Fruit size and shape: Allometry at different taxonomic levels in bird-dispersed plants

Summary The likelihood that a plant's seeds will be dispersed by fruit-eating birds may depend upon the size and shape of its fruits. Assuming that elongate fruits can be swallowed more easily than spherical fruits of equal volume and that plant fitness is enhanced by seed dispersal by many individuals and species of birds, natural selection should favour increasing fruit elongation with increasing fruit size in bird-dispersed plants. According to this view, this allometric pattern would be adaptive. Alternatively, fruit shape in bird-dispersed plants may be constrained by development or phylogeny. To determine whether there was any evidence to support the adaptive allometry hypothesis, we examined allometric relationships between length and diameter in fruits and seeds in a group of neotropical bird-dispersed plant species. Using the major axis technique, we regressed ln(diameter) on ln(length) for fruits and seeds at various taxonomic levels: (1) within individual trees ofOcotea tenera (Lauraceae) (2) among 19 trees within a population ofO. tenera, as well as among pooled fruits from multiple trees within 20 other species in the Lauraceae, (3) among 25 sympatric species within a plant family (Lauraceae) and (4) among 167 species representing 63 angiosperm families within a plant community in Monteverde, Costa Rica. At most taxonomic levels, a tendency for fruit length to increase more rapidly than fruit diameter among fruits (negative allometry) occurred more frequently than expected by chance. Estimated slopes of the regressions of fruit length on fruit diameter were < 1 within 15 of the 19 individualO. tenera trees, among tree means withinO. tenera, among pooled fruits within 16 of the 20 other species in the Lauraceae, among species means within the Lauraceae and among means of all bird-dispersed species in the lower montane forests of Monteverde. Seed allometry showed similar patterns, although for both fruits and seeds the broad confidence intervals of the slopes estimated by major axis regression overlapped 1 in many cases. Among the 63 Monteverde family means, fruit length and diameter scaled isometrically. Based on measurements of ontogenetic changes in fruit shape in a single species,O. viridifolia, we found no evidence that negative allometry in fruit shape within the Lauraceae was an inevitable consequence of developmental constraints. Instead, increasing elongation of fruits and seeds in certain plant taxa is consistent with adaptation to gape-limited avian seed dispersers. Contrary results from vertebrate-dispersed species from Malawi and Spain may reflect differences between the New and Old World in plant taxa, seed dispersers or evolutionary history.     

Exploring sub‐individual variability: role of ontogeny,abiotic environment and seed‐dispersing birds

• Within‐individual trait variation – otherwise known as sub‐individual variation – is an important component of phenotypic variation, with both a genetic and epigenetic basis. We explore its adaptive value and the effects of ontogeny and the environment on sub‐individual variability.
• We conducted a field study to analyse the effects of tree age, soil pH, soil water content and soil nutrients on sub‐individual variability in fruit size of hawthorn (Crataegus monogyna) in three sites in northwest Spain. Additionally, we examined how bird‐mediated selection influences average and sub‐individual variation in fruit size.
• Results show that average and sub‐individual variations in fruit size were related to fitness affecting seed dispersal. Older trees produced larger fruits, but tree age did not affect sub‐individual variation in fruit size. Abiotic environmental factors differently affected sub‐individual variation and average fruit size. Seed‐dispersing birds exerted correlated selection on average and variation in fruit size, favouring trees with larger and less variable fruit size at one site.
• Our work suggests that the fruit size variation within individual trees, the sub‐individual variation, is modified by abiotic environmental factors and, additionally, is an adaptive trait that responds to natural selection.

