Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

Responses of dispersal agents to tree and fruit traits in Virola calophylla (Myristicaceae): implications for selection

Variation in traits affecting seed dispersal in plants has been attributed to selection exerted by dispersal agents. The potential for such selection was investigated in Virola calophylla (Myristicaceae) in Manú National Park, Peru, through identification of seed dispersal agents and of tree and fruit traits significantly affecting the quantity of seeds dispersed. Seventeen bird and one primate species (the spider monkey, Ateles paniscus) dispersed its seeds. Spider monkeys dispersed the majority of seeds (a minimum of 83% of all dispersed seeds). Visitation by dispersal agents depended only on the quantity of ripe fruit available during a tree observation. In contrast, seed removal increased with both greater quantity of ripe fruit and aril: seed ratio. When analyzed separately, seed removal by birds increased only with greater aril: seed ratio, whereas seed removal by spider monkeys was affected by the quantity of ripe fruit and phenological stage. The finding that dispersal agents responded differently to some tree and fruit traits indicates not only that dispersal agents can exert selection on traits affecting seed dispersal, but also that the resulting selection pressures are likely to be inconsistent. This conclusion is supported by the result that the proportion of the seed crop that was dispersed from individual trees, which accounted for cumulative dispersal by all agents, was not influenced by any tree or fruit trait evaluated. Comparing these results with those from studies of V. sebifera and V. nobilis in Panama revealed that the disperser assemblages of these three Virola species were congruent in their similar taxonomic representation. In Panama the proportion of V. nobilis seed crop dispersed was related positively to aril: seed ratio and negatively to seed mass, a result not found for V. calophylla in Peru. The greater importance of dispersal by primates versus birds in V. calophylla, relative to V. nobilis, may explain this difference. Thus, variation in disperser assemblages at regional scales can be another factor contributing to inconsistency in disperser-mediated selection on plant traits.     

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 seed size of wind-dispersed pines (Pinus) on secondary seed dispersal and the caching behavior of rodents

Seeds and nuts dispersed by scatter-hoarding animals are relatively large compared to propagules dispersed by other means. Possible selective forces in the evolution of large seed size include the selectivity of foraging animals and the ways that food-storing animals treat seeds and nuts after harvest. Treatment by rodents, primarily yellow pine chipmunks ( Tamias amoenus ), of four species of pine seeds that vary in size was studied in the Carson Range of western Nevada. The pines, lodgepole pine ( Pinus contorta , 8.7 mg), ponderosa pine ( P. ponderosa , 55 mg), Jeffrey pine ( P. jeffreyi , 157 mg), and sugar pine ( P. lambertiana , 213 mg), produce winged seeds that are initially wind-dispersed but are gathered by rodents and cached in the soil. Radioactive scandium-46 was used to follow the fates of seeds of all fours species placed around three source trees during autumn 1998 to 2000. Rodents gathered the seeds of all four species, but they took fewer of the lodgepole pine seeds and only six lodgepole seed caches (n=2106 total caches) were found during the three years. Among the other three species, number of seeds per cache decreased with increasing seed mass. However, the product of number of seeds per cache and seed mass was similar for all species. Sugar pine seeds were cached slightly deeper than ponderosa and Jeffrey pine seeds. For the species examined, seed size appeared to have had little effect on several other attributes, including mean dispersal distance, substrate choice, and microhabitat choice. Large size decreases wind dispersibility of pine seeds, but secondary dispersal by scatter-hoarding rodents compensates for poor wind dispersal so that total dispersibility of large-seeded pines is not compromised.     

