pastures. Community composition differed considerably between forest fragments from El Salvador and Nicaragua with many species restricted to either the Caribbean or Pacific regions. In contrast, dung beetle community composition and structure were largely the same in the pastures of El Salvador and Nicaragua, regardless of region or original habitat-type, and were similar to published results from pastures in Mexico and elsewhere on the Isthmus. Very small patches of native shrubs and tree stands (<2.5 ha) maintained no forest specialists in Nicaragua, whereas, in El Salvador, some forest specialists occurred even in the smallest stands of trees (ca. 0.25 ha). The study indicates that the expansion of cattle pastures has caused a regional decline in dung beetle diversity. Forest fragments and small isolated patches of native trees and shrubs maintain some of the diversity of the original landscape but their conservation value for dung beetles will depend on the biogeographical history of the sites.     

The effect of habitat fragmentation on the bee visitor assemblages of three Australian tropical rainforest tree species

Tropical forest loss and fragmentation can change bee community dynamics and potentially interrupt plant–pollinator relationships. While bee community responses to forest fragmentation have been investigated in a number of tropical regions, no studies have focused on this topic in Australia. In this study, we examine taxonomic and functional diversity of bees visiting flowers of three tree species across small and large rainforest fragments in Australian tropical landscapes. We found lower taxonomic diversity of bees visiting flowers of trees in small rainforest fragments compared with large forest fragments and show that bee species in small fragments were subsets of species in larger fragments. Bees visiting trees in small fragments also had higher mean body sizes than those in larger fragments, suggesting that small‐sized bees may be less likely to persist in small fragments. Lastly, we found reductions in the abundance of eusocial stingless bees visiting flowers in small fragments compared to large fragments. These results suggest that pollinator visits to native trees living in small tropical forest remnants may be reduced, which may in turn impact on a range of processes, potentially including forest regeneration and diversity maintenance in small forest remnants in Australian tropical countryside landscapes.     

Bat species richness and abundance in tropical rain forest fragments and in agricultural habitats at Los Tuxtlas, Mexico

Faced with the rapid and extensive conversion of tropical rain forests to pasture lands and agricultural fields and with the need to preserve the remaining mammalian fauna, it is imperative to determine how the different species that form the mammalian community have responded to the anthropogenic alterations of their natural habitats To provide data in this direction, we sampled bats m 45 forest islands, m 20 agricultural habitats representing five types of vegetation (cocoa, coffee, mixed, citrus and allspice), in four live-fence sites and in four pasture sites at Los Tuxtlas, Veracruz, Mexico Sampling effort resulted in the capture of 2587 bats representing 35 species In forest habitats we detected 32 species We did not capture any bats at the four pasture sites, but the at the other agricultural habitats studied, we captured 38% of the bats and 77% of the species recorded Thirty-four percent of the species recorded were present at the live-fence habitats Isolating distance was an important variable influencing species richness in forests and in agricultural habitats Only 10% of the species recorded occurred m all the habitats studied, but 77% of the species occurred m a habitat other than ram forest Recaptures of bats indicated inter habitat movements in the fragmented landscape We discuss the conservation value for the bat fauna of agricultural islands of vegetation as elements reducing isolating distances among forest fragments     

Anthropogenic landscape changes and avian diversity at Los Tuxtlas, Mexico

Faced with rapid and extensive conversion of tropical rain forests to a landscape consisting of pasture lands, and with the need to preserve the avian diversity of tropical regions, it is imperative to determine how different species have responded to anthropogenic alterations of their natural habitats. We sampled birds in undisturbed and disturbed forest islands in regenerating forests and in four replicates of each of the following man-made habitats: arboreal agricultural habitats (cacao, coffee, mixed, citrus and allspice), non-arboreal agricultural habitats (corn, jalapen~o chili pepper and bananas), live fences and pastures, at Los Tuxtlas, Veracruz, Mexico. We censused 22145 birds representing 226 species. We detected 79% of the species in forest habitats, 80% in agricultural habitats, 43% in live fences and only 5% in pastures. Isolating distance and continued disturbance by humans of forest fragments were important variables influencing species' richness. Arboreal agricultural habitats and live fences were richer in species and in birds than non-arboreal man-made habitats. Economic surveys showed that some of the crops investigated yield higher returns than cattle ranching based on pastures. We discuss the conservation value for birds of agricultural islands and of live fences as landscape elements that help reduce physical and biotic isolation among remaining configurations of forest fragments in Los Tuxtlas.     

