Incorporating animal behavior into seed dispersal models: implications for seed shadows

Seed dispersal fundamentally influences plant population and community dynamics but is difficult to quantify directly. Consequently, models are frequently used to describe the seed shadow (the seed deposition pattern of a plant population). For vertebrate-dispersed plants, animal behavior is known to influence seed shadows but is poorly integrated in seed dispersal models. Here, we illustrate a modeling approach that incorporates animal behavior and develop a stochastic, spatially explicit simulation model that predicts the seed shadow for a primate-dispersed tree species (Virola calophylla, Myristicaceae) at the forest stand scale. The model was parameterized from field-collected data on fruit production and seed dispersal, behaviors and movement patterns of the key disperser, the spider monkey (Ateles paniscus), densities of dispersed and non-dispersed seeds, and direct estimates of seed dispersal distances. Our model demonstrated that the spatial scale of dispersal for this V. calophylla population was large, as spider monkeys routinely dispersed seeds >100 m, a commonly used threshold for long-distance dispersal. The simulated seed shadow was heterogeneous, with high spatial variance in seed density resulting largely from behaviors and movement patterns of spider monkeys that aggregated seeds (dispersal at their sleeping sites) and that scattered seeds (dispersal during diurnal foraging and resting). The single-distribution dispersal kernels frequently used to model dispersal substantially underestimated this variance and poorly fit the simulated seed-dispersal curve, primarily because of its multimodality, and a mixture distribution always fit the simulated dispersal curve better. Both seed shadow heterogeneity and dispersal curve multimodality arose directly from these different dispersal processes generated by spider monkeys. Compared to models that did not account for disperser behavior, our modeling approach improved prediction of the seed shadow of this V. calophylla population. An important function of seed dispersal models is to use the seed shadows they predict to estimate components of plant demography, particularly seedling population dynamics and distributions. Our model demonstrated that improved seed shadow prediction for animal-dispersed plants can be accomplished by incorporating spatially explicit information on disperser behavior and movements, using scales large enough to capture routine long-distance dispersal, and using dispersal kernels, such as mixture distributions, that account for spatially aggregated dispersal.     

Neighborhood effects on seed dispersal by frugivores: testing theory with a mistletoe-marsupial system in Patagonia

The outcome of the dispersal process in zoochorous plants is largely determined by the behavior of frugivorous animals. Recent simulation studies have found that fruit removal rates and mean dispersal distances are strongly affected by fruiting plant neighborhoods. We empirically tested the effects of conspecific fruiting plant neighborhoods, crop sizes, and plant accessibility on fruit removal rates and seed dispersal distances of a mistletoe species exclusively dispersed by an arboreal marsupial in Northern Patagonia. Moreover, in this study, we overcome technical limitations in the empirical estimation of seed dispersal by using a novel 15N stable isotope enrichment technique together with Bayesian mixing models that allowed us to identify dispersed seeds from focal plants without the need of extensive genotyping. We found that, as predicted by theory, plants in denser neighborhoods had greater fruit removal and shorter mean dispersal distances than more isolated plants. Furthermore, the probability of dispersing seeds farther away decreased with neighborhood density. Larger crop sizes resulted in larger fruit removal rates and smaller probabilities of longer distance dispersal. The interplay between frugivore behavioral decisions and the spatial distribution of plants could have important consequences for plant spatial dynamics.     

