Absence of Recruitment Limitation in Restored Dune Slacks Suggests That Manual Seed Introduction Can Be a Successful Practice for Restoring Orchid Populations

Answering the question whether or not to introduce seeds of target species in restored habitats depends on the relative importance of dispersal and recruitment limitation. Especially in orchid species, recruitment limitation is expected to be important because of their dependence on mycorrhiza for germination. Using a large seed introduction experiment we investigated the relative importance of dispersal versus recruitment limitation in the failure of three rare orchid species to colonize restored habitat patches. For all species, seeds developed successfully into protocorms in both occupied and unoccupied habitats, and no significant differences in the number of protocorms per seed packet were found between occupied and unoccupied habitats. These results show that orchid species do not necessarily suffer from recruitment limitation when introduced in restored habitats, and that manual introduction of orchid seeds can be a successful and necessary ecological restoration practice.     

Biogeography of seed-dispersal syndromes, life-forms and seed sizes among woody rain-forest plants in Australia's subtropics

Aim To enhance our understanding of the evolutionary interactions between seed‐dispersal syndromes, life‐forms, seed size, and habitat characteristics by studying their association with the regional‐scale distributions of subtropical rain‐forest plants in the context of climatic gradients. Location South‐east Queensland, subtropical eastern Australia (152° E, 26° S). Methods We classified 250 rain‐forest sites into six floristic site‐groups based on their woody plant composition. The resulting classification was strongly associated with variation in rainfall. The distribution of species across the floristic site‐groups was used to assign 568 species to seven habitat classes (one class for ‘widespread’ species, with all other species classified according to the site‐group within which they were most frequent). Species were also classified for three other categorical life‐history factors: three dispersal syndromes based on diaspore morphology (fleshy, wind‐assisted, and unadorned); four life‐forms (trees, shrubs and small trees, tall climbers, and short and shrubby climbers); and four seed‐diameter classes (< 3 mm, ≥ 3 and < 4.5 mm, ≥ 4.5 and < 7 mm, and ≥ 7 mm). We used a basic comparative approach augmented by simple phylogenetically constrained comparisons to assess association between dispersal syndrome, seed size, life‐form, and habitat class. Results Across the rain forests of south‐east Queensland, the proportion of species with fleshy diaspores or of large stature increases with rainfall. High‐rainfall sites also have larger average seed sizes, but the difference in average seed size between high‐ and low‐rainfall sites is small compared with variation within sites. Among species, those with fleshy fruit tend to have larger seeds and to favour high‐rainfall sites. Very few small trees produce diaspores adapted for wind‐assisted dispersal. On average, species with unadorned diaspores have smaller seeds than those with fleshy diaspores. However, within sites, species with unadorned and fleshy diaspores have similar average seed sizes, and some species with unadorned diaspores from high‐rainfall habitats have extremely large seeds. Main conclusions Commonly observed associations between fleshy fruit, larger plants, larger seeds, and productive habitats are apparent within the rain‐forest flora of south‐east Queensland. However, these associations are generally weak and involve complex interactions. For example, the strong tendency for species with fleshy fruit to have larger seeds than those with unadorned diaspores concealed a significant group of species from wetter forests that produce extremely large seeds and unadorned diaspores. The most widespread species in this study tend to be large plants (particularly robust lianes) and to produce fleshy fruit, but they tend not to have relatively large seeds. The association between large seeds, large plants, fleshy fruit and productive habitats is discussed as part of an evolutionary strategy favouring fitness in populations close to carrying capacity. We review some problems with focusing on establishment chances per seed as the driver towards association between large seeds, large plants and productive rain‐forest habitats (the difficult‐establishment hypothesis). Instead we suggest that production of large, short‐lived seeds by long‐lived plants in temporally stable, closed habitats may reflect the limited evolutionary potential for strategies enhancing colonization (e.g. producing large numbers of dormant seeds), thus allowing the establishment benefits of large seeds greater selective influence (the slow‐replacement hypothesis). The association of fleshy fruit with large seeds probably reflects the difficulty of dispersing large seeds by other means (the difficult‐dispersal hypothesis).     

