Crop size is more important than neighborhood fruit availability for fruit removal of <Emphasis Type="Italic">Eugenia uniflora</Emphasis> (Myrtaceae) by bird seed dispersers

For a plant with bird-dispersed seeds, the effectiveness of seed dispersal can change with fruit availability at scales ranging from individual plants to neighborhoods, and the scale at which frugivory patterns emerge may be specific for frugivorous species differing in their life-history and behavior. The authors explore the influence of multispecies fruit availability at two local spatial scales on fruit consumption of Eugenia uniflora trees for two functional groups of birds. The authors related visitation and fruit removal by fruit gulpers and pulp mashers to crop size and conspecific and heterospecific fruit abundance to assess the potential roles that facilitative or competitive interactions play on seed dispersal. The same fruiting scenario influenced fruit gulpers (legitimate seed dispersers) and pulp mashers (inefficient dispersers) in different ways. Visits and fruit removal by legitimate seed dispersers were positively related to crop size and slightly related to conspecific, but not to heterospecific fruit neighborhoods. Visits and fruit consumption by pulp mashers was not related to crop size and decreased with heterospecific fruit availability in neighborhoods; however, this might not result in competition for dispersers. The weak evidence for facilitative or competitive processes suggest that interaction of E. uniflora with seed dispersers may depend primarily on crop size or other plant’s attributes susceptible to selection. The results give limited support to the hypothesis that spatial patterns of fruit availability influence fruit consumption by birds, and highlight the importance of considering separately legitimate and inefficient dispersers to explain the mechanisms that lie behind spatial patterns of seed dispersal.     

Geographic variation in seed removal of a myrmecochorous herb: influence of variation in functional guild and species composition of the disperser assemblage through spatial and temporal scales

The ecological influence of changes in the functional guild and species composition of ant assemblages on ant‐dispersal mutualisms is still poorly known. Using a multi‐scale approach over an 800 km range within the Iberian Peninsula, we tested the hypothesis that variation in seed removal rate was related to functional guild rather than to species composition variations of disperser assemblages in the myrmecochore herb Helleborus foetidus. At least two premises must be confirmed to validate this hypothesis: 1) ant assemblages that are dissimilar in species composition but with similar functional guild composition will not differ significantly in seed removal, and 2) assemblages with different functional guild composition will render different seed removal services. We conducted 3328 ant‐visitor censuses on 462 individual plants to identify both the species composition and functional guild variation of the ant‐disperser assemblage, and the plant seed removal rate. Functional guild composition of the ant assemblage was determined by the proportion of visits of ants acting as legitimate dispersers, facultative dispersers or elaiosome predators. Results showed that ant‐seed dispersal success seemed to be more sensitive to species composition changes of the ant assemblage than to functional guild shifts. However, this sensitivity was scale‐dependent. Thus, at the fine, inter‐individual scale, seed removal covaried with the species and functional guild composition of the ant assemblages; at the inter‐populational scale, differences in seed removal tended to be related to the dissimilarity of the assemblage species composition rather than the assemblage functional guild; finally, inter‐regional differences in seed removal were unrelated to dissimilarities of the ant assemblage composition or functional guild. Though differences in seed removal and the relative frequency of the legitimate dispersers tended to be positively correlated, none of the above premises were fully confirmed in this study. Therefore, our results did not support in full the hypothesis that the variation in seed removal was explained by shifts in functional guild composition, rather than shifts in species composition.     