Nonredundancy in the dispersal network of a generalist tropical forest tree

Plant species with generalized dispersal mutualisms are considered to be robust to local frugivore extinctions because of redundancy between dispersal agents. However, real redundancy can only occur if frugivores have similar foraging and ranging patterns and if fruit is a limiting resource. We evaluated the quantitative and qualitative contributions of seed dispersers for an endochorus mast-fruiting species, Prunus javanica (Rosaceae) in Khao Yai National Park, Thailand, to evaluate the potential redundancy of dispersers. Data were collected from tree watches, seed/fruit traps, and seed transects under and away from fruiting trees, feeding and seed deposition by gibbons (Hylobates lar), and evaluations of seed and first-year seedling survival. We identified three clusters of dispersers within the network. Most (>80%) frugivore species observed were small birds and squirrels that were not functional dispersers, dropping most seeds under or very near the tree crown, where seedling survival was ultimately nil. Monkeys (Macaca leonina) were low-quality, short-range dispersers, but they dispersed large numbers of seeds and were responsible for 67% of surviving first-year seedlings. Gibbons and Oriental Pied Hornbills (Anthracoceros albirostris) handled few fruits, but they provided the highest quality service by carrying most seeds away from the canopy to medium and long distances, respectively. Although there was overlap in the deposition patterns of the functional dispersers, they displayed complementary, rather than redundant, roles in seed dispersal. Satiation of all functional dispersers further limited their capacity to "replace" one another. Redundancy must be evaluated at the community level because each type of disperser may shift to different species in the non-masting years of P. javanica. Our results underscore the need for research on broader spatial and temporal scales, which combines studies of dispersal and plant recruitment, to better understand mechanisms that maintain network stability.     

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     

Specialization and interaction strength in a tropical plant-frugivore network differ among forest strata

The degree of interdependence and potential for shared coevolutionary history of frugivorous animals and fleshy-fruited plants are contentious topics. Recently, network analyses revealed that mutualistic relationships between fleshy-fruited plants and frugivores are mostly built upon generalized associations. However, little is known about the determinants of network structure, especially from tropical forests where plants' dependence on animal seed dispersal is particularly high. Here, we present an in-depth analysis of specialization and interaction strength in a plant-frugivore network from a Kenyan rain forest. We recorded fruit removal from 33 plant species in different forest strata (canopy, midstory, understory) and habitats (primary and secondary forest) with a standardized sampling design (3447 interactions in 924 observation hours). We classified the 88 frugivore species into guilds according to dietary specialization (14 obligate, 28 partial, 46 opportunistic frugivores) and forest dependence (50 forest species, 38 visitors). Overall, complementary specialization was similar to that in other plant-frugivore networks. However, the plant-frugivore interactions in the canopy stratum were less specialized than in the mid- and understory, whereas primary and secondary forest did not differ. Plant specialization on frugivores decreased with plant height, and obligate and partial frugivores were less specialized than opportunistic frugivores. The overall impact of a frugivore increased with the number of visits and the specialization on specific plants. Moreover, interaction strength of frugivores differed among forest strata. Obligate frugivores foraged in the canopy where fruit resources were abundant, whereas partial and opportunistic frugivores were more common on mid- and understory plants, respectively. We conclude that the vertical stratification of the frugivore community into obligate and opportunistic feeding guilds structures this plant-frugivore network. The canopy stratum comprises stronger links and generalized associations, whereas the lower strata are composed of weaker links and more specialized interactions. Our results suggest that seed-dispersal relationships of plants in lower forest strata are more prone to disruption than those of canopy trees.     

Disentangling the drivers of reduced long-distance seed dispersal by birds in an experimentally fragmented landscape

Seed dispersal is a crucial component of plant population dynamics. Human landscape modifications, such as habitat destruction and fragmentation, can alter the abundance of fruiting plants and animal dispersers, foraging rates, vector movement, and the composition of the disperser community, all of which can singly or in concert affect seed dispersal. Here, we quantify and tease apart the effects of landscape configuration, namely, fragmentation of primary forest and the composition of the surrounding forest matrix, on individual components of seed dispersal of Heliconia acuminata, an Amazonian understory herb. First we identified the effects of landscape configuration on the abundance of fruiting plants and six bird disperser species. Although highly variable in space and time, densities of fruiting plants were similar in continuous forest and fragments. However, the two largest-bodied avian dispersers were less common or absent in small fragments. Second, we determined whether fragmentation affected foraging rates. Fruit removal rates were similar and very high across the landscape, suggesting that Heliconia fruits are a key resource for small frugivores in this landscape. Third, we used radiotelemetry and statistical models to quantify how landscape configuration influences vector movement patterns. Bird dispersers flew farther and faster, and perched longer in primary relative to secondary forests. One species also altered its movement direction in response to habitat boundaries between primary and secondary forests. Finally, we parameterized a simulation model linking data on fruit density and disperser abundance and behavior with empirical estimates of seed retention times to generate seed dispersal patterns in two hypothetical landscapes. Despite clear changes in bird movement in response to landscape configuration, our simulations demonstrate that these differences had negligible effects on dispersal distances. However, small fragments had reduced densities of Turdus albicollis, the largest-bodied disperser and the only one to both regurgitate and defecate seeds. This change in Turdus abundance acted together with lower numbers of fruiting plants in small fragments to decrease the probability of long-distance dispersal events from small patches. These findings emphasize the importance of foraging style for seed dispersal and highlight the primacy of habitat size relative to spatial configuration in preserving biotic interactions.     