Exploring the interaction of avian frugivory and plant spatial heterogeneity and its effect on seed dispersal kernels using a simulation model

Seed dispersal by avian frugivores is one of the key processes influencing plant spatial patterns, but may fail if there is disruption of plant–frugivore mutualisms, such as decline in abundance of dispersers, fragmentation of habitat, or isolation of individual trees. We used simulation model experiments to examine the interaction between frugivore density and behaviour and the spatial arrangement of fruiting plants and its effect on seed dispersal kernels. We focussed on two New Zealand canopy tree species that produce large fruits and are dispersed predominantly by one avian frugivore (Hemiphaga novaeseelandiae). Although the mean seed dispersal distance decreased when trees became more aggregated, there were more frugivore flights between tree clusters, consequently stretching the tails of the dispersal kernels. Conversely, when trees were less aggregated in the landscape, mean dispersal distances increased because seeds were deposited over larger areas, but the kernels had shorter tails. While there were no statistically meaningful changes in kernel parameters when frugivore density changed, decreases in density did cause a proportional reduction in the total number of dispersed seeds. However, birds were forced to move further when fruit availability and fruit ripening were low. Sensitivity analysis showed that dispersal kernels were primarily influenced by the model parameters relating to disperser behaviour, especially those determining attractiveness based on distance to candidate fruiting trees. Our results suggest that the spatial arrangement of plants plays an important role in seed dispersal processes – although tree aggregation curbed the mean seed dispersal distance, it was accompanied by occasional long distance events, and tree dispersion caused an increase in mean dispersal distance, both potentially increasing the probability of seeds finding suitable habitats for germination and growth. Even though low frugivore densities did not cause dispersal failure, there were negative effects on the quantity of seed dispersal because fewer seeds were dispersed.     

Tree invasion and establishment in old fields at Hutcheson Memorial Forest

In this review I present results from pattern analysis and field experiments in New Jersey, United States concerning tree invasion and establishment in old fields. Hutcheson Memorial Forest Center (HMF) is the New Jersey study site and a logical choice to anchor a review of this topic. HMF contains the longest and largest old field permanent plot study in the USA including extensive data concerning tree invasion and establishment. HMF has also been the site of considerable old field tree experimental research. In addition, I review other investigations of old field tree invasion and establishment of the eastern deciduous biome and contrast them with HMF. I identify critical questions concerning tree invasion and establishment, indicate what progress has been made towards answering these questions and propose future experimentation. I also synthesize results into a conceptual model and investigate the relative importance of seed and seedling processes in determining tree dynamics in old fields. The idea of a possibility or “window” for tree invasion and establishment and its variation form the basis for this review. For example, the temporal variation of windows is reflected in tree establishment patterns at HMF because wind- and bird-dispersed species invaded one year after cessation of cultivation while mammal-dispersed species were not present until year 12. Although analysis of size classes of individual HMF trees imply that windows close quickly, studies are of too short duration to tell definitely. Spatial variation in invasion and establishment pattern includes exponential decline in seed and stem density with distance from forest edge which deteriorates with time as the old field becomes more structurally complex. Additional spatial heterogeneity occurs when seeds and seedlings of bird-dispersed species clump which conversely becomes more pronounced with time. Many of these HMF patterns are seen in other eastern United States old field sites. When reviewing field experiments from my site and others, I focus on the spatial and temporal variation and the difference between species in the effect of processes on tree seeds and seedlings. I do this because this variation relates directly to the dynamic nature of windows whose quantification is necessary for modeling. Combining my pattern analysis with a survey of variation in seed and seedling mortality caused by different successional processes in old fields, I conclude that seed dispersal, conditions necessary for seed germination and seed and seedling predation are the critical processes controlling and constraining tree invasion and establishment in old fields. Competition and indirect effects of litter and the environment, which modify how these processes affect tree dynamics, are also important. Therefore, processes involving the interaction of species with the environment and with other species form the principal filters on and limitations to tree invasion and establishment and not the non-interactive life-history traits of individual tree species such as growth rate or longevity. The interaction of pattern and process and the affect of many of these processes on old field community structure are also discussed.     