POLLINATOR-MEDIATED COMPETITION,REPRODUCTIVE CHARACTER DISPLACEMENT,AND THE EVOLUTION OF SELFING IN ARENARIA UNIFLORA (CARYOPHYLLACEAE)

Ecological factors that reduce the effectiveness of cross-pollination are likely to play a role in the frequent evolution of routine self-fertilization in flowering plants. However, we lack empirical evidence linking the reproductive assurance value of selfing in poor pollination environments to evolutionary shifts in mating system. Here, we investigated the adaptive significance of prior selfing in the polymorphic annual plant Arenaria uniflora (Caryophyllaceae), in which selfer populations occur only in areas of range overlap with congener A. glabra. To examine the hypothesis that secondary contact between the two species contributed to the evolution and maintenance of selfing, we used field competition experiments and controlled hand-pollinations to measure the female fitness consequences of pollinator-mediated interspecific interactions. Uniformly high fruit set by selfers in the naturally pollinated field arrays confirmed the reproductive assurance value of selfing, whereas substantial reductions in outcrosser fruit set (15%) and total seed production (20–35%) in the presence of A. glabra demonstrated that pollinator-mediated interactions can provide strong selection for self-pollination. Heterospecific pollen transfer, rather than competition for pollinator service, appears to be the primary mechanism of pollinator-mediated competition in Arenaria. Premating barriers to hybridization between outcrossers and A. glabra are extremely weak. The production of a few inviable hybrid seeds after heterospecific pollination and intermediate seed set after mixed pollinations indicates that A. glabra pollen can usurp A. uniflora ovules. Thus, any visit to A. uniflora by shared pollinators carries a potential female fitness cost. Moreover, patterns of fruit set and seed set in the competition arrays relative to controls were consistent with the receipt of mixed pollen loads, rather than a lack of pollinator visits. Competition through pollen transfer favors preemptive self-pollination and may be responsible for the evolution of a highly reduced floral morphology in A. uniflora selfers as well as their current geographical distribution.     

Putative local adaptations modulate the interactions of the carnivorous plant Drosera uniflora Willd (1809) (Droseraceae) with cushion and shrub nurse plants

Cushion and shrub plants are typical high mountain nurse plants. In Magellanic tundras of the Cordillera del Sarao on the coast of Chile, the carnivorous plant Drosera uniflora grows in association with cushions of Donatia fascicularis and the shrubs Chusquea montana var. nigricans and Lepidothamnus fonkii. The different microhabitats for recruitment, in addition to the limited gene flow of D. uniflora, enable us to hypothesise that this plant manifests putative local adaptations, expressed in seed germination and population abundance. Our aim was to evaluate the local adaptation of D. uniflora to cushion and shrub microhabitats by estimating germination and abundance. Local adaptation of seed germination was determined by means of reciprocal transplant experiments. Abundance was determined in small plots located inside, on the edge and outside the nurse plants. Seed origin and growth substrate play a decisive role in D. uniflora germination. Seeds that originate from cushions plant habitat and germinate in the same substrate do so in greater numbers than when those which originate from shrubby habitats. Conversely, seeds that originate from shrubby habitat and germinate in the same substrate do not exhibit significant differences. Abundance was always greater inside than outside of the nurse plants. We concluded that availability and quality of different microhabitats for seed germination and recruitment, together with a limited gene flow, would trigger putative local adaptations in D. uniflora, modulating a long history of interactions with the plants that act as nurse plants.     

A 10-year study of changes in forest vegetation on Silhouette island, Seychelles

A 10 year study of forest communities on Silhouette island, Seychelles demonstrates stability of forest composition in most areas over this time-scale. Areas with heavy invasion by alien species were found to be regenerating, particularly with the rapid loss of Clidemia hirta. This is attributed to the abundance of well-adapted native plants allowing competitive exclusion to take place, throughout competition for light. It was noted that invasive plant species tend to be unstable on the rocky slopes covered by native high forest. A high rate of tree fall and limited seed dispersal may reduce the impact of the invasive tree Paraserianthes falcataria in the future. Other species such as Cinnamomum verum and Psidium cattleianum may persist as major invaders due to wider seed dispersal.     

