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.     

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.     

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.     

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.

     

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.     

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.     

Some factors affecting airborne survival of Pseudomonas fluorescens indoors

The effect of relative humidity (RH) on the airborne survival of a Pseudomonas fluorescens strain was studied at 20, 40, 60 and 80% RH indoors. The aero-stable spore of Bacillus subtilis subsp. niger , used as a tracer of physical losses was compared with a light scattering particle counter, as there were doubts about the reliability of the spore as tracer. The Rion counter was validated before use and found to give a good estimate of relative physical losses providing spray suspensions contained between 10⁷ and 10⁹ cfu ml⁻¹ cells and that the humidity was not more than 80% RH. Pseudomonas fluorescens , strain P⁺ S⁺, suspended in distilled water survived best at mid humidities and least at 80% RH. When suspended in 1% glycerol there was an apparent 'increase' in viability after an initial rapid reduction. This was thought to be due to delay in equilibration of glycerol in the cell membrane after concentration on dehydration. The cells were thought to be unstable and sensitive to the stress of rehydration before equilibration occurred. The findings are discussed in relation to Cox's theories of outer membrane damage on aerosolization.     

The escape response of pea aphids to foliar-foraging predators: factors affecting dropping behaviour

1. The effects of predator species, aphid density, aphid age, diel period, and habitat complexity on the dropping behaviour of the pea aphid Acyrthosiphon pisum were assessed in a series of laboratory and field-cage experiments.
2. The presence of foliar-foraging predators significantly increased the proportion of aphids that dropped from alfalfa plants. In the absence of predators, less than 7% of the aphids dropped. Dropping more than doubled (14%) when one of three hemipteran predators , N. americoferus, G. punctipes or O. insidiosus , was present. Nearly 60% of the aphids dropped when the ladybird beetle, Coccinella septempunctata , was present.
3. Adult aphids showed a significantly higher propensity to drop than immature aphids, regardless of the presence or absence of predators. Aphid density had no effect on dropping behaviour.
4. Neither diel period nor habitat complexity had an effect on aphid dropping behaviour. Aphids were significantly more likely to drop in the presence of predators during either the day or night and from either early or late regrowth alfalfa.
5. A review of the factors affecting dropping behaviour, including those elucidated in this study, indicates that the propensity to drop from a plant is influenced by three factors: the risk of predation on the plant, the quality of the resource to be abandoned, and the risk of mortality in the new microhabitat.     

Rodent seed predation: effects on seed survival, recruitment, abundance, and dispersion of bird-dispersed tropical trees

Tropical tree species vary widely in their pattern of spatial dispersion. We focus on how seed predation may modify seed deposition patterns and affect the abundance and dispersion of adult trees in a tropical forest in India. Using plots across a range of seed densities, we examined whether seed predation levels by terrestrial rodents varied across six large-seeded, bird-dispersed tree species. Since inter-specific variation in density-dependent seed mortality may have downstream effects on recruitment and adult tree stages, we determined recruitment patterns close to and away from parent trees, along with adult tree abundance and dispersion patterns. Four species (Canarium resiniferum, Dysoxylum binectariferum, Horsfieldia kingii, and Prunus ceylanica) showed high predation levels (78.5-98.7%) and increased mortality with increasing seed density, while two species, Chisocheton cumingianus and Polyalthia simiarum, showed significantly lower seed predation levels and weak density-dependent mortality. The latter two species also had the highest recruitment near parent trees, with most abundant and aggregated adults. The four species that had high seed mortality had low recruitment under parent trees, were rare, and had more spaced adult tree dispersion. Biotic dispersal may be vital for species that suffer density-dependent mortality factors under parent trees. In tropical forests where large vertebrate seed dispersers but not seed predators are hunted, differences in seed vulnerability to rodent seed predation and density-dependent mortality can affect forest structure and composition.     

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

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

Biotic and abiotic factors affecting survival of early life history intervals of a stream-dwelling sunfish

Synopsis Biotic interactions, such as competition and predation, and abiotic events, such as floods, both have important consequences for organisms inhabiting warmwater streams. Empirical studies can lead to better understanding of how these factors interact to affect distinct life history intervals of stream fishes. Survival of eggs, embryos and larvae in nests of longear sunfishLepomis megalotis was monitored in a section of Jordan Creek for four years. During two years characterized by low, relatively stable flow, nest failures could be attributed to biotic interactions. During two years with more variable flow, most brood losses occurred during floods. Flooding led to nest desertion and loss of offspring regardless of nest location. The number of nests constructed tended to be lower during years with more variable flow. The proportion of nests with successful broods did not differ between colonial and solitary nests. Variation in flow regime modifies the importance of biotic effects on survival of early life history intervals of longear sunfish.     

Social and maternal factors affecting duckling survival in eiders Somateria mollissima

1. With the aid of a novel survivorship model, an 8-year field study of social and maternal factors affecting duckling survival in eiders (Somateria mollissima) revealed that duckling survival probability varies in accordance with maternal brood-rearing strategy. This variability in survival provides compelling evidence of different annual fitness consequences between females that share brood-rearing and those that tend their broods alone. Consequently, as prebreeding survival is often a major source of individual variation in lifetime reproductive success, a female's annual, state-dependent (e.g. condition) choice of a brood-rearing strategy can be a critical fitness decision. 2. Variance in duckling survival among lone tender broods was best explained by a model with significant interannual variability in survival, and survivorship tending to increase with increasing clutch size at hatch. Clutch size was correlated positively with female condition. Hatch date and female body condition together affected duckling survival, but their contributions are confounded. We were unable to identify a relationship between female age or experience and duckling survival. 3. Variance in duckling survival among multifemale brood-rearing coalitions was best explained by a model that included the number of tenders, the number of ducklings and interannual variation in how their ratio affected survivorship. Hatch date did not significantly influence survival. 4. Expected duckling survival is higher in early life for lone tenders when compared with multifemale brood-rearing coalitions. However, as ducklings approach 2-3 weeks of age, two or three females was the optimal number of tenders to maximize daily duckling survival. The survivorship advantage of multifemale brood-rearing coalitions was most evident in years of average survival. 5. The observed frequency distribution of female group sizes corresponds with the distribution of offspring survival probabilities for these groups. Evidence for optimal group sizes in nature is rare, but the most likely candidates may be groups of unrelated animals where entry is controlled by the group members, such as for female eiders. 6. Our study demonstrates that differences in social factors can lead to different predictions of lifetime reproductive success in species with shared parental care of self-feeding young.