Evolutionary trajectory of phenological escape in a flowering plant: Mechanistic insights from bidirectional avoidance of butterfly egg-laying pressure.

Phenological escape, whereby species alter the timing of life-history events to avoid seasonal antagonists, is usually analyzed either as a potential evolutionary outcome given current selection coefficients, or as a realized outcome in response to known enemies. We here gain mechanistic insights into the evolutionary trajectory of phenological escape in the brassicaceous herb Cardamine pratensis, by comparing the flowering schedules of two sympatric ecotypes in different stages of a disruptive response to egg-laying pressure imposed by the pierid butterfly Anthocharis cardamines, whose larvae are pre-dispersal seed predators (reducing realized fecundity by ~70%). When the focal point of highest intensity selection (peak egg-laying) occurs early in the flowering schedule, selection for late flowering dependent on reduced egg-laying combined with selection for early flowering dependent on reduced predator survival results in a symmetrical bimodal flowering curve; when the focal point occurs late, an asymmetrical flowering curve results with a large early flowering mode due to selection for reduced egg-laying augmented by selection for infested plants to outrun larval development and dehisce prior to seed-pod consumption. Unequal selection pressures on high and low fecundity ramets, due to asynchronous flowering and morphologically targeted (size-dependent) egg-laying, constrain phenological escape, with bimodal flowering evolving primarily in response to disruptive selection on high fecundity phenotypes. These results emphasize the importance of analyzing variation in selection coefficients among morphological phenotypes over the entire flowering schedule to predict how populations will evolve in response to altered phenologies resulting from climate change.

Emerging infectious disease triggered a trophic cascade and enhanced recruitment of a masting tree.

There are several mechanisms that allow plants to temporarily escape from top-down control. One of them is trophic cascades triggered by top predators or pathogens. Another is satiation of consumers by mast seeding. These two mechanisms have traditionally been studied in separation. However, their combined action may have a greater effect on plant release than either process alone. In 2015, an outbreak of a disease (African swine fever, ASF) caused a crash in wild boar (Sus scrofa) abundance in Bialowieza Primeval Forest. Wild boar are important consumers of acorns and are difficult to satiate relative to less mobile granivores. We hypothesized that the joint action of the ASF outbreak and masting would enhance regeneration of oaks (Quercus robur). Data from ungulate exclosures demonstrated that ASF led to reduction in acorn predation. Tree seedling data indicated that oak recruitment increased twofold relative to pre-epidemic period. Our results showed that perturbations caused by wildlife disease travel through food webs and influence forest dynamics. The outbreak of ASF acted synergistically with masting and removed herbivore top-down control of oaks by mobile consumers. This illustrates that the ASF epidemic that currently occurs across Europe can have broad effects on forest dynamics.

Acorn Crop, Seed Size and Chemical Defenses Determine the Performance of Specialized Insect Predators and Reproductive Output in a Mediterranean Oak.

Seed predation is an antagonistic interaction that negatively affects the performance of individual plants and can limit plant population dynamics. In animal-dispersed plants, crop size is an important determinant of plant reproductive success through its effect on seed dispersers and predators. Seed traits, such as size or chemical composition, can also increase the tolerance to seed predators or reduce their performance. We investigated the interaction between Quercus faginea and two specialized pre-dispersal insect seed predators (weevils and moths) during two years of contrasting crop size to determine the consequences of oak reproductive investment on seed production and insect performance. Crop size was 44% lower and acorns were 32% smaller in the second year, although acorn predation by insects was proportionally similar between both years at the population level. Individual trees producing larger crops showed a lower incidence of insect predators during the year of abundant acorn production, whereas trees producing bigger acorns experienced higher seed predation rates by insects, and acorns held more insect larvae in the low crop year. Competition between insects increased when acorn production was low, and higher tannin content in acorns further constrained the number of weevil larvae developing together in the same acorn. However, the abundance and size of insect larvae produced per tree were similar between the two crop years, and this was due to larvae often depleting acorn reserves when resources were low. Oak reproductive output increased nearly two-fold during the large crop year. Crop size variation, acorn production in a given year and acorn size and chemical composition seem to be important traits for reducing damage by insect predators in Quercus faginea and improve oak reproductive success.