Spatial and temporal variation in dispersal pattern of an invasive pine

Understanding dispersal ability of an invasive species is crucial for describing its potential spread. Despite this, we still know little about the dispersal potential of many invasive species. We explored dispersal spectra in Pinus strobus, an invasive tree in sandstone areas in Central Europe. We studied dispersal of the species using distribution of self-sown trees in the field. We compared these observed data with theoretical dispersal curves derived using information on wind speed, seed terminal velocity and tree height. Finally, we fitted various empirical dispersal curves to the observed data. All the analyses based on the observed field data were done for the whole dataset, and for the dataset divided by habitat types and age categories of the self-sown trees. P. strobus seeds can disperse up to 757.5 m from the source. The observed data fall within the confidence intervals of the predictions based on a negative exponential model. When comparing different dispersal functions fitted to the data, it was not easy to decide which of the dispersal curves provides the overall best fit. This was because different functions were the best predictors of different parts of the dispersal curve. We suggest that future studies should provide not only empirical fitted dispersal curves but also observed data and provide estimates of confidence intervals. Such information will provide insights into the reliability of the dispersal estimates in general and help to evaluate the predictive power of the different models.     

Vertebrate Fruit Removal and Ant Seed Dispersal in the Neotropical Ginger Renealmia alpinia (Zingiberaceae)*

Plants frequently display fruit characteristics that support multiple seed‐dispersal syndromes. These ambiguous characteristics may reflect the fact that seed dispersal is usually a complex process involving multiple dispersers. This is the case for the Neotropical ginger Renealmia alpinia (Zingiberaceae). It was originally suggested that the aromatic fruits of R. alpinia located at the base of the plant are adapted for terrestrial mammal seed dispersal. However, the dark‐purple coloration of the fruits and bright orange aril surrounding the seeds suggest that birds may play a role in R. alpinia seed dispersal. At La Selva Biological Station, Costa Rica, we used camera traps to record vertebrate visits to infructescences of R. alpinia. Most visitors were toucans and aracaris (Ramphastidae). However fruits were also removed by terrestrial mammals (coatis and armadillos). In addition to vertebrate fruit removal, some of the fruits dehisce and the seeds that fall on the ground are dispersed by ants. Fruitfall traps showed that 77 percent of fruits are removed by vertebrates. However, 15 percent of fruits fall to the base of parent plants to be potentially dispersed by ants. Experiments using a laboratory ant colony showed that ants are effective seed dispersers of R. alpinia. Ant seed manipulation increased germination success and reduced time to germination. In conclusion, primary seed dispersal in the Neotropical ginger R. alpinia is mostly performed by birds, additionally ants are effective dispersers at short distances. Seed dispersal in R. alpinia is a complex process involving a diverse array of dispersal agents.     

Dust seed production and dispersal in Swedish Pyroleae species

Dust seeds are the smallest seeds in angiosperms weighing just about a few micrograms. These seeds are characteristic of most orchids, and several studies have been performed on seed features, fecundity and dispersal of orchid dust seeds. In this study we examine seed features, seed production and seed dispersal in another plant group with dust seeds, the Pyroleae (Monotropoideae, Ericaceae), focusing on six species: Pyrola chlorantha, P. minor, P. rotundifolia, Chimaphila umbellata, Moneses uniflora and Orthilia secunda. Seed production per capsule among these species was in the range between ca 1000 and 7800, and seed production per capsule bearing shoot was in the range between ca 7000 and 60 000. Combining our results with published information on pollen‐ovule ratios suggests that these Pyroleae species have a generally efficient pollination system. The most fecund species was P. minor, the only species among the investigated that is probably largely self‐pollinating. The investigated Pyroleae species have a seed production comparable to the less fecund orchid species. We studied seed dispersal in the field in one of the species, P. chlorantha. Despite the extremely small and potentially buoyant seeds, the vast majority of seeds are deposited close to the seed source, within a few meters. Further studies on the recruitment ecology of the investigated Pyroleae species are currently under way.     

