The role of Rhytidoponera metallica (Hymenoptera,Formicidae) in facilitating post‐fire seed germination of three ant‐dispersed legume species

The fire avoidance hypothesis proposes that a benefit of seed dispersal by ants (myrmecochory) is to protect seeds from being killed during fire and to facilitate post‐fire germination of seeds that require heat shock to break their physical dormancy. The aim of this study was to quantify the effect of fire and seed burial by a predominant seed‐dispersing ant, Rhytidoponera metallica (subfamily: Ectatomminae) on germination levels of three ant‐dispersed legume species (Pultenaea daphnoides, Acacia myrtifolia and Acacia pycnantha). Experimental burial of seeds within aluminium cans at a site prior to being burnt and at an adjacent unburnt site showed that fire increased germination levels, particularly for seeds buried at 1‐ and 2‐cm deep and that overall, germination levels differed among the three plant species. To quantify seed burial depths and post‐fire germination levels facilitated by R. metallica ants, seeds were fed to colonies prior to fire at the burnt and unburnt sites. Of the seeds buried within nests that were recovered, between 45% and 75% occurred within the upper 6 cm of the soil profile, although unexpectedly, greater percentages of seeds were recovered from the upper 0–2 cm of nests in the unburnt site compared with nests in the burnt site. Germination levels of buried seeds associated with R. metallica nests ranged from 21.2% to 29.5% in the burnt site compared with 3.1–14.8% in the unburnt site. While increased seed germination levels were associated with R. metallica nests following fire, most seeds were buried at depths below those where optimal temperatures for breaking seed dormancy occurred during the fire. We suggest that R. metallica ants may provide fire avoidance benefits to myrmecochorous seeds by burying them at a range of depths within a potential germination zone defined by intra‐ and inter‐fire variation in levels of soil heating.     

SEED DISPERSAL IN VIOLA (VIOLACEAE): ADAPTATIONS AND STRATEGIES

Surveys of Viola dispersal mechanisms result in the distinction of two major adaptive syndromes: one purely myrmecochorous for the exploitation of seed-transporting ants, the other partially myrmecochorous (= diplochorous) for the explosive ejection of seeds followed by ant exploitation. Diplochory is exhibited by the majority of Viola species, but myrmecochory is exhibited only by eleven species with limited Eurasian distribution. Morphological and experimental evidence suggests the hypothesis that Viola is basically myrmecochorous but that different selective pressures, especially seed predation, have produced a clear divergence in dispersal systems. The majority of species, the diplochores, have evolved a system which combines ballistic and ant seed dispersal with predator avoidance. Diplochory itself may be a response to predation pressure. The minority of species are purely myrmecochorous, possibly highly coevolved with specific ant species, thus limiting the distribution of the Viola species concerned.     

DUAL FUNCTION OF THE ELAIOSOME OF CORYDALIS AUREA (FUMARIACEAE): ATTRACTION OF DISPERSAL AGENTS AND REPULSION OF PEROMYSCUS MANICULATUS,A SEED PREDATOR

It has been suggested that one of the selective advantages of ant dispersal is the burial of seeds in ant nests where predators such as small rodents cannot find them. The elaiosomes of Corydalis aurea (Fumariaceae) are extremely attractive to ants, which assiduously gather the seeds and take them to nests. However, seed production commonly exceeds the gathering capacity of ants so that seeds accumulate beneath the parent plants. In spite of this, no signs of rodent predation are evident. Experiments with a major seed predator, the deer mouse Peromyscus maniculatus, show that when given the choice of seeds with and without elaiosomes, the mice consume significantly more seeds without elaiosomes. This remains true whether or not the intact seeds bear fresh, moist elaiosomes or dry, withered ones. Our experiments strongly suggest that the elaiosome has a dual function, the attraction of the ant seed dispersers and the repulsion of seed predators.     

