Potential for ants and vertebrate predators to shape seed-dispersal dynamics of the invasive thistles <Emphasis Type="Italic">Cirsium arvense</Emphasis> and <Emphasis Type="Italic">Carduus nutans</Emphasis> in their introduced range (North America)

The population dynamics of invasive plants are influenced by positive and negative associations formed with members of the fauna present in the introduced range. For example, mutualistic associations formed with pollinators or seed dispersers may facilitate invasion, but reduced fitness from attack by native herbivores can also suppress it. Since population expansion depends on effective seed dispersal, interactions with seed dispersers and predators in a plant species introduced range may be of particular importance. We explored the relative contributions of potential seed dispersers (ants) and vertebrate predators (rodents and birds) to seed removal of two diplochorous (i.e., wind- and ant-dispersed), invasive thistles, Cirsium arvense and Carduus nutans, in Colorado, USA. We also conducted behavior trials to explore the potential of different ant species to disperse seeds, and we quantified which potential ant dispersers were prevalent at our study locations. Both ants and vertebrate predators removed significant amounts of C. arvense and C. nutans seed, with the relative proportion of seed removed by each guild varying by location. The behavior trials revealed clear seed preferences among three ant species as well as differences in the foragers’ abilities to move seeds. In addition, two ant species that acted as potential dispersal agents were dominant at the study locations. Since local conditions in part determined whether dispersers or predators removed more seed, it is possible that some thistle populations benefit from a net dispersal effect, while others suffer proportionally more predation. Additionally, because the effectiveness of potential ant dispersers is taxon-specific, changes in ant community composition could affect the seed-dispersal dynamics of these thistles. Until now, most studies describing dispersal dynamics in C. arvense and C. nutans have focused on primary dispersal by wind or pre-dispersal seed predation by insects. Our findings suggest that animal-mediated dispersal and post-dispersal seed predation deserve further consideration.     

White‐tailed deer as the last megafauna dispersing seeds in Neotropical dry forests: the role of fruit and seed traits

Endozoochory is a prominent form of seed dispersal in tropical dry forests. Most extant megafauna that perform such seed dispersal are ungulates, which can also be seed predators. White‐tailed deer (Odocoileus virginianus) is one of the last extant megafauna of Neotropical dry forests, but whether it serves as a legitimate seed disperser is poorly understood. We studied seed dispersal patterns and germination after white‐tailed deer gut passage in a tropical dry forest in southwest Ecuador. Over 23 mo, we recorded ca 2000 seeds of 11 species in 385 fecal samples. Most seeds belonged to four species of Fabaceae: Chloroleucon mangense, Senna mollissima, Piptadenia flava, and Caesalpinia glabrata. Seeds from eight of the 11 species dispersed by white‐tailed deer germinated under controlled conditions. Ingestion did not affect germination of C. mangense and S. mollissima, whereas C. glabrata showed reduced germination. Nevertheless, the removal of fruit pulp resulting from ingestion by white‐tailed deer could have a deinhibition effect on germination due to seed release. Thus, white‐tailed deer play an important role as legitimate seed dispersers of woody species formerly considered autochorous. Our results suggest that more research is needed to fully understand the ecological and evolutionary effects of the remaining extant megafauna on plant regeneration dynamics in the dry Neotropics.     

Role of Brycinus lateralis (Teleostei: Alestidae) in dispersal and germination of Nymphaea nouchali (Angiospermae: Nymphaeaceae) seeds on a seasonal floodplain of the Okavango Delta,Botswana

Seed passage through the gut of vertebrates can be important for seed dispersal, but might influence seed viability. The ability of seeds to germinate after ingestion by seed-eating fish is important for the population dynamics of some plant species, and significant in the evolution of plant–fish interactions. Certain fish in the Okavango Delta, Botswana, are fruit- and seed-eaters and could act as seed dispersers. We sampled 14 fish species in 2013, finding Nymphaea nouchali var. caerulea seeds in the digestive tracts of eight, most commonly in the striped robber Brycinus lateralis. Seeds extracted from the gut of this species had an overall mean germination success of 11.7%. This fish species might well be a legitimate seed disperser, having a positive effect on seed dispersal from parent plants in the Okavango Delta. The current study represents one of the first investigations of the likelihood of seed dispersal by fish on the African continent.     

