Tracking Bat‐Dispersed Seeds Using Fluorescent Pigment1

Tracking the dispersal of seeds by fruit‐eating animals in tropical rain forests is crucial to further our understanding of plant–frugivore interactions and their impacts upon forest regeneration and plant population dynamics. We tested the feasibility of tracking bat‐dispersed seeds in a Philippine lowland rain forest with the help of fluorescent pigment. The powder was mixed with acetone and sprayed to ripe fruits of fig trees, i.e., Ficus septica and F. variegata. During nightly monitoring using a hand‐held ultraviolet lamp bat deposits (seed‐containing spat outs and feces) could successfully be detected. Distances and directions of deposit sites to the focal trees were recorded and seed shadow areas were analyzed. Bats dispersed most of the seeds less than 50 m away from the parent plants resulting in seed shadow areas < 0.30 ha in size. An in situ fruit preference experiment showed that fluorescent powder is unlikely to deter bats from feeding on ripe figs. In conclusion, the technique is simple, inexpensive, noninvasive, applicable to different fields of research and allows one to follow the fate of seeds from known sources.     

The diet of the capped langur (Presbytis pileata) in a moist deciduous forest in Bangladesh

Capped langurs (Presbytis pileata) in Madhupur National Park in north-central Bangladesh have an annual diet that is comprised largely of mature leaves (42% of 20,460 total feeding records). Among colobine monkeys, only some populations of red colobus (Procolobus badius) have a diet richer in mature leaves. New leaves (11%) and fruit (24%) are the other annually important dietary items. Seasonal breakdown of this diet, however, revealed that during the monsoon months of May through September theP. pileata diet is approximately 50% fruit, including pulpy ripe fruit. This is also the period of maximum fruit availability. Data on diet and food availability indicate that while capped langurs subsist on mature leaves during the dry season (80% of diet from November to March), they select fruit and new leaves and switch to these foods whenever they are available. These data support the hypothesis that the colobine feeding strategy is adapted to cope with seasonal food scarcity.     

Advantages of granivory in seasonal environments: feeding ecology of an arboreal seed predator in Amazonian forests

Specialized seed predators are uncommon in arboreal vertebrate assemblages, and the hypothesis that consuming seeds of immature fruits – which may be available for relatively long periods compared to mature fruit – could reduce seasonal food scarcity experienced by generalist frugivores remains largely untested. To test this hypothesis, we examined the diet and feeding ecology of bald‐faced saki monkeys Pithecia irrorata in a largely intact forest mosaic of southeastern Peru based on systematic monitoring of five habituated groups over a three‐year period and compared the relative availability of ripe and unripe fruits in their diet. Plant phenology data from individual tree crowns showed that, compared to ripe fruits, immature fruits were available in more tree species, in greater quantities, and for longer periods. Despite pronounced community‐wide seasonal changes in fruit production at our study area, feeding patterns of bald‐faced saki remained largely invariant: fruits comprised approximately 95% of the species’ monthly diet, with seeds alone accounting for 75%, with no major monthly dietary shifts. The flexible exploitation by this species of a consistently available food supply for which it faces little competition likely reduces foraging effort and consumption of less desirable foods, even during prolonged periods of overall fruit scarcity. The relative rarity of immature fruit specialists in tropical forests may reflect the fact that processing the hard pericarps and neutralizing the toxicities of immature seeds present substantial evolutionary hurdles that few arboreal vertebrate species have overcome.     