     

珍稀植物蒜头果野生植株结实量及果实特征研究

该研究对云南省广南县不同分布点的野生植株大小与结实量,果实、果核性状特征,果皮与果核性状间的关系进行了分析。结果表明:(1)野生成年植株个体间结实量差异大,单株结实量从几十个至几千个,变异系数可达136.38%。结实量与冠幅有正相关关系(R=0.592,P0.01),与胸径和树高无相关关系(P0.05)。(2)扁球型果实平均纵径37.10~40.36 mm,变异系数7.28%~8.65%;平均横径41.15~45.03mm,变异系数6.44%~9.31%;平均果实重量35.77~47.29 g,变异系数18.99%~21.44%。野生蒜头果果实大小差异明显,单个果实重量差别为3.4倍。(3)果核平均纵径27.50~31.69 mm,变异系数7.13%~10.99%;平均横径30.94~34.16 mm,变异系数6.47%~9.41%;平均果核重量14.03~18.77 g,变异系数17.37%~22.68%。单个果核重量差别为3.7倍。(4)平均果皮纵向厚度4.33~4.80 mm,变异系数20.22%~26.91%;平均横向厚度5.10~5.44 mm,变异系数12.92%~20.98%;平均果皮重21.62~28.51 g,变异系数20.01%~24.12%。该研究结果表明野生蒜头果单株结实量、果实和果核大小、果皮厚等表型性状存在广泛的多样性,其资源为人工定向培育和开发利用提供了较为丰富的选择材料。     

Crop Size and Fruit Neighborhood Effects on Bird Visitation to Fruiting Schefflera morototoni Trees in Puerto Rico1

Studies of zoochorous seed dispersal systems often consider crop size, yet seldom consider the kinds and amounts of fruits surrounding parent plants (the fruit neighborhood) when attempting to explain among‐plant variation in fruit removal. We studied avian frugivory at 24 Schefflera morototoni trees from February to May 1998 in central Puerto Rico. The number of fruits removed by avian seed dispersers per visit was similar among focal trees (typically 2–4). In contrast, visitation rate was highly variable (range: 0–71 visits per 4 h). We used multiple regression analyses to evaluate the relative roles of crop size (focal tree ripe fruit abundance) and fruit neighborhood variables (measured within 30 m of focal trees) in affecting visitation to focal trees by avian frugivores. Visitation rate was positively related to crop size (although this variable was only significant in one of four regression models considered) and negatively related to the presence or abundance of conspecific fruits, suggesting that trees competed intraspecifically for dispersers. Relationships between visitation and heterospecific fruits were mixed—some kinds of fruits appeared to enhance visitation to focal trees, while others seemed to reduce visitation. In most regression models, neighborhood variables had larger effects on visitation than focal tree fruit crop size. Our results highlight the important effects of local fruiting environments on the ability of individual plants to attract seed dispersers.     

Intratree Variation in Fruit Production and Implications for Primate Foraging

We tested the hypothesis that fruit quantity and quality vary vertically within trees. We quantified intratree fruit production before exploitation by frugivores at different heights in 89 trees from 17 species fed on by primates in Kibale National Park, Uganda. We also conducted a pilot study to determine if the nutritional value of fruit varied within tree crowns. Depending on the species and crown size, we divided tree canopies into 2 or 3 vertical layers. In 2-layered trees, upper crowns produced fruits that were 9.6–30.1% bigger and 0.52–140 times the densities of those from lower crowns, with one exception. Among 2-layered trees, upper crowns produced a mean of 46.9 fruits/m³ (median 12.1), while lower crowns produced a mean of 14.1 fruits/m³ (median 2.5). Among 3-layered trees, upper crowns produced a mean density of 49.9 fruits/m³ (median 12.5), middle crowns a mean of 16.8 fruits/m³ (median 6.6), and lower crowns a mean of 12.8 fruits/m³ (median 1.8). Dry pulp and moisture were systematically greater per fruit in the highest compared to the lowest canopy layers (22.4% and 16.4% respectively in 2-layered trees, 49.7% and 21.8% respectively in 3-layered trees). In 1 tree of Diospyros abyssinica, a pilot nutritional study showed that upper crown ripe fruit contained 41.9% more sugar, 8.4% more crude proteins, and 1.8 times less of the potentially toxic saponin than lower crown ripe fruit, but the result needs to be verified with more individuals and species of trees. We discuss the consequences of intratree variations in fruit production with respect to competition among frugivorous primates.     