Are all patches worth exploring? Foraging desert birds do not rely on environmental indicators of seed abundance at small scales.

BACKGROUND: Consumers should show strong spatial preferences when foraging in environments where food availability is highly heterogeneous and predictable. Postdispersal granivores face this scenario in most arid areas, where soil seed bank abundance and composition associates persistently with vegetation structure at small scales (decimetres to metres). Those environmental features should be exploited as useful pre-harvest information, at least to avoid patches predicted to be poor. However, we did not find the expected spatial association in the algarrobal of the central Monte desert by observing foraging seed-eating birds, a field technique influenced by how much they exploit visited patches. In this work we tested if the first stage of foraging by granivorous birds (patch visit, encounter or exploration) is positively associated with environmental indicators of patch quality by recording the removal of single seeds from 300 scattered experimental devices during seasonal trials. Spatial selectivity was analysed by comparing the structural characteristics of used vs. available microhabitats, and evaluated against bottom-up and top-down hypotheses based on our previous knowledge on local seed bank abundance, composition and dynamics. Their foraging activity was also explored for spatial autocorrelation and environmental correlates at bigger scales. RESULTS: Postdispersal granivorous birds were less selective in their use of foraging space than expected if microhabitat appearance were providing them relevant information to guide their search for profitable foraging patches. No microhabitat type, as defined by their vegetation structure and soil cover, remained safe from bird exploration. Analyses at bigger temporal and spatial scales proved more important to describe heterogeneity in seed removal. CONCLUSIONS: Closeness to tall trees, probably related to bird territoriality and reproduction or to their perception of predation risk, seemed to determine a first level of habitat selection, constraining explorable space. Then, microhabitat openness (rather than seed abundance) exerted some positive influence on which patches were more frequently visited among those accessible. Selective patterns by birds at small scales were closer to our predictions of a top-down spatial effect, with seed consumption creating or strengthening (and not responding to) the spatial pattern and dynamics of the seed bank.

Seed supply limits seedling recruitment of Eucalyptus miniata: interactions between seed predation by ants and fire in the Australian seasonal tropics.

Seed predation can cause substantial seed losses and influence plant population dynamics, but the impact depends on the extent to which populations are limited by seed availability or favorable microsites for recruitment. Harvester ants are the dominant post-dispersal seed predators in Australia's tropical savannas, and their abundance and foraging efficiency, as well as the availability of seed and microsites, are affected by fire history. We undertook a predator-exclusion experiment to examine the interactive effects of fire history (no fire compared with annual burning over 5 years) and seed predation by ants on seedling establishment of the dominant savanna tree, Eucalyptus miniata, in northern Australia. Despite its large seed size, the rate of removal (~ 20-60%) was similar or higher than typically reported for eucalypts, although it was lower than that recorded for the smaller seeds of the co-occurring E. tetrodonta. Seed predation rates were twice as high in annually burnt compared to unburnt sites, but there was no significant difference in the proportion of seedlings that emerged from the initial seed available. Seedling emergence in both regimes was low, representing < 7% of seed available after harvesting. About one-third of emergent seedlings were still alive during the middle of the following dry season. Our results indicate that seedling recruitment in E. miniata is limited by both seed supply and microsite availability. However, seed predation by ants reduces the likelihood of seedling establishment from low to virtually zero, which suggests that it plays a potentially important role in the population dynamics of savanna eucalypts.

Difference on cone size preferences between two coniferous species by Great Spotted Woodpecker (Dendrocopos major).