Seasonal variation in food availability and relative importance of dietary items in the Gambian epauletted fruit bat (Epomophorus gambianus)

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Regeneration niche partitioning in neotropical pioneers: effects of gap size,seasonal drought and herbivory on growth and survival

Adaptive trade-offs underlie the specialisation that permits habitat partitioning in species rich plant communities. We investigated the influence of the trade-offs that determine differences in growth and survival among six species of neotropical pioneer trees in gaps in semideciduous forest in Panama. Seedlings of Miconia argentea, Cecropia insignis, Luehea seemannii, Trema micrantha, Ochroma pyramidale and Croton bilbergianus were planted into artificial small (25 m²), medium (64 m²) and large (225 m²) gaps in secondary forest in the Barro Colorado Nature Monument. Trema and Ochroma suffered 50% mortality across all gap sizes, while Cecropia had high mortality only during the dry season and in the small gaps, and Miconia and Croton suffered low to zero mortality across all environments. The highest growth rates in large gaps were attained by Cecropia seedlings and in the smaller gaps by Miconia seedlings, although there were indications that Trema and Ochroma required gaps that were larger than any used in this study. Variation in growth and mortality could not be attributed to differences in foliar herbivore damage. Instead, there was strong evidence of a trade-off between maximum growth in the wet season and the ability to survive seasonal drought, particularly in small gaps. We conclude that variation in allocation in response to multiple limiting resources may be as important as allocation to growth and defence in determining the habitat preferences of neotropical pioneers.     

Feeding ecology of mandrills (Mandrillus sphinx) in campo animal reserve,Cameroon

A field study on the ecology of mandrills (Mandrillus sphinx) was carried out for 28 months in Cameroon. Fresh food remnants and large quantities of fresh feces were collected by following the groups. Analyses of these products indicated that fruit (including seeds), monocotyledonous plant leaves and insects (especially ants and termites), were frequently eaten. Mandrills mostly ate the plant and animal foods in the lower forest stratum and on the ground. Fallen seeds and monocotyledonous plant leaves were eaten more frequently in the minor fruiting season than in the major fruiting season presumably to compensate for the shortage of fresh fruit during the former. Daily travel distances were shorter during the minor fruiting season than during the major fruiting season, because in the minor fruiting season mandrills forage for small food items, such as the new leaves and piths of monocotyledons and fallen seeds which are sparsely distributed on the ground, while in the major fruiting season they search for widely distributed food such as fruit. The daily pattern of group movement and a food intake experiment suggest that mandrills move and feed continuously throughout the day. Use of fallen seeds and monocotyledonous plant leaves appears to enable mandrills to maintain a terrestrial life in the tropical rain forest. The feeding and ranging characteristics of mandrills are basically similar to those of other baboon species in open land, though their environments differ extremely.     

Do Meyer’s Parrots Poicephalus meyeri benefit pollination and seed dispersal of plants in the Okavango Delta,Botswana?

Recent studies of new world parrots repeatedly document, with few exceptions, that parrots are wasteful and destructive predispersal seed predators that are unlikely to contribute towards pollination and seed dispersal. Few detailed studies, however, have assessed the contribution of African parrots to forest ecology by quantifying the potential net benefit of seed and flower predation by parrots for most tree species in their diet. Due to the incidence of pollen on the heads of Meyer’s Parrots when feeding on Leguminosae flowers and the dispersal of viable seeds to the ground during seed predation, we compared destruction rates, when feeding on pods, fruits and flowers, with dispersal rates of viable seeds to the ground and frequency of head contact with reproductive apparatus to estimate net benefit from Meyer’s Parrot feeding activity. Meyer’s Parrots were not implicated in endo‐ or epizoochory, but they dropped uneaten fruit pulp and seeds to the ground during feeding bouts, thus providing ripe, undamaged seeds to secondary seed dispersers. This link with forest recruitment was weak, as all tree species utilized by Meyer’s Parrots either had more significant primary dispersal agents or were primarily wind‐dispersed. In most cases, the negative effect of seed predation outweighed any positive effects in terms of dispersal, whereby almost three times more seeds were consumed or destroyed than were dispersed to the ground. Significantly, only Sclerocarya birrea caffra recorded marginal net dispersal benefit from utilization by Meyer’s Parrots. Due to low relative resource abundance and high destruction rate, feeding activity on Berchemia discolor may be significant enough to influence its spatial distribution and abundance. Utilization of flowers of Kigelia africana and Adansonia digitata by parrots likely had a significant negative impact on pollination. Feeding on Acacia nigrescens flowers, however, was potentially advantageous to their pollination. We conclude that Poicephalus parrots are net consumers of ripe, undamaged seeds and flowers, thus having an overall negative impact on forest recruitment in subtropical Africa.     

