Factors affecting post-dispersal seed survival in a tropical forest

Summary Using the subcanopy tree Faramea occidentalis in Panama, I studied post-dispersal seed survival as a function of five characteristics describing seed locations. By simultaneously considering distance from a conspecific adult, size of the nearest conspecific adult, leaf litter quantity, proximity to logs or tree trunks, and whether or not the seed was in a gap, I was able to analyze the influences of individual factors, as well as the interactions among factors. Seed survival was significantly less in treefall gaps than in the forest understory. Seed survival was also influenced by the size of the nearest adult but in a complex interaction with distance to an adult. For seeds beneath adults, survival decreased with increasing tree size, while for seeds away from adults, survival was independent of the size of the nearest conspecific adult. Distance did not directly affect seed survival, nor did the quantity of leaf litter or the proximity to a tree trunk or a log. In a separate analysis, the relationship between distance and seed survival was consistent over four years, suggesting that single cohort studies may provide accurate insights into the consequences of dispersal. In contrast, the spatial locations of surviving seeds were not consistent over the four-year period. Transects with high survival one year did not tend to have high survival in other years, and the locations of surviving seeds in any particular year could not be predicted from the knowledge of where seeds survived in other years. While survival is patchy within a year, the locations of patches shift from year to year.     

Effects of frugivore impoverishment and seed predators on the recruitment of a keystone palm

Many plant species are threatened as a result of extinction of their large-bodied frugivores all over the world. Additionally, introduced herbivores and seed predators may cause severe pressure on early stages of plant recruitment. We studied the seed dispersal and seed predation of the keystone palm Euterpe edulis on a land-bridge island with a highly impoverished frugivore fauna and overabundant seed predators, and in a continuous Atlantic forest in Brazil. While the diversity of avian seed dispersers and predators was higher on the mainland, the abundance of seed dispersers was 4-fold higher on the island. Turdus flavipes was responsible for 72% and 96% of seeds removed in the island and mainland, respectively. However, the higher density of T. flaviceps on the island did not result in higher seed removal. In fact, seed removal rate was 1.7 times lower there than on the mainland, probably due to the aggressive behavior of the major seed disperser who defend palm fruits. Seed predation, on the other hand, was markedly higher on the island, where nearly 100% of seeds were preyed upon, but only 0.3% on the mainland. As a consequence of higher seed predation the population of E. edulis has few numbers of seedlings and saplings on the island. Therefore, management of the seed predator populations on the island is a key priority for recovering the natural population of this keystone palm and the frugivores that depend on its fruits.     

Destination‐based seed dispersal homogenizes genetic structure of a tropical palm

As the dominant seed dispersal agents in many ecosystems, frugivorous animals profoundly impact gene movement and fine‐scale genetic structure of plants. Most frugivores engage in some form of destination‐based dispersal, in that they move seeds towards specific destinations, resulting in clumped distributions of seeds away from the source tree. Molecular analyses of dispersed seeds and seedlings suggest that destination‐based dispersal may often yield clusters of maternal genotypes and lead to pronounced local genetic structure. The long‐wattled umbrellabird Cephalopterus penduliger is a frugivorous bird whose lek mating system creates a species‐specific pattern of seed dispersal that can potentially be distinguished from background dispersal processes. We used this system to test how destination‐based dispersal by umbrellabirds into the lek affects gene movement and genetic structure of one of their preferred food sources Oenocarpus bataua, a canopy palm tree. Relative to background dispersal processes, umbrellabird mating behaviour yielded more diverse seed pools in leks that included on average five times more seed sources and a higher incidence of long‐distance dispersal events. This resulted in markedly lower fine‐scale spatial genetic structure among established seedlings in leks than background areas. These species‐specific impacts of destination‐based dispersal illustrate how detailed knowledge of disperser behaviour can elucidate the mechanistic link driving observed patterns of seed movement and genetic structure.     