Do seed-dispersing birds exert selection on optimal plant trait combinations? Correlated phenotypic selection on the fruit and seed size of hawthorn (<Emphasis Type="Italic">Crataegus monogyna</Emphasis>)

Seed-dispersing birds can be expected to optimize their energy intake through fruit selection and hence exert a positive selection on fruit pulp content, that is to say, favoring big fruits with small seeds. On the other hand they may select both, average fruit size and its variation. We addressed this issue by analyzing the phenotypic selection exerted by the dispersers of Crataegus monogyna on the fruit and seed size of this species. Fruit and seed size were analyzed at two ontogenic plant stages: the initial size of seeds (and their fruits) as future individuals and the mean size of fruits and seeds produced by adult plants. Fruit diameter and its within-individual variation are the actual targets of selection for maternal fruit traits, negatively affecting relative fitness, although total selection acts only on mean fruit size. For individual seeds, size selection is positive and directed at fruit diameter and seed length, while the remaining traits are subject to indirect selection. Birds exerted a correlational selection favoring big fruits with small seeds. Nevertheless the evolutionary consequences of this selection are expected to be limited by several factors. For example, the positive correlation between fruit and seed sizes, the existence of counteracting selective pressures exerted by other mutualistic or antagonist interactions and temporal or spatial changes in the environmental conditions that may alter the selective forces and even the optimum phenotype in each situation.     

Influence of between-year variation in the density of Rhus <Emphasis Type="Italic">trichocarpa</Emphasis> fruits on the removal of fruit by birds

The fruiting phenology and fruit removal patterns of Rhus trichocarpa Miq. (Anacardiaceae) were investigated in a warm-temperate secondary forest in Japan. Mature fruits of this species are dispersed by birds. Effects of fruit display size and canopy openness on fruit removal were investigated in years with different fruit densities (i.e., masting and non-masting years). Moreover, effects of increased canopy openness during winter on fruit removal were also investigated. Seasonal patterns of fruit removal were quite different between masting and non- masting years. In the non-mast year, fruits were removed by birds soon after maturation in the summer. In contrast, in the mast year, fruits were removed gradually by birds from summer to winter. Moreover, the rate of fruit removal was greater for trees with a larger display size in the non-mast year, whereas that was greater for trees with greater canopy openness in the mast year. Canopy openness increased in winter, and fruit removal in winter was enhanced in trees with a more open canopy only in the non-mast year. These observations strongly suggest that avian dispersers became satiated in the mast year, whereas fruit removal was enhanced in the non-mast year. In the mast year, although many fruits were not dispersed until winter, they were neither depredated nor rotten, and the long period of fruit removal by birds may have enhanced fruit dispersal. The large abundance of seedlings and saplings of this species in Japanese secondary forests suggests that this fruiting strategy is beneficial.     

Genetic structure in a population of a tropical tree Ocotea tenera (Lauraceae): influence of avian seed dispersal

We studied the influence of avian seed dispersal on the structuring of genetic diversity in a population of a tropical tree, Ocotea tenera (Lauraceae). The seeds of O. tenera are principally dispersed by four, relatively specialized, fruit-eating bird species (emerald toucanets, keel-billed toucans, resplendent quetzals, and three-wattled bellbirds). We found high genetic diversity within the overall population and significant, nonrandom structuring of that diversity among subpopulations. Subpopulations contained members of several sibling groups, and most saplings within subpopulations were shown not to be the progeny of adult trees within the same subpopulation. Our data indicate that O. tenera subpopulations are founded with several seeds from few maternal families, and that this mode of establishment is an important determinant of population genetic architecture.     

Facultative ripening in Hamelia patens (Rubiaceae): effects of fruit removal and rotting

Summary In Costa Rica individual Hamelia patens trees produce fruit throughout the year and experience dramatic changes in rates of fruit removal and rotting. During some moths, most fruits rot because they are not removed. Rotting fruits increase the probability that other fruits on the same infructescence will rot. When removal rates are high, fruits are taken as soon as their seeds become viable but before the fruit is completely ripe. Experimental removal of fruits produced significantly higher ripening rates than on control infructescences. This response allows Hamelia to ripen more fruit and increase the number of fruits taken when dispersers are abundant (e.g., during migration). The proximate mechanism of this response probably includes reallocation of energy conserved when partially ripe fruits are removed. Responding to fluctuating disperser populations likely increases dispersal success and may function as the ultimate cause.     