Seed banks on Attalea phalerata (Arecaceae) stems in the Pantanal wetland, Brazil

Background and Aims

Seeds can accumulate in the soil or elsewhere, such as on the stems of palms when these are covered by persistent sheaths. These sheaths could act as a safe site for some species. Here, we studied whether persistent sheaths of the palm Attalea phalerata (Arecaceae) are available sites for seed accumulation in the Pantanal wetland of Brazil. We also investigated whether the composition, richness and diversity of species of seeds in the persistent sheaths are determined by habitat (riparian forest and forest patches) and/or season (wet and dry).

Methods

All accumulated material was collected from ten persistent sheaths along the stems of 64 A. phalerata individuals (16 per habitat and 16 per season). The material was then individually inspected under a stereomicroscope to record seed species and number.

Key Results

Of the 640 sheaths sampled, 65 % contained seeds (n = 3468). This seed bank included 75 species belonging to 12 families, and was primarily composed of small, endozoochoric seeds, with a few abundant species (Cecropia pachystachya and Ficus pertusa). Moraceae was the richest family (four species) and Urticaceae the most abundant (1594 seeds). Stems of A. phalerata in the riparian forest had 1·8 times more seeds and 1·3 times more species than those in forest patches. In the wet season we sampled 4·1 times more seeds and 2·2 more species on palm stems than in the dry season. Richness did not differ between habitats, but was higher in the wet season. Abundance was higher in forest patches and in the wet season.

Conclusions

Attalea phalerata stems contain a rich seed bank, comparable to soil seed banks of tropical forests. As most of these seeds are not adapted to grow in flooding conditions, palm stems might be regarded as safe sites for seeds (and seedlings) to escape from the seasonal flooding of the Pantanal.     

Diet composition of chimpanzees inhabiting the montane forest of Kahuzi,Democratic Republic of Congo

The diet of chimpanzees was investigated by direct observations, feeding remains, and fecal analysis from January 1994 to December 2000 in the montane forest of Kahuzi-Biega National Park. A total of 171 food items were identified, among which 156 items were plant materials belonging to 114 species from 57 taxonomic families. Chimpanzees consumed 66 species of fruits (62 species of pulps and four species of seeds). Results of fecal analysis showed that fig fruits were the most frequently eaten. Their seeds occurred in 92% of a total of 7212 chimpanzee fecal samples. The chimpanzees changed their diet according to seasonal and annual variations in both abundance and diversity of fruit species. However, they are very selective frugivores. Only a few pulp-fruit species are regularly identified in their fecal samples. During the rainy season, when ripe fruit was scarce, chimpanzees relied heavily on piths and leaves. They swallowed leaves of two species of Commelinaceae without chewing, probably for medical purposes. Animal foods were eaten infrequently. The montane forest of Kahuzi, where chimpanzees range up to 2600 m above sea level, may be the highest altitudinal limit ever recorded for their distribution. Compared to other chimpanzee habitats, Kahuzi has a low diversity of fruit species and the availability of a few pulp-fruit species may be critical to the survival of Kahuzi chimpanzees.     

Diets of the African Civet Civettictis civetta (Schreber, 1778) in selected coffee forest habitat,south‐western Ethiopia

Feeding ecology of the African civet, Civettictis civetta (Schreber, 1778), was studied in selected coffee forest habitat for two seasons between December 2012 and October 2013. The study was conducted in Limu Seka district, south‐west Ethiopia. Faecal analysis was employed to assess the diet spectrum, seasonal abundance and relative importance of food items. Over 55 different food items were identified from the analysis of 387 scat samples collected during the dry and wet seasons. In the coffee forest habitat, African civets showed omnivorous feeding habit with plant‐to‐animal diet biomass ratio of 1 : 1.36. A slight diet specialization was observed during the wet season (BA = 0.46) favouring animal prey. However, during the dry season, they showed generalist feeding habit (BA = 0.87) with more plant biomass in their diet. With over 64% occurrence and 14.4% biomass, coffee berries significantly contributed to the civets dry season plant diets. The excreted coffee beans, after civets ingested ripe coffee berries, are the tastiest product used by farmers for consumption and market. Seasonal collection of civet coffee from coffee forest floor economically supports the farmers while increasing the importance of civets in the habitat and hence contributing for its conservation.     