Rodents change acorn dispersal behaviour in response to ungulate presence

Small rodents are prominent seed predators, but they also favour plant recruitment as seed dispersers. The direct interactions of ungulates on plants are more one‐sided and negative, as they mainly reduce plant recruitment through predation on seeds and seedlings. The effects of small rodents and ungulates on plant recruitment have been considered and studied as independent episodes within plant regeneration cycles. However, ungulate–rodent interactions and their potential effects on plant regeneration have not been considered so far. A number of studies have recently documented ungulate effects on the abundance, diversity and spatial distribution of small rodents. Here, we hypothesize that ungulates may also affect rodent seed dispersal behaviour. We monitored acorn dispersal by small rodents (Mus spretus and Apodemus sylvaticus) in oak woodlands with and without exclosures for large ungulates, mainly red deer, Cervus elaphus, and wild boar, Sus scrofa. The study was carried out in a typical Mediterranean Holm oak, Quercus ilex, forest throughout the acorn fall season in 2003 and 2004. We found that, in both years, the proportion of acorns cached and not recovered in the short‐term was, on average, lower in the presence (1.4%) than in the absence (19.9%) of ungulates. Acorn dispersal distances were not affected by ungulate presence in either year. However, ungulates had an effect on the spatial distribution of dispersed seeds; rodents apparently avoided shrubs as caching sites in both years. This result was interpreted as a behavioural response to reduce the risk of cache pilferage by conspecifics, which are closely associated with shrubs in presence, but not in absence, of ungulates. Potential effects of different densities of rodents or predators were discarded, as none of them differed between the areas with and without ungulates. The present study found significant interactions between heterospecific seed and seedling consumers that had been considered as independent episodes within tree regeneration cycles. As a result of such interactions, ungulates may have negative indirect effects on oak recruitment by reducing (1) acorn caching frequency, and (2) the proportion of acorns cached under shrubs, key nurse‐plants for the establishment of Holm oak seedlings in Mediterranean areas.     

Context‐dependent fruit–frugivore interactions: partner identities and spatio‐temporal variations

Fruit–frugivore interactions are crucial for the dynamics and regeneration of most forested ecosystems. Still, we lack an understanding of the potential variation in the sign and strength of such interactions in relation to variations in the spatial and temporal ecological context. Here, we evaluated spatial (three sites) and temporal (two fruiting seasons) local variation in the sign (seed predation versus dispersal) and strength (frequency and quantity) of the interactions among six frugivorous mammals and a community of Mediterranean fleshy‐fruited shrubs. We examined mammal faecal samples and quantified frequency of seed occurrence, number of seeds per faecal sample, seed species diversity and quality of seed treatment (i.e. percentage of undamaged seeds). The frequency of seed occurrence and number of seeds per faecal sample strongly varied among dispersers, sites, seasons and fruit species. For instance, fox Vulpes vulpes faeces showed between 6 and 40 times more seeds than wild boar Sus scrofa faeces in seasons or sites in which Rubus and Juniperus seeds were dominant. However, in seasons or sites dominated by Corema seeds, wild boar faeces contained up to seven times more seeds than fox faeces. Mammalian carnivores (fox and badger, Meles meles) treated seeds gently, acting mostly as dispersers, whereas deer (Cervus elaphus and Dama dama) acted mainly as seed predators. Interestingly, rabbit Oryctolagus cuniculus acted as either mostly seed disperser or seed predator depending on the plant species. Our results indicated that the sign of fruit–frugivore interactions depended mainly on the identity of the partners. For a particular fruit–frugivore pair, however, our surrogate of interaction strength largely varied with the spatio‐temporal context (year and habitat), leading to a low specificity across the seed–frugivore network. The high spatio‐temporal variability of seed dispersal (in quantity, quality and seed diversity) by different frugivores would confer resilience against unpredictable environmental conditions, such as those typical of Mediterranean ecosystems.     

