Granivory in the Chilean matorral: extending the information on arid zones of South America

Comparative studies of granivory intensity across different arid and semi-and regions of the world have assumed homogeneity of conditions over large geographical areas, despite the existence of evident heterogeneities at local and regional scales For South America, previous evidence from the Monte desert, Argentina, showed that granivory intensity is low compared with other continents and that seed removal rates by taxon (granivore ranking) are in decreasing order ants > birds = small mammals In this work, we examined the assumed generality of this pattern for another South American region, the Chilean matorral We studied the differences between sparse and dense habitats of the matorral in granivory intensity throughout the year Although total seed removal did not differ between habitats, differences among granivores did occur Ants were the most important seed removers in both habitats, although they did not differ from birds in the sparse habitat Further, ants were the most important seed consumers during summer, but their seed removal rate is almost negligible the rest of the year Birds maintained an intermediate rate of seed consumption through the year Granivory intensity by small mammals was very low in both habitats during the whole year The granivory ranking obtained for the Chilean matorral was ants > birds > mammals, in dense habitat, and ants = birds > mammals, in sparse matorral Am abundance is correlated with seed removal throughout the year This relationship was not found in the other taxa We discuss possible causes for the differences among habitats, taxa, and seasons, as well as distinct granivore rankings from different continents     

Effect of spatial and temporal heterogeneity on granivory in the Monte Desert,Argentina

Abstract An important aspect of arid regions is the spatial heterogeneity resulting from differences among plant communities. There are process differences among different vegetation patches increasing variability in the functioning of the ecosystem. The purpose of this study was to estimate granivory, by studying variation of seed removal rates among patches at the local scale and variation according to seed type. We carried out experiments during four seasons in three plant communities in the Monte Desert, Argentina: ‘peladal’, mesquite forest and creosotebush. Seed trays were offered to ants, birds and rodents. Two types of commercial seeds (sunflower and millet) were used. We found that seed removal rate was significantly different among plant communities, among taxa (birds, ants and murid rodents), and among seasons, and that removal rate was higher for sunflower. Seed removal rates by murid rodents were higher than by birds or ants, and occasionally as high as those found in other deserts. The heterogeneity of seed removal patterns at the local level was as strong as that found between continents. Further studies may show similar local heterogeneity in other deserts of the world.     

Seed predation by birds and small mammals in semiarid Chile

We studied spatial and temporal patterns in foraging activity among diurnal birds and nocturnal mammals at a semiarid site in northern Chile using artificial foraging trays. Small mammals foraged more extensively under shrubs than in open microhabitats, but birds showed no such selection. Moreover, avian foraging was more extensive than that by small mammals in all seasons and both microhabitats. Avian foraging was highly seasonal, as many birds at our site migrate to the Andean prepuna or to Patagonia during the austral summer. Birds have tended to be overshadowed by small mammals and ants in studies of granivory, but this study suggests that their importance may be underestimated in some systems.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

Spatial variation in post-dispersal seed removal in an Atlantic forest: Effects of habitat, location and guilds of seed predators

Studies of post-dispersal seed removal in the Neotropics have rarely examined the magnitude of seed removal by different types of granivores. The relative impact of invertebrates, small rodents, and birds on seed removal was investigated in a 2,178 ha Atlantic forest fragment in southeastern Brazil. We used popcorn kernels (Zea mays—Poaceae) to investigate seed removal in a series of selective exclosure treatments in a replicated, paired design experiment that included forest understory, gaps, and forest edge sites. We recorded the vegetation around the experimental seed stations in detail in order to evaluate the influence of microhabitat traits on seed removal. Vertebrate granivores (rodents and birds) were surveyed to determine whether granivore abundance was correlated with seed removal levels. Seed removal varied spatially and in unpredictable ways at the study site. Seed encounter and seed use varied with treatments, but not with habitat type. However, seed removal by invertebrates was negatively correlated with gap-related traits, which suggested an avoidance of large gaps by granivorous ants. The abundance of small mammals was remarkably low, but granivorous birds (tinamous and doves) were abundant at the study site. Birds were the main seed consumers in open treatments, but there was no correlation between local granivorous bird abundance and seed removal. These results emphasize the stochastic spatial pattern of seed removal, and, contrary to previous studies, highlight the importance of birds as seed predators in forest habitats.     

