Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.     

Mammalian seed dispersal in Cantabrian woodland pastures: Network structure and response to forest loss

We evaluated the role of wild large mammals as dispersers of fleshy-fruited woody plants in woodland pastures of the Cantabrian range (N Spain). By searching for seeds in mammal scats across four localities, we addressed how extensive seed dispersal was in relation to the fleshy-fruited plant community, and applied a network approach to identify the relative role of mammal species in the seed dispersal process. We also tested the response of mammalian dispersers to forest availability at increasing spatial scales. Five carnivores and three ungulates dispersed seeds of eight fleshy-fruited trees and shrubs. Mammalian seed dispersal did not mirror community-wide fruit availability, as abundant fruiting trees were scarce whereas thorny shrubs were over-represented among dispersed species. The dispersal network was dominated by bramble (Rubus ulmifolius/fruticosus), the remaining plants being rarer and showing more restricted disperser coteries. Fox (Vulpes vulpes), badger (Meles meles), and wild boar (Sus scrofa) dispersed mostly bramble, whereas martens (Martes sp.) dispersed mostly wild rose (Rosa sp.). Ungulates occasionally dispersed holly (Ilex aquifolium) and hawthorn (Crataegus monogyna). The empirical network reflected a skewed distribution of interactions and some functional complementarity (as judged from the low levels of connectance and nestedness), but also some degree of specialization. Mammals overused uncovered microsites for seed deposition, and increased their disperser activity in those landscape sectors devoid of forest. Combined with previous findings on avian seed dispersal, this study suggest a strong functional complementarity coming from the low overlap in the main plant types that mammals and birds disperse – thorny shrubs and trees, respectively – and the differential patterns of seed deposition, with mammals mostly dispersing into deforested areas, and birds into forest-rich landscapes.     

Seed Dispersal Patterns by Large Frugivorous Mammals in a Degraded Mosaic Landscape

Seed dispersal by Red fox (Vulpes vulpes), Stone marten (Martes foina), and Wild boar (Sus scrofa) was analyzed in an extensively degraded mosaic landscape in Sierra Nevada (SE Spain). The main objective was to determine whether seed dispersal by mammals was related to habitat degradation within a mosaic of adjacent degraded patches mixed with native forest and thereby to determine the potential role of mammals as seed dispersers in degraded landscape units. For three consecutive years, mammal feces were collected in the fruit production period, extracting all seeds of woody species found therein and analyzing their viability. Feces were collected in three different plots for each of five different landscape units: shrubland, native forest, and dense, cleared, and fenced reforestation stands. Seeds from 16 woody species (which represent more than a half of the total fleshy‐fruited woody species available) were recorded, although some agrarian species are also introduced in a low percentage of the scats. Seeds showed a high viability rate for all dispersed species, irrespective of the mammal disperser. No differences in species composition appeared in the overall landscape units or in the seed density between degraded habitats. Due to the small patch size, the high viability of dispersed seeds, and the large home range of the large mammals, these three animal species act as efficient seed dispersers for a diverse assemblage of woody plant species regardless of the habitat type within this degradation framework. This fact has important consequences for the biodiversity recuperation in these degraded habitats, principally in pine plantations.     

Gibbons (Nomascus gabriellae) provide key seed dispersal for the Pacific walnut (Dracontomelon dao), in Asia's lowland tropical forest

Understanding the mutualisms between frugivores and plants is essential for developing successful forest management and conservation strategies, especially in tropical rainforests where the majority of plants are dispersed by animals. Gibbons are among the most effective seed dispersers in South East Asia's tropical forests, but are also one of the highly threatened arboreal mammals in the region. Here we studied the seed dispersal of the Pacific walnut (Dracontomelon dao), a canopy tree which produces fruit that are common in the diet of the endangered southern yellow-cheeked crested gibbon (Nomascus gabriellae). We found that gibbons were the most effective disperser for this species; they consumed approximately 45% of the fruit crop, which was four times more than that consumed by macaques – the only other legitimate disperser. Gibbons tracked the temporal (but not spatial) abundance of ripe fruits, indicating this fruit was a preferred species for the gibbon. Both gibbons and macaques dispersed the majority (>90%) of the seeds at least 20 m away from parent crowns, with mean dispersal distances by gibbons measuring 179.3 ± 98.0 m (range: 4–425 m). Seeds defecated by gibbons germinated quicker and at greater rates than seeds spat by macaques, or in undispersed fruits. Gibbon-dispersed seeds were also more likely to be removed by unknown seed predators or unknown secondary dispersers. Overall, gibbons play a key role in the regeneration of the Pacific walnut. Our findings have significant implications both for the management of the Pacific walnut tree dominating tropical rainforest as well as the reintroduction program of the Southern yellow-cheeked crested gibbon.     