The number of species that specialize in pre-dispersal seed predation is relatively small. Examples of specialized pre-dispersal seed predators adapted to feeding on closed cones include vertebrate species like Crossbills, Squirrels, Nutcrackers and Woodpeckers. Seed predation selects against certain phenotypic features of cones and favors another phenotypic features. In this study, we document preferences of the Great Spotted Woodpecker (Dendrocopos major) for specific traits in the cones of Norway spruce (Picea abies) and Scots pine (Pinus sylvestris). We found that the Great Spotted Woodpecker prefers to feed on medium sized Norway spruce cones. The results suggest a disruptive selection that favors the extreme cone lengths in Norway spruce. In Scots pine, the woodpeckers avoided cones with large apophyses. Further, the selectivity for the specific characteristics of the cones is probably related to the configuration of the anvil, a place at which woodpeckers extract seeds from the cones. We think that the Great Spotted Woodpecker preferences in relation to the morphological characteristics of cones are a key to the design of the anvil in order to maximize the use of it as a tool for processing cones of both the Norway spruce and the Scots pine.

Contrasting effects of ploidy level on seed production in a diploid tetraploid system.

Previous studies demonstrated the effects of polyploidy on various aspects of plant life. It is, however, difficult to determine which plant characteristics are responsible for fitness differences between cytotypes. We assessed the relationship between polyploidy and seed production. To separate the effects of flowering phenology, flower head size and herbivores from other possible causes, we collected data on these characteristics in single flower heads of diploid and tetraploid Centaurea phrygia in an experimental garden. We used structural equation modelling to identify the main pathways determining seed production. The results showed that the relationship between polyploidy and seed production is mediated by most of the studied factors. The different factors acted in opposing directions. Wider flower heads displayed higher above the ground suggested higher seed production in diploids. In contrast, earlier flowering and a lower abundance of herbivores suggested higher seed production in tetraploids. However, because phenology was the strongest driver of seed production in this system, the sum of all the pathways suggested greater seed production in tetraploids than in diploids. The pathway linking ploidy level directly to seed production, representing unstudied factors, was not significant. This suggests that the factors studied likely are drivers of the between-cytotype differences. Overall, this study demonstrated that tetraploids possess overall higher fitness estimated as seed production. Regardless of the patterns observed here, strong between year fluctuations in the composition and diversity of insect communities have been observed. The direction of the selection may thus vary between years. Consequently, understanding the structure of the interactions is more important for understanding the system than the overall effects of cytotype on a fitness trait in a specific year. Such knowledge can be used to model the evolution of species traits and plant-herbivore and plant-pollinator interactions in diploid-polyploid systems.

Seed preferences by rodents in the agri-environment and implications for biological weed control.

Post-dispersal seed predation and endozoochorous seed dispersal are two antagonistic processes in relation to plant recruitment, but rely on similar preconditions such as feeding behavior of seed consumers and seed traits. In agricultural landscapes, rodents are considered important seed predators, thereby potentially providing regulating ecosystem services in terms of biological weed control. However, their potential to disperse seeds endozoochorously is largely unknown. We exposed seeds of arable plant species with different seed traits (seed weight, nutrient content) and different Red List status in an experimental rye field and assessed seed removal by rodents. In a complementary laboratory experiment, consumption rates, feeding preferences, and potential endozoochory by two vole species (Microtus arvalis and Myodes glareolus) were tested. Seed consumption by rodents after 24 h was 35% in the field and 90% in the laboratory. Both vole species preferred nutrient-rich over nutrient-poor seeds and M. glareolus further preferred light over heavy seeds and seeds of common over those of endangered plants. Endozoochory by voles could be neglected for all tested plant species as no seeds germinated, and only few intact seeds could be retrieved from feces. Synthesis and applications. Our results suggest that voles can provide regulating services in agricultural landscapes by depleting the seed shadow of weeds, rather than facilitating plant recruitment by endozoochory. In the laboratory, endangered arable plants were less preferred by voles than noxious weeds, and thus, our results provide implications for seed choice in restoration approaches. However, other factors such as seed and predator densities need to be taken into account to reliably predict the impact of rodents on the seed fate of arable plants.