Spatial variation in post-dispersal seed removal in an Atlantic forest: Effects of habitat,location and guilds of seed predators

Studies of post-dispersal seed removal in the Neotropics have rarely examined the magnitude of seed removal by different types of granivores. The relative impact of invertebrates, small rodents, and birds on seed removal was investigated in a 2,178 ha Atlantic forest fragment in southeastern Brazil. We used popcorn kernels (Zea mays—Poaceae) to investigate seed removal in a series of selective exclosure treatments in a replicated, paired design experiment that included forest understory, gaps, and forest edge sites. We recorded the vegetation around the experimental seed stations in detail in order to evaluate the influence of microhabitat traits on seed removal. Vertebrate granivores (rodents and birds) were surveyed to determine whether granivore abundance was correlated with seed removal levels. Seed removal varied spatially and in unpredictable ways at the study site. Seed encounter and seed use varied with treatments, but not with habitat type. However, seed removal by invertebrates was negatively correlated with gap-related traits, which suggested an avoidance of large gaps by granivorous ants. The abundance of small mammals was remarkably low, but granivorous birds (tinamous and doves) were abundant at the study site. Birds were the main seed consumers in open treatments, but there was no correlation between local granivorous bird abundance and seed removal. These results emphasize the stochastic spatial pattern of seed removal, and, contrary to previous studies, highlight the importance of birds as seed predators in forest habitats.     

Granivory rates by rodents, insects, and birds at different microsites in the Patagonian steppe

We studied the rates of seed removal by different granivores, in different microsites, at different times, in a Patagonian shrub steppe in South America. Granivory rates of exotic ( Phalaris canariensis ) and native ( Mulinum spinosum ) seed species were an order of magnitude lower than those reported for another cool desert from North America with comparable climatic conditions. Insects and rodents removed the greatest amount of seeds in comparison to birds. In general, there were no differences in seed removal across microsites, except at one sampling time (when rodents and insects removed more seeds from bare soil and beside tussocks in comparison to positions beside shrubs, while birds took similar amounts of seeds from all microsites). The length of the experiment differentially affected the granivory rate of different groups. Removal rates (per day) were significantly greater, and exhibited lower variability, when seeds were left for a longer period of time in the field (a month) than for a few days. Insects were more efficient at finding the seeds rapidly and rodents at depleting them; birds could not find or deplete many seeds in short periods of time. Rates of granivory decreased slightly but significantly as the summer progressed mainly due to a reduction of seed removal by birds and rodents but not by insects. Granivores removed an order of magnitude less native seeds than exotic seeds. These differences seemed to be related to palatability as M. spinosum seeds have more phenols, toxic concentrations of iron and copper, and lower dry matter digestibility, phosphorous, and nitrogen content, in comparison to P. canariensis seeds.     

小果野芭蕉种子散布和不同时空尺度上种子被捕食格局

 以分布在云南西双版纳地区的大型先锋草本植物小果野芭蕉(Musa acuminata)为研究材料,研究其种子初次散布过程和不同时空尺度上种子被 捕食格局。小果野芭蕉的成熟果实有75%在夜间被取食和传播,在白天消失的果实则占25%。蝙蝠是其最主要的种子传播者,鸟类在其种子传播 过程中也起到一定的作用。人工摆放种子试验结果显示小果野芭蕉种子的主要转移者是小型啮齿类(鼠类)和蚁类：在开放处理下3 d后转移率为 86%,排除蚁类(鼠类可进入)处理下种子转移率为69%以及排除鼠类(蚂蚁可进入)处理下种子被转移率为56%。季节、地点和生境均显著影响人工 摆放种子被转移强度：雨季显著高于旱季(p<0.001), 野芭蕉生境显著高于与其相连的自然森林和荒地(p<0.001),在人为干扰较少的补蚌自然 保护区显著低于西双版纳热带植物园和新山,而后两者之间并无显著差异(p>0.05)。同时,地点和生境以及季节、地点和生境都有显著的交互 作用。与相邻的森林和荒地相比,野芭蕉群落中种子被鼠类捕食的强度最大且受蚁类二次转移的比例最少,森林和荒地中种子被鼠类捕食的强 度相对较小且蚁类对种子的二次转移比例较高,从而更好地帮助种子逃避鼠类捕食。因此,依赖于食果动物(主要是蝙蝠, 也包括鸟类)的初次 散布是小果野芭蕉种子逃避捕食的关键。     