Seed Dispersers,Seed Predators,and Browsers Act Synergistically as Biotic Filters in a Mosaic Landscape

In this study, we analize the functional influence of animals on the plants they interact with in a mediterranean mountain. We hypothesise that seed dispersers, seed predators, and browsers can act as biotic filters for plant communities. We analyse the combined effects of mutualistic (seed dispersal) and antagonistic (seed predation, herbivory) animal interactions in a mosaic landscape of Mediterranean mountains, basing our results on observational and experimental field. Most of the dispersed seeds came from tree species, whereas the population of saplings was composed predominantly of zoochorous shrub species. Seed predators preferentially consumed seeds from tree species, whereas seeds from the dominant fleshy-fruited shrubs had a higher probability of escaping these predators. The same pattern was repeated among the different landscape units by browsers, since they browsed selectively and far more intensely on tree-species saplings than on the surrounding shrubs. In synthesis, our work identifies the major biotic processes that appear to be favoring a community dominated by shrubs versus trees because seed dispersers, predators, and herbivores together favored shrub dispersal and establishment versus trees.     

Sowing forests: a synthesis of seed dispersal and predation by agoutis and their influence on plant communities

Granivorous rodents have been traditionally regarded as antagonistic seed predators. Agoutis (Dasyprocta spp.), however, have also been recognized as mutualistic dispersers of plants because of their role as scatter-hoarders of seeds, especially for large-seeded species. A closer look shows that such definitions are too simplistic for these Neotropical animals because agoutis can influence plant communities not only through seed dispersal of large seeds but also through predation of small seeds and seedlings, evidencing their dual role. Herein, we summarize the literature on plant–agouti interactions, decompose agouti seed dispersal into its quantitative and qualitative components, and discuss how environmental factors and plant traits determine whether these interactions result in mutualisms or antagonisms. We also look at the role of agoutis in a community context, assessing their effectiveness as substitutes for extinct megafaunal frugivores and comparing their ecological functions to those of other extant dispersers of large seeds. We also discuss how our conclusions can be extended to the single other genus in the Dasyproctidae family (Myoprocta). Finally, we examine agoutis’ contribution to carbon stocks and summarize current conservation threats and efforts. We recorded 164 interactions between agoutis and plants, which were widespread across the plant phylogeny, confirming that agoutis are generalist frugivores. Seed mass was a main factor determining seed hoarding probability of plant species and agoutis were found to disperse larger seeds than other large-bodied frugivores. Agoutis positively contributed to carbon storage by preying upon seeds of plants with lower carbon biomass and by dispersing species with higher biomass. This synthesis of plant–agouti interactions shows that ecological services provided by agoutis to plant populations and communities go beyond seed dispersal and predation, and we identify still unanswered questions. We hope to emphasise the importance of agoutis in Neotropical forests.     

The effect of limited visibility on vigilance behaviour and speed of predator detection: implications for the conservation of granivorous passerines

Ants have been traditionally considered either as predators or dispersers of seeds, but not both. That is, ant dispersal is restricted to myrmecochorous seeds, while almost all seeds removed by seed‐harvesting ants are eaten. However, harvesting ants might be simultaneously antagonistic and mutualistic towards seeds. This study analyzes the predation–dispersal relationship between seed‐harvesting ants and seeds of Lobularia maritima, a non‐myrmechorous perennial herb, in order to disentangle the dual role of ants as dispersers and predators of L. maritima seeds. The results obtained confirm the role of harvesting ants as both predators and dispersers of the non‐myrmechorous seeds of L. maritima. The removal activity of Messor bouvieri on L. maritima seeds is very important, particularly in autumn, which is the flowering and fruiting peak of this plant. It can be estimated that harvesting ants collect more than 85% of seeds, and almost 70% of them are effectively lost to predation. However, these granivorous ants also have drawbacks as seed dispersers. There is a relatively small percent of seeds collected by ants that escape predation, either because they are dropped on the way to the nest (16.4% of seeds harvested), or because they are mistakenly rejected on the refuse pile (0.9%). Abiotic dispersal of L. maritima seeds in the absence of ants occurs over very short distances from the plant stem. As seeds dispersed by ants reach a considerably greater distance than that obtained by gravity, this might represent a real advantage for the species, because it reduces intraspecific adult competition for seedlings, which directly influences seedling survivorship. These results challenge the generalization that seed removal by ants generally leads to successful seed dispersal if done by legitimate seed dispersers, or seed loss if done by seed consumers that eat them, and confirm that harvesting ants might have a dual role as both predators and dispersers of nonmyrmechorous seeds.     

