Effects of dung and seed size on secondary dispersal,seed predation,and seedling establishment of rain forest trees

Seeds dispersed by tropical, arboreal mammals are usually deposited singly and without dung or in clumps of fecal material. After dispersal through defecation by mammals, most seeds are secondarily dispersed by dung beetles or consumed by rodents. These post-dispersal, plant-animal interactions are likely to interact themselves, as seeds buried by dung beetles are less likely to be found by rodents than unburied seeds. In a series of three experiments with seeds of 15 species in central Amazonia (Brazil), we determined (1) how presence and amount of dung associated with seeds influences long-term seed fate and seedling establishment, (2) how deeply dung beetles bury seeds and how burial depth affects seedling establishment, and (3) how seed size affects the interaction between seeds, dung beetles, and rodents. Our overall goal was to understand how post-dispersal plant-animal interactions determine the link between primary seed dispersal and seedling establishment. On average, 43% of seeds surrounded by dung were buried by dung beetles, compared to 0% of seeds not surrounded by dung (n=2,156). Seeds in dung, however, tended to be more prone than bare seeds to predation by rodents. Of seeds in dung, probability of burial was negatively related to seed size and positively related to amount of dung. Burial of seeds decreased the probability of seed predation by rodents three-fold, and increased the probability of seedling establishment two-fold. Mean burial depth was 4 cm (0.5–20 cm) and was not related to seed size, contrary to previous studies. Probability of seedling establishment was negatively correlated with burial depth and not related to seed size at 5 or 10 cm depths. These results illustrate a complex web of interactions among dung beetles, rodents, and dispersed seeds. These interactions affect the probability of seedling establishment and are themselves strongly tied to how seeds are deposited by primary dispersers. More generally, our results emphasize the importance of looking beyond a single type of plant-animal interaction (e.g., seed dispersal or seed predation) to incorporate potential effects of interacting interactions.     

Tamarins and Dung Beetles: An Efficient Diplochorous Dispersal System in the Peruvian Amazonia

Dung beetles fulfill several key functions in ecosystems but their role as secondary seed dispersers is probably one of the most complex ones. Various factors, such as seed characteristics, dispersal pattern induced by the primary disperser, season, and habitat, can affect the seed–beetle interaction. Particularly little is known about the fate of seeds primarily dispersed in small feces. The aim of this study was to investigate the effects of these factors on the dung beetle community (species composition, number and size of individuals) and its consequences on burial occurrence and depth of seeds primarily dispersed by two tamarin species. We captured dung beetles in a Peruvian rain forest with 299 dung‐baited pitfall traps to characterize the dung beetle community. Seed burial occurrence and depth were assessed by marking in situ 551 dispersed seeds in feces placed in cages. Among these seeds, 22.5 percent were buried by dung beetles after 2 d. We observed a significant effect of the amount of dung, season, time of deposition, and habitat on the number of individuals and species of dung beetles, as well as on seed burial occurrence and depth, while the tamarin species significantly influenced only the number and the size of dung beetles. This seed dispersal loop is particularly important for forest regeneration: small to large seeds dispersed by tamarins in secondary forest can be buried by dung beetles. These seeds can thus benefit from a better protection against predation and a more suitable microenvironment for germination, potentially enhancing seedling recruitment.     

Cheek-pouch dispersal of seeds by Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan

Seed dispersal by Japanese monkeys (Macaca fuscata yakui) via cheek-pouch was studied in a warm temperate evergreen forest on Yakushima Island. Plant list was compiled based on a study during 1986–1995, of which troops of monkeys have been habituated without artificial feeding. We followed the well-habituated monkeys in 1993 and 1994 to observe the feeding behavior and their treatments of fruits and seeds, and collected seeds dispersed by monkeys to record the distance carried from the mother trees. We checked the difference of germination ratio between seeds dispersed via cheek-pouch and seeds taken from mother trees by sowing experiments. Seeds and acorns of 22 species were observed to be dispersed via cheek-pouch of monkeys. Among them, three species with acorns were never dispersed via feces, and 15 species with drupes were seldom dispersed via feces. Plant species of which seeds are dispersed only via cheek-pouch had larger seeds than those of dispersed both via cheek-pouch and via feces, and typically had only one or two seeds in a fruit. As for one of cheek-pouch dispersal species,Persea thunbergii, the mean distance when seeds were carried from the mother trees via cheek-pouch was 19.7 m, and the maximum distance was as long as 105 m although more than 80% of seeds were dispersed within 30 m from mother trees. And 82% of seeds dispersed via cheek-pouch germinated. The easy separation of seeds from other parts of the fruit seems to facilitate cheek-pouch dispersal more than dispersal via feces. Cheek-pouch dispersal by monkeys has possibly enhanced the natural selection for larger seeds which bring forth larger seedlings with high shade-tolerance. In conclusion, cheek-pouch dispersal by monkeys is quite an important mode for trees in the mature stand in a warm temperate evergreen forest on Yakushima Island.     

Effects of dung beetles on seedling emergence from herbaceous seeds in the dung of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) in a temperate Japanese grassland ecosystem

The effects of dung form and condition and of dung beetles on the emergence of seedlings from herbaceous seeds in sika deer dung were examined in a temperate grassland ecosystem dominated by Zoysia japonica and Hydrocotyle maritima. I conducted field experiments to compare seedling emergence between dung exposed to dung beetles and intact dung using both dung pellets and pats during a typical rainy month (June) and the hottest, drier month (August), when large numbers of seeds of the dominant species were present in the dung. The exposed dung was immediately attacked and broken up by dung beetles, whereas dung protected from the beetles remained intact. In June, at least 12 herbaceous species, including Z. japonica, H. maritima, Mazus pumilus, and Plantago asiatica, emerged from the dung, versus at least six species in August. Decomposition rates of the pellets in June and decomposition scores of the pats in June and August were positively correlated with the number of emerging seedlings, suggesting that the acceleration of decomposition by dung beetles can positively affect seed germination. In this system of interactions among sika deer, herbaceous plants, and dung beetles, sika deer dung prevented seeds from germinating, and beetles had an indirect positive effect on seedling emergence by accelerating decomposition of the dung, although the extent of the effect may depend on the dung type, plant species, and environmental factors.     

Larger seeds in tropical floras: consistent patterns independent of growth form and dispersal mode

Many species of tropical moist forests have large seeds compared to those found in temperate floras. This could be attributed to a prevalence of woody growth forms, or adaptation to dispersal by vertebrates, or the dense shade of a closed canopy, rather than to an intrinsic tropical v. temperate difference. We compared tropical v. temperate seed mass data at two geographic scales; firstly within Australia between tropical, subtropical and temperate open woodlands, then between six tropical and temperate datasets from five continents. Within Australia we found that seed mass increased with decreasing latitude in ten out of eleven growth form-dispersal mode combinations: only wind-dispersed graminoids showed no significant effect. While the pattern of generally larger seeds in the tropical flora was associated with tropical families tending to have larger seeds than temperate families, we also found that tropical species had larger seeds than temperate cofamilials. As a preliminary test of the generality of these findings we included a further four tropical and temperate data sets in a second analysis. Average tropical seed masses were larger than temperate seed masses in all testable growth form-dispersal mode combinations, and statistically larger in five out of seven cases: unassisted, vertebrate- and wind-dispersed woody species, and unassisted and vertebrate-dispersed forbs. No difference was found for wind-dispersed forbs or for graminoids lacking dispersal structures. That an effect was found in these combinations rules out explanations based solely on characteristics of tropical v. temperate dispersers, although this may be a contributing factor. Instead we suggest that some aspect of tropical systems, for example higher metabolic costs of seedling production, has selected for a general increase in seed mass among tropical taxa.     

