Biological and environmental effects on fine-scale seed dispersal of an invasive tree in a secondary subtropical forest

Dispersal abilities of invading species emerge from the interaction between the species and some features of the target community. Ligustrum lucidum is a tree species invading different ecosystems. Major spatial patterns of Ligustrum invasions and their ecological consequences have been analyzed, but no study addressed the dispersal process at a fine scale, assessing the effects of different biological and environmental factors. Ligustrum lucidum is an ornithochoric species. The structure of the environment determines bird movements and thus affects seed dispersal. We used inverse modeling to analyze bird-mediated dispersal of L. lucidum seeds in a secondary Yungas forest and surrounding crop-fields. We assessed the effects of egestion mode (regurgitation and defecation) and tree density (as an environment character) on seed dispersal. Seed dispersal presented different spatial patterns depending on the egestion mode. Tree density was positively associated with the number of regurgitated dispersed seeds and negatively associated with the number of defecated dispersed seeds. In both cases, dispersal distance increased in open areas, but absence of perches inhibited seed arrival. Thus, spread of L. lucidum is facilitated in open areas with some trees; inside the native forest, short distance dispersal facilitates the gradual invasion by this exotic species. Our results suggest that processes like crop abandonment and forest succession, which are active in subtropical montane systems, may facilitate L. lucidum invasion. Our seed dispersal models should be combined with actual distribution maps of L. lucidum to identify areas vulnerable to new invasions.     

Directed seed dispersal towards areas with low conspecific tree density by a scatter‐hoarding rodent

Scatter‐hoarding animals spread out cached seeds to reduce density‐dependent theft of their food reserves. This behaviour could lead to directed dispersal into areas with lower densities of conspecific trees, where seed and seedling survival are higher, and could profoundly affect the spatial structure of plant communities. We tested this hypothesis with Central American agoutis and Astrocaryum standleyanum palm seeds on Barro Colorado Island, Panama. We radio‐tracked seeds as they were cached and re‐cached by agoutis, calculated the density of adult Astrocaryum trees surrounding each cache, and tested whether the observed number of trees around seed caches declined more than expected under random dispersal. Seedling establishment success was negatively dependent on seed density, and agoutis carried seeds towards locations with lower conspecific tree densities, thus facilitating the escape of seeds from natural enemies. This behaviour may be a widespread mechanism leading to highly effective seed dispersal by scatter‐hoarding animals.     

Secondary dispersal of forest herb seeds from raccoon dung: contrasting service by multiple vectors

Secondary movement of seeds deposited in dung potentially improves their chances of seedling establishment but has rarely been documented in temperate-zone deciduous forest. We followed mayapple (Podophyllum peltatum) seeds after deposition in raccoon (Procyon lotor) dung to test for secondary dispersal in a long-established deciduous forest in southeastern Ohio, USA. Marked seeds were incorporated into raccoon dung, placed in natural forest sites, and observed at regular intervals until the subsequent Spring. Abiotic seed movement was monitored during natural and simulated rain events. To identify animal vectors and describe their interaction with seeds, motion-triggered wildlife cameras were trained on dung with embedded mayapple seeds. Seed movement was initially limited by adhesion in the viscid dung matrix, but 15% of seeds eventually moved >20 cm from the point of placement, a minimal estimate of a shoot’s competitive neighborhood (maximum movement distance = 130 cm). Significantly greater movement occurred on slopes and in the absence of leaf litter. Rainfall moved seeds as far as 51 cm. White-footed mice (Peromyscus leucopus) and chipmunks (Tamias striatus) began removing seeds within 24 h of dung placement, presumably caching a portion of them outside the 1.5-m radius of observation. At least four other animal species interacted with dung in ways unrelated to seed presence, which nevertheless resulted in seed movement on a scale of centimeters. Thus, multiple secondary processes caused seed movement, each resulting in a distinctive seed distribution. The forest herb potentially benefits through reduction in seed density and distribution to a variety of microsites.     