啮齿类取食的物种偏好与时空格局

通过强烈消耗土壤种子库,动物取食种子对植物种群更新和群落动态产生深远的影响。一般认为种子被食概率的空间格局取决于种子密度和离母树的距离,而环境（如地形）异质性的影响则一直没有得到足够的关注,与此相关的机制及其影响程度亦不清楚。研究设计在野外埋放种子以模拟种子扩散后的情形,监测啮齿类对种子的取食,以检验种子取食受埋藏生境、时间及动物对种子种类的偏好等因素的影响。结果表明,经过1a实验,8种落叶阔叶树种子的累计取食概率为0～48．25％,平均值为20％;山顶部位的取食概率大致是其它部位概率的3倍;埋放在凋落物层中的种子被食概率大约为埋放在土壤层中概率的2倍。利用logistic回归模型进行统计分析表明,种子被食概率变化的45％可以被上述因素解释。其中,物种偏好是影响种子被取食概率的首要因素,其后依次是地形、埋藏时间和深度。啮齿类明显喜好较大的种子;其取食行为在山脊部位明显较其它部位更频繁和剧烈;对埋藏种子的取食从3月份开始加剧,到7月份以后平息下来。种子埋放深度对啮齿类的取食概率有显著影响。     

How intraspecific variation in seed‐dispersing animals matters for plants

Seed dispersal by animals is a complex phenomenon, characterized by multiple mechanisms and variable outcomes. Most researchers approach this complexity by analysing context‐dependency in seed dispersal and investigating extrinsic factors that might influence interactions between plants and seed dispersers. Intrinsic traits of seed dispersers provide an alternative way of making sense of the enormous variation in seed fates. I review causes of intraspecific variability in frugivorous and granivorous animals, discuss their effects on seed dispersal, and outline likely consequences for plant populations and communities. Sources of individual variation in seed‐dispersing animals include sexual dimorphism, changes associated with growth and ageing, individual specialization, and animal personalities. Sexual dimorphism of seed‐dispersing animals influences seed fate through diverse mechanisms that range from effects caused by sex‐specific differences in body size, to influences of male versus female cognitive functions. These differences affect the type of seed treatment (e.g. dispersal versus predation), the number of dispersed seeds, distance of seed dispersal, and likelihood that seeds are left in favourable sites for seeds or seedlings. The best‐documented consequences of individual differences associated with growth and ageing involve quantity of dispersed seeds and the quality of seed treatment in the mouth and gut. Individual specialization on different resources affects the number of dispersed plant species, and therefore the connectivity and architecture of seed‐dispersal networks. Animal personalities might play an important role in shaping interactions between plants and dispersers of their seeds, yet their potential in this regard remains overlooked. In general, intraspecific variation in seed‐dispersing animals often influences plants through effects of these individual differences on the movement ecology of the dispersers. Two conditions are necessary for individual variation to exert a strong influence on seed dispersal. First, the individual differences in traits should translate into differences in crucial characteristics of seed dispersal. Second, individual variation is more likely to be important when the proportions of particular types of individuals fluctuate strongly in a population or vary across space; when proportions are static, it is less likely that intraspecific differences will be responsible for changes in the dynamics and outcomes of plant–animal interactions. In conclusion, focusing on variation among foraging animals rather than on species averages might bring new, mechanistic insights to the phenomenon of seed dispersal. While this shift in perspective is unlikely to replace the traditional approach (based on the assumption that all important variation occurs among species), it provides a complementary alternative to decipher the enormous variation observed in animal‐mediated seed dispersal.     

Small-scale indirect effects determine the outcome of a tripartite plant–disperser–granivore interaction

The microhabitat in which plants grow affects the outcome of their interactions with animals, particularly non-specialist consumers. Nevertheless, as most research on this topic has dealt with either mutualists or antagonists, little is known about the indirect effects of plant microhabitats on the outcome of tripartite interactions involving plants and both mutualists (e.g. seed dispersers) and antagonists (e.g. granivores). During three consecutive years, we analysed small-scale variations in the interaction of a perennial myrmecochore, Helleborus foetidus, with its seed dispersers and consumers as a function of the intensity of plant cover. Most seeds were released during the day and were rapidly removed by ants. Nevertheless, the proportion of ant-removed seeds was higher for plants located in open microhabitats than for plants surrounded by dense vegetation and rocky cover. Ant sampling revealed that seed removers were equally abundant, irrespective of the level of cover. By contrast, a few tiny ant species that feed on the reward without transporting the seeds were more abundant in highly covered microhabitats, irrespective of hellebore diaspore availability. These “cheaters” decrease the chance of removal by removers and increase the probability of seeds remaining on the ground until night, when granivore mice Apodemus sylvaticus become active. Mice also preferred foraging in covered microhabitats, where they consumed a larger proportion of seeds. Therefore, the density of cover indirectly increased seed predation risk by attracting more seed predators and cheater ants that contribute to increase seed availability for seed predators. Our results emphasize the importance of considering the indirect effects of plant microhabitat on their dispersal success. They highlight the indirect effect of cheaters that are likely to interfere in mutualisms and may lead to their collapse unless external factors such as spatio-temporal heterogeneity in seed availability constrain their effect. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Spatiotemporal variation in predispersal seed predation intensity