Seed dispersal potential by wild mallard duck as estimated from digestive tract analysis

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Seed Dispersal by Bats and Birds in Forest and Disturbed Habitats of Chiapas,Mexico1

We examined seed dispersal by bats and birds in four habitats of the Selva Lacandona tropical rain forest region, Chiapas, Mexico. The four habitats represented a disturbance gradient: active cornfield, ten-year-old abandoned cornfield, cacao plantation, and forest. Using seed traps examined before sunrise (0400 h) and before sunset (1800 h), we compared volant vertebrate seed dispersal, assuming that seeds found at the end of the night were dispersed by bats and those found at the end of the day were dispersed by birds. We did not find seeds from other frugivores such as monkeys or opossums. In all habitats bats dispersed more seeds than birds. In most months bats also dispersed more seeds than birds, except in December when no seeds were found in the traps. Bats also consistently dispersed more species of seeds than birds, although a x2 comparison showed differences not to be significant. Fifty percent of the species represented in the dispersed seeds in all habitats were pioneer species. Cecropia seeds represented a high percentage (up to 87% of those dispersed by bats and up to 83% by birds) of dispersed seeds that fell in our traps. The influence of bats and birds on secondary successional processes is likely to be fundamental for the establishment of vegetation. Since bats dispersed more seeds than birds (primarily to disturbed areas and consisting primarily of pioneer species), they are likely to play an important role in successional and restoration processes among habitats as structurally and vegetationally different as cornfields, old fields, cacao plantations, and forest.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Seed dispersal and seedling recruitment of trees at different successional stages in a temperate forest in northeastern China

Aims Spatial distribution of adult trees in a forest community is determined by patterns of both seed dispersal and seedling recruitment. The objectives of our study were to understand the processes of seed dispersal and seedling recruitment of dominant tree species in a temperate forest of northeastern China and to identify the factors constraining seed dispersal and seedling establishment at different stages of forest succession.Methods During three summer and autumn sessions between 2006 and 2008, altogether 113080 seeds from 22 different tree species were collected in three large field plots representing different forest types in the Changbai Mountain region of northeastern China. The spatial distribution of seed abundance was analyzed using a Syrjala test. Regeneration success of nine major tree species was assessed using variables defining 'limitations' in 'seeds' and 'seedling establishment'.Important findings We found that seed production fluctuated between years and varied greatly with forest types. Four tree species, Acer spp., Fraxinus mandshurica, Tilia amurensis and Betula spp., had the greatest seed production and the widest range of seed dispersal, whereas Quercus mongolica showed the most sustained seed production pattern. The spatial patterns of seed abundance differed significantly among forest types and years. The tree species investigated in this study differed in the degree of seed limitation, as well as in limitation of seedling establishment. There were both negative and positive correlations between seed density and seedling density, depending on site and parental tree density. Seeds of 16 tree species were found in the Populus davidiana–Betula platyphylla forest (PBF) plot, 11 in the conifer and broad-leaved mixed forest (CBF) plot but only 8 in the broad-leaved-Korean pine mixed forest (BKF) plot. The number of seed-contributing species was not only greater in the secondary forests (CBF and PBF plots) than in the primary forest (BKF plot) but was also more variable during the 3 years of assessment. Results from the correlations between seed density and seedling occurrence and that between parental tree density or seed weight and dispersal limitation confirm our intuitive expectations, i.e. heavy seeds had greater dispersal limitation but higher establishment success than light seeds.     