Geographic consistency and variation in conflicting selection generated by pollinators and seed predators.

BACKGROUNDS AND AIMS: Floral traits that attract pollinators may also attract seed predators, which, in turn, may generate conflicting natural selection on such traits. Although such selection trade-offs are expected to vary geographically, few studies have investigated selection mediated by pollinators and seed predators across a geographic mosaic of environments and floral variation. METHODS: Floral traits were investigated in 14 populations of the bumble-bee-pollinated herb, Pedicularis rex, in which tubular flowers are subtended by cupular bracts holding rain water. To study potentially conflicting selection on floral traits generated by pollinators and florivores, stigmatic pollen loads, initial seed set, pre-dispersal seed predation and final viable seed production were measured in 12-14 populations in the field. KEY RESULTS: Generalized Linear Model (GLM) analyses indicated that the pollen load on stigmas was positively related to the exsertion of the corolla beyond the cupular bracts and size of the lower corolla lip, but so too was the rate of seed predation, creating conflicting selection on both floral traits. A geographic mosaic of selection mediated by seed predators, but not pollinators, was indicated by significant variation in levels of seed predation and the inclusion of two-, three- and four-way interaction terms between population and seed predation in the best model [lowest corrected Akaike Information Criterion (AICc)] explaining final seed production. CONCLUSIONS: These results indicate opposing selection in operation: pollinators generated selection for greater floral exsertion beyond the bracts, but seed predators generated selection for reduced exsertion above the protective pools of water, although the strength of the latter varied across populations.

Yucca brevifolia fruit production, predispersal seed predation, and fruit removal by rodents during two years of contrasting reproduction.

PREMISE OF THE STUDY: The distribution of Yucca brevifolia, a keystone species of the Mojave Desert, may contract with climate change, yet reproduction and dispersal are poorly understood. We tracked reproduction, seed predation, and fruit dispersal for two years and discuss whether Y. brevifolia is a masting species. METHODS: Fruit maturation, seed predation (larval yucca moths), and fruit dispersal (rodents) were monitored on a random sample of panicles during 2013 and 2014, which were years of high and low reproduction, respectively. Fates of fruits placed on the ground and in canopies were also tracked. Rodents were live-trapped to assess abundance and species composition. KEY RESULTS: In 2013, 66% of inflorescences produced fruit of which 53% escaped larval predation; 19.5% of seeds were destroyed in infested fruits. Total seed production was estimated to be >100 times greater in 2013 than 2014. One-third of the fruit crop fell to the ground and was removed by rodents over the course of 120 d. After ground fruits became scarce, rodents exploited canopy fruits. Rodent numbers were low in 2013, so fruits remained in canopies for 370 d. In 2014, fruit production was approximately 20% lower. Larvae infested the majority of fruits, and almost twice the number of seeds were damaged. Fruits were exploited by rodents within 65 d. CONCLUSIONS: High fertilization, prolific seed production, and low predispersal predation in 2013 suggests that pollinator attraction and satiation of seed predators influence masting in Y. brevifolia. Abundant, prolonged fruit availability to seed-dispersing rodents likely extends recruitment opportunities during mast years.

Reproductive success of Cabralea canjerana (Meliaceae) in Atlantic forest fragments, Brazil / Éxito reproductivo de Cabralea canjerana (Meliaceae) en fragmentos de bosque del Atlántico, Brasil