Do deer and raccoons defecate in the right place? Fitness consequences of vertebrate seed dispersal for a deciduous forest herb

Precision of seed placement in a heterogeneous environment is often assumed to select for the evolution of animal-mediated dispersal systems, but this hypothesis has rarely been tested in a multivariate sense. We quantify the microsite fitness benefits of dispersal by white-tailed deer (Odocoileus virginianus) and raccoons (Procyon lotor) for mayapple (Podophyllum peltatum), a shade-tolerant perennial herb, in deciduous forests of southeastern Ohio, USA. Micro-environmental variables were recorded at dung-deposition microsites, at rooting points of mayapple shoots, and at random (control) points in the forest. Fitness was assessed as the degree of overlap in ordinations of microsites by environmental variables. Mayapple occupied a broad sector (56%) of environment space corresponding to low and mid-slope positions, ravines, and proximity to trees. Deer and raccoon defecation placed dung in 71–74 and 86–95% of environment space, respectively, reaching mayapple microsites in 57–60 and 53–54% of cases. Deer placed dung in mayapple environment space 7–9% more often than predicted by random distribution, and raccoons placed dung in mayapple space 0–5% more often, consistent with only a modest degree of directed dispersal. Thus, the precision hypothesis is only weakly supported. The greatest fitness benefit of vertebrate dispersal appears to be the broad distribution of seeds, thereby increasing the probability of randomly reaching a suitable microsite. Imprecise dispersal suggests that secondary mechanisms of seed movement need to be explored in deciduous forest communities.     

When condition trumps location: seed consumption by fruit‐eating birds removes pathogens and predator attractants

Seed ingestion by frugivorous vertebrates commonly benefits plants by moving seeds to locations with fewer predators and pathogens than under the parent. For plants with high local population densities, however, movement from the parent plant is unlikely to result in ‘escape’ from predators and pathogens. Changes to seed condition caused by gut passage may also provide benefits, yet are rarely evaluated as an alternative. Here, we use a common bird‐dispersed chilli pepper (Capsicum chacoense) to conduct the first experimental comparison of escape‐related benefits to condition‐related benefits of animal‐mediated seed dispersal. Within chilli populations, seeds dispersed far from parent plants gained no advantage from escape alone, but seed consumption by birds increased seed survival by 370% – regardless of dispersal distance – due to removal during gut passage of fungal pathogens and chemical attractants to granivores. These results call into question the pre‐eminence of escape as the primary advantage of dispersal within populations and document two overlooked mechanisms by which frugivores can benefit fruiting plants.     

Diminishing importance of elaiosomes for acacia seed removal in non-native ranges

Myrmecochorous plants produce seeds with lipid-rich appendages (elaiosomes) which act as a reward for seed-dispersing ants. Seed dispersal is important for exotic species, which often need to establish new mutualistic interactions in order to colonize new non-native habitats. However, little is known about the importance of elaiosomes for seed removal in many of their non-native ranges. We studied ant–seed interactions of elaiosome-bearing and elaiosome-removed seeds of the Australian trees Acacia dealbata and Acacia longifolia in order to assess the relative importance of elaiosomes for seed removal between their native (Australia) and non-native (Portugal) ranges. In Portugal, we also studied the co-occurring native plant species with myrmecochorous seeds, Pterospartum tridentatum and Ulex europaeus, across three contiguous levels of acacia invasion: control (i.e. no acacia), low, and high acacia tree density. Acacia seeds were successfully removed by ants in their non-native region by a diversified assemblage of ant species, even in sites where native plants interacted with only one specialized ant species. In the invaded range, diminishing relative importance of elaiosomes was associated with changes in the ant community due to acacia invasion, and for A. dealbata, ant species richness decreased with increasing acacia tree density. Although seed removal was high for both acacia species, the importance of elaiosomes was proportionally lower for A. dealbata in the non-native region. Native plant species experienced significant reductions in seed removal in areas highly invaded by acacia, identifying another mechanism of displacement of native plants by acacias.     