Efficiency of endozoochorous seed dispersal in six dry-fruited species (Cistaceae): from seed ingestion to early seedling establishment

We combined laboratory and nursery experiments to analyse the effectiveness of sheep as endozoochorous seed dispersers of six native shrubby Cistaceae species collected in SE Spain (Helianthemum apenninum (L.) Mill., H. violaceum (Cav.) Pers., Fumana ericoides (Cav.) Grand., F. thymifolia (L.) Spach, Cistus monspeliensis L. and C. laurifolius L.), considering the main stages after seed ingestion, i.e. seed recovery, seed germination, seedling emergence and early seedling establishment. Seed recovery after gut passage was high (around 40%) for all the species, except F. thymifolia (12%). Most seeds (ca. 90%) were recovered within 48 h after ingestion for all the species, although seeds were still recovered up to 96 h after ingestion. Gut passage increased germination up to seven-fold compared to non-ingested seeds. Furthermore, seedling emergence from seeds contained in pellets was overall similar (intact pellets) to or higher (crumbled pellets) than emergence from seeds without dung. Survival of emerged seedlings and mass of seedlings after 20 days were not reduced by dung. Sheep act therefore as effective dispersers of these Cistaceae species by scattering seeds and promoting germination, while faeces do not hamper seedling establishment. We conclude that the interaction between herbivorous ungulates and these dry-fruited species may be considered a mutualism qualitatively similar to the mutualism between frugivorous vertebrates and fleshy-fruited plants.     

Dung beetle activity affects rain forest seed bank dynamics and seedling establishment

Dung beetles relocate vertebrate feces under the soil surface, and this behavior has many ecological consequences. In tropical forests, for example, seeds defecated by mammals that are subsequently buried by dung beetles are less likely to suffer predation. While the effects of dung beetles on the fate of defecated seeds have been relatively well studied, their effect on seeds already buried in the soil has not. To contribute to fill this gap, we designed a study with three objectives: (a) Describe the vertical re‐distribution of soil seeds that occurs due to dung beetle activity; (b) Determine if beetle activity favors establishment of seedlings from the soil seed bank; and (c) Determine if the effect of dung beetles is stronger in sites of recurrent mammal defecation. We carried out three complementary field experiments, one with artificial seeds (plastic beads) of three sizes buried at known depths, one with two species of seeds buried at those same depths, and one with the natural soil seed bank in sites of single vs. recurrent defecation. Buried beads were moved by dung beetles along the vertical axis, both upwards (9.5%) and downwards (11.5%); smaller beads were more frequently moved downwards while the contrary occurred for larger beads. Dung beetle activity caused an increase in seedling establishment, both from experimentally buried seeds and from the natural seed bank. Defecation recurrence had no effect on seedling establishment. We conclude that dung beetle activity affects seed bank dynamics with important consequences for seedling establishment in tropical forests.Abstract in Spanish is available with online material.     

Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination

Canopy gap and chilling requirements for seed germination were assessed in 61 and 44 species, respectively, in the temperate rain forest of southern Chile. Germination assays within canopy gap and understorey were carried out under natural conditions. Germination tests in cold stratified and non-stratified seeds were performed under laboratory conditions. Seeds were collected of common trees, shrubs, vines and perennial herbs of forests in southern Chile. Final percent germination was significantly enhanced under canopy gap conditions in 19 species, and significantly reduced in 11 species. Germination proved indifferent under gap vs. understorey conditions in half the species tested. Cold stratification affected germination in 11 species: significantly increasing final germination of four species, and significantly decreasing final percent germination of seven species. A Principal Components Analysis (PCA) was applied with the purpose of identifying groups of species with similar germination strategies. Four attributes were included in the PCA: final germination percentage in canopy gap, germination rate in the laboratory, and gap-and chilling-dependency indices (EGAP and STRAT, respectively). The first axis separated species mainly on EGAP variation while the second axis separated them mainly according to STRAT variation; the two axes together explaining 73% of the among-species variation. A small group of trees and vines germinating best in the understorey and neutral to chilling could clearly be distinguished from the remaining species analyzed. Multifactorial ANOVAs were used for evaluating the combined effect of successional status, seed mass, dispersal period, life form, phylogenetic categories, and dispersal syndrome on EGAP and STRAT variation. The EGAP value of secondary successional species proved significantly greater than that of primary successional species and species with endozoochorous seed dispersal were significantly less dependent on chilling (according to STRAT values) compared to species with mainly wind-dispersed seeds. The possible implications of these results for seedling establishment are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Reduced germination success of temperate grassland seeds sown in dung: consequences for post‐dispersal seed fate