In Brazil, the Atlantic forest remnants have high biological diversity and a high level of endemism, but very little is known about the reproductive success of native species. Cabralea canjerana is a common tree in the Montane Atlantic forest, and its reproduction is highly dependent on pollinators. In order to contribute with the particular knowledge on this species, we collected data in three fragmented and three continuous forest sites, where the effects of fragmentation on both mutualistic (pollination) and antagonistic (seed predation) interactions were analysed. We determined fruit production and weight of 25 trees per site. The number of seeds and the percentage of predated and aborted seeds were also accessed for seven fruits of 10 trees per site. Pollinator visitation frequencies to flowers were recorded in two forest fragments and in two sites of the continuous forest. Our data showed that plants of C. canjerana produced more fruits (z-value=-8.24; p<0.0001) and seeds per fruit (z-value=-6.58; p=0.002) in the continuous than in the fragmented sites. This was likely due to differences in pollination, because the number of pollinator visits was higher in the continuous forest than in the fragments. Seed abortion (z-value=4.08, p<0.001) and predation (z-value=3.72, p=0.0002), on the other hand, were higher in the fragmented than in the continuous sites. Then, mutualistic and antagonistic interactions were affected by fragmentation, decreasing the reproductive success of the study tree. This study was the first to show a decrease in the reproductive output in forest fragments in an Atlantic forest tree species. This decrease may threaten the population structure and viability of C. canjerana in forest fragments. Rev. Biol. Trop. 63 (2): 515-524. Epub 2015 June 01.

En Brasil, los remanentes de bosque del Atlántico tienen una alta diversidad biológica y un alto nivel de endemismo, pero se conoce muy poco sobre el éxito reproductivo de las especies nativas. Canjerana cabralea es un árbol común en el bosque Atlántico Montano y su reproducción es altamente dependiente de los polinizadores. Con el fin de contribuir con el conocimiento particular de esta especie, se recogieron los datos en tres sitios fragmentados y tres de bosques continuos, donde se analizaron los efectos de la fragmentación tanto en relaciones mutualistas (polinización) como interacciones antagónicas (depredación de semillas). Se determinó la producción de frutos y el peso de 25 árboles. También se calculó el número de semillas y el porcentaje de semillas depredadas y abortadas en 7 frutos de 10 árboles por sitio. Frecuencia de visitas de polinizadores a las flores se registraron en dos fragmentos de bosque y en dos sitios de bosque continuo. Nuestros datos muestran que las plantas de C. canjerana produjeron más frutos (Z=-8.24; p<0.0001) y semillas por fruto (Z=-6.58; p=0.002) en el continuo que en los sitios fragmentados. Esto fue probablemente debido a las diferencias en la polinización, porque el número de visitas de polinizadores fue mayor en el bosque continuo que en los fragmentos. Por otro lado, el aborto de semillas (Z=4.08, p<0.001) y la depredación (Z=3.72, p=0.0002) fueron mayores en los sitios fragmentados que en los continuos. Entonces, las interacciones mutualistas y antagónicas se vieron afectadas por la fragmentación, disminuyendo el éxito reproductivo del árbol estudiado. Este estudio fue el primero en mostrar una disminución en el rendimiento reproductivo en los fragmentos de bosque en una especie de árboles forestales atlánticos. Esta disminución puede poner en peligro la estructura de la población y la viabilidad de C. canjerana en fragmentos de bosque.

Seed predators exert selection on the subindividual variation of seed size.

Subindividual variation among repeated organs in plants constitutes an overlooked level of variation in phenotypic selection studies, despite being a major component of phenotypic variation. Animals that interact with plants could be selective agents on subindividual variation. This study examines selective pressures exerted during post-dispersal seed predation and germination on the subindividual variation of seed size in hawthorn (Crataegus monogyna). With a seed offering experiment and a germination test, we estimated phenotypic selection differentials for average and subindividual variation of seed size due to seed predation and germination. Seed size affects germination, growth rate and the probability of an individual seed of escaping predation. Longer seeds showed higher germination rates, but this did not result in significant selection on phenotypes of the maternal trees. On the other hand, seed predators avoided wider seeds, and by doing so exerted phenotypic selection on adult average and subindividual variation of seed size. The detected selection on subindividual variation suggests that the levels of phenotypic variation within individual plants may be, at least partly, the adaptive consequence of animal-mediated selection.