Uncorrelated mistletoe infection patterns and mating success with local host specialization in <Emphasis Type="Italic">Psittacanthus calyculatus</Emphasis> (Loranthaceae)

Mistletoe infection between conspecific and interspecific hosts can be restricted by seed dispersal, host-mistletoe compatibility and abiotic factors, yet no studies have linked mistletoe infection patterns and pollination together for understanding mistletoe distribution at a local scale. Psittacanthus calyculatus (Loranthaceae) is a hemiparasitic plant with a broad host range across its geographic distribution. The potential for local host adaptation has been shown using cross-inoculation experiments, in which plants of mistletoe seeds collected from a given host are more likely to survive when they are inoculated on conspecific host trees compared with those inoculated on other host provenances. Here we evaluate host adaptation by describing the local patterns of infection (prevalence and intensity) of P. calyculatus mistletoes on three native host tree species (Alnus acuminata, Quercus crassipes, Salix bonplandiana) and one introduced species (Populus alba) and carried out cross-pollination experiments to examine how pollination affects infection patterns of different host species. Mistletoe infection prevalence (proportion of infection) and infection intensity (mean number of mistletoes per tree) were in general disproportional with respect to the availability of native host tree species but higher to that of non-native host tree species. Cross-pollination experiments showed higher mating success on the native host tree species, suggesting higher local adaptation to specially Q. crassipes. The observed spatial distribution of host tree species and mistletoe infection along with the non-random mating could contribute to local genetic structuring of mistletoe populations.     

Forest disturbance and seasonal food availability influence a conditional seed dispersal mutualism

The interaction between granivorous scatterhoarding mammals and plants is a conditional mutualism: scatterhoarders consume seeds (acting as predators), but the movement of seed by scatterhoarders may contribute to dispersal (acting as mutualists). Understanding the ecological factors that shape this relationship is highly relevant in anthropogenically disturbed tropical forests where large‐bodied frugivores are extirpated. In such forests, large‐seeded trees that once depended on these frugivores for dispersal may now only have scatterhoarders as prospective dispersers. We studied Carapa oreophila (Meliaceae) in an Afromontane forest, to test the hypotheses that the proportion of seeds immediately consumed or hoarded (dispersed) would vary over a disturbance gradient. Temporal replication also afforded exploration of how habitat effects might vary with food availability. Using a Bayesian framework, we demonstrate that seeds were more likely to be hoarded in less disturbed forest, irrespective of temporal variation in food abundance. In contrast, forest disturbance only appeared to increase seed predation in temporal replicates that coincided with sustained food availability. These results highlight the potential variability in the dynamics between plants and scatterhoarders over fine temporal scales, elucidating possible ecological scenarios where scatterhoarders might act as mutualists (contributing positively to plant recruitment). Our study also fills important knowledge gaps about the importance of scatterhoarders as dispersers in tropical forests depleted of large‐bodied frugivores, particularly in Africa where scatterhoarding mutualisms have not been extensively studied.     

Role of indehiscent pericarp in formation of soil seed bank in five cold desert Brassicaceae species