The effect of predispersal seed predation by Bruchus atomarius (Bruchidae, Coleoptera) on individual performance and population dynamics of the perennial forest herb, Lathyrus vernus (Leguminosae), was investigated in 11 permanent plots over 4 years. Seed predation and parameters describing intra-specific neighbour distance, plant size, inflorescence size, flowering phenology and current and previous herbivore damage were measured on all plants. In addition, demographic information from all plots was analysed using transition matrix population models in order to estimate the influence of seed predation on population growth rates. Predispersal seed predation rates differed significantly among years. Plot averages ranged from 0 to 83.7%. However, most of the variation occurred among individuals. Within individuals there was no consistency in predation rates among years. Exposure to herbivory, plant size and flowering phenology did not affect predation rates but individuals with larger inflorescences suffered from significantly higher predation. Seed predation in L. vernus was not influenced by neighbour distances of individual plants but it was positively correlated with the average density of seeds within plots, suggesting that seed predation is density dependent at the patch level. The reduction in population growth rate due to seed predation ranged from 0 to 7.6%. The sensitivity of population growth rate to reductions in seed production varied considerably among years and plots. This variation was mainly due to differences in the reproductive value of seeds and seedlings. The intensity of seed predation over the range found was not correlated with changes in population growth rate. The results of this study suggest that the influence of external factors, like seed predation, on population growth rate largely depends on the demographic transition rates in the investigated population.     

Seed dispersal by Galápagos tortoises

Aim Large‐bodied vertebrates often have a dramatic role in ecosystem function through herbivory, trampling, seed dispersal and nutrient cycling. The iconic Galápagos tortoises (Chelonoidis nigra) are the largest extant terrestrial ectotherms, yet their ecology is poorly known. Large body size should confer a generalist diet, benign digestive processes and long‐distance ranging ability, rendering giant tortoises adept seed dispersers. We sought to determine the extent of seed dispersal by Galápagos tortoises and their impact on seed germination for selected species, and to assess potential impacts of tortoise dispersal on the vegetation dynamics of the Galápagos. Location Galápagos, Ecuador. Methods To determine the number of seeds dispersed we identified and counted intact seeds from 120 fresh dung piles in both agricultural and national park land. To estimate the distance over which tortoises move seeds we used estimated digesta retention times from captive tortoises as a proxy for retention times of wild tortoises and tortoise movement data obtained from GPS telemetry. We conducted germination trials for five plant species to determine whether tortoise processing influenced germination success. Results In our dung sample, we found intact seeds from > 45 plant species, of which 11 were from introduced species. Tortoises defecated, on average, 464 (SE 95) seeds and 2.8 (SE 0.2) species per dung pile. Seed numbers were dominated by introduced species, particularly in agricultural land. Tortoises frequently moved seeds over long distances; during mean digesta retention times (12 days) tortoises moved an average of 394 m (SE 34) and a maximum of 4355 m over the longest recorded retention time (28 days). We did not find evidence that tortoise ingestion or the presence of dung influenced seed germination success. Main conclusions Galápagos tortoises are prodigious seed dispersers, regularly moving large quantities of seeds over long distances. This may confer important advantages to tortoise‐dispersed species, including transport of seeds away from the parent plants into sites favourable for germination. More extensive research is needed to quantify germination success, recruitment to adulthood and demography of plants under natural conditions, with and without tortoise dispersal, to determine the seed dispersal effectiveness of Galápagos tortoises.     

