The impact of consumption of fruit by vertebrate and invertebrate frugivores on the germination success of an Australian rainforest seed

Abstract The consumption of fruits by vertebrates and invertebrates can be both advantageous or detrimental to the survival of the seeds they contain. This study investigated the effect of fruit size and consumption of fruit pulp by rodents and beetles on the germination of the seeds of Acmena graveolens, a tropical rainforest canopy tree found in northern Australia. As fruit size increased, germination success and the amount of pulp remaining on the fruits was greater. When beetles were absent, germination success was highest when most of the pulp was removed by rodents, suggesting that they removed an inhibitor of germination. When beetles were present, germination success did not differ significantly across pulp categories, so beetles apparently enhanced germination in seeds with little pulp initially removed, possibly by further removal of fruit pulp. In this study, both rodents and beetles enhanced germination success of A. graveolens seeds by consumption of fruit pulp. Acting as facilitators of germination is a relatively unusual role for both these frugivores that are generally considered to act as seed predators or (in the case of rodents) dispersers.     

The Influence of Temporal Changes in Fruit Availability on Diet Composition and Seed Handling in Blue Monkeys (Cercopithecus mitis doggetti)1

We investigated the relation between temporally varying resources, diet composition, and seed-handling behaviors in a group of blue monkeys (Cercopithecus mitis doggetti) in a tropical montane forest of Rwanda. Changes in diet composition were related to concurrent phenological studies of fruit-producing trees, and density and abundance of tree resources within the monkey's home range. Fruit composed nearly 50 percent of the diet. Over 50 percent of the fruits eaten had juicy fleshy pulp. Observations of seed handling behavior provided insights into the role of these animals as potential seed dispersal agents. The monkeys moved the seeds of 29 species out of parent canopies by defecating seeds intact and by potentially carrying seeds in cheek pouches and dropping them later. Seeds of 18 species were found intact in fecal piles. Our study showed community-level phenology patterns did not indicate a decrease in fruit availability during the study period, but an analysis of the preferred fruits consumed by the monkeys showed distinct periods of low fruit availability. The study period included two dry seasons; only one of these produced a period of fruit scarcity for the animals. The animals employed different strategies during times of preferred fruit scarcity. They increased consumption of leaves and other fleshy fruits, and diet diversity increased, or became mainly seed predators and diet diversity decreased. The variable responses of these monkeys to changes in food availability highlights their dietary plasticity and imposes significant variations in their role as potential seed dispersers.     

Revealing secondary seed removers: results from camera trapping

This paper reports the results of the first study on secondary seed removal of seeds dispersed by Sykes’ monkeys (Cercopithecus albogularis) using camera traps in Africa. Patterns of primary seed dispersal are often superimposed by secondary conveyance, emphasising the need to study these secondary processes carefully. As the agents and mechanisms of seed dispersal are often concealed, being carried out by cryptic or nocturnal animals in dense vegetation, camera trapping was deemed a viable means to investigate secondary removal of seeds disseminated by C. albogularis in the Western Soutpansberg, South Africa. Camera traps were established at preferred feeding trees of the focal Sykes’ monkey group to identify animal species that remove seeds and fruits spat and dropped to the forest floor and seed removal observations were carried out. This method proved to be effective in identifying seed removers and also allowed to get indications about the quantities of seeds removed. Ten animal species were recorded visiting the trees, of which eight were observed removing seeds and fruits. Overall seed and fruit removal rates were high, indicating that the foraging behaviour of C. albogularis attracts many terrestrial frugivores.     

Linking frugivore activity to early recruitment of a bird dispersed tree,Eugenia umbelliflora (Myrtaceae) in the Atlantic rainforest

Seed dispersal by animals is a complex process involving several distinct stages: fruit removal by frugivores, seed delivery in different microhabitats, seed germination, seedling establishment, and adult recruitment. Nevertheless, studies conducted until now have provided scarce information concerning the sequence of stages in a plant's life cycle in its entirety. The main objective of this study was to evaluate the immediate consequences of frugivore activity for Eugenia umbelliflora (Myrtaceae) early recruitment by measuring the relative importance of each fruit‐eating bird species on the establishment of new seedlings in scrub and low restinga vegetation in the Atlantic rainforest, Brazil. We conducted focal tree observations on E. umbelliflora trees recording birds' feeding behaviour and post‐feeding movements. We also recorded the fate of dispersed seeds in scrub and low restinga vegetation. We recorded 17 bird species interacting with fruits in 55 h of observation. Only 30% of the handled fruits were successfully removed. From 108 post flight movements of exit from the fruiting trees, 30.6% were to scrub and 69.4% to low restinga forest. Proportion of seed germination was higher in low restinga than in the scrub vegetation. Incorporating the probabilities of seeds' removal, deposition, and germination in both sites, we found that the relative importance of each frugivorous bird as seed dispersers varies largely among species. Turdus amaurochalinus and Turdus rufiventris were the best dispersers, together representing almost 12% probability of seed germination following removal. Our results show the importance of assessing the overall consequence of seed dispersal within the framework of disperser effectiveness, providing a more comprehensive and realistic evaluation of the relative importance of different seed dispersers on plant population dynamics.     