The dispersal and germination unit of some Brassicaceae species is the fruit, and we hypothesized that it could affect germination phenology and promote formation of a soil seed bank. We determined the effects of the indehiscent pericarp on germination and longevity of buried seeds of five Brassicaceae species native to cold deserts of central Asia. Germination phenology (seedling emergence) was monitored for intact dispersal units and isolated seeds of Chorispora sibirica, Goldbachia laevigata, Spirorrhynchus sabulosus, Tauscheria lasiocarpa (annuals), and Sterigmostemum fuhaiense (perennial) at natural temperatures in watered and non-watered (natural precipitation) soil. Intact dispersal units and isolated seeds were buried under natural conditions and exhumed at regular intervals for 35 months to monitor germination, viability and moisture content of isolated seeds, seeds in dispersal units, and seeds removed from dispersal units after burial. Isolated seeds of Goldbachia, Spirorrhynchus, and Tauscheria germinated only the first autumn and those of Chorispora and Sterigmostemum the first autumn and first spring, with higher germination percentages in all species in watered than in non-watered soil. A high percentage of seeds in buried dispersal units of Chorispora, Goldbachia, and Sterigmostemum was viable after 35 months, and seeds exhibited a 6-month dormancy cycle, being non-dormant only in autumn and spring. Seeds in buried dispersal units of Spirorrhynchus and Tauscheria germinated when exhumed in the first spring, but all non-germinated seeds were dead after 1 year. Thus, the presence of the pericarp allows Chorispora, Goldbachia, and Sterigmostemum to form a persistent seed bank but not Spirorrhynchus and Tauscheria.     

Differences in reward removal efficiency and its consequences for seed germination in the ant-dispersed sedge <Emphasis Type="Italic">Carex tristachya</Emphasis> (Cyperaceae)

Reward removal is an essential step for seed dispersal mutualism because residual rewards inhibit germination. Nevertheless, variation in the reward removal efficiency (RRE) among dispersers and its consequences for germination have rarely been reported. In this study, we compared the RREs of two sympatric seed-dispersing ants, Formica japonica and Pheidole noda, using seeds of the ant-dispersed sedge Carex tristachya. Then, we conducted seed sowing experiments in a non-heated glasshouse to evaluate the effect of RRE on the percentage and speed of germination. The majority (85%) of seeds handled by F. japonica had residual elaiosomes, while elaiosomes were completely removed from all seeds handled by P. noda, demonstrating that P. noda has much higher RRE than F. japonica. The seed sowing experiments revealed that RRE, defined by the presence or absence of residual elaiosomes, was not associated with the percentage germination within a year. However, high RRE seeds with no residual elaiosomes germinated significantly faster than low RRE seeds with residual elaiosomes. Similarly, artificial removal of elaiosomes from C. tristachya seeds accelerated germination speed without affecting germination percentage. These results suggest that RRE is one of the most important parameters determining the effectiveness of a seed dispersal agent.     

Interaction between a threatened endemic plant (<Emphasis Type="Italic">Anchusa crispa</Emphasis>) and the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Seed dispersal mutualisms are essential to ensure the survival of diverse plant species and communities worldwide. Here, we investigated whether the invasive Argentine ant can replace native ants by fulfilling their functional role in the seed dispersal of the rare and threatened endemic myrmecochorous plant, Anchusa crispa, in Corsica (France). Our study addressed the potential of Linepithema humile to disperse elaiosome-bearing seeds of A. crispa, examining L. humile’s effects on (1) the composition of communities of ants removing seeds, (2) the number of seed removals, (3) seed preference, (4) the distance of seed dispersion, and (5) seed germination. We caught seven native species at the control site, but only the Argentine ant at invaded sites. L humile removed A. crispa seeds in greater numbers than did native ants, respectively 66 and 23%, probably due to their higher worker density. The invader was similar to native ants with respect to distance of seed transport. Finally, rates of seed germination were not significantly different between seeds previously in contact with either Argentine ants or not. Taken all together, these results suggest that the Argentine ant is unlikely to pose a threat to A. crispa population. These results have important implications for the management of this rare and threatened endemic plant and provide an example of non-negative interactions between invasive and native species.     