• Endozoochory is one of the main drivers shaping temperate grassland communities by maintaining plant populations of its constituents and enabling plants to colonize new habitats. Successful endozoochorous dispersal implies that seeds not only get consumed and survive the digestive tract but are also able to develop into viable seedlings in a dung environment.
• We experimentally assessed the germination probability and timing of 15 annual and perennial temperate European grassland species in cattle and horse dung and in different climatic conditions (greenhouse and outdoor conditions).
• Interspecific variation in germinability and germination timing are found, while life strategy had only an effect on germination timing. We found adverse effects of both cattle and horse dung on the germination characteristics of all tested grassland species, but the effects of cattle dung were more pronounced. In comparison with the control treatment, fewer seeds emerged in dung and more time was needed to germinate. Also, germination metrics clearly differed between the artificial greenhouse and outdoor conditions, with generally a lower germinability in outdoor conditions.
• According to our results, a large cost seems to be associated with endozoochorous dispersal in this stage of the life cycle, as seed dispersal effectiveness strongly depends on the quality of the deposition site with a lowered survival and germination probability when seeds are deposited in dung.

     

Herbivorous mammals as seed dispersers in a Mediterranean dehesa

Endozoochrous seed dispersal by herbivorous mammals has been verified repeatedly and its possible influence on the structure and function of herbaceous communities has been suggested. Quantitative studies, however, are lacking in the field of seed dispersal via the dung of herbivore guilds in little-altered environments. The present paper analyses seed dispersal via rabbit, fallow deer, red deer and cow dung in a Mediterranean dehesa (open woodland used for hunting and ranching) during the seeding season. Dung seed content was determined by the glasshouse cultivation of eight dung samples from each herbivore, collected fortnightly between February and August. The four herbivores disperse many seeds (spring averages are 6–15 seeds per gram of dry dung and maxima of 25–70) from a large number of species (totals between 52 and 78). Dispersal seems to be mainly determined by seed production of the plant comminity. This is reflected in (i) the dissemination of a high percentage of the species present in the dehesa, (ii) great seasonal variability, related to seed production, in the amount of seeds and number of species dispersed, and (iii) a high semi-quantitative similarity of seed content in the four types of herbivore dung throughout the year. There is also important quantitative variation that depends on animal traits and feeding habits. These results and the characteristics of species found in dung suggest the adaptation of plant species to the dispersal of their seeds via herbivore gut. This process may well have profound implications for vegetation dynamics and the evolution of plant traits.     

Seed Dispersal by Monkeys and the Fate of Dispersed Seeds in a Peruvian Rain Forest1