Depredación pre-dispersiva de semillas en tres poblaciones del árbol Enterolobium contortisiliquum (Fabaceae)

Pre-dispersal seed predation in three populations of the tree Enterolobium contortisiliquum (Fabaceae). Predation is an important selective force that has shaped different strategies related to size, quantity and production time of fruits, seeds and seed reserves. The seeds of E. contortisiliqum are predated by the bruchid Merobruchus bicoloripes. We collected 1 300 fruits from 26 trees in San Salvador de Jujuy (Jujuy province), Castañares, and San Lorenzo (Salta province) Argentina. San Lorenzo is the wettest and highest site, while Castañares is the driest and lowest site. Jujuy is in the middle. In the laboratory, the fruits from each site were weighted and divided in three groups (small, medium and big). We counted the mature and predated seeds per fruit. In San Lorenzo, seed predation was higher in medium sized fruits while in Castañares and Jujuy seed predation was higher in both medium and bigger fruits. However, only in San Lorenzo we found a significant difference in preference for medium size fruits. Rev. Biol. Trop. 57 (3): 781-788. Epub 2009 September 30.

La depredación se ha postulado como una importante fuerza selectiva que ha moldeado distintas estrategias relacionadas con el tamaño, la cantidad y el tiempo de producción de semillas, frutos y el contenido de reservas. Las semillas de Enterolobium contortisiliqum son depredadas por el brúquido Merobruchus bicoloripes. Se estudiaron tres poblaciones de E. contortisiliquum en relación al tamaño de los frutos, a la producción de semillas por fruto, y a la depredación de semillas, comparando la variabilidad dentro y entre poblaciones. Se recolectaron 1300 frutos de 26 árboles en San Salvador de Jujuy (Jujuy), Castañares y San Lorenzo (Salta). Los frutos se pesaron y se contaron las semillas viables y depredadas. San Lorenzo es el sitio más húmedo y de mayor altitud, y Castañares el de menor altitud y más seco. En San Lorenzo encontramos una mayor preferencia de depredación en frutos medianos. En Castañares y San Salvador de Jujuy, a pesar de que los frutos medianos y grandes presentaron mayores niveles de depredación, no encontramos diferencias en la preferencia por alguna clase de tamaño. Los mayores niveles de depredación se encontraron en Castañares y S. S. de Jujuy.

Post-dispersal predation on Pinus sylvestris seeds by Fringilla spp: ground substrate affects selection for seed color.

Background matching might lower the risk of seeds being eaten by seed predators that search visually. In aviary experiments, we analyzed the selection of diff erent-colored seeds by ground-feeding finches (Fringillacoelebs and F.montifringilla) against four naturally occurring forest soil substrates. The substrates were fresh burn (black), 6-year-old burn (brown), mineral soil (pale yellow) and Pleuroziumschreberi feather moss (green). We used color-sorted seeds of Pinussylvestris, a species with a large natural variation in seed color, ranging from pale yellow to black. Although seeds were scattered on the substrates at a density of only 91 seeds m-2, birds removed seeds effectively. Both bird species found more pale than dark seeds on the fresh burn substrate. F. montifringilla also recovered more pale than dark seeds on the old burn, and more dark than pale seeds on mineral soil. In moss, the birds found very few seeds compared to the other substrates, and there was no color discrimination. P.sylvestris is frequently regenerating after fire, suggesting that dark seeds would be favored under selection from visually searching predators. Fire-adapted conifers with serotinous cones, e.g., Pinuscontorta ssp. latifolia, which spread their seeds primarily on freshly burnt surfaces, produce uniformly black or dark brown seeds. However, regeneration of the non-serotinous P.sylvestris is often extended for several years after a fire, during which substrate color and structure change. This may have helped to maintain variation in seed color. When regeneration of a plant species occurs on a substrate of uniform color, we believe that selection by visually searching seed predators will result in the evolution of cryptic seed color.

Effects of experimental manipulation of inflorescence size on pollination and pre-dispersal seed predation in the hummingbird-pollinated plant Ipomopsis aggregata.