Switching roles from antagonist to mutualist: a harvester ant as a key seed disperser of a myrmecochorous plant

1. Harvester ants are major seed predators in arid environments. However, given that many harvester ants are partly omnivorous and therefore potentially attracted to the elaiosomes of myrmecochorous seeds, it is unclear if these ants act as predators or dispersers when removing myrmecochorous seeds. 2. We describe the outcomes of interactions between the harvester ant, Pogonomyrmex naegelii, and myrmecochorous plant, Microstachys serrulata, in a Brazilian savanna. We: (i) evaluated the role of elaiosome in seed removal by P. naegelii; (ii) investigated the fate and viability of removed seeds; (iii) tested if soils associated with P. naegelii nests are nutrient-enriched; (iv) compared seedling survival; and (v) the density of seedling and adult M. serrulata nearby to P. naegelii nests compared with those away from these nests (i.e. controls). 3. Rates of removal of M. serrulata seeds were two-fold higher with elaiosomes than without. The ant attractant oleic acid was the dominant fatty acid in elaiosomes, but it was absent from seeds. Removed seeds are taken into nests, and Tetrazolium tests indicated that 95% of seeds remain viable. Soils associated with P. naegelii nests were not nutrient-enriched, and seedling survival was similar nearby to P. naegelii nests compared with control areas. However, densities of both seedling and adult M. serrulata were higher nearby to P. naegelii nests than in control areas. 4. Our findings show that P. naegelii switches its role from seed predator for most plant species to be the dominant seed disperser for M. serrulata, playing a key role in the distribution of adult plants.     

Unexpected relationships and valuable mistakes: non‐myrmecochorous Prosopis dispersed by messy leafcutting ants in harvesting their seeds

Abstract Ants generally disperse seeds while feeding on fruits or structures attached to the seed. Seed dispersal as a by‐product of seed predation (dyszoochory) was recognized in specialized harvester ants, but not in ants predating seeds opportunistically. Leafcutting ants are the main herbivores in much of the Neotropics, and they have been reported to remove fruits and seeds, but their role as seed predators and dispersers has not been acknowledged. Prosopis flexuosa D.C. (Fabaceae, Mimosoideae) is the most abundant tree species in the central Monte Desert, Argentina, and it is likely to depend on secondary animal dispersal. Mammalian frugivores are usually considered its main dispersers, but the opportunity for dispersal may be small since the removal of fruits and seeds by seed predators is very intense. The objective of this study was to identify which ant species interact with P. flexuosa fruits and to evaluate their relative importance as seed predators and dispersers. In a field experiment, whole and segmented pods were offered and several ant species exploiting the fruits were identified. Additionally, all pod segments remaining around nests of the three ant species able to remove them (the leafcutters Acromyrmex lobicornis Emery and Acromyrmex striatus Roger, and Pheidole bergi Mayr) were examined during and after the P. flexuosa primary dispersal season. Up to 753 pod segments and 90 sound seeds were found accumulated in a circle of 1 m radius over nests of A. lobicornis, and even more in an examined trail. Acromyrmex striatus left a smaller proportion of sound seeds and P. bergi left a smaller number of pod segments. All tendencies were similar during shorter known periods of accumulation. Leafcutting ants are acting as important seed predators, and ‘by mistake’ may be dispersing a key non‐myrmecochorous tree. This is an unexplored path in the seed dispersal cycle of P. flexuosa that challenges the tendency to predict interactions based on classifications made with other goals.     

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

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

Increased seed predation in the second fruiting event during an exceptionally long period of community-level masting in Borneo

In Southeast Asian tropical rainforests, community-level masting (CM) occurs at irregular intervals of 2–10 years. During CM periods, many plant species from various families synchronously flower and subsequently undergo community-level fruiting. Seed predation is a key factor in understanding the ecological and evolutionary factors affecting CM. Masting is proposed to decrease seed mortality due to predation in two ways: by depressing predator abundance through extended and unpredictable absences of seeds; and by satiating predators via mass seed production (predator satiation hypothesis). If the hypothesis is valid in these rainforests, the incidence of seed predation will be higher in a fruiting event that occurs soon after a previous fruiting event, because the intervening period of seed absence would be inadequate to starve the predators. In this study, we examined seed predation by insects, focusing on five dipterocarp species that exceptionally reproduced twice during an extended CM period. All of the five species suffered more intense seed predation in the second fruiting event, consistent with the prediction expected from the predator satiation hypothesis. Weevils, bark beetles and mammals were the main cause of increased seed predation in three, one and one plant species, respectively. However, seed predation intensity did not increase during the second fruiting event in a few combinations of predator and plant species. We discuss the possibility that competition for seeds among predators and/or the interspecific differences in life history traits among predators might affect the varying intensities of seed predation among dipterocarp species by different seed predators.     