Changes in Baboon Feeding Behavior: Maturity-dependent Fruit and Seed Size Selection within a Food Plant Species

Despite considerable inter- and intraindividual variation in fruit and seed size in many plant species, researchers have given little attention to the relevance of the traits for primate fruit choice within a food plant species and its implications for tree regeneration. We studied feeding behavior and selectivity of olive baboons (Papio anubis) in the African locust bean (Parkia biglobosa, Mimosaceae), via direct observations of habituated groups and indirect evidence from leftovers of pods after feeding events. Olive baboons acted as both seed predators and dispersers for Parkia biglobosa. They fed on and destroyed unripe seeds, and swallowed intact ripe seeds when consuming mature fruit pulp. Predation rate was high, and only 10% of the seeds were dispersed. Predation and dispersal of seeds is linked to seed number and size. Digestible unripe seeds accounted for 10% of the unripe fruit mass, while indigestible ripe seeds made up 28% of the mature fruit mass. With these constraints, olive baboons increased food gain per fruit by selecting unripe pods containing a high number of large and heavy seeds. Consequently, only pods with fewer and smaller seeds remained for maturation. Thereafter, baboons fed on mature pods containing the smallest seeds, and exploited pods with more seeds to a greater extent than those with fewer seeds. Thus, fruits with small seeds and an intermediate seed number contributed the most to dispersal by baboons.     

Responses of squirrel monkeys to seasonal changes in food availability in an eastern Amazonian forest

Tropical forests are characterized by marked temporal and spatial variation in productivity, and many primates face foraging problems associated with seasonal shifts in fruit availability. In this study, I examined seasonal changes in diet and foraging behaviors of two groups of squirrel monkeys (Saimiri sciureus), studied for 12 months in Eastern Brazilian Amazonia, an area characterized by seasonal rainfall. Squirrel monkeys were primarily insectivorous (79% of feeding and foraging time), with fruit consumption highest during the rainy season. Although monkeys fed from 68 plant species, fruit of Attalea maripa palms accounted for 28% of annual fruit-feeding records. Dietary shifts in the dry season were correlated with a decline in ripe A. maripa fruits. Despite pronounced seasonal variation in rainfall and fruit abundance, foraging efficiency, travel time, and distance traveled remained stable between seasons. Instead, squirrel monkeys at this Eastern Amazonian site primarily dealt with the seasonal decline in fruit by showing dietary flexibility. Consumption of insects, flowers, and exudates increased during the dry season. In particular, their foraging behavior at this time strongly resembled that of tamarins (Saguinus sp.) and consisted of heavy use of seed-pod exudates and specialized foraging on large-bodied orthopterans near the forest floor. Comparisons with squirrel monkeys at other locations indicate that, across their geographic range, Saimiri use a variety of behavioral tactics during reduced periods of fruit availability.     

Does successful ovule development depend on its position within the pod? Examples from Neotropical Fabaceae

Previous studies have shown a nonrandom pattern of ovule fate probabilities according to ovule position in legume pods. Here, we tested how ovule position within the pods of two Fabaceae affects its fate. We expected higher proportion of well‐formed seeds near the fruit tips and of unfertilized and aborted ovules near fruit bases. We collected pods of Poincianella pyramidalis and Anandenanthera colubrina in a seasonal dry forest in northeastern Brazil and recorded total pod length and ovule number, position, and fate (unfertilized, well formed, aborted, and predated). The proportion of well‐formed ovules at fruit tips was significantly higher than at fruit bases in P. pyramidalis. The opposite pattern was found for unfertilized and aborted ovules, thus corroborating our hypothesis. However, the probability of seed predation in A. colubrina was significantly higher in pod tips, thus providing moderate support for our hypotheses. Interspecific differences in the patterns of ovule fate are likely to be driven by species pollination systems.     