Large floral displays should theoretically provide advantages to plants through increased pollinator visitation and resulting fruit and seed set. However empirical tests of the response of pollinators to floral display size have been limited by a lack of direct experimentation, and the results of such studies have been equivocal. In addition, other selective agents such as pre-dispersal seed predators might modulate effects of floral display on pollination. By artificially altering flower number, we examined the direct effects of floral display in the monocarpic herb, Ipomopsis aggregata (Polemoniaceae), on visitation rates by broad-tailed and rufous hummingbird pollinators, as well destruction of fruits by a pre-dispersal seed predator (Hylemya: Anthomyiidae). In addition, we quantified the ultimate effects of flower number on female reproductive success. Plants with larger floral displays were most likely to be visited first in any given foraging bout (P < 0.01). As expected, plants with more flowers received more total flower visits. However, we found no gain in the proportion of flowers visited for many- versus few-flowered plants, or the total number of approaches/hour. In fact, a significantly greater percentage of flowers were visited on few-flowered plants. Plants did not compensate for our reduction in flowers by increasing investment in the number or proportion of flowers that set fruit, the number of seeds/fruit, or seed weight. Pre-dispersal seed predation was greater for many- than for few-flowered plants (P < 0.001), but this did not offset the potential fitness gains of producing large displays. Our data support the hypothesis that large floral displays function primarily in long-distance attraction of pollinators, and enhance maternal success.

Post-dispersal predation and scatterhoarding of Dipteryx panamensis (Papilionaceae) seeds by rodents in Panama.

In tropical rain forests of Central America, the canopy tree Dipteryx panamensis (Papilionaceae) fruits when overall fruit biomass is low for mammals. Flying and arboreal consumers feed on D. panamensis and drop seeds under the parent or disperse them farther away. Seeds on the ground attract many vertebrate seed-eaters, some of them potential secondary seed dispersers. The fate of seeds artificially distributed to simulate bat dispersal was studied in relation to fruitfall periodicity and the visiting frequency of diurnal rodents at Barro Colorado Island (BCI), Panama. The frequency of visits by agoutis is very high at the beginning of fruitfall, but in the area close (<50 m) to fruiting trees (Dipteryx-rich area) it declines throughout fruiting, whereas it remains unchanged farther (>50 m) away (Dipteryx-poor and Gustavia-rich area). Squirrels were usually observed in the Dipteryx-rich area. Along with intense post-dispersal seed predation by rodents in the Dipteryx-rich area, a significant proportion of seeds were cached by rodents in the Dipteryx-poor area. Post-dispersal seed predation rate was inversely related to hoarding rate. A significantly greater proportion of seeds was cached in March, especially more than 100 m from the nearest fruiting tree. This correlates with the mid-fruiting period, i.e. during the height of D. panamensis fruiting, when rodents seem to be temporarily satiated with the food supply at parent trees. Hoarding remained high toward April, i.e. late in the fruiting season of D. panamensis. Low survival of scatterhoarded seeds suggests that the alternative food supply over the animal's home-ranges in May-June 1990 was too low to promote survival of cached seeds. Seedlings are assumed to establish in the less-used area of the rodents' home-range when overall food supply is sufficient to satiate post-dispersal predators.

Oviposition choices by a pre-dispersal seed predator (Hylemya sp.) : I. Correspondence with hummingbird pollinators, and the role of plant size, density and floral morphology.

Although the importance of pollinators has most often been examined in the evolution of floral characters, seed predators may also play a role in shaping floral evolution. In this study, I examined the role of interplant distance, plant size, and flower morphology on Ipomopsis aggregatás (Polemoniaceae) attractiveness to a pre-dispersal seed predator, Hylemya sp. (Anthomyiidae) and to hummingbird pollinators. The attractiveness of I. aggregata individuals to Hylemya was nonlinearly related to interplant distance in experimental arrays. Clumped and highly dispersed plants were preyed upon more frequently than those at intermediate distances. I found no relationship between interplant distance and visitation rates by hummingbird pollinators in these experimental arrays. However, in natural populations studied, clumped plants were more frequently approached by hummingbirds than those growing more widely dispersed. Display size was unrelated to visitation by Hylemya on inflorescences I clipped and maintained as "large", "small" and "control". Display size was also unrelated to the total number of visits by hummingbird pollinators to each of these experimental plants, however "large" display plants were more likely to be visited first in any given visitation sequence. Of various morphological measurements, corolla length showed the strongest positive correlation with Hylemya egg presence. To the extent that plant spacing and morphology is correlated with pollinator visits and ultimate seed set, Hylemya could be choosing flowers optimally, and playing a role in the evolution of floral traits.