A multi-species assessment of post-dispersal seed predation in the central Chilean Andes

BACKGROUND AND AIMS: Post-dispersal seed predation in alpine communities has received little attention despite evidence that seeds removed by granivores can decrease plant recruitment into ecosystems. Moreover, few studies have assessed the effects of removal of seeds of a range of species after dispersal on the seeds remaining in ecosystems. A comparison was made of the magnitude of seed removal by ants and birds of nine different shrubby-, herbaceous- and cushion-plant species in the central Chilean Andes in order to assess the interactions between birds, ants and wind, and the types of seeds. METHODS: A total of 324 soil-covered plates, each containing 50 seeds of one species, were placed in the field at an altitude of 2700 m and assigned to one of four treatments: control, exclusion of ants, birds, and both. The design also allowed the effects of wind to be assessed. Seed removal from plates was monitored over 20 d. KEY RESULTS: Mean accumulative seed removal by granivores averaged over all nine species combined was 25%. However, large differences between species were evident, with limited seed removal (3-11%) in three herbaceous species (Alstroemeria pallida, Sisyrinchium arenarium, Pozoa coriacea), moderate (18-33%) in five species, including a shrub (Chuquiraga oppositifolia), two herbs (Taraxacum officinale, Rhodophiala rhodolirion), and two cushion-plants (Laretia acaulis, Azorella monantha), and substantial (78%) in the shrub Anarthrophyllum cumingii. The magnitudes of losses caused by birds compared with ants did not differ for the majority of species, although removal by birds was greater than by ants in A. cumingii, and smaller for C. oppositifolia. CONCLUSIONS: Post-dispersal seed removal is shown to be an important cause of decreased potential plant species recruitment into alpine ecosystems. The substantial differences in the magnitude of seed losses to ants and birds demonstrate the need for evaluation of seed removal on a wide range of species in any given ecosystem.     

The Bushmeat Harvest Alters Seedling Banks by Favoring Lianas, Large Seeds, and Seeds Dispersed by Bats, Birds, and Wind

We evaluated predictions that hunters favor lianas, large seeds, and seeds dispersed by bats, small birds, and mechanical means for seedling banks in central Panama. We censused 3201 trees in 20 1-ha plots and 38,250 seedlings in the central 64 m² of each plot. We found significant differences in the species composition of the seedling bank between nine protected sites in the Barro Colorado Nature Monument and 11 hunted sites in the contiguous Parque Nacional Soberanía. Lianas, species with large seeds, and species with seeds dispersed by bats, small birds, and mechanical means were all overrepresented at hunted sites. The latter two findings could also be evaluated relative to the species composition of reproductively mature adults for canopy trees. The tree species present in the seedling bank had significantly heavier seeds than the tree species present as adults at hunted sites but not at protected sites. The representation of seed dispersal modes among the species present in the seedling bank did not reflect pre-existing differences in the local species composition of adults. We hypothesize that hunting large seed predators favors large seeds by reducing predation and increasing survival. We also hypothesize that the harvest of large birds and mammals that disperse many seeds favors other species whose seeds are dispersed by bats, small birds, and mechanical means. This process also favors lianas because the seeds of disproportionate numbers of liana species are dispersed by wind.     