Small fruit problem in Citrus trees

Summary Physiological causes of the small fruit problem which occurs in certain trees of orange [Citrus sinensis (L.) Osbeck cv. Valencia] were investigated in terms of water relations and gas exchange of fruits during early fruit development as well as tree carbohydrate reserves. These data from cv. Valencia trees with and without a small fruit potential were compared with those of the large fruited cv. Navel. Neither fruit water potential nor fruit transpiration nor tree carbohydrate reserves appeared to be a cause of the small fruit. Yield records showed the small fruit to be assocaited with a large number of fruit per tree. However, fruits from cv. Valencia trees with a small fruit potential respired faster than either fruits of the same cultivar and size from trees without the physiological disorder or fruits of the same size of cv. Navel and also exceeded the dark respiration of the respective leaves. Hence, the small fruit problem in cv. Valencia was partly attributed to inefficient fruit photosynthesis, causing excessive respiration of each of a larger number of fruits compared to fruits of a tree of the same cultivar but without the physiological disorder. Fruits of cv. Valencia respired more in their 2 months longer lifetime on the tree relative to those of cv. Navel. It is concluded that orchard management methods will have to be investigated to balance the fruit load of the cv. Valencia tree utilizing the carbon available for fruit growth and to minimise stress during the early fruit development.     

Exploring the limiting causes of fruit production in the tropical tree Ipomoea wolcottiana Rose (Convolvulaceae)

Path analysis was used to evaluate a group of variables related to fruit production in the tropical tree Ipomoea wolcottiana (Convolvulaceae). The variables were tree size (trunk diameter), available phosphorus in the soil, floral phenotypic sex expression (i.e. gynoecium biomass allocation), flower production, fruits initiated, and length of the corolla and diameter of its opening. The path model explained 79.9% of the total variation, and the production of mature fruits was related in large part by the number of fruits initiated and by phenotypic sex expression. In turn, fruits initiated were related to corolla tube diameter and to a lesser extent to corolla tube length. The model could explain a low percentage of the variation (18.5%) of the number of fruits initiated, but once a fruit was initiated it had a high probability of maturing. Phenotypic sex expression related negatively to the number of flowers produced and positively, to almost the same degree, to available phosphorus in the soil. Tree size had a positive effect on flower production but a negligible effect on fruits initiated and matured. Soil phosphorus affected positively, although weakly all variables. The relationships between tree size, number of flowers and phenotypic sex expression suggested that abundant flowering in this species functions more for pollen donation than its reception.     

The effect of spatio-temporal variation in understanding the fruit crop size hypothesis

In order to evaluate the 'fruit crop size' hypothesis, we analyzed the effect of fruit availability on the number of visits by birds and on the proportion of removed fruits, and on how much of the variation in the proportion of removed fruits could be explained by spatio-temporal variability. Fieldwork was done on the coast of the state of Veracruz, Mexico, using the small tree Bursera fagaroides (Burseraceae) and its frugivorous feeding assemblage, on four hills during four years. The number of fruits produced, the number of visits by birds, and the proportion of removed fruits varied between years with more than half an order of magnitude. The number of available fruits was associated to both the number of visits by birds and the proportion of fruits removed from each plant individual, and supports the fruit crop size hypothesis. However, the hypothesis varies in strength depending on total fruit production by the population per year and per hill. The spatio-temporal variation of the system was the effect of variation in the intensity of fructification between years and hills; the inclusion of spatio-temporal variability helped to explain most of the variation found in our results. The latter approach may explain the explain the conflicting results found by other authors in studies done in only one year or at one site, where variation is greatly reduced. Considering the number of published reports, a meta-analysis seems to be in order to determine the existence of a general effect between fruit production, the number of visits by birds and the proportion of fruits removed. If this is possible, we hypothezise that the association between fruit production and the number of visits by birds should be of low intensity.     