Seed predation limits post-fire recruitment in the waratah (<Emphasis Type="Italic">Telopea speciosissima</Emphasis>)

Seed predation may reduce recruitment in populations that are limited by the availability of seeds rather than microsites. Fires increase the availability of both seeds and microsites, but in plants that lack a soil- or canopy-stored seed bank, post-fire recruitment is often delayed compared to the majority of species. Pyrogenic flowering species, such as Telopea speciosissima, release their non-dormant seeds more than 1 year after fire, by which time seed predation and the availability of microsites may differ from that experienced by plants recruiting soon after fire. I assessed the role of post-dispersal seed predation in limiting seedling establishment after fire in T. speciosissima, in southeastern Australia. Using a seed-planting experiment, I manipulated vertebrate access to seeds and the combined cover of litter and vegetation within experimental microsites in the 2 years of natural seed fall after a fire. Losses to vertebrate and invertebrate seed predators were rapid and substantial, with 50% of seeds consumed after 2 months in exposed locations and after 5 months when vertebrates were excluded. After 7 months, only 6% of seeds or seedlings survived, even where vertebrates were excluded. Removing litter and vegetation increased the likelihood of seed predation by vertebrates, but had little influence on losses due to invertebrates. Microsites with high-density vegetation and litter cover were more likely to have seed survival or germination than microsites with low-density cover. Recruitment in pyrogenic flowering species may depend upon the release of seeds into locations where dense cover may allow them to escape from vertebrate predators. Even here, conditions suitable for germination must occur soon after seed release for seeds to escape from invertebrate predators. Seed production will also affect recruitment after any one fire, while the ability of some juvenile and most adult plants to resprout after fire buffers populations against rapid declines when there is little successful recruitment.     

The effect of limited visibility on vigilance behaviour and speed of predator detection: implications for the conservation of granivorous passerines

Ants have been traditionally considered either as predators or dispersers of seeds, but not both. That is, ant dispersal is restricted to myrmecochorous seeds, while almost all seeds removed by seed‐harvesting ants are eaten. However, harvesting ants might be simultaneously antagonistic and mutualistic towards seeds. This study analyzes the predation–dispersal relationship between seed‐harvesting ants and seeds of Lobularia maritima, a non‐myrmechorous perennial herb, in order to disentangle the dual role of ants as dispersers and predators of L. maritima seeds. The results obtained confirm the role of harvesting ants as both predators and dispersers of the non‐myrmechorous seeds of L. maritima. The removal activity of Messor bouvieri on L. maritima seeds is very important, particularly in autumn, which is the flowering and fruiting peak of this plant. It can be estimated that harvesting ants collect more than 85% of seeds, and almost 70% of them are effectively lost to predation. However, these granivorous ants also have drawbacks as seed dispersers. There is a relatively small percent of seeds collected by ants that escape predation, either because they are dropped on the way to the nest (16.4% of seeds harvested), or because they are mistakenly rejected on the refuse pile (0.9%). Abiotic dispersal of L. maritima seeds in the absence of ants occurs over very short distances from the plant stem. As seeds dispersed by ants reach a considerably greater distance than that obtained by gravity, this might represent a real advantage for the species, because it reduces intraspecific adult competition for seedlings, which directly influences seedling survivorship. These results challenge the generalization that seed removal by ants generally leads to successful seed dispersal if done by legitimate seed dispersers, or seed loss if done by seed consumers that eat them, and confirm that harvesting ants might have a dual role as both predators and dispersers of nonmyrmechorous seeds.     