Diplochory in western chokecherry: you can��t judge a fruit by its mesocarp

Western chokecherry (Prunus virginiana var. demissa, Rosaceae) is dispersed by frugivorous birds and carnivores, but it has large seeds that are potentially attractive to rodents that could act as seed predators and dispersers. Here, we quantify the benefits of primary dispersal by birds and secondary dispersal by scatter-hoarding rodents. In the fall, avian frugivores (mostly American robins, Turdus migratorius, and cedar waxwings, Bombycilla cedrorum) consumed 87% of the fruit crop and dispersed 67% of the fruit crop away from parent plants. Rodents removed 89% of seeds that simulated bird-dispersed seed rain from transects in riparian zones and 58% from transects in upland habitats. Rodents scatter-hoarded 91.6% of the seeds they removed, burying most in small caches (two to eight seeds) 8?C25?mm deep. About 39% of the seeds in spring caches produced seedlings. Inside rodent-proof exclosures, 52.1% of seeds buried to simulate rodent caches produced seedlings, 29.7% of which were still alive after 1?year. In contrast, only 3.8% of seeds placed on the soil surface, simulating dispersal by avian frugivores, produced seedlings. Seed dispersal by frugivorous birds likely contributes to colonization of unoccupied habitat through long-range dispersal and to escape from distance-dependent seed mortality near the parent plant. Despite seed losses, rodents offer short-range seed dispersal and bury seeds in more favorable sites for germination, improving seedling emergence and establishment. The combined mechanisms of seed dispersal significantly enhanced chokecherry seedling recruitment by providing more dispersal-related benefits than either frugivorous bird or scatter-hoarding rodents could provide alone.     

Ant interactions with native and exotic seeds in the Patagonian steppe: Influence of seed traits,disturbance levels and ant assemblage

Invasive plants may establish strong interactions with species in their new range which could limit or enhance their establishment and spread. These interactions depend upon traits of the invader and the recipient community, and may alter interactions among native species. In the Patagonian steppe we studied interactions of native ant assemblages with seeds of native and exotic plants, and asked whether ant–seed interactions differ with seed types and disturbance levels and whether the amount and type of ant–seed interactions can be predicted if both ant and seed traits are known. To characterize and quantify ant–seed interactions, we offered baits with large seeds of Pappostipa speciosa (native) and medium-sized elaiosome-bearing seeds of Carduus thoermeri (exotic), near and far from a road (high vs. low disturbed areas), and compared ant abundance and composition between areas. Interaction frequency was the highest for C. thoermeri seeds far from the road. Composition of ants interacting with C. thoermeri in these areas differed from that near the road and from that interacting with native seeds. Ant composition and abundance were similar between areas, but some species interacted more with exotic seeds in low disturbed areas. Ant foraging type predicted ant–seed interactions since the abundance of seed harvesters was positively correlated to interactions with P. speciosa, and that of generalists and predators, with interactions with C. thoermeri. The high interaction of ants with exotic seeds in low invaded areas suggests that ant activity could influence plant invasion, either by predating or dispersing seeds of invasive plants.     

Seed Dispersal by Primates and Implications for the Conservation of a Biodiversity Hotspot,the Atlantic Forest of South America

Primates play a fundamental role as seed dispersers, particularly in tropical rainforests. Because defaunation and fragmentation are leading several primate species to local extinction, it is fundamental to understand the role of primates as effective seed dispersers. Here we present a systematic review of studies of seed dispersal by primates in a biodiversity hotspot, the Atlantic Forest of South America, to 1) highlight gaps in our knowledge, 2) determine species richness and proportion of seed species dispersed, and 3) test the relationship between primate body size and the size of dispersed species. Our review found 79 studies of the diet of six ecospecies (Callithrix, Leontopithecus, Callicebus, Sapajus, Alouatta, Brachyteles) but only 20 of these report information on seed dispersal, and none of these are on Callithrix or Callicebus. We found a strong bias in the distribution of species and regions, with most of the studies concentrated in southeastern Brazil. All ecospecies dispersed a large proportion of the seed species they handled (72.1–93.6%). Brachyteles dispersed the highest diversity of plants (N = 73), followed by Sapajus (N = 66), Leontopithecus (N = 49), and Alouatta (N = 26). Although we found no significant relationship between primate body size and the size of seeds dispersed, Brachyteles disperse a higher diversity of large-seeded species than smaller-bodied primates. These results suggest that the local extinction of large primate species may lead to dramatic changes in the plant community, as many large-seeded plants are inaccessible to smaller arboreal frugivores. We propose guidelines for future research on primate seed dispersal to enable the evaluation of seed dispersal effectiveness and to improve our understanding of the fundamental role of primates in this key ecosystem process.     