The role of frugivorous birds in fruit removal and seed germination of the invasive alien Cotoneaster franchetii in central Argentina

Many invasive plant species have fleshy fruits that are eaten by native frugivorous birds which disperse their seeds and may facilitate their germination, playing an important role in plant invasion success. The fleshy‐fruited shrub Cotoneaster franchetii (Rosaceae) is an important invasive alien in the mountainous regions of central Argentina. To determine the role of avian frugivorous in fruit removal of this species, we conducted a frugivore exclusion experiment including bagged and unbagged branches in 75 plants of C. franchetii. At the end of the dispersal period, we compared the percentage of missing fruits (removed by birds + naturally dropped) in unbagged branches with the percentage of naturally dropped fruits in bagged branches. To assess whether any mechanism acting on seeds during their passage through bird guts (de‐inhibition by pulp removal and/or seed scarification) affects seed germination of this species, we compared percentage and speed of germination among seeds obtained from faeces of the native frugivorous Turdus chiguanco, from manually de‐pulped fruits, and from intact fruits. The percentage of missing fruits per shrub in unbagged branches was significantly higher than the percentage of naturally dropped fruits in bagged branches, suggesting that frugivorous birds play an important role in fruit removal of C. franchetii in the study area. Seeds from bird faeces and from manually de‐pulped fruits germinated in higher percentage and faster than seeds from intact fruits. Germination percentage and speed of seeds from manually de‐pulped fruits were significantly higher than those of gut‐passed seeds. These results indicate that T. chiguanco increases and accelerates seed germination of C. franchetii through pulp removal, but not through seed scarification. Overall, our findings indicate that native frugivorous birds facilitate the dispersal and germination success of C. franchetii, likely playing an important role in its invasion throughout the mountainous region of central Argentina.     

Seed Dispersal and Demography of Pioneer Trees: The Case of Hortia arborea

Abstract: This study investigates the relationships among seed dispersal, patterns of seedling recruitment and the spatial distribution of a pioneer tree (Hortia arborea, Rutaceae) in the Brazilian Atlantic forest. The study was carried out at Dois Irmãos Reserve, a 387.4 ha reserve in northeastern Brazil. Fruit shadow concentrated beneath parental trees (74.6 % < 6 m distant) and fruit removal averaged 5.6 % (0 ‐ 25.5 %). Agoutis (Dasyprocta prymnolopha, Rodentia) ate fruits in the forest and in captivity and were the only vertebrates recorded feeding on Hortia fruits. Agoutis, however, destroyed 86.7 % of the seeds they ingested. In addition, only one seed from an expected number of 1980 germinated in a treefall gap after it passed through the digestive tract of agoutis. Both seed germination and seedling recruitment were restricted to gaps, and occurred among seeds manually released from the pulp or among those from fruits that naturally rotted in gaps. Moreover, exposed seedlings were taller than those covered by plants in a gap. H. arborea appears to self‐maintain populations in the same patches of forest, which are delimited by a pool of old and newly created gaps. More specifically, there is successful recruitment in patches occupied by parents, resulting in dense clumps of H. arborea.     

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.     

Red-Tailed Guenons (Cercopithecus ascanius) and Strychnos mitis: Evidence for Plant Benefits Beyond Seed Dispersal