Seed predation of Copaifera langsdorffii (Fabaceae): a tropical tree with supra‐annual fruiting

Seed predation is an important ecological and evolutionary force that directly affects the distribution of plant species. Copaifera langsdorffii is a tropical tree species with supra‐annual fruiting, which has its seeds predated by a specialist endogenous insect (Rynochenus brevicollis: Curculionidae) in the Brazilian savanna. Three hypotheses were addressed: (i) the predator satiation hypothesis, (ii) the resource concentration hypothesis and (iii) the larger seed predation hypothesis. A total of 112 individual C. langsdorffii were monitored monthly from January to August during four consecutive years (from 2008 to 2011) to determine the presence of fruits on each plant. All trees produced fruits in the year 2008, whereas none of them produced flowers or fruits in 2009 or 2010. Moreover, only 65 individuals (58%) marked in 2008 produced fruits in 2011. The number of fruits per plant was approximately 21% greater in 2008 than in 2011, while the percentage of seed predation was 76% greater in 2011, thereby supporting the predator satiation hypothesis. The percentage of seeds predated was not affected by the number of fruits per plant. Therefore, our data did not support the resource concentration hypothesis. Plants producing large seeds experienced more seed predation by R. brevicollis, supporting the larger seed predation hypothesis. In addition, we also observed a positive relationship between seed volume and adult R. brevicollis weight. This study demonstrates the importance of supra‐annual fruiting for increasing survivorship of C. langsdorffii seeds both at the individual and the population level, and suggests that seed predators select plants producing large seeds as a way of increasing the number of offspring.     

Insect Seed Predators in Erythrina falcata (Fabaceae): Identification of Predatory Species and Ecological Consequences of Asynchronous Flowering

Seed predation by insects exerts negative effects on plant reproduction by limiting the supply of seeds and preventing germination. Seed predators of the family Fabaceae are usually generalists, which increases the rate of predation. One strategy to minimize seed predation, developed by plants from temperate regions, is “escape in time,” i.e., flowering before or after the peak of predation. For tropical species, few studies have investigated the strategies used by plants to minimize seed predation. Here, using Erythrina falcata, a tropical species of Fabaceae, we test three main hypotheses: (i) escape in time is a mechanism used by E. falcata to minimize seed predation, (ii) the predators of E. falcata seeds are generalists, and (iii) the biometric variables of the pods can influence seed predation. In order to test these hypotheses, we determined the flowering time of E. falcata, rate of seed predation, the predators insects, and biometric variables of the pods. The analyzed trees were grouped into three classes: “early,” “peak,” and “late” flowering. The average seed predation rates on trees in the early and late classes were 65% and 50%, respectively, and in the peak class, 80%; thus, our first hypothesis can be accepted. Three species of Lepidoptera and two of Coleoptera were found preying on E. falcata seeds. These species were observed to be generalist predators; thus, our second hypothesis can be accepted. The biometric variables of the pods cannot influence seed predation rate. The ecological consequences of asynchronous flowering on plants and insects are discussed.     

Spatial relationships and mechanisms of coexistence between dominant and subordinate top predators

Most forest ecosystems contain a diverse community of top‐level predators. How these predator species interact, and how their interactions influence their spatial distribution is still poorly understood. Here we studied interactions among top predators in a guild of diurnal forest raptors in order to test the hypothesis that predation among competing predators (intraguild predation) significantly affects the spatial distribution of predator species, causing subordinate species to nest farther away from the dominant ones. The study analyzed a guild in southwestern Europe comprising three raptor species. For 8 years we studied the spatial distribution of used nests, breeding phenology, intraguild predation, territory occupancy, and nest‐builder species and subsequent nest‐user species. The subordinate species (sparrowhawk Accipiter nisus) nested farther away from the dominant species (goshawk A. gentilis), which preyed on sparrowhawks but not on buzzards Buteo buteo, and closer to buzzards, with which sparrowhawks do not share many common prey. This presumably reflects an effort to seek protection from goshawks. This potential positive effect of buzzards on sparrowhawks may be reciprocal, because buzzards benefit from old sparrowhawk nests, which buzzards used as a base for their nests, and from used sparrowhawk nests, from which buzzards stole prey. Buzzards occasionally occupied old goshawk nests. These results support our initial hypothesis that interspecific interactions within the raptor guild influence the spatial distribution of predator species in forest ecosystems, with intraguild predation as a key driver. We discuss several mechanisms that may promote the coexistence of subordinate and dominant predators and the spatial assembly of this raptor guild: spatial refuges, different breeding phenology, spatial avoidance, low territory occupancy between neighboring nesting territories, nest concealment and protection, and diet segregation.     