Fruit production is influenced by tree size and size‐asymmetric crowding in a wet tropical forest

In tropical forest communities, seedling recruitment can be limited by the number of fruit produced by adults. Fruit production tends to be highly unequal among trees of the same species, which may be due to environmental factors. We observed fruit production for ~2,000 trees of 17 species across 3 years in a wet tropical forest in Costa Rica. Fruit production was modeled as a function of tree size, nutrient availability, and neighborhood crowding. Following model selection, tree size and neighborhood crowding predicted both the probability of reproduction and the number of fruit produced. Nutrient availability only predicted only the probability of reproduction. In all species, larger trees were more likely to be reproductive and produce more fruit. In addition, number of fruit was strongly negatively related to presence of larger neighboring trees in 13 species; presence of all neighboring trees had a weak‐to‐moderate negative influence on reproductive status in 16 species. Among various metrics of soil nutrient availability, only sum of base cations was positively associated with reproductive status, and for only four species. Synthesis Overall, these results suggest that direct influences on fruit production tend to be mediated through tree size and crowding from neighboring trees, rather than soil nutrients. However, we found variation in the effects of neighbors and nutrients among species; mechanistic studies of allocation to fruit production are needed to explain these differences.     

Bird Assemblage and Visitation Pattern at Fruiting Elmerrillia tsiampaca (Magnoliaceae) Trees in Papua New Guinea

Most tropical trees produce fleshy fruits that attract frugivores that disperse their seeds. Early demography and distribution for these tree species depend on the effects of frugivores and their behavior. Anthropogenic changes that affect frugivore communities could ultimately result in changes in tree distribution and population demography. We studied the frugivore assemblage at 38 fruiting Elmerrillia tsiampaca, a rain forest canopy tree species in Papua New Guinea. Elmerrillia tsiampaca is an important resource for frugivorous birds at our study site because it produces abundant lipid-rich fruits at a time of low fruit availability. We classified avian frugivores into functional disperser groups and quantified visitation rates and behavior at trees during 56 canopy and 35 ground observation periods. We tested predictions derived from other studies of plant–frugivore interactions with this little-studied frugivore assemblage in an undisturbed rain forest. Elmerrillia tsiampaca fruits were consumed by 26 bird species, but most seeds were removed by eight species. The most important visitors (Columbidae, Paradisaeidae and Rhyticeros plicatus) were of a larger size than predicted based on diaspore size. Columbidae efficiently exploited the structurally protected fruit, which was inconsistent with other studies in New Guinea where structurally protected fruits were predominantly consumed by Paradisaeidae. Birds vulnerable to predation foraged for short time periods, consistent with the hypothesis that predator avoidance enhances seed dispersal. We identified seven functional disperser groups, indicating there is little redundancy in disperser groups among the regular and frequent visitors to this tropical rain forest tree species.     

Does individual variation in fruit profitability override color differences in avian choice of red or white <Emphasis Type="Italic">Ilex serrata</Emphasis> fruits?

Although avian color preferences have been studied and documented in controlled experiments, they have not been demonstrated under natural conditions in most cases. We hypothesized that avian fruit choice reflects intraspecific variation in fruit characteristics other than color, rather than fruit color differences. By planting one Ilex serrata Thunb. (red form) and one I. serrata forma leucocarpa Beissner (white form), which produce red and white fruits, respectively, at each of five points, we examined the proportion of fruits removed per tree and fruit choice by three avian species based on fruit color and other fruit characteristics. The proportion of fruits removed increased with pulpy sugar concentration and fruit diameter, but it did not differ between fruit colors. The main foragers, resident brown-eared bulbuls Hypsypetes amaurotis, consumed fruits regardless of color, but correspondingly to fruit removal, and appeared to base their fruit choice on pulpy sugar concentration and fruit diameter rather than on color. In contrast, the minor foragers, migrant Daurian redstarts Phoenicurus auroreus (Pallas) and Siberian bluechats Tarsiger cyanurus (Pallas), tended to choose red fruits and were possibly attracted by them. In conclusion, fruit removal per tree reflected individual variation in fruit profitability more strongly than differences in fruit color, even though the individual variation was not remarkable. The importance of color in fruit choice differed based on species, residency status, and major/minor foragers.     