I report data collected on red-tailed guenon (Cercopithecus ascanius schmidti) fruit processing behaviors between June 1993 and April 1994. Red-tailed guenons consumed the fruit of Strychnos mitis in 542 of 2,930 fruit-eating events (FEEs). The monkeys spat out cleaned seeds of Strychnos mitis in a majority of these records (477/542; 88%); seeds were occasionally swallowed whole, but only when pulp was unripe (69/542; 12%). In 83% of the FEEs on Strychnos, the red-tailed guenons spat out seeds within 10 m of the removal site; they typically stayed in the same tree while processing fruit, and in 56% of the FEEs, they moved <1 m before spitting seeds. I monitored spat seeds to evaluate the impact of monkey fruit processing on seed fate. Results indicate that 83% of seeds spat out by the red-tailed guenons germinated, while only 12% of unprocessed seeds survived to germination (p < 0.01). Of the processed seeds that germinated, 60% survived to germination and seedling establishment, while only 5% of unprocessed seeds survived to seedling establishment (p < 0.01). Unprocessed seeds were also more likely to be attacked by seed predators (p < 0.01) and fungus (p < 0.01). Although there is generally high mortality in seeds/seedlings, mature trees of Strychnos mitis are found in groves of adults, under which dense populations of seedlings and saplings can occur. These data suggest that Strychnos mitis does not conform to expectations of the Janzen-Connell model of seed escape from parent trees. Instead, I suggest that by removing pulp, a process that results in a reduction of fungal pathogen attack, red-tailed guenons positively effect the seed survivorship of Strychnos mitis. Although this effect has been observed in pulp-cleaning ant species, it is a hitherto undescribed effect of primates on their fruit resources.     

Interactions between Ants and Non‐myrmecochorous Fruits in Miconia (Melastomataceae) in a Neotropical Savanna

The aim of this study was to evaluate the role of ants as secondary seed dispersers of six primarily bird‐dispersed Miconia species in the cerrados of southeastern Brazil. Vertebrate exclosure and seed germination experiments were performed for M. albicans, M. alborufescens, M. corallina, M. ferruginata, M. ibaguensis, and M. irwinii. Excluding vertebrates did not significantly alter fruit removal rate for any of the Miconia species relative to open controls. Fruits on stalks and fallen fruits were removed and transported to nests mainly by species of Atta, Acromyrmex, and Ectatomma (dispersal distance ranging from 0.1 to 45.2 m), while Camponotus ants tended to be observed removing the fruit pulp (seed cleaning) where the fruits were found. Seed manipulation by Atta decreased germination of M. irwinii, but not M. ferruginata. Germination did not occur in intact fruits, and thus seed cleaning was an important service provided by the ants. Ant nest soils did not inhibit germination of any of the Miconia species, suggesting they are a good substrate for long‐lived Miconia seeds. We conclude that ant activity could have important effects on the fate of Miconia seeds adapted for bird dispersal.     

Ant seed removal in a non-myrmecochorous Neotropical shrub: Implications for seed dispersal

This study investigated ant seed removal of Piper sancti-felicis, an early successional Neotropical shrub. Neotropical Piper are a classic example of bat-dispersed plants, but we suggest that ants are underappreciated dispersal agents. We identified eleven ant species from the genera Aphaenogaster, Ectatomma, Paratrechina, Pheidole, Trachymyrmex, and Wasmannia recruiting to and harvesting P. sancti-felicis seeds in forest edge and secondary forest sites at La Selva, Costa Rica. We also tested for differences in ant recruitment to five states in which ants can commonly encounter seeds: unripe fruit, ripe fruit, overripe fruit, bat feces, and cleaned seeds. Overall, ants harvested more seeds from ripe and overripe fruits than other states, but this varied among species. To better understand the mechanisms behind ant preferences for ripe/overripe fruit, we also studied how alkenylphenols, secondary metabolites found in high concentrations in P. sancti-felicis fruits, affected foraging behavior in one genus of potential ant dispersers, Ectatomma. We found no effects of alkenylphenols on recruitment of Ectatomma to fruits, and thus, these compounds are unlikely to explain differences in ant recruitment among fruits of different maturity. Considering that P. sancti-felicis seeds have no apparent adaptations for ant dispersal, and few ants removed seeds that were cleaned of pulp, we hypothesize that most ants are harvesting its seeds for the nutritional rewards in the attached pulp. This study emphasizes the importance of ants as important additional dispersers of P. sancti-felicis and suggests that other non-myrmecochorous, vertebrate-dispersed plants may similarly benefit from the recruitment to fruit by ants.     