Effects of seed passage through slugs on germination

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

The Role of Arboreal Seed Dispersal Groups on the Seed Rain of a Lowland Tropical Forest1

Most tropical plants produce fleshy fruits that are dispersed primarily by vertebrate frugivores. Behavioral disparities among vertebrate seed dispersers could influence patterns of seed distribution and thus forest structure. This study investigated the relative importance of arboreal seed dispersers and seed predators on the initial stage of forest organization–seed deposition. We asked the following questions: (1) To what degree do arboreal seed dispersers influence the species richness and abundance of the seed rain? and (2) Based on the plant species and strata of the forest for which they provide dispersal services, do arboreal seed dispersers represent similar or distinct functional groups? To answer these questions, seed rain was sampled for 12 months in the Dja Reserve, Cameroon. Seed traps representing five percent of the crown area were erected below the canopies of 90 trees belonging to nine focal tree species: 3 dispersed by monkeys, 3 dispersed by large frugivorous birds, and 3 wind‐dispersed species. Seeds disseminated by arboreal seed dispersers accounted for ca 12 percent of the seeds and 68 percent of the seed species identified in seed traps. Monkeys dispersed more than twice the number of seed species than large frugivorous birds, but birds dispersed more individual seeds. We identified two distinct functional dispersal groups, one composed of large frugivorous birds and one composed of monkeys, drop dispersers, and seed predators. These groups dispersed plants found in different canopy strata and exhibited low overlap in the seed species they disseminated. We conclude it is unlikely that seed dispersal services provided by monkeys could be compensated for by frugivorous birds in the event of their extirpation from Afrotropical forests.     

Are possums important dispersers of large-seeded fruit?

The claim by Dungan et al. (2002) that “in many areas possums may be the only potential dispersal vector for large-seeded native species” is unsubstantiated. There is little evidence possums excrete viable seed of large-seeded fruit greater than 10 mm diameter, and seeds up to this size are dispersed by a suite of bird species. Nowhere in New Zealand are there likely to be possums in the absence of this suite of bird species.     

Effects of Seed Dispersal by Three Ateline Monkey Species on Seed Germination at Tinigua National Park, Colombia

We examined the effect of seed ingestion by three ateline primates: woolly monkeys, Lagothrix lagothricha; spider monkeys, Ateles belzebuth; and, red howler, Alouatta seniculus on germination rates and latency periods of seeds of several plant species in Tinigua National Park, Colombia. We collected dispersed seeds from feces and control seeds from the parental trees and washed them for germination trials. For the majority of plants, dispersed seeds germinated as well or better than control seeds did. Although spider monkeys depend more heavily on fruits than the other monkey species do, they were not more efficient than howlers or woolly monkeys at improving germination rates. A considerable proportion of the seeds dispersed by howlers and woolly monkeys showed reduced latency periods to germination, but spider monkeys showed less effect on reducing germination time. This result may be related to longer gut retention times, but such a trend has not been observed in other primate species. We conclude that, like many other primates, ateline monkeys are effective seed dispersers in terms of their effects on the seeds they swallow because they rarely decrease their germination rates. We discuss problems that make interspecific comparisons difficult, such as inappropriate control seeds and differences associated with germination substrates, and we stress the importance of studying other components of seed dispersal effectiveness.     

Macaques as Seed Dispersal Agents in Asian Forests: A Review

The role of primates in seed dispersal is well recognized. Macaques (Macaca spp.) are major primate seed dispersers in Asia, and recent studies have revealed their role as seed dispersal agents in this region. Here, we review present knowledge of the traits that define the role of macaques as seed dispersers. The size of seeds in fruit influences whether macaques swallow (0.5–17.1 mm; median: 3.0), spit (1–37 mm; median: 7.6), or drop (8.2–57.7 mm; median: 20.5) them. Dispersal distances via defecation are several hundreds of meters (median: 259 m, range: 0–1300 m), shorter than those achieved by some mammals and birds in tropical and temperate regions. However, macaques disperse seeds by defecation at comparable distances to omnivorous carnivores, and further than passerines. Seed dispersal distance by spitting is much shorter (median: 20 m, range: 0–405 m) than by defecation. Among Asian primates, seed dispersal distances resulting from macaque defecation are shorter than those for gibbons and longer than those for langurs. The effects of seed ingestion on the percentage and speed of germination vary among both plant and macaque species. The degree of frugivory, fruit/seed handling methods, seed dispersal distance, microhabitats of dispersed seeds, and effects of dispersal on seed germination vary seasonally and interannually, and long-term studies of the ecological role of macaques are needed. Researchers have begun to assess the effectiveness of seed dispersal by macaques, secondary dispersal of seeds originally dispersed by macaques, and the effects of provisioning on seed dispersal. Future studies should also test the effects of social factors (such as age and rank), which have received little attention in studies of seed dispersal.     