Non-redundancy in seed dispersal and germination by native and introduced frugivorous birds: implications of invasive bird impact on native plant communities

Seed dispersal by vertebrate animals is important for the establishment of many fleshy-fruited plant species. Different frugivorous species can provide different seed dispersal services according to their specific dietary preferences as well as behaviour and body traits (e.g. body size and beak size of birds). Our aim was to study redundancies and complementarities in seed dispersal and germination between the two main native seed disperser birds and the introduced silver pheasant Lophura nycthemera in the temperate Patagonian forests. For this, we collected fresh droppings from the studied species and analyzed seed content. We conducted germination trials for four plant species common in bird droppings; two native species (Aristotelia chilensis and Rhaphithamnus spinosus) and two invasive non-native species (Rubus ulmifolius and Rosa rubiginosa). Both native frugivorous birds and the silver pheasant dispersed fruits of non- native fleshy-fruited plants, but their roles were non-redundant in terms of species dispersed and effect on seed germination. The silver pheasant dispersed a proportionally high number of non-native seeds, while native birds dispersed a high number of native seeds. In addition, the effect of gut treatment in seed germination differed between seed dispersers. Native birds promoted the germination for the two native plant species studied, while the silver pheasant promoted the germination of one non-native plant. This suggests that seed dispersal by the silver pheasant may contribute to the spread of some invasive fleshy-fruited plants in the ecosystems that otherwise would not be dispersed by any other bird. The understanding of redundancies and complementarities on seed dispersal and germination between native and introduced birds will allow improving the management of fleshy-fruited non-native plants.     

Seed dispersal of Diospyros virginiana in the past and the present: Evidence for a generalist evolutionary strategy

Several North American trees are hypothesized to have lost their co‐evolved seed disperser during the late‐Pleistocene extinction and are therefore considered anachronistic. We tested this hypothesis for the American persimmon (Diospyros virginiana) by studying the effects of gut passage of proposed seed dispersers on seedling survival and growth, natural fruiting characteristics, and modern animal consumption patterns. We tested gut passage effects on persimmon seeds using three native living species, the raccoon (Procyon lotor), Virginia opossum (Didelphis virginiana), and coyote (Canis latrans), and two Pleistocene analogs; the Asian elephant (Elephas maximus) and alpaca (Vicugna pacos). Persimmon seeds excreted by raccoons, coyotes, and elephants survived gut transit. Gut passage did not affect sprouting success, but did tend to decrease time to sprout and increase seedling quality. Under field conditions, persimmon fruits were palatable on the parent tree and on the ground for an equal duration, but most fruits were consumed on the ground. Seven vertebrate species fed upon persimmon fruits, with the white‐tailed deer (Odocoileus virginianus)—a species not capable of dispersing persimmon seeds—comprising over 90% of detections. Conversely, potential living seed dispersers were rarely detected. Our results suggest the American persimmon evolved to attract a variety of seed dispersers and thus is not anachronistic. However, human‐induced changes in mammal communities could be affecting successful seed dispersal. We argue that changes in the relative abundance of mammals during the Anthropocene may be modifying seed dispersal patterns, leading to potential changes in forest community composition.     

Persistence of the effect of frugivore identity on post‐dispersal seed fate: consequences for the assessment of functional redundancy

