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

Summary 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.     

Predispersal seed predation on five Piper species in tropical rainforest

Absolute number of seeds lost to predispersal seed predators and proportion of total seeds lost per infructescence were compared among five Costa Rican Piper species of different annual fecundities. Mean seed number and mean seed size in the five species were negatively correlated. The impact of predation on these species was inversely related to the number of seeds they produced. The two early successional species had very high fecundities, a combination of many seeds per infructescence, many infructescences per plant, and, in one species, year-round reproduction. Although seed predators destroyed as many or more seeds of these early successional species than they did of the less fecund, late successional species, this loss accounted for a relatively minor proportion (9 and 12%) of the seeds of the early successional species. In contrast, late successional species produced fewer, larger seeds in a smaller number of infructescences and were not continually in fruit. One of these species, which produced intermediate numbers of intermediately sized seeds, lost 30% of the seeds in each infructescence on average. Seed predators destroyed a larger proportion (65 and 76%) of the seeds per infructescence in the two species with fewest seeds per infructescence. High levels of insect damage in these late successional species caused many of their infructescences to abort prematurely. Taken together these factors resulted in annual fecundities several orders of magnitude smaller in shade-tolerant Piper species than the annual fecundities of shade-intolerant, early successional species. Seedlings of the two early successional species were common in large gaps and other sunny clearings and seedlings of the species with 30% seed loss were occasional, whereas no seedlings were seen of the two species with the highest proportional seed loss, suggesting that seed predation on the latter species may limit seedling recruitment.     

Satiation of predispersal seed predators: the importance of considering both plant and seed levels

Plants can reduce the fitness costs of granivory by satiating seed predators. The most common satiation mechanism is the production of large crops, which ensures that a proportion of the seeds survive predation. Nevertheless, satiation of small granivores at the seed level may also exist. Larger seeds would satiate more efficiently, enhancing the probability of seed survival after having been attacked. However, a larger seed size could compromise the efficiency of satiation by means of large crops if there were a negative relationship between seed size and the number of seeds produced by an individual plant. We analyze both types of satiation in the interaction between the holm oak Quercus ilex and the chestnut weevil Curculio elephas. Both crop size and acorn size differed strongly in a sample of 32 trees. Larger crop sizes satiated weevils, and higher proportions of the seeds were not attacked as crop size increased. Larger seeds also satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival. Seedling size was strongly related to acorn size and was reduced by weevil attack, but seedlings coming from large weeviled acorns were still larger. The number and the size of the acorns produced by individual trees were negatively related. Larger proportions of the crop were infested in oaks producing less numerous crops of larger acorns. However, contrary to expectations, these trees did not satiate more effectively at the seed level either. Effective satiation by larger acorns was precluded by larger multi-infestation rates associated to smaller seed crops, in such a way that the proportion of attacked seeds that survived did not vary among trees with different acorn sizes. These results highlight the need of considering satiation by means of large crops and large seeds in studies of predispersal seed predation. Long-term monitoring on individual oaks will help to assess whether there is a trade-off between the number and the size of the acorns and, if it existed, how it could condition the fitness consequences of both types of satiation.     

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

Summary 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.     

Shell strength and primate seed predation of nontoxic species in eastern and southern Africa

The large African primates that eat fruit destroy the seeds of a number of fruiting species. This paper addresses several questions about seed-eating: What is the nature of the dietary niche provided by the nonpoisonous seeds of eastern and southern Africa? How well are these seeds mechanically protected? What other means of reducing seed predation are employed by the plants? and is the niche ecologically stable? Measurements of seed shell strength on 37 species from 17 families reveal a range of values, from <100-kg (numerous species) to over 2000-kg (palm nuts) breaking load. Primates crack open with their teeth seed shells from species exhibiting test strengths less than 600 kg. Variation in shell strength appears to increase dramatically for average species strengths above 100 kg. Plant species are not characterized by specific shell strengths but instead, display envelopes of shell strength overlapping broadly with other species. Taking this into account, adult male baboons (Papio spp.) appear to be dentally capable of preying upon most of the seed species of eastern and southern Africa. The possibility for predation of nonpoisonous seeds exists primarily because the plants periodically produce large crops in synchrony and the hard-shelled seeds are effectively dispersed, sometimes explosively but more often by means of edible fruits. The concomitant primate seed predation is a facultative specialization, of little apparent threat to the community of plants that support it.     