Seed germination in Chrysophyllum sp. nov., a large-seeded rainforest species in north Queensland: effects of seed size,litter depth and seed position

Leaf litter affects seed germination in many ways and past studies have shown greater impacts on relatively small seeds, both within and among species. In this shade-house experiment I examined the impact of forest litter on seed germination in Chrysophyllum sp. nov. (Sapotaceae), a large-seeded (2.4 g) rainforest tree from north Queensland. Seed mass varies more than 30-fold in this species, making it useful for studying the role of litter as a possible selective pressure in the evolution of seed size in large-seeded species. Seeds of varying size (small, medium, large) were sown in planting boxes containing one of three litter levels (low, medium, high) and placed either below or on top of the litter. Seed size and litter biomass had no significant impact on the number of germinating seeds or the time to germination but seeds placed below the litter germinated around twice as frequently, and 20% sooner, than seeds placed on top of the litter. There were no significant interactions between any of the three factors. This shade-house experiment suggests that leaf litter is not an important selective pressure in the evolution of seed size in this species. However, if litter disturbance under field conditions differentially affects the probability of seed germination in relation to seed size and/or litter biomass, then litter could still act as a selective pressure in the evolution of seed size in Chrysophyllum and other large-seeded species.     

Predicting dispersal of auto‐gyrating fruit in tropical trees: a case study from the Dipterocarpaceae

Seed dispersal governs the distribution of plant propagules in the landscape and hence forms the template on which density‐dependent processes act. Dispersal is therefore a vital component of many species coexistence and forest dynamics models and is of applied value in understanding forest regeneration. Research on the processes that facilitate forest regeneration and restoration is given further weight in the context of widespread loss and degradation of tropical forests, and provides impetus to improve estimates of seed dispersal for tropical forest trees. South‐East Asian lowland rainforests, which have been subject to severe degradation, are dominated by trees of the Dipterocarpaceae family which constitute over 40% of forest biomass. Dipterocarp dispersal is generally considered to be poor given their large, gyration‐dispersed fruits. However, there is wide variability in fruit size and morphology which we hypothesize mechanistically underpins dispersal potential through the lift provided to seeds mediated by the wings. We explored experimentally how the ratio of fruit wing area to mass (“inverse wing loading,” IWL) explains variation in seed dispersal kernels among 13 dipterocarp species by releasing fruit from a canopy tower. Horizontal seed dispersal distances increased with IWL, especially at high wind speeds. Seed dispersal of all species was predominantly local, with 90% of seed dispersing <10 m, although maximum dispersal distances varied widely among species. We present a generic seed dispersal model for dipterocarps based on attributes of seed morphology and provide modeled seed dispersal kernels for all dipterocarp species with IWLs of 1–50, representing 75% of species in Borneo.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

Seeds photoblastism and its relationship with some plant traits in 136 cacti taxa

Seed germination triggered by light exposure (positive photoblastism) has been determined in quantitative studies for numerous plant families and species. For Cactaceae, positive photoblastism is thought to be associated with life form and seed mass, but this association has never been evaluated. To explore hypotheses on associations between seed mass, seed dispersal, seed dormancy, life form, taxa and plant height with Relative Light Germination (RLG) in Cactaceae, we evaluated the effect of light on seed germination of 136 taxa. The taxa studied are native to several countries: México, Chile, Argentina, Brazil, Perú, USA, and Venezuela. Seed traits contrasted with RLG were life form, seed mass, seed dispersal, seed dormancy, adult plant height and taxon. We found some differences between RLG among taxa; Cacteae, Pachycereeae and Trichocereeae had higher RLG than Notocacteae. RLG was lower for seeds from taller than for shorter taxa, and lower for taxa with heavier seeds than for taxa with lighter seeds. Dispersal syndrome groups varied with RLG. RLG did not differ between cylindrical and globose taxa. Trends found here were in agreement with expectations for small-seeded species to have a light requirement to germinate more often than large-seeded species. This is the first time that cactus height is related to photoblastism. It is possible that seeds from tall plants are larger and thus have the capacity to produce taller seedlings than those from small plants, and that seedlings from large seeds with more resources have the ability to emerge from greater soil depths than those from small seeds.     