Roles of non‐native species in large‐scale regeneration of moist tropical forests on anthropogenic grassland

This paper presents a new synthesis of the role of native and non‐native species in diverse pathways and processes that influence forest regeneration on anthropogenic grassland in the moist tropics. Because of altered species composition, abiotic conditions and landscape habitat mosaics, together with human interventions, these successional pathways differ from those seen in pre‐clearing forests. However, representation of different functional life forms of plant (tree, vine, grass, herb and fern) and animal (frugivorous seed disperser, granivorous seed predator, seedling herbivore and carnivore) shows consistent global variation among areas of pasture, intact forest, and post‐grassland regrowth. Biotic webs of interaction involve complex indirect influences and feedbacks, which can account for wide observed variation in regeneration trajectories over time. Important processes include: limitation of tree establishment by dense grasses; recruitment and growth of pioneer pasture trees (shading grasses and facilitating bird‐assisted seed dispersal); and smothering of trees by vines. In these interactions, species’ functional roles are more important than their biogeographic origins. Case studies in eastern Australia show native rain forest plant species diversity in all life forms increasing over time when pioneer trees are non‐native (e.g., Cinnamomum camphora, Solanum mauritianum), concurrent with decreased grass and fern cover and increased abundance of trees and vine tangles. The global literature shows both native and non‐native species facilitating and inhibiting regeneration. However conservation goals are often targeted at removing non‐native species. Achieving large‐scale tropical forest restoration will require increased recognition of their multiple roles, and compromises about allocating resources to their removal.     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

Reproductive ecology of tropical forest trees in logged and fragmented habitats in Thailand and Costa Rica

Invertebrates mediate several important ecological processes, including pollination and seed predation, and events that affect invertebrate diversity or behaviour can potentially disrupt forest regeneration processes. This study investigates the impact of logging in Thailand and forest fragmentation in Costa Rica on the pollination and seed production of two self-incompatible forest trees. Logging in a dry deciduous dipterocarp forest in Thailand resulted in reduced densities of the common dipterocarp treeShorea siamensis and variably isolated individual trees. The number of flower visits to S. siamensis by pollinating Trigona bees was not affected by logging disturbance. However, pollinators did spend longer periods of time foraging in the canopies of isolated trees which were more prevalent in logged areas where tree density had been reduced. Consequently, at the logged site few cross-pollinations were effected and fruit set of S. siamensis was considerably lower than at nearby unlogged sites where distances between flowering conspecifics were smaller. Reduced fruit set has long-term implications for the recovery of S. siamensis populations in disturbed areas, and local population genetic structure is likely to be affected as reduced outcrossing rates among trees in disturbed regions results in relatively inbred seed. In Costa Rica forest fragmentation has restricted the once widespread tree Anacardium excelsum to forest patches located in an agriculturally-dominated landscape. As with S. siamensis, the abundance of pollinators, also Trigona bees, in the canopies of A. excelsum was largely unaffected by fragment size. Nevertheless, pollination success and seed production was positively correlated with fragment size. We propose that small bees rarely move between forest fragments and gene exchange through pollination occurs predominantly among trees within fragments and, together with likely low genetic variability in small fragments, that this contributes to the observed reduced fertilisation and seed set of A. excelsum. Thus increased tree isolation tree through selective logging or habitat fragmentation by forest clearance can result in reduced seed set due to changes in the foraging patterns of poorly mobile pollinators. Even if population sizes of the pollinators are maintained following environmental perturbation, this study shows that disturbance may disrupt pollination processes through changes in pollinator foraging behaviour. More attention needs to be focussed on changes in the behaviour of species involved in key ecological interactions following disturbance events in tropical forests.     

Tropical rain forest fragmentation and wild populations of primates at Los Tuxtlas,Mexico

In view of the extensive destruction, fragmentation, and conversion of primate habitats to anthropogenic vegetation, information on Neotropical primate ability to use a landscape consisting of forest fragments and agricultural habitats is necessary to understand the ecological flexibility of the species involved and it is of relevance to the design of conservation scenarios at the landscape level. We censused howlers and spider monkeys in 126 forest fragments and in 44 agricultural sites at Los Tuxtlas, southern Veracruz, Mexico, and used the IDRISI Geographic Information System to assess the extent of primate habitat remaining. We conducted economic surveys to assess the productivity of several systems of land management, including cattle ranching. Seventy- five percent of native habitat has been lost at Los Tuxtlas, 20% remains in the form of isolated fragments of vegetation, and only 5% consists of contiguous rain forest at high elevations (> 800 m). Howlers and spider monkeys were present in only 60 and 8% of the forest sites investigated, respectively, attesting to extensive local extinction. Populations of both species are small and found in a fragmented and isolated condition throughout the landscape. A large number of howlers were detected in artifactual habitats such as cacao, coffee and mixed (cacao and coffee) plantations shaded by rain forest trees. Residual rain forest vegetation along streams, rivers, and lakes facilitated the interfragment and interhabitat movement of howlers. Economic surveys showed that some of the agricultural habitats were more productive than cattle ranching. Conservation of spider monkeys requires large areas of contiguous forest, but only small areas are present at high elevations. Howlers are restricted to elevations < 800 m, where most of the forest has disappeared, but can apparently exist in a matrix of forest fragments, arboreal agricultural habitats, and pasture lands. Structural aspects of the vegetation and connectivity among habitat islands may be indispensable components of both landscape scenarios.     