Primary seed dispersal by two species of monkeys and the effects of rodents and dung beetles on the fate of dispersed seeds are described for a rain forest in southeastern Perú. During the six-month study period (June–November 1992) spider monkeys (Ateles paniscus) dispersed the seeds of 71 plant species, whereas howler monkeys (Alouatta seniculus) dispersed seeds of 14 species. Spider and howler monkeys also differed greatly in their ranging behavior and defecation patterns, and as a consequence, produced different seed rain patterns. Monkey defecations were visited by 27 species of dung beetles (Scarabaeidae). Dung beetles buried 41 percent of the seeds in the dung, but the number of seeds buried varied greatly, according to seed size. Removal rates of unburied seeds by rodents varied between 63–97 percent after 30 d for 8 plant species. The presence of fecal material increased the percentage of seeds removed by seed predators, but this effect became insignificant with time. Although seed predators found some seeds buried in dung balls (mimicking burial by dung beetles), depth of burial significantly affected the fate of these seeds. Less than 35 percent of Brosimum lactescens seeds buried inside dung balls at a depth of 1 cm remained undiscovered by rodents, whereas at least 75 percent of the seeds escaped rodent detection at a depth of 3 cm and 96 percent escaped at 5 cm. Both dung beetles and rodents greatly affected the fate of seeds dispersed by monkeys. It is thus important to consider postdispersal factors affecting the fate of seeds when assessing the effectiveness of frugivores as seed dispersers.     

Potential and realised contribution of endozoochory to seedling establishment

Many plant species have been shown to be dispersed by large herbivores through endozoochory, but there have been few studies that have demonstrated the ecological significance of this dispersal method. Known quantities of seeds were fed to sheep and the emergence of seeds germinating from dung in the field and glasshouse were compared with emergence in the field after direct sowing. All 12 species studied showed potential endozoochory as they germinated and grew from dung in the glasshouse. Six species showed no emergence in the field from either dung or direct sowing. Of the remaining, three did not emerge from dung in the field but did from direct sowing indicating a very high cost of dispersal (one other species showed this pattern but not significantly), one emerged in the field from dung but at comparatively low rates and one showed no cost to endozoochory as it emerged equally well from dung or after direct sowing. The results indicate that potential endozoochory, as measured by germination from dung in the glasshouse, over-estimates likely rates of establishment in the field. The results also indicate that there are potentially high costs to endozoochory that have to be balanced against the benefits of long-distance dispersal by large herbivores.     

Seedling establishment after endozoochory in disturbed and undisturbed grasslands

Local plant community composition and structure may be largely influenced by germination and seedling establishment from seeds dispersed in animal dung, through seed input, gap creation and nutrient enrichment. With an experimental approach we assessed (1) what the effect is of dung deposition on the number of seedlings in the plant community 3 months and 1 year after dung deposition, (2) what the effect is of this seedling establishment on the local plant community characteristics such as species richness and (3) if this effect interacts with large-scale soil disturbance which removes the close canopy, such as sod-cutting. Viable seeds of monocotylous species were abundantly present in the dung, and dung deposition led to a higher number of monocotylous seedlings after 3 months. However, this effect was no longer significant after 1 year. Moreover, the proportion of viable monocotylous seeds that effectively established in the field after 3 months was less than 5%. A lower number of viable seeds of the less-dominant dicotylous species was dispersed in the dung but they had a higher cover and species richness after 1 year. This resulted in an increased total small-scale species richness and diversity after dung deposition through a decreasing dominance of monocotylous species. Sod-cutting had a pronounced effect on seedling emergence: viable seeds dispersed by dung had a higher probability of successful establishment when the dung was deposited in large gaps. This indicates that an increase of safe sites associated with disturbance strengthens the effects of seed dispersal and gap creation by dung deposition.     