Large frugivores play an important role as seed dispersers and their extinction may affect plant regeneration. The consequences of such extinctions depend on the likelihood of other species being functionally redundant and on how post‐dispersal events are affected. We assess the functional redundancy of two seed dispersers of the Atlantic Forest, the muriqui (Brachyteles arachnoides) and the tapir (Tapirus terrestris) through the comparison of their seed dispersal quality, taking into account post‐dispersal events. We compare tapirs and muriquis for: (1) the dung beetle community associated with their feces; (2) the seed burial probability and burial depth by dung beetles; and (3) the seed mortality due to predators or other causes according to burial depth. We determine how seed burial affects seed dispersal effectiveness (SDE) and compare the dispersal quality of four plant species dispersed by these frugivores. Muriqui feces attract 16‐fold more dung beetles per gram of fecal matter and seeds experience 10.5‐fold more burial than seeds in tapir feces. In both feces types, seed mortality due to predation decreases with burial depth but seed mortality due to other causes increases. Total seed mortality differ within plant species according to the primary disperser. Therefore, the effect of seed burial on SDE varies according to the plant species, burial depth, and primary disperser. As tapirs and muriquis differently affect the seed fate, they are not functionally redundant. Since the effect of the primary disperser persists into the post‐dispersal events, we should consider the cascading effects of these processes when assessing functional redundancy.     

Podarcis lilfordi from the Balearic islands as a potential disperser of the rare Mediterranean plant Withania frutescens

This study examines whether the lizard Podarcis lilfordi is a legitimate disperser of the rare Mediterranean plant Withania frutescens by using the biochemical test of triphenyl-2H-tetrazolium chloride for testing seed viability. This lizard eats the fresh fruits of the plants and defecates intact seeds which have been retained 1 to 3 d in their gut. Viability of seeds recovered from faeces was very high and comparable to the viability of fresh seeds. Seed dispersal by this lizard in the Balearic islands may facilitate population expansion of this rare plant in the Balearics.     

Integration of exotic seeds into an Azorean seed dispersal network

Seed dispersal plays a central role in plant ecology. Among animals, birds are particularly important seed dispersers, often incorporating exotic plants into their diets and facilitating their integration in the communities. Network theory offers a highly informative framework to study the structural and functional attributes of complex interactions networks. We used information from bird fecal samples to build a quantitative seed dispersal network for the last fragment of native laurel forest in the island of São Miguel—Azores with three specific objectives: (1) to assess the integration of exotic seeds into seed dispersal; (2) to evaluate the impact of exotic plants in network structure; (3) to test the potential of an exotic species to reduce the seed dispersal of a co-occurring native, via competition for seed dispersers. The seed dispersal network was based on the analysis of 1,121 droppings and described 74 unique interactions between 41 plant species and 7 bird species. Exotic seeds deeply infiltrated into the seed dispersal network forming the majority (59 %) of seeds in the droppings and including those of three globally invasive plants. Overall, birds depended equally on native and exotic fruits despite the lower abundance of the latter in the study area. In an experiment, birds did not show a preference for fruits of the exotic Leycesteria formosa over the native Vaccinium cylindraceum consuming them equally. However, the presence of the exotic plant negatively affected the number of native seeds dispersed, by diverting some of the consumers of the native fruits. Taken altogether the results reveal an alarming invasion level of seed dispersal systems in one of the last remnant native forests of the Azores.     

The role of seed dispersal,pollination and historical effects on genetic patterns of an insular plant that has lost its only seed disperser

Aim The genetic structure of many plant species is heavily dependent on their pollinators and seed dispersers, and can thus be altered if either of the associated mutualistic interactions is disrupted. In this study we assess the genetic diversity and structure and infer the seed/pollen gene‐flow patterns among insular populations of Daphne rodriguezii, a shrub pollinated and dispersed by animals that has lost its only disperser (the lizard Podarcis lilfordi) in most of its populations. Location The island of Menorca and the islet of Colom (Balearic Islands, Western Mediterranean). Methods To assess the contribution of gene flow via pollen and seeds to the genetic structure of D. rodriguezii we used amplified fragment length polymorphisms (AFLPs; seeds and pollen) and plastid DNA sequences (cpDNA; seeds). We sampled individuals from all population nuclei of the species (12–19 adults per population): one population in Colom, where the plant–lizard interaction persists, and four in Menorca, where the seed dispersal mutualism disappeared with the extinction of the lizard. Results The highest heterozygosity values were found in Colom and in its closest population (Favàritx), whereas values were lower in the smallest Menorcan populations, which also had higher relatedness among individuals. We found distinct genetic signals between AFLP and cpDNA analyses. While AFLP markers showed low differentiation between populations, cpDNA showed a clear differentiation between them. Main conclusions Our results point to negative impacts of the disperser loss on genetic diversity and relatedness in the smaller and more isolated populations. They also suggest an old isolation by seeds, probably occurring well before the extinction of the lizard (c. 2000 years ago). Gene flow was maintained via pollination; however, the seed disperser loss may ultimately hinder pollinator‐mediated gene flow, as a result of reduced probabilities of effective pollination among increasingly distant and scarce individuals.     