Are subordinate ants the best seed dispersers? Linking dominance hierarchies and seed dispersal ability in myrmecochory interactions

True myrmecochory involves the dispersal of elaiosome-bearing seeds by ants. Between the guild of ants that are attracted to these seeds, only a few of them will act as effective dispersers, that is, transporting the seeds to suitable sites (the nests) for germination and plant establishment. Ant communities are known to be highly hierarchical, and subordinate ants quickly deliver resources to their nest rather than consuming it on-site, thereby avoiding encounters with more dominant species. As a result of a series of studies that were carried out during summer in semi-arid Northwest Argentina, we have found that the most important seed disperser of the myrmecochorous plant Jatropha excisa Griseb. (Euphorbiaceae), the ant Pogonomyrmex cunicularius pencosensis Forel, was the most subordinate species during interspecific interactions. The daily timing of release of the J. excisa seeds through ballistic dispersal increased their probability of being removed by the highly thermophilic P. cunicularius pencosensis. Foraging during the warmest hours of the day allowed P. cunicularius pencosensis ants to avoid the risk of interference competition with dominant species, which also behaved as elaiosome predators. As a conclusion, subordinance behaviour appears to be integral to successful myrmecochory, and also the timing of seed release plays a key role in shaping the dynamics of myrmecochorous interactions. Therefore, ant-dispersed plants should not only favour their discovery by subordinate ants, but also should present their seeds at those times of the day when the behaviourally dominant ants are less active.     

Interactions among phytophagous insect species colonizing cones of white fir (Abies concolor)

Summary The insect complex colonizing white fir (Abies concolor [Gord. and Glend.] Lindl.) cones is composed of eleven species that can be separated into three feeding guilds: the seed-mining guild, Megastigmus pinus Parfitt, M. rafni Hoffmeyer, Earomyia abietum McAlpine; the cone-and seed-mining guild, Dioryctria abietivorella Grote, Eucosma prob. siskiyouana (Kearfoot), Cydia prob. bracteatana (Fernald), Barbara sp.; the scale-and bract-feeding guild, Asynapta hokinsi (Felt), Dasineura prob. abiesemia Foote, Ressiliella conicola (Foote), Lasiomma abietis Huckett). In three of four study sites the cone crop decreased from one year to the next. As cone crop size decreased there was a concomitant increase in the percent of cones with more than one species. In addition, there was a shift toward an increased co-occurrence of members of different guilds within a single cone. Both positive and negative interactions were detected between selected species-pairs. Present-day competition was only inferred between species-pairs belonging to the cone and seed-mining guild. Decreasing resources over time, combined with increasing insect populations and the absence of acceptable alternate hosts appeared to be important factors for setting conditions conducive to interspecific competition. It is hypothesized, that the aperiodicity of white fir cone crops was important in keeping insect populations below levels which would result in interspecific competition. The possible elimination of aperiodicity in cone crops of white fir, such as might occur in managed seed orchards, may lead to decreased species diversity via competitive exclusion and thereby simplify development of IPM programs.     

Hydrodynamics and seed dispersal in the lower Amazon

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Coexistence of plant species in a biodiversity hotspot is stabilized by competition but not by seed predation

Understanding the mechanisms of species coexistence is a key task for ecology. Recent theory predicts that both competition and predation (which causes apparent competition among prey) can either promote or limit species coexistence. Both mechanisms cause negative interactions between individuals, and each mechanism promotes stable coexistence if it causes negative interactions to be stronger between conspecifics than between heterospecifics. However, the relative importance of competition and predation for coexistence in natural communities is poorly known. Here, we study how competition and apparent competition via pre‐dispersal seed predators affect the long‐term fecundity of Protea shrubs in the fire‐prone Fynbos biome (South Africa). These shrubs store all viable seeds produced since the last fire in fire‐proof cones. Competitive effects on cone number and pre‐dispersal seed predation reduce their fecundity and can thus limit recruitment after the next fire. In 27 communities comprising 49 990 shrubs of 22 Protea species, we measured cone number and per‐cone seed predation rate of 2154 and 1755 focal individuals, respectively. Neighbourhood analyses related these measures to individual‐based community maps. We found that conspecific neighbours had stronger competitive effects on cone number than heterospecific neighbours. In contrast, apparent competition via seed predators was comparable between conspecifics and heterospecifics. This indicates that competition stabilizes coexistence of Protea species, whereas pre‐dispersal seed predation does not. Larger neighbours had stronger competitive effects and neighbours with large seed crops exerted stronger apparent competition. For 97% of the focal plants, competition reduced fecundity more than apparent competition. Our results show that even in communities of closely related and ecologically similar species, intraspecific competition can be stronger than interspecific competition. On the other hand, apparent competition through seed predators need not have such a stabilizing effect. These findings illustrate the potential of ‘community demography’, the demographic study of multiple interacting species, for understanding plant coexistence.     