Roles of cottony hairs in directed seed dispersal in riparian willows

Willows usually establish on wet substrates with fine sediments at sites that are created by large disturbances, but suitable microsites are spatially and temporally limited. Thus, we hypothesized that willow seeds are selectively dispersed to suitable microsites, such as those with a wet substrate, rather than unsuitable microsites, such as those with a dry substrate, with seedling establishment mediated by the cottony hairs attached to seeds (directed dispersal). To test our hypothesis, we compared several recruitment-related traits, including buoyancy, germination, and trapping at favorable microsites, in seeds of the riparian willows Salix sachalinensis and S. integra with and without cottony hairs in laboratory and field experiments. In both field and laboratory experiments, more seeds with cottony hairs were trapped in water and wet sand than in dry sand, in which no seeds of either species germinated. These results indicate that cottony hairs facilitate the recruitment of seeds to microsites favorable for seed germination and help seeds avoid unfavorable microsites. On the water surface, 17.6% of S. sachalinensis seeds and 68.0% S. integra seeds with cottony hairs floated for more than 6 days, whereas all seeds without cottony hairs sank immediately after being placed on the water surface. These results suggest that cottony hairs facilitate long-distance dispersal via flowing water and also help avoid germination under water, where willow seedlings fail to establish. Seeds of the two willow species were released from the cottony hairs and germinated immediately after the seeds were placed on wet sand, but not after placement on water or dry sand. These results suggest that the seeds are released from the cottony hairs when the hairs become wet and the seeds are striking to a suitable microsite for seedling establishment, such as wet sand. In riparian willows, the cottony hairs promote directed dispersal by moving seeds to discrete and predictable microsites where the seedling establishment is disproportionately high.     

Aggregated seed dispersal by wreathed hornbills at a roost site in a moist evergreen forest of Thailand

Hornbills (Bucerotidae) are widely regarded as important seed dispersers in tropical forests in Africa and Asia. We investigated how the roosting behavior of wreathed hornbills (Aceros undulatus) influences seed deposition and seedling survival at a roost site in a moist evergreen forest of Khao Yai National Park, Thailand. Fallen fruits and seeds were collected in traps that were placed around a roosting site for 14 months, and seedlings were monitored in adjacent quadrats for 3 years. Seedfall and seedlings of species represented in the hornbill diet occurred at significantly higher densities in the traps and quadrats located beneath the crown of the roosting tree than in those located beyond the crown. With the exception of Cinnamomum subavenium, the seeds and seedlings of most diet species rarely survived beyond the first year. The quality of hornbill dispersal to this roosting site may be poor due to the highly concentrated seedfall, which results in high seed and seedling mortality. However, the number of seeds deposited by each hornbill each day at roosting sites is relatively low. Wreathed hornbills are primarily scatter dispersers during the day and probably serve as agents of seed dispersal in the moist evergreen forest of Khao Yai.     

Germination and water dispersal of seeds from a threatened plant species Penthorum chinense

To conserve a threatened plant species (Penthorum chinense Pursh) in Japan, seed germination responses to pretreatment (imbibition and/or chilled), temperature and light, and seed dispersal by water were examined. The seeds collected from abandoned paddy fields in a warm temperate region, central Japan, germinated in light (14 h photoperiod; light 22°C, dark 21°C) after a moist-chilled treatment. After this pretreatment, the seeds germinated well at 10–25°C (optimum temperature 15°C), but did not germinate in darkness even at the optimum temperature. Most of the seeds floated on distilled water, but 20–60% of the seeds that were collected from several populations sank in distilled water, indicating dimorphism in seed dispersal by water. The floating and sunken seeds did not show significant differences in weight and germination rate within a population. The addition of a surface-active agent in distilled water submerged the seeds, indicating that the buoyancy of the seeds is attributable to an oil coating on the seed surface that enhances the interfacial tension on the seeds. Three times the number of seeds sank in river water collected from a rural area than in distilled water. A greater number of seeds also sank in water that had increasing concentrations of linear alkylbenzenesulfonate, which is a major component of synthetic detergents. This suggests that the water dispersal of this species is suppressed by surface-active agents, including detergents, in river water.     