Planting Design Effects on Avian Seed Dispersers in a Tropical Forest Restoration Experiment

Seed dispersal often limits tropical forest regeneration and animals disperse most rainforest tree seeds. This presents two important questions for restoration ecologists: (1) which animals are common seed dispersers? and (2) which restoration techniques attract them? Fourteen restoration sites were planted with four tree species in three designs, (1) controls (no planting, natural regeneration) (2) islands (trees planted in small patches), and (3) plantations (trees planted continuously over a large patch). We sampled birds in November, February, and April 2007–2008 with mist nets, in February and July 2009 with observations, and in July 2008 with both techniques. We documented 30 seed species from fecal samples of captured birds. All identified seed species were early‐successional forms. Four tanager species, three thrushes, two saltators, two flycatchers, and one finch were categorized as common seed dispersers, based on their high likelihood of dispersing seeds. Common dispersers were generalist species with small gape widths (<15 mm). Common dispersers were captured significantly more often in plantations than controls in most seasons and more often in plantations than islands during one season. Common disperser observations were significantly greater in plantations than controls during two periods and in plantations compared with islands in one period. Results indicate that plantation‐style planting is the conservative strategy to maximize attractiveness to common dispersers in tropical restoration sites. Island planting is an alternative when resources are limited although disperser activity may be lower in some seasons than in plantations. Additional research should investigate how to attract large, forest‐associated dispersers.     

Intra-specific downsizing of frugivores affects seed germination of fleshy-fruited plant species

The loss of largest-bodied individuals within species of frugivorous animals is one of the major consequences of defaunation. The gradual disappearance of large-bodied frugivores is expected to entail a parallel deterioration in seed dispersal functionality if the remaining smaller-sized individuals are not so effective as seed dispersers. While the multiple impacts of the extinction of large bodied species have been relatively well studied, the impact of intraspecific downsizing (i.e. the extinction of large individuals within species) on seed dispersal has rarely been evaluated. Here we experimentally assessed the impact of body-size reduction in the frugivorous lizard Gallotia galloti (Lacertidae), an endemic species of the Canary Islands, on the seed germination patterns of two fleshy-fruited plant species (Rubia fruticosa and Withania aristata). Seed germination curves and the proportions of germinated seeds were compared for both plant species after being defecated by large-sized individuals and small-sized individuals. The data show that seeds of W. aristata defecated by larger-sized lizards germinated faster and in a higher percentage than those defecated by small-sized lizards, while no differences were found for R. fruticosa seeds. Our results suggest that disappearance of the largest individuals of frugivorous species may impair recruitment of some plant species by worsening seed germination. They also warn us of a potential cryptic loss of seed dispersal functionality on defaunated ecosystems, even when frugivorous species remain abundant.     

Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

Conflicting selection on Cneorum tricoccon (Rutaceae) seed size caused by native and alien seed dispersers

The disappearance of native seed dispersers due to anthropogenic activities is often accompanied by the introduction of alien species, which may to some extent replace the ecological service provided by the extinct ones. Yet, little empirical evidence exists demonstrating the evolutionary consequences of such alien “replacement.” Here, we document the conflicting selection exerted on seed size by two native lizards (Podarcis lilfordi and P. pityusensis) and an alien mammal species (Martes martes), all acting as legitimate seed dispersers of the Mediterranean relict Cneorum tricoccon. While lizards mostly exerted a negative directional selection on seed diameter, especially P. pityusensis, the much larger pine marten exerted positive selection on seed size. Our findings suggest that this among‐disperser variation in the selection regimes, together with the occurrence of spatial variation in the presence of each seed disperser, help to create the geographical variation observed for seed size of C. tricoccon. To our knowledge, this is the first empirical evidence showing opposing selective pressures between native and alien species in the seed dispersal process in an invaded ecosystem.     