Effects of Local Disturbance of Tropical Forests on Frugivores and Seed Removal of a Small-Seeded Afrotropical Tree

Abstract:  Small-scale, local disturbance of tropical forests, for example from selective logging, is widespread, but its effects on biodiversity and ecosystem function have rarely been studied. In 3 East African tropical rainforests, we investigated the effect of different levels of local forest disturbance on the frugivore community and on tree visitation and fruit removal rates of the small-seeded tree Celtis durandii. We quantified birds and primates in little and heavily disturbed sites, distinguishing between forest specialists, forest generalists, and forest visitors. We quantified frugivorous tree visitors and seed removal rates of C. durandii trees in the same sites. Forest disturbance reduced the species richness and density of the frugivore community and of forest specialists. Frugivorous species and individuals visiting the study trees were reduced significantly, which led to a marginally significant decline in fruit removal by all frugivores and a significant reduction in removal by forest specialists. Reduction in fruit removal by forest specialists was not compensated for by increases in removal by forest generalists or visitors. Results did not differ among the 3 rainforests, which suggests they were consistent at a regional scale. So local forest disturbance led to a loss of frugivores and their seed removal services. This suggests that large-seeded tree species and trees with small fruits are losing seed dispersers. Thus, local forest disturbance appears to have a more general negative impact on frugivores and their seed dispersal services than anticipated previously.     

The memory of spatial patterns: changes in local abundance and aggregation in a tropical forest

The current spatial pattern of a population is the result of previous individual birth, death, and dispersal events. We present a simple model followed by a comparative analysis for a species-rich plant community to show how the current spatial aggregation of a population may hold information about recent population dynamics. Previous research has shown how locally restricted seed dispersal often leads to stronger aggregation in less abundant populations than it does in more abundant populations. In contrast, little is known about how changes in the local abundance of a species may affect the spatial distribution of individuals. If the level of aggregation within a species depends to some extent on the abundance of the species, then changes in abundance should lead to subsequent changes in aggregation. However, an overall change of spatial pattern relies on many individual birth and death events, and a surplus of deaths or births may have short-term effects on aggregation that are opposite to the long-term change predicted by the change in abundance. The change in aggregation may therefore lag behind the change in abundance, and consequently, the current aggregation may hold information about recent population dynamics. Using an individual-based simulation model with local dispersal and density-dependent competition, we show that, on average, recently growing populations should be more aggregated than shrinking populations of the same current local abundance. We tested this hypothesis using spatial data on individuals from a long-term tropical rain forest plot, and find support for this relationship in canopy trees, but not in understory and shrub species. On this basis we argue that current spatial aggregation is an important characteristic that contains information on recent changes in local abundance, and may be applied to taxonomic groups where dispersal is limited and within-species aggregation is observed.     

Seed dispersal by pulp consumers, not "legitimate" seed dispersers, increases Guettarda viburnoides population growth

We examined the effect of seed dispersal by Purplish Jays (Cyanocorax cyanomelas; pulp consumers) and the Chestnut-eared Ara?ari (Pteroglossus castanotis; "legitimate" seed dispersers) on population growth of the small tree Guettarda viburnoides (Rubiaceae) in northeastern Bolivian savannas. Because each bird species differs with respect to feeding and post-feeding behavior, we hypothesized that seed dispersal by each species will contribute differently to the rate of increase of G. viburnoides, but that seed dispersal by either species will increase population growth when compared to a scenario with no seed dispersal. To examine the effects of individual dispersers on the future population size of G. viburnoides, we projected population growth rate using demographic models for G. viburnoides that explicitly incorporate data on quantitative and qualitative aspects of seed dispersal by each frugivore species. Our model suggests that seed dispersal by C. cyanomelas leads to positive population growth of G. viburnoides, whereas seed dispersal by P. castanotis has a detrimental effect on the population growth of this species. To our knowledge, this is the first study to report negative effects of a "legitimate" seed disperser on the population dynamics of the plant it consumes. Our results stress the importance of incorporating frugivore effects into population projection matrices, to allow a comprehensive analysis of the effectiveness of different dispersers for plant population dynamics.     