Negative density dependence of seed dispersal and seedling recruitment in a Neotropical palm

Negative density dependence (NDD) of recruitment is pervasive in tropical tree species. We tested the hypotheses that seed dispersal is NDD, due to intraspecific competition for dispersers, and that this contributes to NDD of recruitment. We compared dispersal in the palm Attalea butyracea across a wide range of population density on Barro Colorado Island in Panama and assessed its consequences for seed distributions. We found that frugivore visitation, seed removal and dispersal distance all declined with population density of A. butyracea, demonstrating NDD of seed dispersal due to competition for dispersers. Furthermore, as population density increased, the distances of seeds from the nearest adult decreased, conspecific seed crowding increased and seedling recruitment success decreased, all patterns expected under poorer dispersal. Unexpectedly, however, our analyses showed that NDD of dispersal did not contribute substantially to these changes in the quality of the seed distribution; patterns with population density were dominated by effects due solely to increasing adult and seed density.     

Potential for ants and vertebrate predators to shape seed-dispersal dynamics of the invasive thistles <Emphasis Type="Italic">Cirsium arvense</Emphasis> and <Emphasis Type="Italic">Carduus nutans</Emphasis> in their introduced range (North America)

The population dynamics of invasive plants are influenced by positive and negative associations formed with members of the fauna present in the introduced range. For example, mutualistic associations formed with pollinators or seed dispersers may facilitate invasion, but reduced fitness from attack by native herbivores can also suppress it. Since population expansion depends on effective seed dispersal, interactions with seed dispersers and predators in a plant species introduced range may be of particular importance. We explored the relative contributions of potential seed dispersers (ants) and vertebrate predators (rodents and birds) to seed removal of two diplochorous (i.e., wind- and ant-dispersed), invasive thistles, Cirsium arvense and Carduus nutans, in Colorado, USA. We also conducted behavior trials to explore the potential of different ant species to disperse seeds, and we quantified which potential ant dispersers were prevalent at our study locations. Both ants and vertebrate predators removed significant amounts of C. arvense and C. nutans seed, with the relative proportion of seed removed by each guild varying by location. The behavior trials revealed clear seed preferences among three ant species as well as differences in the foragers’ abilities to move seeds. In addition, two ant species that acted as potential dispersal agents were dominant at the study locations. Since local conditions in part determined whether dispersers or predators removed more seed, it is possible that some thistle populations benefit from a net dispersal effect, while others suffer proportionally more predation. Additionally, because the effectiveness of potential ant dispersers is taxon-specific, changes in ant community composition could affect the seed-dispersal dynamics of these thistles. Until now, most studies describing dispersal dynamics in C. arvense and C. nutans have focused on primary dispersal by wind or pre-dispersal seed predation by insects. Our findings suggest that animal-mediated dispersal and post-dispersal seed predation deserve further consideration.     

Interference effects of Cassia uniflora Mill on Parthenium hysterophorus L.