Fire survival of Cape Proteaceae-influence of fire season and seed predators

Many Cape Proteaceae store seed reserves in closed cones on the plant and rely entirely on these reserves for episodic recruitment after fires. Population size is sensitive to intervals between fires but also to fire season. Populations can be nearly eliminated by successive winter or spring fires. Three hypotheses explaining seasonal variation in recruitment were tested: (a) seeds germinate immediately after fire but seedlings die from summer drought; (b) seeds remain dormant over summer but the longer the delay between seed release after fire and germination 1) the greater the competition between seedlings and resprouts, or 2) the greater the seed losses to predators and/or decay before germination.Drought-avoiding dormancy occurred in 9 of 11 Cape Proteaceae studied, all of which delayed germination to autumn or winter. Seedling emergence and survival was not significantly increased after removal of competitors by methyl bromide poisoning. Seed predation, measured by exclosures, however, significantly reduced seed reserves before germination and also number of seedlings emerging. Post emergence seedling predation was negligible in the burn in contrast to adjacent mature vegetation where seedling predation was very heavy.The role of germination cues and rodent behaviour in controlling population recruitment is discussed and it is concluded that a knowledge of both is essential for predicting vegetation dynamics in this system.Acknowledgements: I am grateful to J. Vlok and R. America for field assistance and J. Breytenbach, F. Kruger, P. Slingsby and J. Yensch for observation and comment. This work was supported by the Directorate of Forestry, South Africa as part of their conservation research programme.     

Tri-trophic interactions affect density dependence of seed fate in a tropical forest palm

Natural enemies, especially host-specific enemies, are hypothesised to facilitate the coexistence of plant species by disproportionately inflicting more damage at increasing host abundance. However, few studies have assessed such Janzen-Connell mechanisms on a scale relevant for coexistence and no study has evaluated potential top-down influences on the specialized pests. We quantified seed predation by specialist invertebrates and generalist vertebrates, as well as larval predation on these invertebrates, for the Neotropical palm Attalea butyracea across ten 4-ha plots spanning 20-fold variation in palm density. As palm density increased, seed attack by bruchid beetles increased, whereas seed predation by rodents held constant. But because rodent predation on bruchid larvae increased disproportionately with increasing palm density, bruchid emergence rates and total seed predation by rodents and bruchids combined were both density-independent. Our results demonstrate that top-down effects can limit the potential of host-specific insects to induce negative-density dependence in plant populations.     

Seed size in tropical trees: a comparative study of factors affecting seed size in Peruvian angiosperms

Evolutionary comparative methods taking into account the potential effects of relatedness reveal that, among 202 species of animal-dispersed tropical woody angiosperms from Peru, large seeds may be an adaptation to: (1) dispersal by mammals rather than by birds and (2) greater plant height. Using the most powerful techniques currently available, appropriate comparisons were made between related species or species groups at all taxonomic levels, thus allowing use of all species in the data set. Even so, the results show no evidence of adaptation (i.e., a functional relationship) of seed mass to successional syndrome or to growth form (trees or vines). In previous cross-species analyses, seed mass has been linked to both of these ecological variables. The lack of relationship between seed mass and habitat use or growth form may be due to targeting the wrong trait to predict these two variables, or because seed mass is subject to exaptation, i.e., a particular seed mass has arisen for other reasons, but allows invasion into a habitat and currently serves to maintain that species in the habitat. Differentiation between these two explanations for the lack of correlation between seed mass and successional syndrome or growth form would entail species-by-species examination of the function of seed mass under natural conditions. The effect of relatedness on the ecological correlates of seed mass and plant height is determined for each of three taxonomic schemes. It is concluded that the three schemes do not differ greatly in the results obtained.     

A strong conditional mutualism limits and enhances seed dispersal and germination of a tropical palm

Seed predation and seed dispersal can have strong effects on early life history stages of plants. These processes have often been studied as individual effects, but the degree to which their relative importance co-varies with seed predator abundance and how this influences seed germination rates is poorly understood. Therefore, we used a combination of observations and field experiments to determine the degree to which germination rates of the palm Astrocaryum mexicanum varied with abundance of a small mammal seed predator/disperser, Heteromys desmarestianus, in a lowland tropical forest. Patterns of abundance of the two species were strongly related; density of H. desmarestianus was low in sites with low density of A. mexicanum and vice versa. Rates of predation and dispersal of A. mexicanum seeds depended on abundance of H. desmarestianus; sites with high densities of H. desmarestianus had the highest rates of seed predation and lowest rates of seed germination, but a greater total number of seeds were dispersed and there was greater density of seedlings, saplings, and adults of A. mexicanum in these sites. When abundance of H. desmarestianus was experimentally reduced, rates of seed predation decreased, but so did dispersal of A. mexicanum seeds. Critically, rates of germination of dispersed seeds were 5 times greater than undispersed seeds. The results suggest that the relationship between A. mexicanum and H. desmarestianus is a conditional mutualism that results in a strong local effect on the abundance of each species. However, the magnitude and direction of these effects are determined by the relative strength of opposing, but related, mechanisms. A. mexicanum nuts provide H. desmarestianus with a critical food resource, and while seed predation on A. mexicanum nuts by H. desmarestianus is very intense, A. mexicanum ultimately benefits because of the relatively high germination rates of its seeds that are dispersed by H. desmarestianus.     