Distribution patterns of plants explained by human movement behaviour

Distribution patterns of plants are affected by human activities such as creation, destruction or modification of habitats. However, another important question is to what extent humans shape plant distributions by acting as dispersal vectors. In order to answer this question we developed a simulation model for the spread of plant species between human settlements. This was done on the basis of extensive sociological and ecological data on a regional scale. With regard to the sociological data, human movement behaviour defined the amount of exchange between the settlements. Gardening types represented the potential habitat in our model. The ecological data was derived from a vegetation survey carried out in 2003, which was a repeat of a survey between 1974 and 1981 along the same transects. From these surveys, we studied the distributions of 13 species in 67 settlements. In our model, the earlier survey provided the data for the initial distribution. The simulated pattern was consequently compared with the distribution pattern in 2003. In the model, dispersal kernels based on patterns of human movement between settlements led to a better match with the distribution patterns than a null model simulating pure distance dependent dispersal for all species. This was statistically significant for seven of the thirteen species. A striking result was that alien species seem to benefit more from human dispersal than native species. We emphasize the importance of the sociological data on human movement behaviour in parameterizing our regional scale model. This study provides quantitative evidence on the impact of human movement behaviour on the distribution of plant species in suburban areas.     

Effects of howler monkey reintroduction on ecological interactions and processes

Rewilding has been an increasingly popular tool to restore plant–animal interactions and ecological processes impaired by defaunation. However, the reestablishment of such processes has seldom been assessed. We investigated the restoration of ecological interactions following the reintroduction of the brown howler monkey (Alouatta guariba) to a defaunated Atlantic forest site. We expected the reintroduction to restore plant–animal interactions and interactions between howlers and dung beetles, which promote secondary seed dispersal. We estimated the number of interactions expected to be restored by the reintroduction to provide the baseline interaction richness that could be restored. We followed the reintroduced howler monkeys twice a week for 24 months (337 hours total) to assess their diet. We used howler monkey dung in secondary seed dispersal experiments with 2484 seed mimics to estimate the removal rates by dung beetles and collected the beetles to assess community attributes. We compared the potential future contribution of howler monkeys and other frugivores to seed dispersal based on the seed sizes they disperse in other areas where they occur. In 2 years, howler monkeys consumed 60 animal-dispersed plant species out of the 330 estimated. Twenty-one dung beetle species were attracted to experimentally provided dung; most of them were tunnelers, nocturnal, and large-sized (>10 mm). On average 30% (range 0–100%) of the large seed mimics (14 mm) were moved by dung beetles. About 91% of the species consumed by howlers (size range 0.3–34.3 mm) overlapped in seed size with those removed by dung beetles. In our study area, howler monkeys may consume more large-seeded fruit species than most other frugivores, highlighting their potential to affect forest regeneration. Our results show reintroductions may effectively restore ecological links and enhance ecological processes.     

Soil vs. canopy seed storage and plant species coexistence in species-rich Australian shrublands