Evidence of a component Allee effect driven by predispersal seed predation in a plant (Pedicularis rex,Orobanchaceae)

A small or sparse population may suffer a reduction in fitness owing to Allee effects. Here, we explored effects of plant density on pollination, reproduction and predation in the alpine herb Pedicularis rex over two years. We did not detect a significant difference in the pollination rate or fecundity (fruit set and the initial seed set) before predation between sparse and dense patches in either year, indicating no pollination-driven Allee effect. However, dense patches experienced significantly fewer attacks by predispersal seed predators in both years, resulting in a significantly decreased realized fecundity (final seed set), suggesting a component Allee effect driven by predispersal seed predation. Predation-driven Allee effects have been predicted by many models and demonstrated for a range of animals, but there is scant evidence for such effects in plants. Our study provides strong evidence of a component Allee effect driven by predation in a plant species.     

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.     

Allozyme diversity in relation to geographic distribution and population size inLathyrus vernus (L.) Bernh. (Fabaceae)

Lathyrus vernus (L.) Bernh. is a diploid, long-lived perennial and insect-pollinated herb with no special adaptation to long-distance dispersal. It occurs on neutral soil in deciduous forests throughout western Eurasia. Due to specific habitat preferences,L. vernus has a fragmented distribution with isolated populations. We investigated allozyme variation at eleven loci in 20 populations ofL. vernus from one geographically central region (the Czech Republic and the Slovak Republic) and two geographically marginal regions (southern and central Sweden) in the species present-day distribution. There was a clear differentiation between the three regions and the genetic distance between the populations was highly correlated with geographic distance. The total genetic diversity (H_T) was 0.354. The proportion of genetic diversity due to differentiation between regions, and to differentiation between populations within regions, accounted for 10% each. There was no difference in level of genetic diversity between the three regions. No significant difference in level of genetic diversity was found between small and large populations. The genetic diversity inL. vernus may either be a result of the long generation-time of the species or peculiarities in the post-glacial migration species, e.g. survival only in refugia far east of the sampled populations and/or migration as a continuous process not involving founder-events.     

Masting,seed dispersal and seed predation in the cycad Macrozamia communis

Summary Little is known about the adaptive value of mast seeding, a common phenomenon in temperate trees and shrubs. Masting is likely to affect both seed dispersal and seed predation. In systems where similar taxa of animals are involved in these two processes, the consequences of mast seeding are likely to be particularly complicated. This study examined the effects of mast seeding in a cycad, Macrozamia communis, on the dispersal of seeds, the pattern of dispersion of seeds and post-dispersal predation on seeds. Dispersal of seeds by possums was poorer from source plants in a masting population than from source plants in an adjacent, non-masting population. This resulted in fewer seeds per seeding female plant in the masting plot being dispersed to favourable sites. We conclude that this is caused by the feeding behaviour and movements of possums in the masting site. The abundance of seeds in this site did not satiate the post-dispersal predators, native rats. In fact, more seeds in this site were eaten than in the nonmasting site. We suggest that the mast seeding observed in M. communis may not be adaptive, but is more likely a consequence of other factors which synchromize flowering within local populations.     

Ecological and evolutionary consequences of spatial and temporal variation in pre-dispersal seed predation

Pre-dispersal seed predation may have important effects on population dynamics and trait evolution in plants. In this review, we first present a conceptual framework of the strength of pre-dispersal seed predation and its variation in space and time. We consider the interaction between plants and their seed predators to be “strong” when it affects plant population dynamics or causes changes in plant trait–fitness relationships, and “weak” when it has no such effects, and propose ways of how to adequately assess these effects. Second, we review the ecological literature between 1991 and 2005 to evaluate documented effects of pre-dispersal seed predation on plants and draw five major conclusions. (1) Pre-dispersal seed predation rates are usually low but sometimes high, and show a considerable variation in space and time. (2) Direct evidence suggests that pre-dispersal seed predation can have a significant effect on recruitment and plant population growth rate. Accumulating evidence of seed-limited recruitment suggests that such effects are common. (3) Pre-dispersal seed predation affects selection on several plant traits, such as flowering phenology and flower number, which are usually interpreted mainly in the context of plant–pollinator interactions. (4) The patterns of variation in the interactions between plants and pre-dispersal seed predators suggest that geographic selection mosaics may be common. (5) Although there are numerous studies estimating seed predation, there are still rather few studies that have aimed at examining the interaction explicitly in terms of effects on plant population dynamics and trait selection. From these we know that seed predators can have important, and often variable, effects on plant population dynamics and trait evolution. However, it still remains to assess how important they are across study systems and relative to other aspects of the plant's biotic and abiotic environment.     