Some factors affecting the composition of tropical ichneumonid faunas

An increasing accumulation of data shows that tropical ichneumonid faunas are no more species-rich than extra-tropical ones, despite the fact that most of their host groups show increased tropical species-richness. This lack of increase in ichneumonid species-richness can be attributed to the absence of groups whose hosts are not present (e.g. Ctenopelmatinae) and poor tropical representation by many groups of diurnal koinobionts (e.g. Campopleginae). Low host density has been postulated as a barrier to tropical koinobiont species-richness, but it is here suggested that this is not the only limiting factor as groups of nocturnal koinobionts, such as the Ophioninae, show increased tropical species-richness. It is postulated that koinobionts have the capability of being able to locate sparse hosts, but as they host-search in flight, a prolonged daytime host-searching period in the tropics would expose them to a high level of predation pressure. By being active at night koinobionts can avoid diurnally active predators. It is also postulated that sparse hosts may be located more easily at night and more habitats may be climatically suitable for ichneumonid activity when they are not subject to direct sunlight. Idiobionts, such as the Mesostenini and Pimplini, are more species-rich and morphologically diverse in the tropics than they are in extra-tropical regions. It is suggested that this results from the fact that tropical idiobionts can be active during the whole of the diapausing period, when their hosts are available, whereas activity by temperate idiobionts is prevented by inclement weather. Although many idiobionts are probably less exposed to predators than koinobionts, many have evolved obvious protective devices.     

Contemporary and historic factors influence differently genetic differentiation and diversity in a tropical palm

Population genetics theory predicts loss in genetic variability because of drift and inbreeding in isolated plant populations; however, it has been argued that long-distance pollination and seed dispersal may be able to maintain gene flow, even in highly fragmented landscapes. We tested how historical effective population size, historical migration and contemporary landscape structure, such as forest cover, patch isolation and matrix resistance, affect genetic variability and differentiation of seedlings in a tropical palm (Euterpe edulis) in a human-modified rainforest. We sampled 16 sites within five landscapes in the Brazilian Atlantic forest and assessed genetic variability and differentiation using eight microsatellite loci. Using a model selection approach, none of the covariates explained the variation observed in inbreeding coefficients among populations. The variation in genetic diversity among sites was best explained by historical effective population size. Allelic richness was best explained by historical effective population size and matrix resistance, whereas genetic differentiation was explained by matrix resistance. Coalescence analysis revealed high historical migration between sites within landscapes and constant historical population sizes, showing that the genetic differentiation is most likely due to recent changes caused by habitat loss and fragmentation. Overall, recent landscape changes have a greater influence on among-population genetic variation than historical gene flow process. As immediate restoration actions in landscapes with low forest amount, the development of more permeable matrices to allow the movement of pollinators and seed dispersers may be an effective strategy to maintain microevolutionary processes.     

Assessing the potential for indirect interactions between tropical tree species via shared insect seed predators

Natural enemies of plants have the potential to influence the dynamics of plant populations and the structure of plant communities. In diverse tropical forests, research on the effects of plant enemies has largely focused on the diversity-enhancing effects of highly specialized enemies, while the community-level effects of enemies with broader diets have rarely been considered. We investigated the community of insect seed predators interacting with seven tree species in the family Lauraceae on Barro Colorado Island (Panama). We present one of the first quantitative food webs for pre-dispersal insect seed predators and their host plants, and use the information in the web to assess the potential for indirect interactions between the tree species. Our data suggest that there is high potential for indirect interactions between Lauraceae species via their shared seed predators. The strength and direction of these interactions are largely unrelated to the phylogenetic distance and trait similarity between species but are likely governed by the volume of fruit produced by each tree species.     

A highly resolved food web for insect seed predators in a species‐rich tropical forest

The top‐down and indirect effects of insects on plant communities depend on patterns of host use, which are often poorly documented, particularly in species‐rich tropical forests. At Barro Colorado Island, Panama, we compiled the first food web quantifying trophic interactions between the majority of co‐occurring woody plant species and their internally feeding insect seed predators. Our study is based on more than 200 000 fruits representing 478 plant species, associated with 369 insect species. Insect host‐specificity was remarkably high: only 20% of seed predator species were associated with more than one plant species, while each tree species experienced seed predation from a median of two insect species. Phylogeny, but not plant traits, explained patterns of seed predator attack. These data suggest that seed predators are unlikely to mediate indirect interactions such as apparent competition between plant species, but are consistent with their proposed contribution to maintaining plant diversity via the Janzen–Connell mechanism.     