Savanna fires increase rates and distances of seed dispersal by ants

Myrmecochory (seed dispersal by ants) is a prominent dispersal mechanism in many environments, and can play a key role in local vegetation dynamics. Here we investigate its interaction with another key process in vegetation dynamics—fire. We examine ant dispersal of seeds immediately before and after experimental burning in an Australian tropical savanna, one of the world’s most fire-prone ecosystems. Specifically, our study addressed the effects of burning on: (1) the composition of ants removing seeds, (2) number of seed removals, and (3) distance of seed dispersal. Fire led to higher rates of seed removal post-fire when compared with unburnt habitat, and markedly altered dispersal distance, with mean dispersal distance increasing more than twofold (from 1.6 to 3.8 m), and many distance dispersal events greater than the pre-fire maximum (7.55 m) being recorded. These changes were due primarily to longer foraging ranges of species of Iridomyrmex, most likely in response to the simplification of their foraging landscape. The significance of enhanced seed-removal rates and distance dispersal for seedling establishment is unclear because the benefits to plants in having their seeds dispersed by ants in northern Australia are poorly known. However, an enhanced removal rate would enhance any benefit of reduced predation by rodents. Similarly, the broader range of dispersal distances would appear to benefit plants in terms of reduced parent–offspring conflict and sibling competition, and the location of favourable seedling microsites. Given the high frequency of fire in Australian tropical savannas, enhanced benefits of seed dispersal by ants would apply for much of the year.     

Seed dispersal by the Florida box turtle (<Emphasis Type="Italic">Terrapene carolina bauri</Emphasis>) in pine rockland forests of the lower Florida Keys,United States

Seed dispersal by animals is one of the most important plant-animal mutualisms, but saurochory, the dispersal of seeds by reptiles, has received little attention. We investigated the role of the Florida box turtle (Terrapene carolina bauri) as a seed dispersal agent in pine rockland forests of the lower Florida Keys and examined the effect of turtle digestion on seed germination. We obtained seeds of 11 species with fleshy fruits and 2 species with non-fleshy fruits (a grass and legume) from the feces of 145 box turtles collected on Key Deer National Wildlife Refuge from 1999 to 2000. We planted the seeds of nine species and germination percentage (percentage of seeds that germinated during the experiment) varied from 10% to 80%. Comparative germination experiments were conducted with Thrinax morrissii, Serenoa repens, and Byrsonima lucida. We compared the germination percentage and germination rate (number of days from planting to seedling emergence) of seeds from three treatments (seeds recovered from feces, control seeds with pulp, and control seeds without pulp) and continued these experiments for up to 2 years. Passage through the box turtle digestive tract greatly enhanced the germination percentage and germination rate of S. repens, but decreased the germination percentage of B. lucida and T. morrissii, and decreased germination rate for T. morrissii. Subsequent destructive seed viability tests revealed that many ungerminated T. morrissii seeds remained viable, suggesting long-term seed dormancy may occur, even after passage through the turtle digestive system. In addition, the proportion of ungerminated seeds which remained viable was greater for seeds recovered from turtle feces than from control seeds with pulp. Furthermore, removal of fleshy pulp either manually or by the turtle digestive system may allow T. morrissii to escape insect predation.     

Fruit removal and postdispersal survivorship in the tropical dry forest shrub Erythroxylum havanense: ecological and evolutionary implications

We studied the relationship between the removal rate and the spatiotemporal availability of ripe fruits of the tropical deciduous shrub Erythroxylum havanense in western Mexico. We also evaluated the effects of dispersal on seed survival during the first stages of establishment. Fast and early dispersal should be favored in E. havanense, since propagules have more time to grow and accumulate resources before the beginning of the severe dry season. In general, high rates of fruit removal imply faster and earlier dispersal. Thus, plants producing large crops should benefit from high removal rates, which will increase the probability of successful establishment by their progeny. To characterize both individual and population fruiting patterns, we made daily counts of fruits on 51 plants arranged in six clumps of different sizes. The daily number of fruits removed per plant was higher for plants with larger initial crop sizes and larger numbers of ripe fruits on a given day, but decreased as clump size increased. Additionally, we monitored postdispersal survival and germination in an experiment manipulating seed density, distance from adult plants, and seed predation. Early establishment was independent of density or distance, and vertebrate seed predation was the main agent of seed mortality. Our results indicate that the critical variable with respect to fruit removal is the number of fruits a plant produces, large plants having higher dispersal rates. Large plants are also more likely to have more seeds escaping postdispersal seed predation.