Different contributions of birds and mammals to seed dispersal of a fleshy-fruited tree

Birds and mammals are important seed dispersers of fleshy-fruited plants. Although their behaviors are different, they frequently consume the same species. Thus to understand the dispersal of fleshy-fruited plants, the contribution of birds and mammals to seed dispersal has to be evaluated. Besides, within birds or mammals, some species may functionally different from others. In this study, we examined seed dispersal of the fleshy-fruited tree Swida controversa focusing on the difference between two frugivore groups (birds and mammals), and differences between species within groups. Collected seeds and S. controversa trees were identified using simple sequence repeat (SSR) genotyping, thus enabling to determine the distance between mother tree and dispersed seeds. The avian species were identified by DNA barcoding of feces, whereas the mammalian species were identified by the shape and smell of feces. Most seeds that fell near or under the maternal trees were dispersed by birds, resulting in short seed dispersal distances (average, 13 m). DNA barcoding detected five taxa of avian dispersers. No differences were detected in seed dispersal distance by different avian taxa (i.e., the distance between dispersed seeds and their maternal trees within the research plot); however the rate of seed immigration from outside the research plot by some avian taxa varied significantly. The seed dispersal distance by mammals was significantly further (127 m; min > 50 m) than that by birds. Additionally, immigrated seeds accounted for approximately two-thirds of mammal-dispersed seeds, indicating that these seeds were from outside the research plot, and that mammals significantly contributed to the long-distance seed dispersal of S. controversa. No differences in seed dispersal distance were detected between different mammalian taxa. Overall, this study revealed that birds and mammals show clearly different seed dispersal patterns, and thus, they play different roles in the regeneration of S. controversa.     

Predicting the Long-Term Effects of Hunting on Plant Species Composition and Diversity in Tropical Forests

Hunting can change abundances of vertebrate seed predators and seed dispersers, causing species‐specific changes in seed dispersal and seed predation and altering seedling communities. What are the consequences of these changes for the adult plant community in the next generation and beyond? Here, I derive equations showing how reduced seed dispersal reduces plant reproduction by intensifying kin competition, increasing vulnerability to natural enemies, and reducing the proportion of seeds passing through disperser guts. I parameterize these equations with available empirical data to estimate the likely effects on next‐generation abundances. I then consider the indirect effects and longer‐term feedbacks of changed seed or adult abundances on reproductive rates due to density‐dependent interactions with natural enemies and mutualists, as well as niche differentiation with competitors, and discuss their likely qualitative effects. The factors limiting seed disperser and seed predator populations in natural and hunted forests emerge as critical for determining the long‐term effects of hunting on rates of seed dispersal and seed predation. For example, where seed dispersers are held to a constant abundance by hunters, decreases in the availability of their preferred food plants are expected to lead to increased per‐seed dispersal probabilities, potentially to the point of compensating for the initial disperser decline. I close by discussing the likely reversibility of hunting‐induced changes in tropical forests and key questions and directions for future research.     

Introduction of mammalian seed predators and the loss of an endemic flightless bird impair seed dispersal of the New Zealand tree Elaeocarpus dentatus

Understanding the mutualistic services provided by species is critical when considering both the consequences of their loss or the benefits of their reintroduction. Like many other Pacific islands, New Zealand seed dispersal networks have been changed by both significant losses of large frugivorous birds and the introduction of invasive mammals. These changes are particularly concerning when important dispersers remain unidentified. We tested the impact of frugivore declines and invasive seed predators on seed dispersal for an endemic tree, hinau Elaeocarpus dentatus, by comparing seed dispersal and predation rates on the mainland of New Zealand with offshore sanctuary islands with higher bird and lower mammal numbers. We used cameras and seed traps to measure predation and dispersal from the ground and canopy, respectively. We found that canopy fruit handling rates (an index of dispersal quantity) were poor even on island sanctuaries (only 14% of seeds captured below parent trees on islands had passed through a bird), which suggests that hinau may be adapted for ground‐based dispersal by flightless birds. Ground‐based dispersal of hinau was low on the New Zealand mainland compared to sanctuary islands (4% of seeds dispersed on the mainland vs. 76% dispersed on islands), due to low frugivore numbers. A flightless endemic rail (Gallirallus australis) conducted the majority of ground‐based fruit removal on islands. Despite being threatened, this rail is controversial in restoration projects because of its predatory impacts on native fauna. Our study demonstrates the importance of testing which species perform important mutualistic services, rather than simply relying on logical assumptions.     