Some factors affecting the detection period of aphid remains in predators using ELISA

Quantitative enzyme-linked immunosorbent assay (ELSIA) was used to detect antigens of the aphid Sitobion averae (F.) in the guts of Linyphiidae, Carabidae and Staphylinidae. The effects of temperature, both constant and variable, and size of meal on the detection period and antigen decay rate were studied in the laboratory. Predators fed freshly-killed aphids were subsequently kept at one of several temperature regimes for a period from 0 to 13 days before being assayed for aphid remains. The proportion of prey remaining at intervals after feeding was measured. Curves were fitted to transformed data and the detection period estimated. The rate of decline in detectable remains was temperature-related, with the rate increasing as temperature increased. Prey remains in Staphylinidae declined much faster than in either Carabidae or Linyphiidae. In all but one case the decline was exponential with time. Variable temperature regimes produced results very similar to those obtained under conditions of constant temperature. Meal size produced a considerable difference in the amount of aphid remains detectable but little difference in the rate of decline or the estimated detection period. Data of the above types are a prerequisite for postmortem quantification of predator meals.

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Influence de certains facteurs sur la détection par ELISA de vestiges de pucerons à l'intérieur des prédateurs

Résumé La recherche d'antigènes du puceron Sitobion avenae F. a été effectuée dans les tractus digestifs de Linyphiidae, Carabidae et Staphylinidae par ELISA (adsorption des antigènes sur l'anticorps fixé et dosage par l'anticorps enzymatiquement marqué). Les influences de la température, —soit constante, soit périodique —, de l'importance du repas, du moment de la détection et de la vitesse de disparition des antigènes ont été examinées au laboratoire. Des prédateurs alimentés en pucerons tués depuis peu ont été conservés de un à 13 jours à defférents régimes de températures avant la recherche de vestiges de pucerons. Une courbe d'atalonnage a été établie pour permettre la conversion de la valeur de la densité optique observée en mg de vestiges de pucerons en fonction du temps écoulé depuis le repas. Les données ont été soumises à une transformation angulaire et les courbes ajustées pour estimer la période de détection.La vitesse de disparition de vestiges détectables a augmenté avec la température. Les vestiges ont disparu beaucoup plus vite dans les Staphylinidae que dans les Carabidae ou les Linyphiidae. A l'exception d'un cas, la disparition était une expontentielle du temps. Les thermopériodes ont produit les mêmes effets que les températures constantes. La taille du repas a provoqué des différences considérables sur la quantité de pucerons décelable, mais peu sur la vitesse de la disparition ou la durée de la période où la détection est possible. L'obtention d'informations de ce type est une condition préalable à toute quantification post-mortem des repas de prédateurs.

     

Mineral cycling in a tropical palm forest

Nutrient cycling and biomass characteristics of a tropical palm forest dominated byOrbignya cohune were found to be different from thsoe of hardwood dominated forests. The cohune palm forest had a high proportion of biomass in leaves (5%), a reduced sapling layer, a large amount of standing forest litter and an exceptionally low decomposition rate factor (0.1 year^–1). Mineral concentrations in palm leaves were generally lower than in hardwood species with the exception of Na, which was exceptionally high inOrbignya cohune. Biomass was estimated at 226 tons ha^–1 containing 1173 kg ha^–1 N; 126 kg ha^–1 P; 437 kg ha^–1 K; 1869 kg ha^–1 Mg; 125 kg ha^–1 Ca, and 2177 kg ha^–1 Na. Soils of cohune association did not differ significantly from those of neighbouring hardwood dominated associations with the exception of Na which occurred in higher concentration because of bioaccumulation in the dominant. The results suggest that the growth habits and physiology of a dominant can strongly influence some of the ecological parameters used to describe aforest association.     

Some factors affecting the airborne survival of bacteria outdoors

Airborne survival of two pseudomonads and a reference strain of Escherichia coli (strain MRE 162) was studied outdoors using a modified microthread technique. When cells of E. coli were-suspended as clusters, survival was much greater than single cells, particularly outdoors. Culture age had a highly significant effect on survival of Pseudomonas maltophila with survival of 24 h cultures being more than 100-fold higher than 48 h cultures. Survival of Pseudomonas fluorescens was variable and depended also upon the method of culture. Survival of E. coli and Ps. maltophila was studied at three locations differing in air quality and was found to be significantly reduced outdoors, particularly when held in direct daylight. Outdoor survival was not significantly different at the three locations but was reduced at increasing temperatures. There was no apparent effect of wind direction or air quality. Results are discussed with reference to the release of genetically-modified micro-organisms.     