The fire-prone shrublands of southwestern Australia are renowned for their high plant species diversity and prominence of canopy seed storage (serotiny). We compared species richness, abundance, and life history attributes for soil and canopy seed banks in relation to extant vegetation among four sites with different substrate conditions and high species turnover (50-80%) to identify whether this unusual community-level organization of seed storage might contribute to maintenance of high species richness. Soil seed bank (SSB) densities were low to moderate (233-1435 seeds/m2) compared with densities for other Mediterranean-type vegetation and were lowest for sites with highest canopy seed bank (CSB) species richness and lowest nutrient availability, but not richness or abundance of resprouters. Annuals were infrequent in the lowest nutrient sites, but there was no evidence that small SSB size was due to low seed inputs or a trade-off between seed production/storage and seed size in response to low nutrient availability. Sorensen's similarity between SSB and extant vegetation was 26-43% but increased to 54-57% when the CSB was included, representing levels higher than reported for most other ecosystems. Resprouting species were well represented in both the SSB and CSB, and there was no evidence for lower seed production in resprouters than in non-sprouters overall. The SSB and CSB held no species in common and were characterized by markedly different seed dispersal attributes, with winged or small seeds in the CSB and seeds dispersed by ants, birds, and wind (though none with wings) in the SSB. There was no evidence of spatial differentiation in the distribution of seeds of SSB species between vegetated and open microsites that might facilitate species coexistence, but most woody non-sprouters showed aggregation at scales of 1-2 m, implying limited seed dispersal. High similarity between overall seed bank (SSB + CSB) and extant species composition, high number of resprouting species, and seed dispersal processes before (SSB) and after fire (CSB) leading to differential spatial aggregation of post-fire recruits from the two seed bank types may buffer species composition against rapid change and provide a mechanism for maintaining species coexistence at the local scale.     

A method for building spatial model of annual weed seed dispersal from experimental data and its application to simulating Bromus sterilis population dispersal

Spatial model of annual weed seed dispersal, in this article, was theoretically derived. According to the requirements of building the spatial model, we designed and done an indoor experiment of weed seed dispersal by wind. In the experiment, the seeds of Bromus sterilis were released at 100 cm height under different wind velocity conditions. Based on the experimental data, the spatial models of seed dispersal of the weed species were built, which were divided into three types according to the coefficient β < 0, β = 0, β > 0. The results showed that dispersal of annual weed seed in any direction obeyed an approximate Gaussian distribution; under the experimental conditions, spatial distribution type of weed seed dispersal changed with variation of wind velocity. Well-known Howard et al.'s model (Howard et al., 1991) of Bromus sterilis seed dispersal is an especial example of the model built in this article. The result of model analysis indicated that the distribution type described by Howard's model was similar to that of seed dispersal of the weed species at the height of 100 cm under the condition of lower wind velocity (about 2.18 m/s). Using CA simulation analysis we found that mean control agent applying to a cell with weed should have a decrease with an increase of wind velocity to prevent weed with the initial configuration from spreading, which implicated less herbicide needs spraying in every cell with weed on average when wind velocity increases.     

The impact of fragmentation and density regulation on forest succession in the Atlantic rain forest

The Atlantic Rain Forest, an important biodiversity hot spot, has faced severe habitat loss since the last century which has resulted in a highly fragmented landscape with a large number of small forest patches (<100 ha). For conservation planning it is essential to understand how current and future forest regeneration depends on ecological processes, fragment size and the connection to the regional seed pool. We have investigated the following questions by applying the forest growth simulation model FORMIND to the situation of the Atlantic Forest in the state of São Paulo, SE Brazil: (1) which set of parameters describing the local regeneration and level of density regulation can reproduce the biomass distribution and stem density of an old growth forest in a reserve? (2) Which additional processes apart from those describing the dynamics of an old growth forest, drive forest succession of small isolated fragments? (3) Which role does external seed input play during succession? Therefore, more than 300 tree species have been classified into nine plant functional types (PFTs), which are characterized by maximum potential height and shade tolerance. We differentiate between two seed dispersal modes: (i) local dispersal, i.e. all seedlings originated from fertile trees within the simulated area and (ii) external seed rain. Local seed dispersal has been parameterized following the pattern oriented approach, using biomass estimates of old growth forest. We have found that moderate density regulation is essential to achieve coexistence for a broad range of regeneration parameters. Considering the expected uncertainty and variability in the regeneration processes it is important that the forest dynamics are robust to variations in the regeneration parameters. Furthermore, edge effects such as increased mortality at the border and external seed rain have been necessary to reproduce the patterns for small isolated fragments. Overall, simulated biomass is much lower in the fragments compared to the continuous forest, whereas shade tolerant species are affected most strongly by fragmentation. Our simulations can supplement empirical studies by extrapolating local knowledge on edge effects of fragments to larger temporal and spatial scales. In particular our results show the importance of external seed rain and therefore highlight the importance of structural connectivity between regenerating fragments and mature forest stands.     