Microsatellite analysis of seed dispersal and parentage of saplings in bur oak, Quercus macrocarpa

Microsatellite analysis was used to examine parentage and spatial distributions of 62 adult bur oaks Quercus macrocarpa, and 100 saplings in a single stand. Using genotypes scored by PCR products at four microsatellite loci, we determined that 94 saplings matched at least one parent in the stand. Saplings often occur as dense clusters of half-sibs around the presumed maternal parent, and only four adults were seed parents to a large proportion of the saplings sampled. A stump apparently was the seed parent of the largest cluster of half-sibs, which occupied a sizeable light gap opened up by the death of their maternal tree. Approximately half of the saplings appeared to have grown from seeds that had not been removed after falling from the tree, and half from seeds that were dispersed beyond the crown of their maternal parent. Long-distance seed dispersal may be more common than has been previously reported. Extremely high levels of long-distance pollination were indicated, and pollen donors within the stand were generally distributed randomly around maternal trees. More than half of the saplings had paternal parents outside of the stand. This study demonstrates the utility of microsatellite analysis for studying mating systems, seed dispersal and seedling establishment in natural plant populations.     

Evolution of recruitment features in plants: a comparative study of species in the Rosaceae

Theory predicts that life history traits that reduce the impact of environmental variation show patterns of negative covariation (trade‐offs). In plants, seed size, seed dormancy and dispersal (in space) interact to reduce risk in a temporally and spatially variable environment. Dispersal in time and space permits escape from unfavourable conditions, whereas a large seed size may improve establishment under unfavourable conditions. However, large seeds may suffer a high rate of loss due to seed predators. The objective of this study was to examine relationships between seed size, seedling establishment and seed dormancy by combining data from field experiments with information of phylogenetic relationships among 11 species in the subfamily Rosoideae (Rosaceae). The predicted relationship between seed size and seed predation was also examined. Emergence from seed, survival and recruitment of seedlings, effects of seed predators (post‐dispersal predation), and seed dormancy were studied experimentally in the field. All species generated seedlings in the experimental plots. Overall, the emergence of seedlings was promoted by disturbance. Several species possessed seeds that were able to germinate after being buried in the soil column for three years. Despite a small data set (ten phylogenetically independent contrasts), the comparative analysis confirmed an expected positive effect of seed size on establishment ability. The emergence of seedlings increased with seed size. The results did not demonstrate the hypothesised positive association between seed size and predation risk, or negative association between seed size and seed dormancy. However, the contrast analysis showed a significant negative relationship between seed dormancy and survivorship of seedlings, i.e. between quantitative data of dormancy and establishment capacity of the species obtained under natural conditions in the field. When not controlling for phylogeny (without contrasts), the only significant effect was a negative association between seed size and seed dormancy.     

What are the consequences of ant–seed interactions on the abundance of two dry-fruited shrubs in a Mediterranean scrub?

Strong interactions between dry-fruited shrubs and seed-harvesting ants are expected in early successional scrubs, where both groups have a major presence. We have analysed the implications of the seed characteristics of two dry-fruited shrub species (Coronilla minima and Dorycnium pentaphyllum) on seed predation and dispersal mediated by harvester ants and the consequences of these processes on spatio-temporal patterns of plant abundance in a heterogeneous environment. We found that large C. minima seeds were collected much more (39%) than small D. pentaphyllum seeds (2%). However, not all of the removed seeds of these plant species were consumed, and 12.8% of the seeds were lost along the trails, which increased dispersal distances compared with abiotic dispersal alone. Seed dropping occurred among all microhabitats of the two plant species, but especially in open microhabitats, which are the most suitable ones for plant establishment. The two plant species increased their presence in the study area during the study period: C. minima in open microhabitats and D. pentaphyllum in high vegetation. The large size of C. minima seeds probably limited the primary seed dispersal of this species, but may have allowed strong interaction with ants. Thus, seed dispersal by ants resulted in C. minima seeds reaching more suitable microhabitats by means of increasing dispersal distance and redistribution among microhabitats. In contrast, the smaller size of D. pentaphyllum seeds arguably allows abiotic seed dispersal over longer distances and colonization of all types of microhabitats, although it probably also limits their interaction with ants and, consequently, their redistribution in suitable microhabitats. We suggest that dyszoochory could contribute to the success of plant species with different seed characteristics in scrub habitats where seeds are abundantly collected by seed-harvesting ants.     