Pre-dispersal seed predation in Central AmericanAcacia farnesiana: factors affecting the abundance of co-occurring bruchid beetles.

Pre-dispersal seed predation of the leguminousAcacia farnesiana byMimosestes nubigens andM. mimosae (Coleoptera: Bruchidae) was investigated in Santa Rosa National Park, northwestern Costa Rica. The purpose of the study was to determine the patterns of resource utilization by the seed predators and the mechanisms causing such patterns. Immature, mature, and fallen fruits were monitored during the dry seasons of 1987 and 1988 from different shrubs and areas. Parameters describing plant size, fecundity, and relative plant isolation were measured on each shrub. No evidence of spatial or temporal segregation was found between the two species and the intensity of seed predation was independent of the variables measured from each plant.M. mimosae was scarcer and always occurred withM. nubigens. Both were present in areas with low and high densities of the host plant, and the frequency distributions of their emergences from the fruits overlapped through the fruiting season.urosigalphus sp., a hymenopteran parasitoid, represented ca. 40% of all insect emergences in 1987 and ca. 30% in 1988. This wasp attacked a greater proportion of bruchid eggs on pods on the shrub than on pods beneath it, and more on green than on mature fruits. Parasitism thus appears to select against bruchid females that oviposit at an early stage of pod maturation. The harsh conditions of the dry season, namely heat and desiccation, also accounted for a high level of bruchid pre-emergence mortality, especially in fallen fruits, where survival from egg to adult was only about 18%. Beneath the shrubs, bruchid females exhibited selectivity, ovipositing more on pods in the shade than on those exposed to direct sunlight. In contrast to parasitoids, abiotic factors probably impose a selective force against those bruchid females that oviposit at a late stage of maturation or on pods already dropped. Both bruchid species can have more than one generation during the fruiting period. The intensity of seed predation did not, however, change during the season. The data obtained in this study suggest that factors like natural enemies and severe weather are more likely to limit the bruchid population densities than intra- or interspecific competition.

Post-dispersal predation of Acacia farnesiana seeds by Stator vachelliae (Bruchidae) in Central America.

Post-dispersal seed predation by the bruchid beetle Stator vachelliae was investigated in Santa Rosa National Park, Costa Rica. This insect finds the seeds of the leguminous Acacia farnesiana in the feces of horses, deer, and ctenosaur lizards, the current major dispersers. Patterns of oviposition and pre-adult survival of beetles in the seeds were investigated in a series of experiments using fresh horse dung. S. vachelliae never minded into the dung balls, attacking only those seeds located on the surface. Fresh horse dung did not attract insects more readily than dry dung. The proportion of seeds attacked was not related to their density in a defecation, and was similar in three areas with different densities of the host plant. In a fourth area with no fruiting A. farnesiana shrubs all seeds survived insect predation. Bruchids attacked a greater proportion of seeds at 1 m than at 5 m from the edge of the shrub's crown. Seeds were mainly removed from horse dung by rodents with similar intensity in all areas and at both distances; this seed removal interfered with bruchid oviposition and probably with bruchid survival. S. vachelliae oviposited less frequently on seeds in dung fully exposed to sun. When oviposition on a dung pile was high, the distribution of eggs on the seeds was clumped, suggesting that some seeds were preferred to others. By the end of the dry season, bruchids stopped attacking the seeds. The results show that the fate of both seeds and bruchids is greatly influenced by the location and time of defecation.