Seed re-dispersal of four myrmecochorous plants by a keystone ant in central China

Seed re-dispersal following initial harvesting by ants may have important implications for the distribution and fate of myrmecochorous seeds. However, the probability of seed re-dispersed by ants and the effect it may have on subsequent survival appear variable, the functional role of diaspore, disperser and seed predator to the fate of discarded seeds remain unclear. To clarify the ecology, we compared the consequences of seed re-dispersal by a keystone seed-dispersing ant (Myrmica ruginodis Nylander) for four sympatric myrmecochorous plants common to the temperate deciduous forests in Qinling Mountains, central China. Plants varied in the probability of re-dispersal and in elaiosome condition. Ants preferred seeds with residual elaiosomes, while rodents only consumed the two larger-seeded species, regardless of the elaiosome presence. The scattered distribution of discarded seeds increased the probability of ant re-harvesting and, to some extent, reduced rodent predation. Thus, difference in the probability of seed re-dispersal and its subsequent effect on seed fate in relation to ants and rodents was attributed primarily to the elaiosome condition, seed size and seed spatial pattern. The results imply that seed re-dispersal could affect the fitness of plants and ultimately influence the plant abundance and distribution pattern. This highlights the necessity to incorporate re-dispersal into myrmecochory to advance our understanding of the benefits of myrmecochory to plants.     

Ignorant seed predators and factors affecting the seed survival of a tropical palm

In addition to acting as seed predators, some terrestrial mammals bury seeds via scatter hoarding. This study system used two permanent plots in examining the interaction between small rodents and the seeds of the palm Astrocaryum mexicanum. We tested how experimental burial, and fruiting status of the parent, distance to the parent, seed size, and microsite characteristics affect the survival of these seeds. Up to 34% of the buried seeds that were exposed only to ignorant rodent foragers (individuals not responsible for burial) survived. In comparison, less than 1% of seeds buried by scatter hoarding rodents survived in previous studies, a percentage that is comparable to the low survival of unburied seeds in this study (<2%). Although unburied seeds had very low survival, increasing distance and/or seed density positively affected survival of unburied seeds. Distance to parent had no effect on buried seed survival.
Buried seed survival was most strongly and significantly determined by the fruiting status of the trees under which they occurred. Seeds experienced significantly greater predation if buried under “parent” trees that fruited during the experiment. Buried seed survival was also negatively affected by germination, as germination may signal the presence of a seed to foraging rodents. There was some indication of a positive effect of tree density on seed survival between the two plots, whereas differences in rodent abundance appear to have no effect on seed survival. Seed size and microsite characteristics had no significant effect on buried seed survival, likely due to the greater proportional effects of other factors and the longevity of A. mexicanum seeds. The results of this study were used to generate a hypothetical causal network showing how comparatively low recovery of buried seeds by ignorant foragers – combined with processes determining the removal of scatter hoarding foragers from their scattered seed caches – may affect seedling recruitment in A. mexicanum.     

The role of ants and mammals in dispersal and post-dispersal seed predation of the shrubs Grevillea (Proteaceae)

The role seed predators play in influencing the dynamics of plant populations has been little studied in Australia. The interaction of ant dispersal and seed predation on the soil seedbank in six shrubby species of Grevillea from the Sydney region of southeastern Australia was examined in selective exclusion experiments, seed array trials and placement of single seeds on the ground.Two distinct seed types in Grevillea were examined and different seed dispersal and post-dispersal seed predation patterns were associated with each: (a) seeds lacking an elaiosome were not attractive to ants and annual seed losses of between 82 and 95% were found in vegetation unburnt for greater than 8 years. Native rodents, Rattus fuscipes, and macropods, Wallabia bicolor, were responsible for these seed losses; (b) seeds with an elaiosome were rapidly handled by ants. Two functional types of ants were recognised. Most encounters were by ants that were small (Local) relative to seed size and these ants simply removed the elaiosome in situ or moved seeds only small distances (<20 cm). Some 0–24% of ant/seed encounters were by large (Removalist) ant species that were capable of moving seeds back to nests. In addition, Rattus fuscipes and Wallabia bicolor consumed at least 32–68% of seeds of Grevillea species with an elaiosome.Ants may reduce the overall levels of seed predation where seeds moved by Removalist ant species escape predation and are deposited in safe sites, hence allowing more seeds to reach the persistent soil seedbank. Mammals do not consume all seeds when ants are excluded, allowing for the potential for some seed escape from predation after seeds are discarded by Local ant species.     