Fish dispersal in fragmented landscapes: a modeling framework for quantifying the permeability of structural barriers

Dispersal is a key determinant of the spatial distribution and abundance of populations, but human-made fragmentation can create barriers that hinder dispersal and reduce population viability. This study presents a modeling framework based on dispersal kernels (modified Laplace distributions) that describe stream fish dispersal in the presence of obstacles to passage. We used mark-recapture trials to quantify summer dispersal of brook trout (Salvelinus fontinalis) in four streams crossed by a highway. The analysis identified population heterogeneity in dispersal behavior, as revealed by the presence of a dominant sedentary component (48-72% of all individuals) characterized by short mean dispersal distance (<10 m), and a secondary mobile component characterized by longer mean dispersal distance (56-1086 m). We did not detect evidence of barrier effects on dispersal through highway crossings. Simulation of various plausible scenarios indicated that detectability of barrier effects was strongly dependent on features of sampling design, such as spatial configuration of the sampling area, barrier extent, and sample size. The proposed modeling framework extends conventional dispersal kernels by incorporating structural barriers. A major strength of the approach is that ecological process (dispersal model) and sampling design (observation model) are incorporated simultaneously into the analysis. This feature can facilitate the use of prior knowledge to improve sampling efficiency of mark-recapture trials in movement studies. Model-based estimation of barrier permeability and its associated uncertainty provides a rigorous approach for quantifying the effect of barriers on stream fish dispersal and assessing population dynamics of stream fish in fragmented landscapes.     

Using CA model to obtain insight into mechanism of plant population spread in a controllable system: annual weeds as an example

Using cellular automata (CA) model, in this article, a mechanistic approach has been conducted to help to insight into spread process of plant population such as annual weed population. A controllable CA model with 25 neighbourhood cells has been built, in which seed dispersal, as a key process of annual weed population dynamics, has been described by Gaussian distribution. The hypothesis that initial configurations affect annual weed population dynamics and control strategies has been tested by simulating. And the results support strongly the hypothesis. Patch, pattern and configuration are important concepts in weed patch management, which have been mathematically defined in this article. The aggregation effects have been found out while simulating, which is subject to patch size, patch density and distribution function of seed dispersal. Perhaps it could partly explain the reason why weeds often distribute in patch. Accordingly, true-patch and generalised patch have been distinguished from each other, which would be practically applicable to weed patch management. Based on this approach to the particular weed-crop system, a new hypothesis has been proposed: in a real controllable weed-crop system, the mean law might hold. If so, algorithm of patch control for long-term weed management could be simplified to control a true-patch only according to its mean density, it is unnecessary to control it according to the density in its each cell.     

Effects of Growth Form and Functional Traits on Response of Woody Plants to Clearing and Fragmentation of Subtropical Rainforest

The conservation implications of large‐scale rainforest clearing and fragmentation on the persistence of functional and taxonomic diversity remain poorly understood. If traits represent adaptive strategies of plant species to particular circumstances, the expectation is that the effect of forest clearing and fragmentation will be affected by species functional traits, particularly those related to dispersal. We used species occurrence data for woody plants in 46 rainforest patches across 75,000 ha largely cleared of forest by the early 1900s to determine the combined effects of area reduction, fragmentation, and patch size on the taxonomic structure and functional diversity of subtropical rainforest. We compiled species trait values for leaf area, seed dry mass, wood density, and maximum height and calculated species niche breadths. Taxonomic structure, trait values (means, ranges), and the functional diversity of assemblages of climbing and free‐standing plants in remnant patches were quantified. Larger rainforest patches had higher species richness. Species in smaller patches were taxonomically less related than species in larger patches. Free‐standing plants had a high percentage of frugivore dispersed seeds; climbers had a high proportion of small wind‐dispersed seeds. Connections between the patchy spatial distribution of free‐standing species, larger seed sizes, and dispersal syndrome were weak. Assemblages of free‐standing plants in patches showed more taxonomic and spatial structuring than climbing plants. Smaller isolated patches retained relatively high functional diversity and similar taxonomic structure to larger tracts of forest despite lower species richness. The response of woody plants to clearing and fragmentation of subtropical rainforest differed between climbers and slow‐growing mature‐phase forest trees but not between climbers and pioneer trees. Quantifying taxonomic structure and functional diversity provides an improved basis for conservation planning and management by elucidating the effects of forest‐area reduction and fragmentation. Efectos de la Forma de Crecimiento y Atributos Funcionales en la Respuesta de Plantas Leñosas al Desmonte y Fragmentación de Bosque Lluvioso Subtropical     