Effects of microhabitat on palm seed predation in two forest fragments in southeast Brazil

The establishment of plants depends crucially on where seeds are deposited in the environment. Some authors suggest that in forest understory seed predation is lower than in gaps, and higher than at the forest edge. However, most studies have been carried out in large forest patches and very little is known about the effects of microhabitat conditions on seed predation in forest fragments. We evaluated the effects of three microhabitats (gaps, forest edge, and understory) on seed predation of two palm species (Euterpe edulis and Syagrus romanzoffiana) in two semi-deciduous forest fragments (230 and 2100 ha) in southeast Brazil. Our objective was to test two hypotheses: (1) Low rodent abundance in small fragments as a result of meso-predator action levels leads to lower seed predation in small fragments. (2) Most mammal species in small fragments are generalists with respect to diet and habitat, so that seed predation is similar in different microhabitats (gaps, forest edge and understory) in the small fragment, but not in the larger one. The study community of small fragments is usually composed of generalist species (in diet and habitat aspects), so we expected the same rate of seed predation among microhabitats (gaps, forest edge and understory) in the tested smaller fragment. The experiment was carried out in the dry season (for E. edulis) and in the wet season (for S. romanzoffiana) in 1999. We conclude that post-dispersal seed predation in forest fragments can be directly connected with mammal communities, reflecting their historical and ecological aspects.     

Chaoborus predation and the function of phenotypic variation in Daphnia

To investigate the role of helmet formation in defense against predation, laboratory experiments were used to analyze the effects of morphological changes in Daphnia on susceptibility to Chaoborus predation. Behavioral observations of Chaoborus preying on helmeted and non-helmeted Daphnia suggest pre-contact advantages for helmeted prey but post-contact advantages for non-helmeted prey. Helmeted Daphnia are better at evading capture by Chaoborus but may also be more easily handled by the predator. Swimming behavior of the prey, which is influenced by the presence of a tailspine, may affect Chaoborus strike distance. These results re-emphasize the potential hydromechanical importance of body shape changes in defense against predation.     

Impact of selective predation by Mesocyclops pehpeiensis on a zooplankton community: experimental analysis using mesocosms

Predation by cyclopoid copepods is an important factor affecting zooplankton communities in freshwater habitats. Experiments provide strong evidence of the role of selective predation by cyclopoid copepods in structuring zooplankton communities. To assess the predation impact of a cyclopoid copepod, Mesocyclops pehpeiensis, we conducted a mesocosm experiment using 20-l polyethylene tanks in which the density of the predator and the food available to herbivorous zooplankton varied. M. pehpeiensis had a notable but selective effect on the zooplankton community. The population of a small cladoceran, Bosmina fatalis was affected negatively, but M. pehpeiensis did not have any apparent impact on the population dynamics of another Bosmina species, B. longirostris. On the other hand, the population of small rotifers responded positively to the presence of M. pehpeiensis, and their densities increased in mesocosms with a high density of M. pehpeiensis. It seems that suppression of B. fatalis by M. pehpeiensis predation indirectly affected rotifers by releasing them from competition with B. fatalis. The results suggest that copepod predation is a powerful factor regulating zooplankton communities directly and indirectly.     

Pre-dispersal seed predation reduces the reproductive compensatory advantage of thrum individuals in Erythroxylum havanense (Erythroxylaceae)

Male-sterility mutations in hermaphroditic species represent the first step in the evolution of gender specialization. Male-sterile individuals commonly compensate the loss of the male function by increasing the number or quality of seeds. Because the magnitude of compensation determines the maintenance of females within populations and the evolution of sexual specialization, plant-animal interactions may affect these evolutionary processes if animals are sensitive to such reproductive asymmetries. Here we explore the effect of seed predation on the reproductive compensation of Erythroxylum havanense, a distylous shrub with morph-biased partial male sterility, during two consecutive years. Seed predation reduced the compensatory advantage of thrums in 1987, but not in 1988. Annual differences in the intensity of seed predation seem to be accounted for differences in the onset and synchrony of flowering. Thus, although seed predators may reduce the compensatory advantage of thrums, their impact is modulated by the environmental cues triggering flowering and insect emergence.     