In some parts of Karnataka and Maharashtra states (India), a leguminous undershrub, Cassia uniflora has effectively replaced the problematic weed Parthenium hysterophorus (parthenium). The studies revealed that the phenolic leachates from different parts of C. uniflora, specially from germinating seeds significantly inhibited the germination of parthenium seeds and retarded the growth of seedlings from the successfully germinated parthenium seeds. At field level, seedlings of C. uniflora outcompeted the seedlings of summer generation of parthenium causing reduction in height, dry weight and number of inflorescences of the latter which ultimately resulted in reduction of their seed output. Because of its longer life span, C. uniflora effectively prevented the establishment of parthenium plants of the winter generation in its area of growth. Results indicate that C. uniflora replaces parthenium through interference.     

Incorporating patterns of disperser behaviour into models of seed dispersal and its effects on estimated dispersal curves

The processes determining where seeds fall relative to their parent plant influence the spatial structure and dynamics of plant populations and communities. For animal dispersed species the factors influencing seed shadows are poorly understood. In this paper we test the hypothesis that the daily temporal distribution of disperser behaviours, for example, foraging and movement, influences dispersal outcomes, in particular the shape and scale of dispersal curves. To do this, we describe frugivory and the dispersal curves produced by the southern cassowary, Casuarius casuarius, the only large-bodied disperser in Australia’s rainforests. We found C. casuarius consumed fruits of 238 species and of all fleshy-fruit types. In feeding trials, seeds of 11 species were retained on average for 309 min (±256 SD). Sampling radio-telemetry data randomly, that is, assuming foraging occurs at random times during the day, gives an estimated average dispersal distance of 239 m (±207 SD) for seeds consumed by C. casuarius. Approximately 4% of seeds were dispersed further than 1,000 m. However, observation of wild birds indicated that foraging and movement occur more frequently early and late in the day. Seeds consumed early in the day were estimated to receive dispersal distances 1.4 times the ‘random’ average estimate, while afternoon consumed seeds received estimated mean dispersal distances of 0.46 times the ‘random’ estimate. Sampling movement data according to the daily distribution of C. casuarius foraging gives an estimated mean dispersal distance of 337 m (±194 SD). Most animals’ behaviour has a non-random temporal distribution. Consequently such effects should be common and need to be incorporated into seed shadow estimation. Our results point to dispersal curves being an emergent property of the plant–disperser interaction rather than being a property of a plant or species.     

Limits and effects of invasion by the nonindigenous wetland plant <Emphasis Type="Italic">Lythrum salicaria</Emphasis> (purple loosestrife): a seed bank analysis

We used seed bank analyses to investigate the role of dispersal in limiting invasion by Eurasian Lythrum salicaria within and among North American wetlands, and the changes in seed bank diversity associated with this invader. We compared the number and species composition of seedlings emerging from soil sampled in 11 uninvaded wetlands and paired uninvaded and invaded sites within 10 invaded wetlands under both seedling competition and noncompetitive conditions. Almost no L. salicaria emerged in samples from uninvaded wetlands, indicating dispersal limitation despite prodigious seed production in nearby wetlands. However L. salicaria emerged in all samples from uninvaded sites in invaded wetlands, suggesting environmental limits on establishment within invaded wetlands. Conditions that provided opportunities for seedlings to compete reduced survival of Typha spp. but not L. salicaria seedlings. However, this was due to species-specific differences in post-emergence mortality rather than response to competition. Competition did reduce seedling mass, but this effect did not differ among species. Species richness of emerging seedlings was lower for invaded than uninvaded wetlands. Lower seed bank richness may be a cause or consequence of L. salicaria invasion. Efforts to reduce seed dispersal to uninvaded wetlands would likely slow the spread of this invader.     