Predation of annual grass weed seeds in arable field margins

Seeds of three species of annual grass weeds (Alopecurus myosuroides, Avena fatua and Bromus sterilis) were placed in field margins around arable fields in a randomised block split-plot design experiment. The field margin vegetation was either sown or naturally regenerated and either cut or uncut. The seeds were either placed in cages designed to exclude small mammals and birds or were uncaged. The proportion of seeds removed was monitored on 10 occasions and mean seed loss was analysed. In general, a greater proportion of weed seeds was removed from uncaged trays in uncut swards, suggesting predation by small mammals, which inhabit tall grass. This effect was mainly due to removal of seeds of the two large-seeded species (A. fatua and B. sterilis), with A. fatua being especially favoured. It is therefore likely that small mammals play a role in the population dynamics of major crop weeds by feeding on their seeds in field margins, especially when these are dense and uncut.     

Rates of seed removal by ants at heath and woodland sites in southeastern Australia

Ants often appear to be important post-dispersal seed predators, particularly in Australia where they are exceptionally abundant and apparently can remove large quantities of seeds from the ground. Rates of seed removal by ants usually are measured by recording removal from artificial seed baits, but the reliability of this approach has not been tested, nor have there been many attempts to integrate the results with the activity of seed-eating ants. This paper describes the rates of seed removal, estimated using a baiting technique that is tested for its reliability, by the seed-eating ants in adjacent heath and woodland sites at Wilson's Promontory, Victoria. Ants removed up to 100% of seeds, but rates varied according to seed species, size of seed clumps, season, time of exposure, and other aspects of the baiting technique. Methodological guidelines are provided to make baiting conditions approximately those likely to occur in nature. Seed-eating ants, particularly species of Rhytidoponera, Chelaner and Pheidole, were by far the most important post-dispersal seed predators, and patterns of seed removal were directly related to their composition, abundance and foraging behaviour. Lygaeid bugs were also observed eating seeds, but there was no evidence of seed predation by rodents or birds. The results suggest that seed predation by ants can substantially deplete seed reserves: however, its actual effect on seedling recruitment is likely to depend on many factors including seed size, crop size, weather, timing and location of seed fall, availability of alternative food sources, patterns of seedling mortality, and fire, none of which have been adequately investigated.     

Seed dispersal in an African fig tree: birds as high quantity,low quality dispersers?

Abstract. Ficus burtt-davyi is a shrub or small tree found in the south and east of South Africa. Based on studies carried out in an area where the plant grows mainly as a rock-splitter, we first describe the nature and timing of the fruit resources it offers to potential dispersal agents, and then the animals that feed on the fruits. The figs are eaten by a diverse avian disperser assemblage, although just two species comprised about half of the recorded visits to the trees. Germination trials with seeds defecated by the birds found that they germinated more quickly than control seeds. Small terrestrial mammals and ants were also found to have a role in fig seed dispersal which may be disproportionate to the number of seeds they transport.     

Seed dispersal by ants in the semi-arid Caatinga of North-East Brazil

BACKGROUND AND AIMS: Myrmecochory is a conspicuous feature of several sclerophyll ecosystems around the world but it has received little attention in the semi-arid areas of South America. This study addresses the importance of seed dispersal by ants in a 2500-km(2) area of the Caatinga ecosystem (north-east Brazil) and investigates ant-derived benefits to the plant through myrmecochory. METHODS: Seed manipulation and dispersal by ants was investigated during a 3-year period in the Xingó region. Both plant and ant assemblages involved in seed dispersal were described and ant behaviour was characterized. True myrmecochorous seeds of seven Euphorbiaceae species (i.e. elaiosome-bearing seeds) were used in experiments designed to: (1) quantify the rates of seed cleaning/removal and the influence of both seed size and elaiosome presence on seed removal; (2) identify the fate of seeds dispersed by ants; and (3) document the benefits of seed dispersal by ants in terms of seed germination and seedling growth. KEY RESULTS: Seed dispersal by ants involved one-quarter of the woody flora inhabiting the Xingó region, but true myrmecochory was restricted to 12.8 % of the woody plant species. Myrmecochorous seeds manipulated by ants faced high levels of seed removal (38-84 %) and 83 % of removed seeds were discarded on ant nests. Moreover, seed removal positively correlated with the presence of elaiosome, and elaiosome removal increased germination success by at least 30 %. Finally, some Euphorbiaceae species presented both increased germination and seedling growth on ant-nest soils. CONCLUSIONS: Myrmecochory is a relevant seed dispersal mode in the Caatinga ecosystem, and is particularly frequent among Euphorbiaceae trees and shrubs. The fact that seeds reach micro-sites suitable for establishment (ant nests) supports the directed dispersal hypothesis as a possible force favouring myrmecochory in this ecosystem. Ecosystems with a high frequency of myrmecochorous plants appear not to be restricted to regions of nutrient-impoverished soil or to fire-prone regions.     