Mechanisms affecting the fate of Prosopis flexuosa (Fabaceae,Mimosoideae) seeds during early secondary dispersal in the Monte Desert,Argentina

Abstract The fate of seeds during secondary dispersal is largely unknown for most species in most ecosystems. This paper deals with sources of seed output of Prosopis flexuosa D.C. (Fabaceae, Mimosoideae) from the surface soil seed‐bank. Prosopis flexuosa is the main tree species in the central Monte Desert, Argentina. In spite of occasional high fruit production, P. flexuosa seeds are not usually found in the soil, suggesting that this species does not form a persistent soil seed‐bank. The magnitude of removal by animals and germination of P. flexuosa seeds was experimentally analysed during the first stage of secondary dispersal (early autumn). The proportion of seeds removed by granivores was assessed by offering different types of diaspores: free seeds, seeds inside intact endocarps, pod segments consisting of 2–3 seeds, and seeds from faeces of one herbivorous hystricognath rodent, the mara (Dolichotis patagonum). The proportion of seeds lost through germination was measured for seeds inside intact endocarps, seeds inside artificially broken endocarps, and free seeds. Removal by ants and mammals is the main factor limiting the formation of a persistent soil seed‐bank of P. flexuosa: >90% of the offered seeds were removed within 24 h of exposure to granivores in three of four treatments. Seeds from the faeces of maras, on the other hand, were less vulnerable to granivory than were other types of diaspores. These results suggest that herbivory might be an indirect mechanism promoting seed longevity in the soil (and likely germination) by discouraging granivore attack. On the other hand, germination did not seem to have an important postdispersal impact on the persistence of P. flexuosa seeds in the soil. Both direct and indirect interactions between vertebrate herbivores and plants may foster P. flexuosa's seed germination in some South American arid zones.     

Chimpanzee (Pan troglodytes) Seed Dispersal in an Afromontane Forest: Microhabitat Influences on the Postdispersal Fate of Large Seeds1

We examined the postdispersal fate of large seeds (≥5 mm) dispersed by chimpanzees in an afromontane forest to evaluate aspects of the effectiveness of seed dispersal by chimpanzees, Pan troglodytes. We assessed the influence of six microhabitat characteristics on seed persistence and germination in seeds dispersed in chimpanzee feces and “wadges.” A total of 257 fecal samples and 56 wadges were located over a 4‐mo period by tracking a semi‐habituated chimpanzee community on day follows. Forty‐nine (19.1%) of the fecal samples contained large seeds from five different tree species. The majority of fecal samples with seeds contained seeds from the mature forest tree Olea capensis (Oleaceae) (83.7%). Forty‐two wadges (75%) contained seeds from the mature forest tree Syzygium guineense (Myrtaceae). Seeds were monitored at their deposition site for removal and germination up to 49 d following deposition. We collected data on the microhabitat surrounding each fecal and wadge sample. Multivariate analyses indicated that while fecal and wadge samples were not clustered into particular microhabitats, there was little overlap in the microhabitats in which wadges and fecal samples were deposited. Significantly more seeds persisted over 49 d in wadges (67.9%) than in feces (30.3%). Elevation was the only microhabitat variable determined to have a significant influence on seed persistence, whereas slope was determined to have a significant influence on germination.     

Consequences of southern cassowary (Casuarius casuarius,L.) gut passage and deposition pattern on the germination of rainforest seeds

In Australia's tropical rainforests the endangered southern cassowary, Casuarius casuarius, L., is the largest native frugivore and many plant species, because of the size of their fruits or seeds, are thought to be largely dependent on cassowaries for their dispersal. In this study we asked whether the contribution of cassowaries to plant recruitment extends beyond removing seeds from the vicinity of the parent. To do this we conducted germination trials involving 17 rainforest plant species to test whether cassowary consumption and seed deposition pattern alter germination probability or time to germination. Twenty‐four per cent of species showed changed germination probabilities, with one species showing no germination without cassowary consumption, and 35% showed changed time to germination. However these differences did not translate into any significant effects when considered across all species. We examined gut scarification, fruit pulp removal (de‐inhibition) and deposition in faecal material as mechanisms for changing germination success; each contributed to the changed success of individual species. There was a negative effect of seed clumping on five species. We conclude that cassowary consumption can modify germination performance in a minority of rainforest plants and that the effect is generally positive. Although the effect on large seeded species was small it is most likely to be important as the cassowary is the only animal in Australia able to internally process large numbers of these seeds.     