Chimpanzee seed dispersal quantity in a tropical montane forest of Rwanda

We describe chimpanzee seed dispersal in the tropical montane forest of Nyungwe National Park (NNP), Rwanda, for a total of three years from January 1998 through May 2000 and May 2006 through March 2007. Relatively few studies have examined chimpanzee seed dispersal in montane communities where there are generally fewer fruiting tree species than in lowland forests. Such studies may reveal new insights into chimpanzee seed dispersal behaviors and the role that they play in forest regeneration processes. Chimpanzees are large‐bodied, highly frugivorous, and tend to deposit the seeds of both large‐ and small‐seeded fruits they consume in a viable state. We found that chimpanzees dispersed a total of 37 fruiting species (20 families) in their feces, 35% of which were large‐seeded trees (≥0.5 cm). A single large‐seeded tree, Syzygium guineense, was the only species to be dispersed in both wadges and feces. Based on phenological patterns of the top five large‐seeded tree species found in chimpanzee feces, our results indicate that chimpanzees do not choose fruits based on their availability. There was, however, a positive relationship between the presence of Ekebergia capensis seeds in chimpanzee feces and S. guineense seeds in chimpanzee wadges and their respective fruit availabilities. Our data reveal that proportionately fewer chimpanzee fecal samples at NNP contained seeds than that reported in two other communities in the Albertine Rift including one at mid‐elevation and one in montane forest. As in other chimpanzee communities, seeds of Ficus spp. were the most common genus in NNP chimpanzee feces. Our data do not support previous studies that describe Ficus spp. as a fallback food for chimpanzees and highlights an intriguing relationship between chimpanzees and the large‐seeded tree species, S. guineense. Am. J. Primatol. 71:901–911, 2009. © 2009 Wiley‐Liss, Inc.     

Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

Unravelling seed dispersal through fragmented landscapes: Frugivore species operate unevenly as mobile links

Seed dispersal constitutes a pivotal process in an increasingly fragmented world, promoting population connectivity, colonization and range shifts in plants. Unveiling how multiple frugivore species disperse seeds through fragmented landscapes, operating as mobile links, has remained elusive owing to methodological constraints for monitoring seed dispersal events. We combine for the first time DNA barcoding and DNA microsatellites to identify, respectively, the frugivore species and the source trees of animal‐dispersed seeds in forest and matrix of a fragmented landscape. We found a high functional complementarity among frugivores in terms of seed deposition at different habitats (forest vs. matrix), perches (isolated trees vs. electricity pylons) and matrix sectors (close vs. far from the forest edge), cross‐habitat seed fluxes, dispersal distances and canopy‐cover dependency. Seed rain at the landscape‐scale, from forest to distant matrix sectors, was characterized by turnovers in the contribution of frugivores and source‐tree habitats: open‐habitat frugivores replaced forest‐dependent frugivores, whereas matrix trees replaced forest trees. As a result of such turnovers, the magnitude of seed rain was evenly distributed between habitats and landscape sectors. We thus uncover key mechanisms behind “biodiversity–ecosystem function” relationships, in this case, the relationship between frugivore diversity and landscape‐scale seed dispersal. Our results reveal the importance of open‐habitat frugivores, isolated fruiting trees and anthropogenic perching sites (infrastructures) in generating seed dispersal events far from the remnant forest, highlighting their potential to drive regeneration dynamics through the matrix. This study helps to broaden the “mobile‐link” concept in seed dispersal studies by providing a comprehensive and integrative view of the way in which multiple frugivore species disseminate seeds through real‐world landscapes.     