Contagious seed dispersal beneath heterospecific fruiting trees and its consequences

An hypothesized advantage of seed dispersal is avoidance of high per capita mortality (i.e. density-dependent mortality) associated with dense populations of seeds and seedlings beneath parent trees. This hypothesis, inherent in nearly all seed dispersal studies, assumes that density effects are species-specific. Yet because many tree species exhibit overlapping fruiting phenologies and share dispersers, seeds may be deposited preferentially under synchronously fruiting heterospecific trees, another location where they may be particularly vulnerable to mortality, in this case by generalist seed predators. We demonstrate that frugivores disperse higher densities of Cornus florida seeds under fruiting (female) Ilex opaca trees than under non-fruiting (male) Ilex trees in temperate hardwood forest settings in South Carolina, USA. To determine if density of Cornus and/or Ilex seeds influences survivorship of dispersed Cornus seeds, we followed the fates of experimentally dispersed Cornus seeds in neighborhoods of differing, manipulated background densities of Cornus and Ilex seeds. We found that the probability of predation on dispersed Cornus seeds was a function of both Cornus and Ilex background seed densities. Higher densities of Ilex seeds negatively affected Cornus seed survivorship, and this was particularly evident as background densities of dispersed Cornus seeds increased. These results illustrate the importance of viewing seed dispersal and predation in a community context, as the pattern and intensity of density-dependent mortality may not be solely a function of conspecific densities.     

Herbivore-induced expansion of <Emphasis Type="Italic">Helianthemum nummularium</Emphasis> in grassland–scrub mosaic vegetation: circumstantial evidence for zoochory and indirect grazing impact

Extensive grazing often has a strong influence on the structure and composition of herbaceous plant communities with increasing population sizes for some species and decreasing presence in others. Herbivores affect plant communities directly by selective grazing of plant species, and indirectly by either epizoochory or endozoochory. Helianthemum nummularium is considered an increasing species because its distribution increased after the introduction of large, free-ranging grazers in at least two coastal dune grassland areas in Belgium. However, its seeds lack any obvious adaptations for epizoochory, and direct observations of plant/seed consumption are scarce. Through field and lab experiments, we assessed the dispersal ability of H. nummularium via endozoochory and epizoochory. In a differentiated grazer exclusion experiment, evidence was found that plants are grazed by large domestic ungulates and small wild herbivores although these incidences were rare. Direct endozoochory evidence remained scarce. No seeds were found germinating in field-collected dung, and only few seedlings emerged following a seed feeding experiment. However, once deposited, we found higher growth rates when seeds were mixed with dung and decreased establishment success when seeds were sown in combination with competitively superior species. Epizoochory was plausible because both fur and hooves of cattle and horses were potentially capable of contributing to the transport of H. nummularium seeds. We conclude that herbivores play a role in seed dispersal, while their selective grazing behaviour most probably creates an appropriate environment for Helianthemum establishment and maintenance.     

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.     

Implications of Behavior and Gut Passage for Seed Dispersal Quality: The Case of Black and Gold Howler Monkeys