Seed handling in chimpanzees (Pan troglodytes) and redtail monkeys (Cercopithecus ascanius): Implications for understanding hominoid and cercopithecine fruit-processing strategies and seed dispersal

Primates are confronted with an array of constraints in feeding on fruit, including the removal of adhesive, energy-rich pulp from seeds. In this paper, I discuss how primates meet this challenge and present data on the fruit-processing and seed-handling behavior of chimpanzees and redtail monkeys in Kibale National Park, Uganda. These data are then related to these species' services as seed dispersers. Particular attention was paid to the methods by which primates removed pulp from seeds, the density of seed clumps that they deposited (by spitting, dropping, or defecating) to the forest floor, and the distance seeds were moved from parent trees. Distance and density differences in chimpanzee and redtail seed dispersal resulted from distinct fruit-processing and seed-handling methods. It was observed, in general, that redtail monkeys engaged in fine oral processing and were seed spitters: most seeds were dispersed in close proximity to parent trees (84% of spat seeds <10 m of parent tree), and deposited singly (100% seeds spat singly). In contrast, chimpanzees were coarse fruit processors and seed swallowers: seeds were defecated in denser clumps (e.g., a mean of 149 large seeds/dung sample and hundreds of small seeds/dung sample), far from parent trees. I evaluate the factors that shape patterns of fruit processing in hominoids and cercopithecines, and argue that the observed seed handling differences can be attributed to differences in digestive retention times, oral anatomy, and alternative mechanisms by which to avoid the cost of seed ballast. Am J Phys Anthropol 109:365–386, 1999. © 1999 Wiley-Liss, Inc.     

Effects of bird ingestion on seed germination of Sorbus commixta

To determine the effects of ingestion by birds on seed germination, we performed germination experiments in the field and laboratory with Sorbus commixta. The germination of four groups of seeds was compared: ingested seeds, seeds defecated in feces after feeding of fruits to birds; extracted seeds, seeds deliberately extracted from the fruit pulp; juiced seeds, seeds plus the juice of the pulp after seeds had been deliberately extracted from the pulp; intact seeds, seeds in untreated intact fruits. In the laboratory, intact and juiced seeds hardly germinated, but ingested and extracted seeds germinated. Thus, the pulp and its juice appeared to inhibit germination, but seeds could germinate without ingestion by birds once the seeds had been manually extracted from the pulp. In the field, intact fruits did not germinate in the first spring, because the seed was still covered with pulp. The pulp of intact seeds decomposed during the first summer, and thus, the seeds had the potential to germinate during the second spring. In fact, most intact seeds do not germinate during the second spring either, since they lose their viability during the first summer. Thus, under natural conditions, most seeds of Sorbus commixta cannot germinate without bird ingestion. Received: 5 July 1997 / Accepted: 7 November 1997     

Alternative seed-handling strategies in primates: seed-spitting by long-tailed macaques (Macaca fascicularis)

Summary The seeds in fruits consumed by primates may be chewed and digested, swallowed and defecated intact, or separated from the flesh and spat out. We show by a combination of close field observations and experiments with caged animals, that long-tailed macaques (Macaca fascicularis) have a remarkably low threshold of 3–4 mm for swallowing seeds and also that wild macaques rarely break them. The seeds of 69% of the ripe fruit species eaten are spat out intact or cleaned outside the mouth and dropped. Seed-spitting significantly reduces the swallowed food bulk and may lessen the risk of releasing seed toxins during mastication. However, it requires that even small fruits are processed in the mouth one or a few at a time. We suggest that fruit storage in the cheek pouches of cercopithecine monkeys allows them to spit seeds individually without excessively slowing fruit intake while feeding on patchily distributed fruit. In contrast, Apes and New World monkeys apparently swallow and defecate most ripe seeds in their diet and colobine monkeys break and digest them, detoxifying seed defenses by bacterial fermentation.     

Rapid Assessment of Seed Removal and Frugivore Activity as a Tool for Monitoring the Health Status of Tropical Forests

Ecological processes in tropical forests are being affected at unprecedented rates by human activities. Yet, the continuity of ecological functions like seed dispersal is crucial for forest regeneration. It thus becomes increasingly urgent to be able to rapidly assess the health status of these processes in order to take appropriate management measures. We tested a method to rapidly evaluate seed removal rates on two animal‐dispersed tree species, Virola kwatae and V. michelii (Myristicaceae), at three sites in French Guiana with increasing levels of anthropogenic disturbance. We counted fallen fruits, fruit valves, and seeds of each focal fruiting tree in a single 1 m² quadrat, and calculated two indices: the proportion of seeds removed and the proportion of fruits opened by mammals. They both provide an indirect and rapid assessment of frugivore activity. Our results showed a significant decrease in the proportion of removed seeds (16%) and fruits opened (19%) at the most impacted site in comparison with the other two sites (79% for seeds, 60% and 35% for fruits). This testifies to an increased impoverishment of the primate and toucan communities at the disturbed sites. This standardized protocol provides fast information about the health status of the community of seed dispersers and predators and of their seed removal services. It is time‐ and cost‐effective and is not species‐specific, allowing comparisons among sites or over time. We suggest using it with the pantropical Myristicaceae and any other capsule‐producing family to rapidly assess the health status of seed removal processes across the tropics.     