Secondary seed dispersal by ants in Neotropical cerrado savanna: species‐specific effects on seeds and seedlings of Siparuna guianensis (Siparunaceae)

1. Most woody plant species in tropical habitats are primarily vertebrate‐dispersed, but interactions between ants and fallen seeds and fruits are frequent. This study assesses the species‐specific services provided by ants to fallen arillate seeds of Siparuna guianensis, a primarily bird‐dispersed tree in cerrado savanna. The questions of which species interact with fallen seeds, their relative contribution (versus vertebrates) to seed removal, and the potential effects on seedling establishment are investigated. 2. Seeds are removed in similar quantities in caged and control treatments, suggesting that ants are the main dispersers on the ground. Five ant species attended seeds. Pheidole megacephala (≈0.4 cm) cooperatively transported seeds, whereas the smaller Pheidole sp. removed the seed aril on spot. Large (> 1.0 cm) Odontomachus chelifer, Pachycondyla striata, and Ectatomma edentatum individually carried seeds up to 4 m. Bits of aril are fed to larvae and intact seeds are discarded near the nest entrance. 3. Overall, greater numbers of seedlings were recorded near ant nests than in control plots without nests. This effect, however, was only detected near P. megacephala and P. striata nests, where soil penetrability was greater compared with controls. Soil nutrients did not differ between paired plots. 4. This study confirms the prevalence of ant–seed interactions in cerrado and shows that ant‐derived benefits are species‐specific. Ant services range from seed cleaning on the spot to seed displacement promoting non‐random spatial seedling recruitment. Although seed dispersal distances by ants are likely to be shorter than those by birds, our study of S. guianensis shows that fine‐scale ant‐induced seed movements may ultimately enhance plant regeneration in cerrado.     

Effects of seed passage through slugs on germination

Seed dispersal by invertebrates is mostly external (e.g. by ants). However, internal dispersal (endozoochory) seems to be uncommon due to size limitations. Slugs are generalist herbivores and increasing evidence suggests that they often disperse seeds. Nevertheless, we know very little on the consequences for plant recruitment. Here, we assess the effect of slug seed passage on germination and early seedling establishment of a set of nine common plant species with limited dispersal capabilities (unassisted or ant dispersed). Germination trials were performed under natural conditions (NW of Spain). Size was a clear limitation for seed ingestion: smaller seeds were eaten more readily, whereas only the largest slugs could swallow the largest seeds. All ingested seeds were voided undamaged. Only not-ingested seeds (the biggest) were damaged, with chewing marks in the surface and 0.8% (N = 250) were broken. In some cases, slugs ate the elaiosomes but discarded the seeds. Slug seed passage had an overall positive effect on total germination and sped it up in two species. Elaiosome removal by slugs had also a positive effect in Ulex europaeus seeds (+160%). However, slugs had no effect on early seedling survival and growth of the study species. Our results show that slugs are effective dispersers in terms of seed treatment in the digestive track. However, according to their reduced movement range (in the range of tens of metres; as shown by other studies), they can only provide rather limited dispersal as compared to vertebrate dispersers. This can be nonetheless significant for species with limited dispersal, e.g. unassisted and ant-dispersed plants, for which they act as non-standard dispersers.     

Effectiveness of Spider Monkeys (<Emphasis Type="Italic">Ateles geoffroyi vellerosus</Emphasis>) as Seed Dispersers in Continuous and Fragmented Rain Forests in Southern Mexico

Seed dispersal is considered a key process determining spatial structure and dynamics of plant populations, and has crucial implications for forest regeneration. We evaluated the effectiveness of spider monkeys (Ateles geoffroyi) as seed dispersers in continuous and fragmented habitats to test if this interaction is altered in forest fragments. We documented fruit and seed handling, defecation patterns, diversity and composition of seeds in feces, and seed germination of defecated and control seeds in the Lacandona rainforest, Mexico. For most species contributing to 80% of total fruit feeding time, monkeys swallowed and spat seeds, but swallowing was the most frequent seed handling category in continuous and fragmented forests. However, the proportion of feeding records of swallowed seeds was higher in continuous forest (0.59) than in fragments (0.46), whereas the opposite was true for proportion of dropped seeds (0.16 vs. 0.31). This pattern was reflected in the number of fecal samples containing seeds, which was greater in continuous (95.5%) than in fragmented forests (82.5%). Seeds in fecal samples included a total of 71 species from 23 plant families. The numbers of defecated seed species were similar between forest conditions, and in both cases most seeds (>86%) were undamaged. Defecated seeds showed greater germination percentages than control seeds in all of the five species evaluated. Although we identified some differences in seed handling and the percentage of feces with seeds between continuous forest and fragments, our results indicate that, in general terms, spider monkeys are effective seed dispersers in both forest conditions.     