Scatter-hoarding rodents use different foraging strategies for seeds from different plant species

Seed predation and dispersal by scatter-hoarding rodents are key processes that determine seed survival, and thus, plant regeneration within forests. For decades, there has been much debate on the important effects of seed size (one of the most important seed traits) on rodent foraging preference. Furthermore, the possible selective forces in the evolution of seed size may be influenced by primary selectivity and how rodents treat seeds after harvesting. In this study, different-sized seeds from four species (Pinus armandii, Pinus densata, Abies sp., and Viburnum sp.) harvested by scatter-hoarding rodents were studied in an alpine forest in Southwestern China for two consecutive years. Our results showed that seed size influenced rodent foraging preferences, with bigger seeds being preferred over smaller seeds, within and across species. Rodents only removed and cached the larger seeds of P. armandii, and ate the seeds of the other three species in situ. Rodents are purely seed predators for these three species. For the cached seeds of P. armandii, significantly positive correlations were observed between seed size and dispersal distance among both primary and secondary cached seeds in 2006, but not in 2005. Our results indicate that among many coexisting species with widely different-sized seeds, scatter-hoarding rodents played important roles in the seed dispersal of the big-seeded species alone. This caching behavior could offset the limited seed dispersal of large-seeded and wingless species (P. armandii), in comparison with that of small winged seed species (P. densata and Abies sp.) and frugivore-dispersed species (Viburnum sp.).     

Diversity of seeds captured by interception exceeds diversity of seeds deposited in traps

Seed dispersal, a key process in terrestrial landscapes, is increasingly important in the face of habitat fragmentation and global climate change. Seed dispersal is also notoriously difficult to characterize, especially in species rich and spatially complex tropical forests. We contrasted assemblages of biotically dispersed seeds collected from four sites using two methods: deposition into seed traps and interception by the capture of frugivorous birds. We also compared seed deposition and interception with local fruit production. Species accumulation curves for seeds deposited in seed traps began to level off sooner than curves for seeds collected from birds captured in mist nets, and extrapolation showed significantly greater estimated species richness for seeds collected from birds than for those deposited in traps. Assemblages from birds and from traps at each site were quite different, with an abundance‐based similarity index of 0.64; this dissimilarity increases if bat‐dispersed seeds are included in the analysis. Common bird‐dispersed species were retrieved from both mist‐netted birds and from seed traps, but numerous locally fruiting understory species were recovered only from birds. We conclude that the sampling of seeds carried by birds provides a valuable complement to other methods of studying seed dispersal in species‐rich tropical forests by revealing relationships between specific dispersers and their seed plants and by creating a more complete account of species diversity of seeds being transported at a given site.     

Seed Swallowing in Tamarins: Evidence of a Curative Function or Enhanced Foraging Efficiency?

Nonhuman primates represent a major component of the frugivore biomass in several rain-forest communities. Although there is considerable evidence that prosimians, monkeys, and apes serve as dispersal agents for many tropical trees, little attention has been paid to the more basic questions of why certain species of primates swallow and void seeds, and what, if any, are the advantages to an animal of having a large, hard, bolus pass through its digestive tract. We examine patterns of fruit-eating and seed-swallowing in two species of free-ranging tamarins: Saguinus mystax and Saguinus geoffroyi. Fruits commonly eaten by tamarins contain large seeds surrounded by a fibrous and adhesive pulp or arilate seed coat. They generally swallow seeds and pulp together. Intact seeds are voided over a 1- to 3-h period. Measurements of 132 seeds naturally voided by Panamanian tamarins average 11.2 mm in length and 0.3 g. The greatest number of large seeds contained in the digestive tract of a single animal at one time was 13. In the case of moustached tamarins, we collected 220 seeds. Average seed length is 11.9 mm and average seed weight is 0.3 g. At the time of capture, one animal had 26 seeds in its digestive tract. In both tamarin species, there is evidence of sex-based differences in feeding behavior. Adult female moustached and Panamanian tamarins swallowed and voided seeds of larger size than adult males did. Seed size is positively correlated with pulp weight (p <. 001), therefore females were selecting food items with higher nutritional rewards than adult males did. Given their small body size and relatively short digestive tract, why do tamarins swallow such large seeds? Although several explanations are possible, we propose that the large number and size of undigested seeds continuously passing through the tamarin gut serve a curative role in mechanically dislodging and expelling intestinal parasites—Ancanthocephala (spiny-headed worms)—from their digestive tracts.     