Seed removal patterns of pioneer trees in an agricultural landscape

The seeds of most tropical plants are dispersed by animals, many of which also act as seed predators. Shifts in animal community composition, such as those driven by the clearing of native vegetation, are therefore likely to drive changes in plant recruitment. We used manipulative experiments excluding ants, small rodents, and birds to quantify the relative impacts of these granivores on animal-dispersed pioneer trees (ADPT) in fragments of savanna vegetation and adjacent soy plantations in Brazil’s Cerrado. We found that ants were the main consumers of ADPT seeds, that the rates of seed removal varied with seed size, and that removal rates were higher in savanna fragments than in soy plantations. However, we also found significant interactions between habitat type, seed species, and the type of seed predator being excluded. Our results underscore how challenging it can be to predict the influence of human disturbances on the interactions between plant and animal communities. Because ants, rodents, and birds are Cerrado’s the main seed dispersers and granivores, seedling recruitment in Cerrado landscape mosaics will depend on how these distinct but related processes are each influenced by species-specific patterns of seed size and seed abundance.     

Comparison of seed removal by ants in vegetation on fertile and infertile soils

Plants which have seeds adapted for dispersal by ants (myrmecochores) are more common in some vegetation types than others. In Australia, myrmecochores are common in sclerophyll shrublands which are associated with infertile soils but not in forests on fertile soils. One hypothesis to explain this difference is that ants are less abundant or less active in seed removal on fertile soils. A test of this hypothesis was carried out on four sites in the Royal National Park, south of Sydney, by measuring the number of myrmecochorous seeds removed by ants in a 24 h period. It was found that the removal rates of seeds were slightly higher on fertile sites although this difference was not significant. The hypothesis that a lower removal rate on fertile soils could account for the lower percentage myrmecochory was therefore rejected. Apart from the gross removal rate of seeds by ants, other differences in ant behaviour might be important. It is possible that a greater proportion of ant species unfertile soils are seed predators or ineffective seed dispersers, and evidence obtained indicates that this is worthy of further research.     

Frugivory and seed dispersal by vertebrates in the Oriental (Indomalayan) Region

Current knowledge of frugivory and seed dispersal by vertebrates in the Oriental Region is summarized. Some degree of frugivory has been reported for many fish and reptile species, almost half the genera of non-marine mammals and more than 40% of bird genera in the region. Highly frugivorous species, for which fruit dominates the diet for at least part of the year, occur in at least two families of reptiles, 12 families of mammals and 17 families of birds. Predation on seeds in fleshy fruits is much less widespread taxonomically: the major seed predators are colobine monkeys and rodents among the mammals, and parrots, some pigeons, and finches among the birds. Most seeds in the Oriental Region, except near its northern margins, are dispersed by vertebrate families which are endemic to the region or to the Old World. Small fruits and large, soft fruits with many small seeds are consumed by a wide range of potential seed dispersal agents, including species which thrive in small forest fragments and degraded landscapes. Larger, bigger-seeded fruits are consumed by progressively fewer dispersers, and the largest depend on a few species of mammals and birds which are highly vulnerable to hunting, fragmentation and habitat loss.     

Are red imported fire ants facilitators of native seed dispersal?