Patterns of fruit and seed production inBauhinia ungulata (Leguminosae)

Patterns of seed and fruit production ofBauhinia ungulata, a small tree legume indigenous in tropical America, were studied in Costa Rica. Only about 8% of flowers produced fruits. The average pod had 19 ovules and about two thirds of these began seed development, with mature pods containing an average of 9.7 mature undamaged seeds. About half of the mature pods were damaged by herbivores and within these, 27% of ovules or seeds had been eaten. Among trees there was no significant variation in pod production, but the number of ovules per pod and seed production per pod varied significantly. Within infructescences most pods were retained at middle positions. Within pods, the probability of an ovule developing into a seed increased toward the distal end. The pattern of seed and fruit production in this species agrees well in general with that reported for other neotropical legumes. The abortion of seeds and fruits can be regarded as a way of controlling maternal investment, and as a response to herbivory.     

Vertebrate frugivory and seed dispersal of a Chihuahuan Desert cactus

Vertebrate frugivory of fleshy-fruited plants may be very important for the recruitment of sexually derived seedlings if it represents the main mechanism of primary seed-dispersal.Opuntia rastrera produces fleshy fruits rich in water and sugars that are attractive to vertebrate frugivores. However, there is a very low rate of seedling recruitment in natural conditions. One of the causes that can influence this low recruitment is an insufficient seed dispersal due to a low fruit removal even under different resource (fruits) availability. To test this prediction, we studied the production and consumption of fruits in two consecutive years in two vegetation types: nopaleras (dense Opuntia-dominated scrublands) and grasslands with sparse populations of O. rastrera. Plant cover, fruit production and removal, and frugivore identity were recorded within each vegetation type in four randomly selected circular plots (7854 m²). Fruit production per area was higher in nopaleras in both years whereas per cladode production did not differ between vegetation types but differed between years in response to variation in precipitation. Fruit consumption by vertebrates was high (100%) and independent of spatial and temporal fruit availability. The intensity of fruit removal was inversely related to resource availability: it was faster in the less dense community (grassland) and in the driest year. Contrary to other studies with similar cacti, fruit removal by small mammals was insignificant whereas main consumers were birds and large mammals. Vertebrate frugivory represents the only mechanism of primary dispersal of seeds as all fruits are removed in about one month. Despite the high quantity of viable seeds (more than one million per ha in nopaleras and a tenth of that in grasslands) that are dispersed by frugivores after the consumption of about 300 kg of fruit per ha in nopaleras and a tenth of that in grasslands, the rare establishment of seedlings (about one seedling per three million of seeds produced) reported in the literature indicates that the interaction between O. rastrera and the disperser guild is indeed very asymmetrical. We speculate that the harsh conditions for cactus establishment found in this ecosystem demand a high investment in disperser rewards (fleshy fruits) to allow a very modest rate of sexually-derived seedling establishment.     

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

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

Seed predation of two alien Opuntia species invading Mediterranean communities

Invasibility depends on the interaction of the introduced species with the abiotic and biotic factors of the recipient community. In particular, the biotic resistance posed by native herbivores has been claimed to be of great importance in controlling plant invasion. We investigated fruit and seed predation of two exotic Opuntia species within and between Mediterranean communities in order to determine how patterns of predation matched patterns of invasion. Predators were small mammals, presumably mice, which could consume more than 50% of the seeds produced. Predators could be equally effective in consuming fruit and single seeds. O. maxima fruits were slightly preferred to O. stricta fruits, but predators did not distinguish between seeds. Seed predation was more intense in invaded than in non-invaded communities. However, there was a high spatial variation in seed predation that did not always match patterns of invasion, suggesting that seed predation alone is not a good predictor of community invasibility to Opuntia. According to these results invasibility to Opuntia is limited in some (but not all) communities by native mice. Seed losses by predation were high for both species. However, we estimated that more than 75% of seeds dispersed by birds to non-invaded areas are not predated.