Impact on fruit removal and seed predation of a secondary metabolite, emodin, in Rhamnus alaternus fruit pulp

There are two contradictory approaches to explaining the presence of secondary metabolites in ripe fruits. One holds that they evolved toward enhancing dispersal success (adaptive approach); the other claims that they evolved primarily to deter herbivores from eating leaves and seeds and that their presence in ripe fruits is a byproduct of that function (non‐adaptive approach). We tested the validity of three hypotheses of the adaptive approach that explain the presence of secondary metabolites in ripe fruits. We explored the current function of a secondary metabolite, emodin, in Mediterranean buckthorn (Rhamnus alaternus, Rhamnaceae) fruits by relating intraspecific variation and seasonal patterns of concentration to fruit removal and seed damage and by conducting feeding trials with captive birds presented artificial fruits that varied in emodin concentration. The concentration of emodin in the pulp of 10–13 Rhamnus plants from the same population was determined by HPLC every month during two fruiting seasons. Fruit removal by birds and seed predation by invertebrates and microbes were determined for the same plants. Emodin concentration rose during the first stages of ripening, reaching a peak before the fruits were ripe, and then decreased to a minimum when the fruits were ripe. No significant correlation between emodin concentration and ripe fruit removal rate among trees was observed in the first year, whereas in the second year the correlation was positive and significant. Thus, the impact of emodin on fruit selection varied between years, suggesting that emodin concentration does not solely govern fruit selection. A significant negative correlation was found in the first year between emodin concentration and seed predation during the first fruiting month. The yellow‐vented bulbul (Pycnonotus xanthopygos), a seed dispersing bird, distinguished between artificial foods that differed in emodin concentration (control, 0.001% and 0.002%), always preferring the lower concentration. In contrast, house sparrows, (Passer domesticus), a seed predator, did not detect such differences in emodin concentration but did distinguish between control foods and food with 0.005% and 0.001% emodin. We suggest that emodin has an ecological role, preventing seed predation by invertebrates and microbes without decreasing fruit removal by avian dispersers.     

Dispersal of a Human-Cultivated Crop by Wild Chimpanzees (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) in a Forest–Farm Matrix

With the conversion of natural habitats to farmland, nonhuman primates (hereafter primates) are increasingly exposed to agricultural crops. Although frugivorous primates are important seed dispersers that sometimes feed on agricultural fruits, evidence for dispersal of crops by primates is lacking. Here, we examine flexible feeding on cacao (Theobroma cacao) fruit and seed dispersal patterns by chimpanzees (Pan troglodytes verus) at Bossou in Guinea, and consequent cacao germination and survival. From direct observations, we confirm that cacao fruit is not an important food to chimpanzees, representing 0.23 % of focal animal feeding time. Chimpanzees ingest cacao pulp and either spit out the large seeds intact from unripe cacao fruit or swallow the seeds from ripe cacao fruits, which are consequently deposited in feces. From ecological surveys we show that chimpanzees distributed cacao extensively throughout their home range, at a mean distance of 407 m?±?SE 0.6 (N?=?90 clusters, range: 4–1130 m) from cacao plantations. As distance from the cacao plantation increased, cacao plants were more likely to survive. Other factors, including number of cacao plants in a cluster, plant height, and openness of the understory did not predict short-term cacao survival. Cacao plants within the forest did not produce fruit. By contrast, when chimpanzees deposited seeds in a plantation, cacao plants produced fruits as a result of farmers’ maintenance of the area. Our local-scale findings emphasize the complex behavioral and ecological interconnections between coexisting humans and primates in agricultural landscapes and generate interesting questions regarding primate niche construction and crop “ownership” related to who “plants” the crop.     

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.     