Spit it out: Monkeys disperse the unorthodox and toxic seeds of Clivia miniata (Amaryllidaceae)

Seeds of many Amaryllidaceae are unorthodox (recalcitrant) and toxic, and cannot survive ingestion, yet are packaged in brightly colored fruits suggestive of zoochory. Seed dispersal and germination of the understory amaryllid, Clivia miniata, were investigated in KwaZulu‐Natal, South Africa. Motion‐activated cameras revealed that samango monkeys (Cercopithecus mitis labiatus) are the primary disperser of C. miniata seeds. They eat the mesocarp and, to a lesser extent, the exocarp, and spit the large (13 mm diameter) seeds whole and cleaned onto the forest floor. Most seeds were dispersed farther than 1 m from the parent. Experimental removal of the fruit pulp had a small positive effect on the rate of seed germination, but did not affect subsequent seedling growth rates. The main advantages of monkey dispersal of Clivia seeds appear to be short‐distance dispersal away from the dense foliage of clumped parent plants and occasional long‐distance dispersal through cheek‐pouching behavior.     

Patterns of frugivory in the columnar cactus Pilosocereus leucocephalus

In the frugivory networks of many arid and semi‐arid Mesoamerican ecosystems, columnar cacti act as keystone species that produce fruits with a high content of water and nutrients attractive to numerous vertebrates. The aim of this investigation was to assess the fruit removal patterns of two guilds of frugivores on the fruits of the woolly torch Pilosocereus leucocephalus. We assessed fruit pulp removal in two ways: by estimating the consumption of seeds given the amount of pulp removed per visit and by estimating the percentage of pulp removal over time. We put exclosures on unripe, intact fruits to keep frugivores from removing material. Once ripe, we removed the exclosures and tracked animal visitation of 69 fruits using camera traps. We obtained a total of 2,162 hr of footage (14:47 hours of them with effective pulp removal). The highest number of visitors is that of diurnal species (n = 12, all birds) versus only four nocturnal (three bats, one rodent). The most effective species in pulp removal are birds. Bats play a modest role in frugivory of this cactus. The significance of this work is twofold: (a) birds and bats consume the fruit pulp of this cactus and likely disperse its seeds, and (b) although bats rank high in pulp removal effectiveness, birds as a guild far outweigh their importance in this system, as they are not only more frequent but also remove more pulp and seeds. Both groups are known to be important in cacti seed dispersal, and our findings are essential in understanding the population dynamics of the woolly torch and in elucidating its seed dispersal ecology.     

Interaction between ants and fruits of<Emphasis Type="Italic"> Guapira opposita</Emphasis> (Nyctaginaceae) in a Brazilian sandy plain rainforest: ant effects on seeds and seedlings

This study examines the dispersal system of Guapira opposita in a tropical sandy rainforest in southeast Brazil. Guapira trees produce small fruits with a high protein content (28.4%) and low lipid content (0.3%), and the plant is primarily dispersed by birds. Mature fruits of G. opposita can fall spontaneously with the pulp intact, or be dropped by birds with bits of pulp attached. In either case, ground-dwelling ants rapidly remove the fruits to their nest (93% after 12 h). The ponerine ants Odontomachus chelifer and Pachycondyla striata are the main seed vectors among the ants, and together account for 56% (20 of 36) of the ant-fruit interactions recorded on the forest floor. Individual workers of O. chelifer and P. striata transport single fruits to their nests. Bits of pulp are fed to larvae and worker nestmates, and intact seeds are discarded outside the nest. Germination success in Guapira is higher for cleaned seeds (pulp removed) than for seeds coated by pulp. Guapira seedlings and juveniles are more frequent close to Odontomachus nests than at sites without such nests. Soil samples from Odontomachus nests had greater penetrability, and higher concentrations of P, K, and Ca than random soil samples. Field experiments suggest that the association between G. opposita seedlings and O. chelifer nests can potentially render the plant some protection against herbivores. Results indicate that fruit displacement by ponerine ants play an important role in the biology of G. opposita seeds and seedlings in the sandy forest, and illustrate the complex nature of the dispersal ecology of tropical tree species.