Comparing the Dispersal of Large-seeded Tree Species by Frugivore Assemblagesin Tropical Montane Forest in Africa

We examined frugivore visitation and seed dispersal of five large-seeded (≥ 5 mm) tree species in tropical montane forest based on their occurrence in frugivorous primate diets: Ekebergia capensis, Olea capensis, Parinari excelsa, Prunus africana , and Syzygium guineense. A total of 21 frugivores in five assemblages ( i.e. , chimpanzees, cercopithecines, large-bodied birds, small-bodied birds, and squirrels) were observed over the study period (August 2006 and October–April 2007). We observed seed dispersal in four of five tree species studied; no dispersal was observed for P. excelsa . Frugivore assemblages did not visit tree species equally. Primates spent the most time in trees and had the largest group size. Large-bodied birds (LB) and chimpanzees dispersed the highest number of seeds per minute. LB and cercopithecines potentially dispersed the greatest number of seeds for E. capensis , and chimpanzees for S. guineense . Our analyses indicated that the mean fruiting duration of the focal tree, time in the tree, and number of species present are important predictor variables for seed dispersal by small- and large-bodied birds, and cercopithecines. The number of fruiting trees in the immediate vicinity of the focal tree further predicted seed dispersal for small-bodied birds (SB). Large-bodied birdseed dispersal also was predicted by time in tree by SB, and the number of individuals for SB and cercopithecines. Cercopithecines (CS) were further explained by the time in tree and number of species (SB & LB), and number of individuals for CS. Our study highlights the complexity of describing the relative importance of a frugivore assemblage to the dispersal of a tree species seeds.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.     

The capuchin, the howler, and the Caatinga: seed dispersal by monkeys in a threatened Brazilian forest

Regeneration of the Brazilian Caatinga forest may be restricted by the naturally low diversity and density of fruit-eating animals, which has been aggravated by local faunal extinction induced by human activities. We made a preliminary evaluation of the potential seed-dispersal role of capuchin (Cebus apella libidinosus) and howler monkeys (Alouatta caraya) in Serra da Capivara National Park. The monkeys dispersed at least 26 species. Alouatta dispersed larger seeds than Cebus, and the two species apparently dispersed seeds in different local habitats. Seed dispersal by monkeys potentially makes a significant contribution to Caatinga regeneration.     

Pulp–seed attachment is a dominant variable explaining legitimate seed dispersal: a case study on woolly monkeys

Seed dispersal is a mutualistic interaction in which frugivores gain nutrients and plants gain when seeds are transported to adequate places for establishment. However, this relationship is prone to deceit, for example, when frugivores spit-out seeds in the proximity of parental trees. Still, few hypotheses have offered explanations on why deceiving strategies are not widespread. In this study, I explore the importance of how difficult it is to mechanically separate the nutritious pulp from the seeds, as a factor that can explain the chance a seed has of being dispersed by woolly monkeys (Lagothrix lagothricha). I completed 1,440 h of focal animal follows during 2 years in order to quantify the chances plant species have of being dispersed. To do so, I evaluated the residuals from the relationship between the number of seeds manipulated and seeds dispersed by a population of woolly monkeys in Tinigua Park (Colombia). For 74 fruit species, I estimated how difficult it is to separate pulp from seeds as the time needed to separate the parts. An exponential model showed that this variable was able to predict 38% of the variation on dispersal probability, demonstrating that, when it is difficult to separate the pulp from the seeds, the probability of legitimate dispersal increases. However, when fruit parts were easy to separate, there was more variation in the outcome. My results suggest that many plants have evolved mechanisms (e.g., small seeds embedded in pulp, strong attachment, irregular seed surface, and thin pulp layer) that preclude deceit by frugivores.