Effects of two types of herbivores on the population dynamics of a perennial herb

The joint effects of multiple herbivores on their shared host plant have received increasing interest recently. The influence of herbivores on population dynamics of their host plants, especially the relative roles of different types of damage, is, however, still poorly understood. Here, we present a modelling approach, including both deterministic and stochastic matrix modelling, to be used in estimating fitness effects of multiple herbivores on perennial plants. We examined the effects and relative roles of two specialist herbivores, a pre-dispersal seed predator, Euphranta connexa, and a leaf-feeding moth, Abrostola asclepiadis, on the population dynamics and long-term fitness of their shared host plant, a long-lived perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). We collected demographic data during 3 years and combined these data with the effects of natural levels of herbivory measured from the same individuals. We found that both seed predation and leaf herbivory reduced population growth of V. hirundinaria, but only very high damage levels changed the growth trend of the vigorously growing study populations from positive to negative. Demographic modelling indicated that seed predation had a greater impact on plant population growth than leaf herbivory. The effect of leaf herbivory was weaker and diminished with increasing level of seed predation. Evaluation of individual fitness components, however, suggested that leaf herbivory contributed more strongly to host plant fitness than seed predation. Our results emphasize that understanding the effects of a particular herbivore on plant population dynamics requires also knowledge on other herbivores present in the system, because the effect of a particular type of herbivory on plant population dynamics is likely to vary according to the intensity of other types of herbivory. Furthermore, evaluating herbivore impact from using individual fitness components does not necessarily reflect the long-term effects on total plant fitness.     

High regeneration capacity helps tropical seeds to counter rodent predation

Rapid germination of non-dormant seeds is one adaptation plants have evolved to counter seed predation by rodents. Some rodent species have evolved behaviors that prevent or slow the seed germination process through seed embryo removal or seed pruning; however, no plant species is known to have successfully escaped embryo removal or seed pruning by rodents. Here, we report that the non-dormant seeds of Pittosporopsis kerrii Craib in tropical rain forests in China have a high regeneration capacity to counter seed pruning by rodents. We found seed pruning, instead of embryo removal, was commonly used by rodents to increase food storage time by slowing down the seed germination process, but that P. kerrii seeds have a high regeneration capacity to escape seed predation by rodents: all pruned seeds, pruned roots and embryo-removed seeds by rodents or people retain the ability to develop into seedlings. Seeds of P. kerrii also have other capacities (i.e. rapid seed decomposition and indigestible dormant taproots) to escape predation by reducing the plant’s attractiveness to rodents. The association between seed pruning behavior in rodents and high regeneration capacity of pruned seeds or roots in P. kerrii seeds are likely novel adaptation strategies adopted by seeds and rodents, respectively.     

Seed dispersal effectiveness in a plant–lizard interaction and its consequences for plant regeneration after disperser loss

Mutualistic disruptions, such as those promoted by the loss of seed dispersers, can have negative effects on the plant regeneration of those species that strongly depend upon them. In order to adequately assess how plant communities are affected by such disruptions, we need to know the importance of the dispersal phase, both in its quantitative and qualitative components. We examined this in the narrow interaction between the shrub Daphne rodriguezii and its (only) disperser, the lizard Podarcis lilfordi. We quantified fruit removal and the effect of fruit/seed-size selection, seed treatment in the disperser’s guts and seed deposition patterns on seedling emergence and survival. In the only locality in which lizards persist, they removed most fruits and showed preference for larger ones in one of the two study years. Seed treatment in lizard’s guts had no effect on germination, although it tended to reduce the effect of seed size on germination (differences between large vs. small seeds in seed germination were higher for non-ingested seeds). Probability of seedling emergence, but not survival, was higher in the locality with lizards. Dispersed seeds under heterospecific shrubs showed higher seedling survival than those under conspecifics in all localities, especially the year with higher rainfall. Our findings support that the movement of seeds to nurse shrubs by lizards is the most important component of the seed dispersal process in the only remaining locality where both species coexist.