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.     

Tropical tree species diversity: a test of the Janzen-Connell model

To test the premises and predictions of the Janzen-Connell model (Janzen's spacing mechanism), seeds of the rainforest canopy tree, Brosimum alicastrum, were placed at different distances from the parent tree and their removal observed over 3 weeks. The number and density of naturally occurring seeds at different distances from the parent tree were also estimated. Predation was not greater near the parent tree, except on the very small spatial scale: the proportion of experimental seeds removed was greater 1 m from the trunk than it was 5–25 m from the trunk. Predation was negatively correlated with seed density, not positively as the Janzen-Connell model assumes-presumably due to predator satiation. The density of seeds after predation peaked 5 m from the tree trunk, but this is well within the crown radius of the parent tree. There is a peak in the number of potential recruits at a distance of 10 m from the parent tree, due to the peaked initial distribution of seeds. This peak is caused by the interaction between the seed density curve and the increasing area of an annulus around the parent tree at increasing distances, not by the product of the density curve and the predation curve. However, it is important to realize that it is not the presence of a peak in recruitment away from the parent that is essential to maintaining tropical tree species diversity, but frequency-dependent recruitment induced by poor recruitment near conspecifics. Predator satiation seems to be an important factor in the survival of B. alicastrum seeds, possibly at several spatial scales. The number of seeds produced by the tree is negatively correlated with the loss to predators, and trees that have a fruiting conspecific nearby also suffer lower levels of predation. Seed predation increases as one moves from the forest edge into the interior, creating an edge effect that may have long-term effects on the forest composition and tree species diversity. More studies are needed, for other species, other localities, and larger spatial and temporal scales, on both the Janzen-Connell mechanism and this edge effect.     

Seed and microsite limitations of recruitment and the impacts of post-dispersal seed predation at the within population level

Seed predation may cause important seed losses in plant populations, but its impact on the dynamics of populations will depend on the degree of seed or microsite limitations for recruitment. Seed losses will only affect recruitment if it is seed limited. The nature of recruitment limitation (seeds or microsites) is usually ascribed to whole plant populations but it may vary within populations among microhabitats and habitats. Thus, the potential impact of seed predation will also vary within the population, being highest where recruitment is seed limited. The impact to the whole population will depend on the spatial concordance between the intensity of seed predation and that of seed limitation. Recruitment limitations (with seed addition experiments), seed predation (with seed removal experiments), and the dynamics of seed availability in the soil (with soil samples taken both after seed dispersal and before the following dispersal event) of the shrub Corema album (Empetraceae) were investigated in dunes in NW Spain, at microhabitats ‘open ground’, ‘underneath C. album ♀’, and ‘underneath C. album ♂’ at two habitats, sparse and dense scrub. The nature of recruitment limitation (seeds vs. microsites) varied within the population. It was seed limited in the microhabitat ‘open ground’ and microsite limited under shrub cover. The spatial patterns of seedling recruitment were unrelated to seed availability but strongly affected by germination requirements. The spatial discordance between seed availability and recruitment implies a crucial constraint for processes affecting seed availability (seed predation but also e.g., dispersal) to impact recruitment. They will not affect its spatial pattern but only its quantity as long as they act in those sites selected by seeds to germinate. Seed predation was highest underneath mother plants and lowest in open ground. Thus, its potential impact is low, as it is centred where recruitment is not seed limited. This study shows that the analysis of seed predation in relation to recruitment limitations at smaller spatial scales within the population provides more insight to understand its impact.     

Offspring spatial patterns in Picconia excelsa (Oleaceae) in the Canarian laurel forest

We studied the spatial patterns of seedlings and seeds in isolated Picconia excelsa (Oleaceae) trees in the laurel forest of Anaga, Tenerife (Canary Islands). By finding isolated trees we assessed the correlation of seed and seedling bank traits and parent trees by removing the confounding effects of proximity (<100 m radius) of conspecific fruiting trees. We counted all the seedlings per age (height) class within its parental range, and sampled the seed number along transects departing from beneath the parent canopy at regular intervals. We mapped all seedlings per age class and plotted seed and seedling profiles in relation to distance to parent trees. Older Picconia seedlings tended to clump significantly further from parent trees than younger seedlings, which clumped just beneath the parents. We found significant differences among distances to parent tree in numbers of seedlings per age class. The seedling bank area was significantly correlated with maximum distance of seedlings to parent trees. The majority of seeds were deposited within the first 4 m below the parent crown. Seedlings amount at further distances from the trees is larger than seeds/fruits as counted on the ground. Our results suggest that disseminated, older seedlings have occupied germination sites far from the parent tree because there is probably lower seedling–seedling and parent–seedling competition for resources, and perhaps no intraspecific allelopathy and predation/disease.     

Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities

Animals aid population growth and fitness in tropical forest communities through dispersal and negatively impact populations through seed predation. The interaction between dispersal and seed predation can produce distance- or density-dependence; powerful mechanisms for maintaining species diversity incorporated in the Janzen–Connell model. Large mammals, the highest biomass seed predators of intact Amazonian communities and at risk due to human disturbance, are potentially central to these interactions. This study tests the Janzen–Connell model and investigates the impact of mammalian seed predators on seedling recruitment and maintenance of tree diversity. Patterns of both vertebrate and invertebrate seed predation and seedling recruitment were studied in the two most abundant canopy tree species in western Amazonia (Arecaceae: Astrocaryum murumuru and Iriartea deltoidea). We specifically examined effects of both spatial and temporal variation of the highest biomass seed predator in southwest Amazonian forests, the white-lipped peccary (Tayassu pecari), on recruitment through disturbed and undisturbed sites and through a fortuitous 12 year natural extinction and recolonization event of T. pecari. Distance-dependent seedling recruitment was found in Astrocaryum and Iriartea at both sites. However, the median distance of seedlings was ~1.5× farther from reproductive adults in both palms at the undisturbed site. The number of Iriartea seeds escaping predation increased 6,000% in both space and time due to the decline of T. pecari abundance. The results demonstrate that Janzen–Connell effects are stronger in intact ecosystems and tie these mechanistically to changes in seed predator abundance. This study shows that anthropogenic changes in mammal communities decrease the magnitude of Janzen–Connell effects in Amazonian forests and may result in decreases in tree diversity.     

A test of the escape and colonization hypotheses for zoochorous tree species in a Western Amazonian forest

In order to assess the importance of seed dispersal (escape and colonization hypotheses), I used transplant experiments for seeds and seedlings of 5–11 plant species with fleshy fruits in a lowland tropical forest (Tinigua National Park, Colombia). I controlled seed density, distance to parental tree, and habitat type. I monitored seed removal, seedling survival, and seedling growth during the first year of development for an average of 554 seeds and 169 seedlings for each species. I supplemented the experimental results with measurements of natural recruitment. I found little support for the escape hypothesis during the seed and seedling stages. For six species that showed differences in seed removal associated with distance, five showed highest removal away from, than close to parent trees, suggesting predator satiation. Seedling survival during the first year was not consistently associated with low densities and long distances from parent trees. For the majority of species, seedlings did not survive flooding in low basins, and there was growth advantage for most plant species in canopy gaps. These differences imply advantages for seed dispersal to adequate habitats, as predicted by the colonization hypothesis. In contrast to experiments, strong negative distance-dependent effects were evident when analyzing natural recruitment patterns. The ratio between saplings and seedlings was higher away from parent trees for the species with enough recruitment to be analyzed and this suggests that a negative distance-dependent effect may also occur after seedling establishment. This pattern is suspected for several other species, but an analysis with some of the other most common trees showed a variety of negative, neutral, and positive distance dependent effects. This study emphasizes the importance of long-term studies to asses the role of seed dispersal.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

Poachers and Forest Fragmentation Alter Seed Dispersal,Seed Survival,and Seedling Recruitment in the Palm Attalea butyraceae,with Implications for Tropical Tree Diversity1

We examined the interaction between a palm and two bruchid beetles along with several mammal species to explore how poachers and habitat fragmentation may indirectly alter the spatial pattern of seed dispersal, seed predation, and seedling recruitment in central Panama. The large, stony endocarps of Attalea butyraceae decay slowly and bear distinctive scars when opened by rodents or beetles. We determined the final distance between endocarps and reproductive trees (which we call an ecologically effective dispersal distance), the predation status of each endocarp, and the distance between seedlings and reproductive trees. The 68 focal trees were divided among 14 sites and four levels of anthropogenic disturbance. Levels of disturbance included full protection from poachers, light and heavy pressure from poachers, and small island habitat fragments. Ecologically effective seed dispersal distances were greatest for protected sites, intermediate for lightly poached sites, and shortest for heavily poached sites and habitat fragments. Seed predation by rodents increased with distance to the nearest reproductive Attalea and was greatest for fully protected sites, intermediate for lightly poached sites, and least for heavily poached sites and habitat fragments. Seed predation by beetles reversed the patterns described for seed predation by rodents. Total seed predation by beetles and rodents combined was independent of distance, greatest for fully protected sites, and lower for poached sites and habitat fragments. Seedling densities were always greatest close to reproductive trees; however, the increase in seedling densities close to reproductive trees was minimal for fully protected sites, clearly evident for poached sites, and pronounced for habitat fragments. Increased seedling recruitment near conspecific trees may in time reduce tree diversity where humans disrupt mammal communities.     