Implications of waterbird ecology for the dispersal of aquatic organisms

In this paper, we review some potential implications of waterbird ecology for their role as dispersers of aquatic plants and invertebrates. We focus particularly on internal transport (endozoochory) by the Anatidae (mainly ducks) and shorebirds, groups especially important for dispersal processes owing to their abundance, migratory habitats and diets. We conduct a literature review to assess the seasonal patterns shown by Anatidae in consumption of seeds and plankton, the interspecific patterns in such consumption (including the effects of body size, bill morphology, etc.), and differences in habitat use (e.g., shoreline vs. open water specialists) and migration patterns between species (e.g., true migrants vs. nomads). We show that many shorebirds are important consumers of seeds as well as plankton, and suggest that their role in plant dispersal has been underestimated. This review confirms that Anatidae, shorebirds and other waterbirds have great potential as dispersers of aquatic organisms, but illustrates how closely related, sympatric bird species can have very different roles in dispersal of specific aquatic organisms. Furthermore, great spatial and temporal variation is likely in dispersal patterns realized by a given bird population. We present evidence suggesting that northbound dispersal of aquatic propagules by endozoochory during spring migration is a frequent process in the northern hemisphere. Much more systematic fieldwork and reanalysis of the existing data sets (e.g., from diet studies) are needed before the relative roles of various waterbird species as dispersers can be fully assessed.     

Small traits with big consequences: how seed traits of nitrogen‐fixing plants might influence ecosystem nutrient cycling

Symbiotic nitrogen (N)‐fixing plants have important effects on the biogeochemical processes of the sites they inhabit, but their ability to reach these sites is determined by the dispersal of their seeds. Differences in seed size and dispersal vectors of N‐fixing and non‐fixing plants could influence the spatial and temporal distributions of N fixers, and thus could have important impacts on biogeochemical cycling. Using seed mass, dispersal vector, and biome data retrieved from online public databases, we ask if there are systematic differences in seed mass and dispersal vectors between N‐fixing and non‐fixing plants. We demonstrate that rhizobial N fixers tend to have larger seeds that are more likely to be biotically dispersed than seeds of non‐fixers, whereas actinorhizal N‐fixing trees tend to have small, abiotically dispersed seeds. We then synthesize existing evidence from the literature to draw links between these dispersal traits and the spatio–temporal patterns of N fixers, as well as their biogeochemical effects on terrestrial ecosystems. Using this literature, we argue that the spatio–temporal distributions of N fixers are influenced by their seed dispersal characteristics, and that these distribution patterns have important effects on the total amount of N fixed at a site and the timing of N inputs during processes such as succession.     

The effects of host-plant distribution and local abundance on the species richness of agromyzid flies attacking British umbellifers

Abstract. 1. The number of agromyzid species (Diptera: Agromyzidae) attacking British Umbelliferae generally increases with the size of the geographic range of the host, measured as occupied 10 km squares in the Atlas of the British Flora (Lawton & Price, 1979). 2. In the present study we tried to explain the large, residual variation in this species—area relationship using two new variables, namely the local abundance of the host plant, and the number of habitats within which it grows. 3. Local abundance was estimated from eight floras that map plant distributions within English countries by tetrads (2 times 2 km squares). Local abundance was defined as: Total number of occupied tetrads Total number of available tetrads within occupied 10 km squares 4. The number of habitats occupied by each host plant was taken from the only county flora to record plant habitats objectively, that for Warwickshire. 5. We expected to find a correlation between local abundance and the residuals from the national species—area relationship, with locally scarce plants having fewer agromyzids than expected from the sizes of their national ranges, and vice versa. 6. What we found was that size of geographic range and local abundance were highly correlated; hence their relative contributions to agromyzid species richness were difficult to disentangle. Residuals from the national species—area relationship were positively correlated with local abundance, but the relationship marginally failed to achieve statistical significance (P= 0.06). 7. In contrast, the number of habitats occupied by each species of umbellifer in Warwickshire had a marked effect upon agrornyzid species richness, with plants that grow in more habitats supporting more species of insects. Not surprisingly, local abundance and number of habitats occupied were highly correlated. 8. Lawton & Price's observation that aquatic umbellifers are faunally impoverished now emerges as part of the general effect of number of habitats occupied by the host plants on agromyzid species richness. 9. Once the number of habitats occupied by each host plant in Warwickshire was entered into a multiple regression, the effect of size of host geographic range on agromyzid species richness was no longer statistically significant. 10. A combination of the number of habitats occupied, and leaf-form of the host (the latter taken from Lawton & Price, 1979), explains 61% of the variation in agromyzid species richness on British Umbelliferae.     