Seed dispersal effectiveness increases with body size in New Zealand alpine scree weta (Deinacrida connectens)

Weta are giant, flightless orthopterans that are endemic to New Zealand. Although they are known to consume fleshy fruits and disperse seeds after gut passage, which is unusual among insects, their effectiveness as seed dispersal mutualists is debated. We conducted a series of laboratory experiments on alpine scree weta (Deinacrida connectens) and mountain snowberries (Gaultheria depressa) to investigate how fruit consumption rates, the proportion of ingested seeds dispersed intact and weta movement patterns vary with weta body sizes. On average weta dispersed 252 snowberry seeds nightly and travelled at a rate of 4 m min^?1. However, seed dispersal effectiveness varied over three orders of magnitude and was strongly associated with body sizes. Smaller weta consumed few snowberry seeds and acted primarily as seed predators. On the other hand, the largest weta consumed and dispersed thousands of seeds each night and appear to be capable of transporting seeds over large distances. Overall results indicate that scree weta shift from being weakly interacting seed predators to strongly interacting, effective seed dispersers as they increase in size.     

Fruit removal of Coprosma quadrifida (Rubiaceae) by birds in south-eastern Australia

Abstract Removal of the fleshy fruits of Coprosma quadrifida by birds was examined in relation to fruit crop size, neighbouring fruit crop size and time in the season in montane wet sclerophyll forest in south-eastern Australia. On average, 84% of individual fruit crops were removed. Frugivores removed fruit in proportion to its abundance, indicating that they continually assess availability rather than respond to a critical abundance. The size of neighbouring fruit crops did not influence fruit removal rates, suggesting that there is little competition among neighbours for dispersers. The ripe fruits of C. quadrifida were eaten by 14 bird species, comprising 50% of the bird species trapped. Silvereyes (Zosterops lateralis) were the most abundant consumers of fruits. With the exception of the parrot, Platycercus elegans, all birds defecated seeds intact and had no influence on seed viability.     

Fruit Availability and Habitat Use by Chimpanzees in the Kalinzu Forest,Uganda: Examination of Fallback Foods

We studied seasonal change in habitat use by chimpanzees in the Kalinzu Forest, Uganda. The forest comprises various types of vegetation. For each vegetation type, we compared number of chimpanzees (per km²) that used the vegetation with fruit availability in different census periods. We estimated the number of chimpanzees by nest count and fruit availability via density of fallen fruit. The mixed mature forest contained a large amount of fruit during the high-fruiting season, but it decreased rapidly in the low-fruiting season. The number of chimpanzees also decreased in mixed mature forest in approximate proportion with fruit availability. In the Parinari-dominated mature and secondary forests, both fruit availability and number of chimpanzees were very low throughout the study. In the Musanga-dominated secondary forest, the number of chimpanzees increased toward the low-fruiting season, though the fruit availability decreased slightly. A multiple regression analysis showed that various fruits had significant effects on the number of chimpanzees during the high-fruiting season, while only Musanga leo-errerae had a significant effect during the low-fruiting season. The results suggest that the fruit of Musanga leo-errerae functions as a fallback food, and a combination of different vegetation types supports the chimpanzees in the Kalinzu Forest.     

Effects of bird ingestion on seed germination of two Prunus species with different fruit-ripening seasons

To evaluate the effects of ingestion by birds on seed germination under natural conditions, we carried out germination experiments in the field using seeds of two Prunus species that have different fruit-ripening seasons. Germination of seeds with the following three treatments was compared: ingested seeds, seeds excreted after feeding of fruits to birds; extracted seeds, seeds deliberately extracted from the fruit pulp; and intact fruit, seeds in untreated intact fruit. Many ingested and extracted seeds of both Prunus species germinated during the first spring, and the difference in germination percentage between ingested and extracted seeds was not significant. Many seeds in intact fruit of Prunus sargentii also germinated during the first spring, but those of Prunus ssiori did not germinate until the second spring. Pulp removal through bird ingestion enabled rapid germination for the autumn-fruiting P. ssiori, whose fruit pulp was not likely to be decomposed until the first spring. In contrast, the effects of ingestion were not striking for the summer-fruiting P. sargentii, whose fruit pulp is quickly decomposed.