Deceptive fruits reduce seed predation by insects inPistacia terebinthus L. (Anacardiaceae)

Summary The production of seedless fruits (parthenocarpy) is not yet understood from an evolutionary viewpoint, even though it is taxonomically widespread. Here I present a case in which parthenocarpy reduces the incidence of seed predation by insects. At least the first generation of chalcidoid wasps that oviposit in the fruits ofPistacia terebinthus L. (Anacardiaceae) cannot discriminate among viable and inviable fruits, allocating energy and time to oviposition on fruits that are not suitable for larval development.     

Seed predation,pathogen infection and life-history traits in Brassica rapa

Herbivory and disease can shape the evolution of plant populations, but their joint effects are rarely investigated. Families of plants of Brassica rapa (Brassicaceae) were grown from seeds collected in two naturalized populations in an experimental garden. We examined leaf infection by the fungus Alternaria, seed predation by a gall midge (Cecidomyiidae) and plant life-history traits. Plants from one population had heavier seeds, were more likely to flower, had less fungal infection, had more seed predation and were more fecund. Fungal infection score and seed predation rate increased with plant size, but large plants still had the greatest number of undamaged fruits. Spatial heterogeneity in the experimental garden was significant; seed predation rate and fecundity varied among blocks. An apparent tradeoff existed between susceptibility to disease and seed predation: plants with the highest fungal infection score had the lowest seed predation rate. Alternaria infection varied between populations, but the disease had no effect on fecundity. Seed predation did reduce fecundity. Damaged fruits had 31.4% fewer intact seeds. However, evidence for additive genetic variation in resistance to seed predation was weak. Therefore, neither disease nor seed predation was likely to be a strong agent of genetically based fecundity selection.     

Removal and predation of Quercus liaotungensis acorns by animals

Two types of acorns (fresh, current-year acorns and old, previous-year acorns) of Quercus liaotungensis were used under two different situations (night and day) to examine the removal and predation by animals in the field. The acorns disappeared very fast. During the day, all of the 280 this-year acorns (TA) and all of the 140 last-year acorns (LA) were removed by animals within 6 h and 5 h, respectively, after acorns were put in place. At night, all of the 280 TA acorns were removed by animals within 10 h after acorns were distributed, and 71 out of 140 LA acorns were removed within 10 h. Animals consumed a small proportion of acorns in situ (11.8%). The number of TA acorns consumed in situ at night was higher than that during the day. During the day, the number of LA acorns consumed in situ was significantly higher than TA acorns. This result suggests that the most important harvesters of Q. liaotungensis during the day were visually orientated diurnal animals, and the most important harvesters at night were olfactorily orientated nocturnal animals. Furthermore, the latter are more efficient than the former at finding and harvesting acorns, because they spend less energy on harvesting the same number of sound acorns.     

When good bugs go bad: intraguild predation by Jalysus wickhami on the parasitoid, Cotesia congregata

We report a case of direct intraguild predation involving an insect predator and parasitoid in an agricultural system. The spined stilt bug, Jalysus wickhami Van Duzee, feeds on eggs of the tobacco hornworm, Manduca sexta L., and also on prepupal and pupal stages of the gregarious hornworm parasitoid, Cotesia congregata (Say). In two separate trials, mean mortality of attached parasitoids was significantly lower (66%, 73%) than that of their detached siblings (97%, 96%) after a 3 day exposure to stilt bugs, demonstrating that attachment to the host offered some protection against predation. In no-choice experiments, prepupal parasitoids suffered greater mortality (0 day-old=61%, 1 day-old=65%) than pupal parasitoids (2 day-old=50%, 3 day-old=14%). When offered in combination with 0 or 2 day-old hornworm eggs, respective mortality of 0, 1, 2 and 3-day-old pupal parasitoids showed a similar pattern (67%, 63%, 33% and 23%). In another experiment, mortality of 0-day-old pupal parasitoids (64%) was greater than that of 3 day-old pupal parasitoids (38%). Mortality of pupal parasitoids was not affected by the availability of hornworm eggs, a highly acceptable food. Younger pupal parasitoids (=prepupae) probably suffered greater mortality because they were more easily fed on by stily bugs than older (pupated) ones. Because C. congregata overwinters in the prepupal stage, it may be particularly vulnerable to attack late in the season when stilt bug populations are large and hornworm eggs are relatively uncommon.