The Circe principle explains how resource-rich land can waylay pollinators in fragmented landscapes

Global declines in pollinators, associated with land-use change [1-6] and fragmentation [7-10], constitute a serious threat to crop production and biodiversity [11]. Models investigating impacts of habitat fragmentation on pollen flow have categorized landscapes simply in terms of habitat and nonhabitat. We show that pollen flow depends strongly on types of land use between habitat fragments. We used paternity analysis of seeds and a combination of circuit and general linear models to analyze pollen flow for the endangered tree Gomortega keule (Gomortegaceae) [12] in the fragmented Central Chile Biodiversity Hotspot [13]. Pollination probability was highest over pine plantation, moderate over low-intensity agriculture and native forest, and lowest over clearfells. Changing the proportions of the land uses over one kilometer altered pollination probability up to 7-fold. We explain our results by the novel "Circe principle." In contrast to models where land uses similar to native habitat promote pollinator movement, pollinators may actually be waylaid in resource-rich areas between habitat patches. Moreover, pollinators may move with higher probability between habitat patches separated by some resource-poor land uses. Pollination research in fragmented landscapes requires explicit recognition of the nature of the nonhabitat matrix, rather than applying simple binary landscape models.     

The relative importance of birds and bees in the pollination of Metrosideros excelsa (Myrtaceae)

Exclusion experiments were used to assess the effect of different pollinator groups on outcrossing and seed production in Metrosideros excelsa. The main study site was Little Barrier Island, New Zealand where indigenous bird and native solitary bees are the main flower visitors. Our results showed that native birds were more important pollinators of M. excelsa than native bees. Seed production was much higher in open pollination than in two exclusion experiments where either birds were excluded and native bees only had access to flowers, or where all pollinators had been excluded. The number of fertile seeds per capsule was 45% higher after open pollination than in treatments with bee visitation only and 28% higher than in treatments where all flower visitors were excluded. Estimated outcrossing rates were significantly higher (t_m = 0.71) for open pollination in the upper canopy (>4 m above‐ground level) where bird visitation is presumed to be more frequent than for a treatment with native bee access only (t_m = 0.40). Our results also suggest that a large proportion of seeds (66%) arise from autonomous self‐pollination when all pollinators are excluded. In four trees of a modified mainland population with predominantly introduced birds and a mixture of introduced and native bees there was no decrease in seed production for the treatment allowing bee access only, indicating that – in contrast to native bees – honeybees may be more efficient pollinators of M. excelsa. Observation of the foraging behaviour of both groups of bees showed that native bees contact the stigma of flowers less frequently than honeybees. This is likely to be a consequence of their smaller body size relative to honeybees.     

Tropical Montane Forest Restoration in Costa Rica: Overcoming Barriers to Dispersal and Establishment

Tropical forests are being cleared at an alarming rate although our understanding of their ecology is limited. It is therefore essential to design restoration experiments that both further our basic knowledge of tropical ecology and inform management strategies to facilitate recovery of these ecosystems. Here we synthesize the results of research on tropical montane forest recovery in abandoned pasture in Costa Rica to address the following questions: (1) What factors limit tropical forest recovery in abandoned pasture? and (2) How can we use this information to design strategies to facilitate ecosystem recovery? Our results indicate that a number of factors impede tropical forest recovery in abandoned pasture land. The most important barriers are lack of dispersal of forest seeds and seedling competition with pasture grasses. High seed predation, low seed germination, lack of nutrients, high light intensity, and rabbit herbivory also affect recovery. Successful strategies to facilitate recovery in abandoned pastures must simultaneously overcome numerous obstacles. Our research shows that establishment of woody species, either native tree seedlings or early‐successional shrubs, can be successful in facilitating recovery, by enhancing seed dispersal and shading out pasture grasses. On the contrary, bird perching structures alone are not an effective strategy, because they only serve to enhance seed dispersal but do not reduce grass cover. Remnant pasture trees can serve as foci of natural recovery and may enhance growth of planted seedlings. Our results highlight the importance of: (1) understanding the basic biology of an ecosystem to design effective restoration strategies; (2) comparing results across a range of sites to determine which restoration strategies are most generally useful; and (3) considering where best to allocate efforts in large‐scale restoration projects.