Invasive ants threaten native communities, in part, through their potential to disrupt mutualisms, yet invasive species may also facilitate native species. The red imported fire ant (Solenopsis invicta) is one of the most conspicuous invasive ants in North America and its high densities, combined with its potential to displace native ants, have led to concerns that it may disrupt ant-plant seed dispersal mutualisms. We examined the potential of fire ants to disperse seeds in the longleaf pine ecosystem by comparing the removal of elaiosome-bearing seeds by fire ants versus native ants. A total of 14 ant species were observed removing seeds, with fire ants responsible for more than half of all removals. While fire ants were the dominant seed remover in this system, they did not remove significantly more seeds than would be expected based on their population density (46% of ground-dwelling ants). Moreover, red imported fire ants were similar to native ants with respect to distance of seed movement and frequency of moving seeds back to the nest. Areas of higher fire ant densities were found to have greater rates of seed removal by ants without a subsequent drop in seed dispersal by native ants, suggesting that fire ant-invaded areas may experience overall higher levels of seed dispersal. Thus, fire ants may actually facilitate dispersal of elaiosome-bearing plant species in the longleaf pine ecosystem.     

Comparing methods of weed seed exposure to predators

Several methods of seed exposure are used in seed predation studies, but how these methods influence the results remains poorly studied. In this article, two commonly used methods of seed exposure – seed cards and plasticine trays – were compared in the field and in the laboratory using three species of weed seeds. In the field, cards or trays with seeds were exposed either with or without cages to keep vertebrates out and either with or without impermeable roofs to provide protection from the weather. The overall seed retrieval from the control stands varied significantly between the methods of seed exposure, roof treatment and the species of seeds, and the scatter in the seed retrieval increased or decreased monotonically with the temperature or precipitation for some species of seeds. This indicates that the controls were more or less relevant depending on the weather conditions and species. The seed removal from cards varied between the species of seeds. The effect of exclosure cages indicated that invertebrates were the dominant seed predators of Capsella bursa‐pastoris, while in Poa annua and Stellaria media both vertebrates and invertebrates were important. Higher seed removal from plasticine trays compared to seed cards was found for all three species of seeds, and placing roofs over the seeds affected seed removal in C. bursa‐pastoris and P. annua. In the laboratory, seed consumption varied with the method of exposure in 10 out of 12 combinations of seed species and predator (two species of carabids and two species of isopods). The overall tendency was the opposite of the field observations: seed consumption was higher for seeds on filter paper and cards compared to seeds on tin trays. We conclude that seed cards are more convenient for short term studies in the field, while tin trays are useful in laboratory multi‐choice experiments. To measure the realistic consumption of invertebrate predators in the laboratory, filter paper seems to be the best option as it does not present an obstacle to predators eating the seed. Using roofs introduced further bias regarding the estimates of seed removal in the field and thus should be avoided.     

Seed removal,seed dispersers,and the allocation of tissues in Myrtaceae seeds

To avoid seed predation, plants may invest in protective seed tissues. Often related to seed size, allocation in seeds' physical defenses can also be influenced by dispersers. We explore the relationships between seed traits (seed mass and hardness) and seed removal in 22 Myrtaceae species of the Brazilian Atlantic Forest, a dominant and diverse fleshy-fruited taxon dispersed by birds, rodents, and other mammals. Our goal is to understand how seed traits influence seed removal rates, and whether dispersers can affect tissue allocation in the seed coat. Seeds were exposed to field removal experiments. In the laboratory, total seed mass and seed coat mass were obtained. To evaluate the influence of seed traits on removal, we performed Kruskal–Wallis and Simple Linear Regression tests. We assessed seed coat and seed mass covariation through standardized major-axis allometric regressions. Harder seeds were larger than softer ones. Seed traits affect removal rates, as tougher and heavier seeds had lower removal. Seed mass significantly predicts seed coat proportion in seven of the 14 species tested. Bird-dispersed species tend to exhibit lower proportions of seed coat as seed mass increases, whereas rodent-dispersed species apparently present the opposite trend, with seed coat proportion increasing with seed mass. Such difference may be caused by the contrasting seed predation pressure represented by birds and rodents. Energy allocation for defense, expressed in seed coat proportion, is greater in large seeds, as these are mostly dispersed by rodents whose propensity to cache and disperse seeds is greater for large and well-protected seeds.