The most generalized mechanism of seed dispersal in tropical and subtropical forests is dispersal by vertebrates. The pattern of seed dispersal and germination capacity of dispersed seeds can determine the seed disperser quality because both affect the postdispersal fate of dispersed seeds. Animals' behavior is an important determinant of seed deposition pattern. One of most abundant frugivores of Neotropical forests are howler monkeys Alouatta spp., so my aim was to evaluate the potential quality of Alouatta caraya as a seed disperser, obtaining details of: (1) seed deposition pattern, (2) behavioral context of defecations, and (3) germination capacity of dispersed seeds. During 2 years the seed shadow created by howlers and the behavioral context of depositions was examined in the Paraná flooded forest, and germination tests were conducted in the laboratory. Black and gold howler monkeys consumed fruits of five canopy trees, three understory trees, one shrub, and two vines. Howlers dispersed seeds in a complex pattern: 35 percent (337 scats) of scats were deposited individually, and 65 percent were deposited in big latrines associated with the main sleeping trees and in small latrines associated with secondary sleeping trees and confrontations at territory edges. From 261 dispersal distances recorded, 20 and 40 m were the mode, but 57 percent of distances were >50 m and 31 percent were >100 m. Germination speed increased significantly in ingested seeds of Banara arguta, Ocotea diospyrifolia and Eugenia punicifolia . The seed deposition pattern generated by A. caraya may produce a diversification of environmental conditions for dispersed seeds that should be considered in future evaluations.     

Composition of harvested seeds and seed selection by the invasive tropical fire ant, <Emphasis Type="Italic">Solenopsis geminata</Emphasis> (Hymenoptera: Formicidae) in Taiwan

Solenopsis geminata (F.) is an invasive ant that is widely distributed in weedy areas and agricultural fields in Taiwan. Previous studies have found that S. geminata harvests the seeds of numerous plants. In the present study, we further investigated the composition of harvested seeds in ant nests and seed selection by workers. The seed caches in S. geminata nests sampled in eight areas in Taiwan suggest that the seeds harvested by workers were diverse and belonged to 52 plant species in 17 plant families. Twenty-three species (44%) belong to the family Gramineae, and most of the seeds weighed from 0.02 to 2.29 mg, which might suggest that these are the main seeds harvested by S. geminata from their habitat. Ten common species with similar seed weights were used to compare the seed preferences of workers from two experimental sites. The results suggest that the seed preference was different between the two experimental sites. The seeds of Casuarina equisetifolia showed the most obvious difference in seed removal speed, which might suggest that S. geminata potentially prefers the encountered seed species in the habitat. The various plant species in the ant nests and seed preference suggest that fire ants easily accept newly encountered plant species. As more than half of the plant species (52%) and the total seed number (63%) belonged to exotic species, the role of S. geminata might be negative because it tends to harvest exotic seeds and has a high opportunity to improve the establishment of exotic seedlings.     

Effects of invasive Green Iguanas (<Emphasis Type="Italic">Iguana iguana</Emphasis>) on seed germination and seed dispersal potential in southeastern Puerto Rico

Green Iguanas (Iguana iguana) are invasive in Puerto Rico due to a variety of negative economic effects, yet we know very little about their ecological impacts. Because they are herbivorous, defecate intact seeds, move through the forest, and have long gut-passage times, Green Iguanas may affect seed germination and seed dispersal. In summer 2013, a total of 258 Green Iguana scat samples were collected at the Humacao Natural Reserve in southeastern Puerto Rico. Seeds extracted from scat and collected from fruit were planted under common garden conditions using experimental treatments designed to tease apart the effects of feces, fruit, and ingestion on seed germination. Green Iguanas decreased the time for seeds to germinate in Ficus spp. by removing fruit pulp, but had no effect on germination of native Annona glabra seeds. For non-native P. pterocarpus and Pterocarpus spp., Green Iguanas produced conflicting results, decreasing the percentage of seeds germinating, but at the same time, reducing the time for seeds to germinate. Green Iguanas likely disperse most seeds beyond the canopies of parental tree at our site. Government and economic resources are being used to eradicate Green Iguana populations in Puerto Rico, but the lack of consistent effects of Green Iguanas on seed germination for the plant species consumed at our site complicates generalizing about their ecological effects and developing management plans that minimize negative effects for native plant communities. We recommend additional studies that target both species of particular concern, such as threatened native or invasive species, as well as studies of sensitive habitats in Puerto Rico.