Scatterhoarding of Manchurian walnut <Emphasis Type="Italic">Juglans mandshurica</Emphasis> by small mammals: response to seed familiarity and seed size

Dispersal patterns can be affected by seed familiarity and seed traits, including size, mass, and nutritional value, but these factors have not been intensively studied in the context of seed dispersal processes. Our aim was to study how small rodents respond to seed size and seed familiarity in their pattern of Manchurian walnut (Juglans mandshurica) seeds in two different habitats in temperate forests of northeast China. Our results demonstrated that Apodemus penisulae acts as the most important disperser for Manchurian walnut seeds. Inexperienced small rodents did not reject seeds of the Manchurian walnut and show similar seed removal rates as compared with experienced rodents. Both experienced and naïve rodents actively participated in seed scatterhoarding of Manchurian walnut seeds. Consecutive survey showed that seeds with large size/mass were removed faster than those with small size/mass, indicating a preference for large seeds. However, small seeds scatter-hoarded by small rodents were transported farther than large ones, failing to support the traditional optimization models for various tree species. Small seeds of Manchurian walnut in caches were less likely to be recovered than large ones and showed greater cache survival rates, indicating that small seeds would be more advantageous for regeneration than large seeds in small rodent-dominated forests.     

Tree recovery and seed dispersal by birds: Comparing forest, agroforestry and abandoned agroforestry in coastal Ecuador

We used a highly replicated study to examine vegetation characteristics between patches of intervened forest, abandoned agroforestry systems with coffee and actively managed agroforestry systems with coffee in a tropical landscape. In all habitats, plant structural characteristics, individual abundance, species richness and composition were recorded for the three plant size classes: adult trees, saplings and seedlings. Furthermore, bird species richness and composition, and seeds dispersed by birds were recorded. Tree abundance was higher in forest habitats while saplings and seedlings were more abundant in abandoned coffee sites. Although species richness of adult trees was similar in the three habitats, species richness of saplings and seedlings was much higher in forest and abandoned coffee than in managed coffee sites. However, in spite of their relatively low species richness, managed coffee sites are an important refuge for tree species common to the almost disappeared mature forest in the area. Floristic similarity for adult trees was relatively low between land use types, but clearly higher for seedlings, indicating homogenizing processes at the landscape level. More than half of the saplings and seedling were not represented by adults in the canopy layer, suggesting the importance of seed dispersal by birds between habitats. Our results show that each of the studied ecosystems plays a unique and complementary role as seed source and as habitat for tree recovery and tree diversity.     

Extremely long-distance seed dispersal by an overfished Amazonian frugivore

Throughout Amazonia, overfishing has decimated populations of fruit-eating fishes, especially the large-bodied characid, Colossoma macropomum. During lengthy annual floods, frugivorous fishes enter vast Amazonian floodplains, consume massive quantities of fallen fruits and egest viable seeds. Many tree and liana species are clearly specialized for icthyochory, and seed dispersal by fish may be crucial for the maintenance of Amazonian wetland forests. Unlike frugivorous mammals and birds, little is known about seed dispersal effectiveness of fishes. Extensive mobility of frugivorous fish could result in extremely effective, multi-directional, long-distance seed dispersal. Over three annual flood seasons, we tracked fine-scale movement patterns and habitat use of wild Colossoma, and seed retention in the digestive tracts of captive individuals. Our mechanistic model predicts that Colossoma disperses seeds extremely long distances to favourable habitats. Modelled mean dispersal distances of 337-552 m and maximum of 5495 m are among the longest ever reported. At least 5 per cent of seeds are predicted to disperse 1700-2110 m, farther than dispersal by almost all other frugivores reported in the literature. Additionally, seed dispersal distances increased with fish size, but overfishing has biased Colossoma populations to smaller individuals. Thus, overexploitation probably disrupts an ancient coevolutionary relationship between Colossoma and Amazonian plants.