Geographical and interspecific variation and the nutrient‐enrichment hypothesis as an adaptive advantage of myrmecochory

In myrmecochory, the relocation of diaspores to ant nests may lead to the enhancement of plant fitness because ant nests and their middens are often richer in essential nutrients than surrounding areas. This idea is the basis of the nutrient‐enrichment hypothesis (NEH), which suggests that nutrient enrichment may be a major selective influence in the evolution of myrmecochory. However, there is little evidence regarding whether the greater plant performance and fitness enhancement in ant nests is due to nutrient enrichment or other benefits of directed dispersal. Here, we present the results of a large‐scale seed‐sowing experiment that tests the NEH in the ant‐dispersed perennial herb Helleborus foetidus, exploring geographical and inter‐ant taxa variation. Experiments were conducted in three well‐separated regions of the Iberian Peninsula, targeting the nests of major and minor local ant dispersers (nine ant species in total) and the soil beneath maternal plants as seed destinations. Seedling emergence, survival and early establishment rates, as well as variation in soil characteristics, were obtained for each seed destination at each region. Our results do not fully support the NEH in our study system. Instead, we found that the advantage of ant nest soil for establishment in H. foetidus was conditional. Differences in soil fertility and concomitant differences in seedling establishment between ant nests and beneath the canopy of maternal plants were observed in some regions and for some ant species, but not in others. Thus, the conditional outcomes arise from inconsistencies among regions, between stages of seedling regeneration and among ant species in the advantages of being dispersed to nests. Because variation in the guilds of ant dispersers of myrmecochore plants across their ranges is common, this study illustrates the need to consider geographic and inter‐ant taxa variation for a complete evaluation of the NEH.     

Habitat effects on second-order predation of the seed predator Harpalus rufipes and implications for weed seedbank management

Seed predators provide a valuable ecosystem service to farmers by reducing densities of weed seeds, and, in turn, densities of weed seedlings they must manage. The predominant invertebrate weed seed predator in Maine, USA, agroecosystems is the carabid beetle Harpalus rufipes DeGeer. Pitfall trapping has shown that H. rufipes prefers sites with vegetative cover to fallow sites, preference speculated to be driven by predator avoidance behavior. To test this hypothesis, ‘second-order predation assays’ were developed, in which live H. rufipes prey were presented to second-order predators. Field experiments were conducted to determine foremost if H. rufipes was subject to second-order predation, and secondly, whether (a) vegetative cover affords H. rufipes protection from second-order predators, and (b) high rates of second-order predation correspond with decreased invertebrate seed predation rates. Two 72-h experiments were conducted (mid August and September 2012) at crop and non-crop sites across a 28 ha diversified farm in Stillwater, ME, USA.Second-order predation was 2.8% per day. Based on images from motion-sensing cameras, H. rufipes’ predators included birds and small mammals. Neither a relationship between second-order predation and vegetative treatment, nor an empirical relationship between second-order predation and invertebrate seed predation were detected. However, a simulation model predicted that 2.8% per day second-order predation could increase the number of seeds entering the seedbank by more than 17% annually. Additionally, complex habitats supported higher rates of second-order predation than did simple habitats.     