Antelope mating strategies facilitate invasion of grasslands by a woody weed

Intra and interspecific variation in frugivore behaviour can have important consequences for seed dispersal outcomes. However, most information comes from among‐species comparisons, and within‐species variation is relatively poorly understood. We examined how large intraspecific differences in the behaviour of a native disperser, blackbuck antelope Antilope cervicapra, influence dispersal of a woody invasive, Prosopis juliflora, in a grassland ecosystem. Blackbuck disperse P. juliflora seeds through their dung. In lekking blackbuck populations, males defend clustered or dispersed mating territories. Territorial male movement is restricted, and within their territories males defecate on dung‐piles. In contrast, mixed‐sex herds range over large areas and do not create dung‐piles. We expected territorial males to shape seed dispersal patterns, and seed deposition and seedling recruitment to be spatially localized. Territorial males had a disproportionately large influence on seed dispersal. Adult males removed twice as much fruit as females, and seed arrival was disproportionately high on territories. Also, because lek‐territories are clustered, seed arrival was spatially highly concentrated. Seedling recruitment was also substantially higher on territories compared with random sites, indicating that the local concentration of seeds created by territorial males continued into high local recruitment of seedlings. Territorial male behaviour may, thus, result in a distinct spatial pattern of invasion of grasslands by the woody P. juliflora. An ex situ experiment showed no beneficial effect of dung and a negative effect of light on seed germination. We conclude that large intraspecific behavioural differences within frugivore populations can result in significant variation in their effectiveness as seed dispersers. Mating strategies in a disperser could shape seed dispersal, seedling recruitment and potentially plant distribution patterns. These mating strategies may aid in the spread of invasives, such as P. juliflora, which could, in turn, negatively influence the behaviour and ecology of native dispersers.     

Dispersal of fig seeds in the Cook Islands: introduced frugivores are no substitutes for natives

Across the Pacific, island vegetation is altering in response to changes in seed disperser assemblages brought about by extinctions and introductions of birds and other animals. On the Cook Islands in the South Pacific, the Pacific Banyan (Ficus prolixa, Moraceae) is undergoing little if any recruitment, possibly linked to a lack of dispersal agents. On Rarotonga, where F. prolixa is found in semi-urban and agricultural environments, there is no recent recruitment in contrast to the situation on Atiu where the tree is common in native forest. We examined the quality and quantity of seed dispersal offered to F. prolixa by the available frugivores on these islands, comparing the effectiveness of extant native and introduced species. The native Cook Islands fauna, particularly birds and bats, appear to be the most effective seed dispersers of F. prolixa, both in terms of quantity and quality. Whilst these are relatively numerous on Atiu, they rarely visit F. prolixa on Rarotonga. The native Chocolate hermit crab Coenobita cavipes is a previously unreported additional native seed disperser, conferring low quantity, but high quality dispersal. Introduced birds and mammals are the most numerous F. prolixa frugivores on Rarotonga and in non-forest environments on Atiu, but they act mainly as seed predators. Consequently, the losses and rarity of remaining native frugivores have not been compensated for by introduced species on Rarotonga which may be contributing to the absence of recruitment there.     

The Bruce effect revisited: is pregnancy termination in female rodents an adaptation to ensure breeding success after male turnover in low densities?

Animal-mediated seed dispersal (zoochory) is considered to be an important mechanism regulating biological processes at larger spatial scales. To date, intra-specific variation in seed disperser quality within seed-dispersing animals has not been studied. Here, I employed seed feeding trials to quantify individual differences in disperser quality within the common carp (Cyprinus carpio) using seeds of two aquatic plants: unbranched bur-reed (Sparganium emersum, Sparganiaceae) and arrowhead (Sagittaria sagittifolia, Alismataceae). I found substantial variation among carp individuals in their propensity to ingest seeds and their ability to digest them, resulting in up to 31-fold differences in the probability of seed dispersal. In addition, there were significant differences in the time that seeds are retained in their digestive systems, generating a twofold difference in the maximum distance over which they can potentially disperse seeds. I propose that seed-eating animal species consist of individuals that display continuous variation in disperser quality, with at one end of the continuum individuals that are likely to eat seeds, pass them unharmed through their digestive tract and transport them over large distances to new locations (i.e. high-quality seed dispersers) and at the other end individuals that rarely eat seeds, destroy most of the ones they ingest and transport the few surviving seeds over relatively short distances (low-quality seed dispersers). Although individual differences in seed dispersal quality could be the result of a variety of factors, these results underline the ecological and evolutionary potential of such variation for both plants and animals.