The relative contributions of seed and pollen dispersal to gene flow and genetic diversity in seedlings of a tropical palm

Seed and pollen dispersal shape patterns of gene flow and genetic diversity in plants. Pollen is generally thought to travel longer distances than seeds, but seeds determine the ultimate location of gametes. Resolving how interactions between these two dispersal processes shape microevolutionary processes is a long‐standing research priority. We unambiguously isolated the separate and combined contributions of these two dispersal processes in seedlings of the animal‐dispersed palm Oenocarpus bataua to address two questions. First, what is the spatial extent of pollen versus seed movement in a system characterized by long‐distance seed dispersal? Second, how does seed dispersal mediate seedling genetic diversity? Despite evidence of frequent long‐distance seed dispersal, we found that pollen moves much further than seeds. Nonetheless, seed dispersal ultimately mediates genetic diversity and fine‐scale spatial genetic structure. Compared to undispersed seedlings, seedlings dispersed by vertebrates were characterized by higher female gametic and diploid seedling diversity and weaker fine‐scale spatial genetic structure for female gametes, male gametes and diploid seedlings. Interestingly, the diversity of maternal seed sources at seed deposition sites (N _em) was associated with higher effective number of pollen sources (N _ep), higher effective number of parents (N _e) and weaker spatial genetic structure, whereas seed dispersal distance had little impact on these or other parameters we measured. These findings highlight the importance maternal seed source diversity (N _em) at frugivore seed deposition sites in driving emergent patterns of fine‐scale genetic diversity and structure.     

Indirect interactions among tropical tree species through shared rodent seed predators: a novel mechanism of tree species coexistence

The coexistence of numerous tree species in tropical forests is commonly explained by negative dependence of recruitment on the conspecific seed and tree density due to specialist natural enemies that attack seeds and seedlings (‘Janzen–Connell’ effects). Less known is whether guilds of shared seed predators can induce a negative dependence of recruitment on the density of different species of the same plant functional group. We studied 54 plots in tropical forest on Barro Colorado Island, Panama, with contrasting mature tree densities of three coexisting large seeded tree species with shared seed predators. Levels of seed predation were far better explained by incorporating seed densities of all three focal species than by conspecific seed density alone. Both positive and negative density dependencies were observed for different species combinations. Thus, indirect interactions via shared seed predators can either promote or reduce the coexistence of different plant functional groups in tropical forest.     

Indirect interactions between browsers and seed predators affect the seed bank dynamics of a chaparral shrub

Interactions between herbivores and seed predators may have long-term consequences for plant populations that rely on persistent seed banks for recovery after unpredictable fires. We assessed the effects of browsing by deer and seed predation by rodents, ants and birds on the densities of seeds entering the seed bank of Ceanothus cuneatus var. rigidus, a maritime chaparral shrub in coastal California. Ceanothus produced many more seeds when protected from browsers in long-term experimental exclosures than did browsed plants, but the seed densities in the soil beneath browsed and unbrowsed Ceanothus were the same at the start of an intensive one-year study. The density of seeds in the soil initially increased in both treatments following summer seed drop: while densities returned to pre-drop levels within a few weeks under browsed plants, soil seed densities remained high for 5–8 months beneath unbrowsed plants. Rodent abundance (especially deer mice) was higher near unbrowsed plants than >30 m away, and rodents removed Ceanothus seeds from dishes in the experimental plots. At least in the short term, rodent density and rates of seed removal were inversely related to the intensity of browsing. Our data have management implications for maintaining viable Ceanothus populations by regulating the intensity of browsing and the timing, intensity and frequency of fires.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Some factors affecting survival and oxygen uptake in a subtropical beach flea

The survival and oxygen uptake of the supralittoral amphipod Chroestia lota Marsden & Fenwick were investigated in humid air and sea water between 15 and 35°C. Seven-day exposure experiments were made on three size groups of amphipods at 6 constant temperatures (15, 20, 25, 30, 35, 40 °C) and three cyclic temperatures (15–25, 20–30, 25–35°C) in air and in sea water at 34 and 17%. salinity. Neither size, treatment nor temperature affected survival between 15 and 30°C. Mortality increased > 30°C with large individuals being consistently less tolerant than medium and small amphipods. While amphipods exposed to cyclic temperatures during submersion had reduced survival compared with constant temperatures, those individuals exposed to cyclic conditions in humid air showed the greatest resistance. Oxygen uptake of Chroestia increased with dry body wt and, over the range 15–35°C, this semi-terrestrial beach flea could maintain its aerial VO₂ following submersion. Oxygen uptake increased directly in proportion to gill area and the weight specific gill area was low, consistent with the need to reduce desiccation. It is suggested that total gill area does not limit oxygen uptake in Chroestia and that cutaneous respiration may be important especially in aquatic conditions.