On the density-dependence of seed predation in<Emphasis Type="Italic"> Dipteryx micrantha</Emphasis>, a bat-dispersed rain forest tree

We studied the effect of seed density on seed predation by following the fate of bat-dispersed Dipteryx micrantha (Leguminosae) seeds deposited under bat feeding roosts. The study was conducted in Cocha Cashu biological station, Amazonian Peru, during the fruiting period of Dipteryx. Predation of Dipteryx seeds in the area is mainly by large to medium-sized rodents. Seed deposits beneath bat feeding roosts were monitored for a 13-week period in an 18-ha study area. A total of 210 seed deposits were found, and on average, seed predators encountered 22% of them during any one week. About one-third of the seed deposits escaped predation, and those deposits that had relatively few seeds were more likely to go unnoticed by rodents than were deposits with many seeds. The mean seed destruction rate was 8% per week; deposits with many seeds tended to lose a smaller proportion of their seeds to seed predators than did deposits with few seeds. Regression tests for the weekly data showed that, at the beginning of the observation period, seed predation was not density-dependent. Later, when the total seed crop beneath roosts was high, the number of seeds predated per deposit was positively density-dependent, while the proportion of seeds predated was negatively density-dependent, indicating predator satiation. Seed deposits that had been visited by seed predators once had a higher probability of being revisited the week after, especially if they contained many seeds when first encountered. This indicates that the foraging behavior of rodents may be affected by their remembering the location of seed-rich patches.     

Responses of pre‐dispersal seed predators to sequential flowering of Dipterocarps in Malaysia

Many species of Dipterocarpaceae and other plant families reproduce synchronously at irregular, multi‐year intervals in Southeast Asian forests. These community‐wide general flowering events are thought to facilitate seed survival through satiation of generalist seed predators. During a general flowering event, closely related Shorea species (Dipterocarpaceae) stagger their flowering times by several weeks, which may minimize cross pollination and interspecific competition for pollinators. Generalist, pre‐dispersal seed predators might also track flowering hosts and influence predator satiation. We addressed the question of whether pre‐dispersal seed predation differed between early and late flowering Shorea species by monitoring flowering, fruiting and seed predation intensity over two general flowering events at the Pasoh Research Forest, Malaysia. Pre‐dispersal insect seed predators killed up to 63 percent of developing seeds, with Nanophyes shoreae, a weevil that feeds on immature seeds being the most important predator for all Shorea species. This weevil caused significantly greater pre‐dispersal seed predation in earlier flowering species. Long larval development time precluded oviposition by adults that emerged from the earliest flowering Shorea on the final flowering Shorea. In contrast, larvae of weevils that feed on mature seeds before seed dispersal (Alcidodes spp.), appeared in seeds of all Shorea species almost simultaneously. We conclude that general flowering events have the potential to satiate post‐dispersal seed predators and pre‐dispersal seed predators of mature fruit, but are less effective at satiating pre‐dispersal predators of immature fruit attacking early flowering species.     

REPRODUCTIVE ECOLOGY OF JEFFERSONIA DIPHYLLA (BERBERIDACEAE)

The reproductive ecology of Jeffersonia diphylla (L.) Pers. (Berberidaceae) was investigated by studying its breeding system, ovule production, seed set, seed dispersal by ants and seed predation by rodents. This species flowers early in the spring and is facultatively autogamous. In a typical year fruit and seed set is high (90%), however, freezing temperatures from late spring frosts in 1983 and 1985 resulted in low fruit set (7% and 20%, respectively), and reduced seed set in those flowers that produced fruit. No differences in seed set between selfed and outcrossed flowers were observed over a two-yr period (1983–84). Ovule number per capsule increased with plant size as measured by leaf number. Seed set and seed wt were unaffected by leaf number unless leaves were removed after flowering was initiated. Jeffersonia diphylla is myrmecochorous. Ants removed seeds faster when seeds were placed in areas where J. diphylla plants were absent, suggesting that dispersal within J. diphylla populations is ant limited. Moreover, fresh (1 day old) seeds were removed by ants faster than 3 day old seeds. Seed predation by rodents prior to dehiscence from capsules is heavy in large populations (85–90%), and apparently negligible in small populations. Predation of seeds that are released from capsules is heavy (approx. 66%), particularly at night. Overall, seed predators consume about 96% of the seed crop in well established populations, but probably much less in small young populations. Hence, seedling recruitment is likely to be higher in small populations, whereas ramet production from rhizomes is the primary mode of propagation in large ones. The evolution of autogamy, early flowering, and myrmecochory are discussed in light of the results of this study.