Seasonally different modes of seed dispersal in the prostrate annual, <Emphasis Type="Italic">Chamaesyce maculata</Emphasis> (L.) Small (Euphorbiaceae), with multiple overlapping generations

The modes of seed dispersal in the prostrate annual, Chamaesyce maculata, with multiple overlapping generations were investigated. We found that C. maculata has two modes of seed dispersal; autochory in the summer and myrmecochory in the autumn. Seasonally different modes of seed dispersal have not been known in other plant species. The large proportion of seeds produced in the summer was positioned further than the expanse of the parent plants by automatic mechanical seed dispersal. Therefore, autochory would be effective for avoiding competition between parent and offspring plants. No autochory occurred in the seeds produced in the autumn. The seeds of C. maculata without an elaiosome were dispersed by seed-collecting ants in the autumn. Although 18 ant species in total visited the plants of C. maculata at the 50 sites investigated, only two ant species, Tetramorium tsushimae and Pheidole noda frequently carried the seeds of C. maculata. The low frequency of seeds carried out of the nest by P. noda suggests that the workers of P. noda carry the seeds as food into their nest. So, P. noda might be a less effective seed disperser for C. maculata, corresponding to the effectiveness of seed dispersal by harvester ants. However, T. tsushimae ants frequently carried the seeds into and out of their nest, suggesting that T. tsushimae do not regard the seeds of C. maculata as a food resource. Thus, T. tsushimae may be an effective seed disperser for C. maculata.     

Trophic and non-trophic pathways mediate apparent competition through post-dispersal seed predation in a Patagonian mixed forest

Post‐dispersal predation can be a major source of seed loss in temperate forests. Little is known, however, about how predator‐mediated indirect interactions such as apparent competition alter survival patterns of canopy tree seeds. Understorey plants may enhance tree seed predation by providing sheltered habitat to granivores (non‐trophic pathway). In addition, occurrence of different tree seeds in mixed patches may lead to short‐term apparent competition between seed types, because of the granivores’ foraging response to changes in food patch quality (trophic pathway). We hypothesised that understorey bamboo cover and mixing of seed species in food patches would both increase tree seed predation in a Nothofagus dombeyi?Austrocedrus chilensis forest in northern Patagonia, Argentina. Seed removal experiments were conducted for three consecutive years (2000–2002) differing in overall granivory rates. Seed patch encounter and seed removal rates were consistently higher for the larger and more nutritious Austrocedrus seeds than for the smaller Nothofagus seeds. Seed removal was greater beneath bamboo than in open areas. This apparent competition pathway was stronger in a low‐predation year (2000) than in high‐predation years (2001–2002), suggesting a shift in microhabitat use by rodents. Patch composition had a significant, though weaker, impact on seed survival across study years, whereas seed density per patch enhanced encounter rates but did not influence seed removal. Removal of the less‐preferred Nothofagus seeds increased in the presence of Austrocedrus seeds, but the reciprocal indirect effect was not observed. However, this non‐reciprocal apparent competition between seed species was only significant in the high‐predation years. Our study shows that granivore‐mediated indirect effects can arise through different interaction pathways, affecting seed survival patterns according to the predator's preference for alternative seed types. Moreover, results indicate that the occurrence and relative strength of trophic vs non‐trophic pathways of apparent competition may change under contrasting predation scenarios.     

Seed dispersal in Erythronium dens-canis L. (Liliaceae): variation among habitats in a myrmecochorous plant

Erythronium dens-canis is a geophyte which produces a single flower each season. The fruits produce small seeds with relatively large elaiosomes. We performed experiments to investigate primary and secondary seed dispersal mechanisms of this species in different habitats in the western part of the Cantabrian Range in northwest Spain. Sticky traps were used to measure primary dispersal of seeds up to 0.5 m from mother plants. Seed cafeteria experiments were performed in different habitats to examine the role of ants and rodents in secondary seed transport and seed predation. Our results indicate that: (a) primary seed dispersal is positively skewed (99% of seeds fall within 20 cm of the mother plant) and seed dispersal distances vary significantly among plants; (b) secondary dispersal is exclusively by myrmecochory, although the proportion of seeds removed by ants differs significantly among habitats; (c) ant species composition and abundances vary among habitats; and (d) freshly dropped seeds are more likely to be removed than seeds that have begun to dry out. We conclude that secondary dispersal of seeds is greatly influenced by habitat but not by small-scale microhabitat. This revised version was published online in June 2006 with corrections to the Cover Date.