Experimental manipulation of seed shadows of an Afrotropical tree determines drivers of recruitment

The loss of animals in tropical forests may alter seed dispersal patterns and reduce seedling recruitment of tree species, but direct experimental evidence is scarce. We manipulated dispersal patterns of Manilkara mabokeensis, a monkey-dispersed tree, to assess the extent to which spatial distributions of seeds drive seedling recruitment. Based on the natural seed shadow, we created seed distributions with seeds deposited under the canopy ("no dispersal"), with declining density from the tree ("natural dispersal"), and at uniform densities ("good dispersal"). These distributions mimicked dispersal patterns that could occur with the extirpation of monkeys, low levels of hunting, and high rates of seed dispersal. We monitored seedling emergence and survival for 18 months and recorded the number of leaves and damage to leaves. "Good dispersal" increased seedling survival by 26%, and "no dispersal" decreased survival by 78%, relative to "natural dispersal." Using a mixed-effects survival model, we decoupled the distance and density components of the seed shadow: seedling survival depended on the seed density, but not on the distance from the tree. Although community seedling diversity tended to decrease with longer dispersal distances, we found no conclusive evidence that patterns of seed dispersal influence the diversity of the seedling community. Local seed dispersal does affect seedling recruitment and survival, with better dispersal resulting in higher seedling recruitment; hence the loss of dispersal services that comes with the reduction or extirpation of seed dispersers will decrease regeneration of some tree species.     

Animal dispersal modelling: Handling landscape features and related animal choices

Animal dispersal in a fragmented landscape depends on the complex interaction between landscape structure and animal behavior. To better understand how individuals disperse, it is important to explicitly represent the properties of organisms and the landscape in which they move. A common approach to modelling dispersal includes representing the landscape as a grid of equal sized cells and then simulating individual movement as a correlated random walk. This approach uses a priori scale of resolution, which limits the representation of all landscape features and how different dispersal abilities are modelled.We develop a vector-based landscape model coupled with an object-oriented model for animal dispersal. In this spatially explicit dispersal model, landscape features are defined based on their geographic and thematic properties and dispersal is modelled through consideration of an organism's behavior, movement rules and searching strategies (such as visual cues). We present the model's underlying concepts, its ability to adequately represent landscape features and provide simulation of dispersal according to different dispersal abilities. We demonstrate the potential of the model by simulating two virtual species in a real Swiss landscape. This illustrates the model's ability to simulate complex dispersal processes and provides information about dispersal such as colonization probability and spatial distribution of the organism's path.     

Local population size in a flightless insect: importance of patch structure-dependent mortality

In spatially heterogeneous systems, utilizing population models to integrate the effects of multiple population rates can yield powerful insights into the relative importance of the component rates. The relative importance of demographic rates and dispersal in shaping the distribution of the western tussock moth (Orgyia vetusta) among patches of its host plant was explored using stage-structured population models. Tussock moth dispersal occurs passively in first-instar larvae and is poor or absent in all other life stages. Spatial surveys suggested, however, that moth distribution is not well explained by passive dispersal; moth populations were greater on small patches and on isolated ones. Further analysis showed that several local demographic rates varied significantly with patch characteristics. Two mortality factors in particular may explain the observed patterns. First, crawler mortality both increased with patch size and was density-dependent. A single-patch difference equation model showed mortality related to patch size is strong enough to overcome the homogenizing effect of density dependence; greater equilibrium densities were predicted for smaller patches. Second, although three rates were found to vary with local patch density, only pupal parasitism by a chalcid wasp could potentially account for higher moth abundances on isolated patches. A spatially explicit simulation model of the multiple-patch system showed that spatial variation in pupal parasitism is indeed strong enough to generate such a pattern. These results demonstrate that habitat spatial structure can affect multiple population processes simultaneously, and even relatively low attack rates imposed on a reproductively valuable life stage of the host can have a dominant effect on population distribution among habitat patches.