Dispersal and recruitment dynamics in the fleshy‐fruited Persoonia lanceolata (Proteaceae)

Question: What is the role of dispersal, persistent soil seed banks and seedling recruitment in population persistence of fleshy‐fruited obligate seeding plant species in fire‐prone habitats? Location: Southeastern Australia. Methods: We used a long‐term study of a shrubby, fleshy‐fruited Persoonia species (Proteaceae) to examine (1) seed removal from beneath the canopy of adult plants; (2) seedling recruitment after fire; (3) the magnitude and location of the residual soil seed bank; and (4) the implications for fire management of obligate seeding species. We used demographic sampling techniques combined with Generalised Linear Modelling and regression to quantify population changes over time. Results: Most of the mature fruits (90%) on the ground below the canopy of plants were removed by Wallabia bicolor (Swamp wallaby) with 88% of seeds extracted from W. bicolor scats viable and dormant. Wallabies play an important role in moving seeds away from parent plants. Their role in occasional long distance dispersal events remains unknown. We detected almost no seed predation in situ under canopies (< 1%). Seedling recruitment was cued to fire, with post‐fire seedling densities 6‐7 times pre‐fire adult densities. After fire, a residual soil seed bank was present, as many seeds (77‐100%) remained dormant and viable at a soil depth where successful future seedling emergence is possible (0‐5 cm). Seedling survival was high (> 80%) with most mortality within 2 years of emergence. Plant growth averaged 17 cm per year. The primary juvenile period of plants was 7–8 years, within the period of likely return fire intervals in the study area. We predicted that the study population increased some five‐fold after the wildfire at the site. Conclusions: Residual soil seed banks are important, especially in species with long primary juvenile periods, to buffer the populations against the impact of a second fire occurring before the seed bank is replenished.     

Does predation contribute to tree diversity?

Seed and seedling predation may differentially affect competitively superior tree species to increase the relative recruitment success of poor competitors and contribute to the coexistence of tree species. We examined the effect of seed and seedling predation on the seedling recruitment of three tree species, Acer rubrum (red maple), Liriodendron tulipifera (yellow poplar), and Quercus rubra (northern red oak), over three years by manipulating seed and seedling exposure to predators under contrasting microsite conditions of shrub cover, leaf litter, and overstory canopy. Species rankings of seedling emergence were constant across microsites, regardless of exposure to seed predators, but varied across years. A. rubrum had the highest emergence probabilities across microsites in 1997, but Q. rubra had the highest emergence probabilities in 1999. Predators decreased seedling survival uniformly across species, but did not affect relative growth rates (RGRs). Q. rubra had the highest seedling survivorship across microsites, while L. tulipifera had the highest RGRs. Our results suggest that annual variability in recruitment success contributes more to seedling diversity than differential predation across microsites. We synthesized our results from separate seedling emergence and survival experiments to project seedling bank composition. With equal fecundity assumed across species, Q. rubra dominated the seedling bank, capturing 90% of the regeneration sites on average, followed by A. rubrum (8% of sites) and L. tulipifera (2% of sites). When seed abundance was weighted by species-specific fecundity, seedling bank composition was more diverse; L. tulipifera captured 62% of the regeneration sites, followed by A. rubrum (21% of sites) and Q. rubra (17% of sites). Tradeoffs between seedling performance and fecundity may promote the diversity of seedling regeneration by increasing the probability of inferior competitors capturing regeneration sites.     

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.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

Combining distances of ballistic and myrmecochorous seed dispersal in Adriana quadripartita (Euphorbiaceae)

The separate contributions of different vectors to net seed dispersal curves of diplochorous systems have rarely been characterised. In Australia, myrmecochory is a common seed dispersal syndrome and in the majority of such systems, seeds are initially dispersed ballistically. We measured ballistic and myrmecochorous seed dispersal distances in relation to canopies of Adriana quadripartita (Euphorbiaceae) and used a simulation model to estimate the net dispersal curve. We also compared seed removal rates and ant abundances under, and outside, plant canopies to examine how foraging patterns by ants may affect net dispersal.Overall ant abundance did not show a significant numerical response to seedfall; however, the abundance of the main seed dispersing ant, Rhytidoponera ‘metallica’ did. Despite this, seed removal rates did not differ significantly between canopy and open locations. Rhytidoponera ‘metallica’ account for 93% of observed seed dispersal events. On average, the ants dispersed seeds 1.54 m and in doing so, moved seed a mean radial distance of 0.76 m away from canopy edges. This contribution to net dispersal distance by ants is considerable since ballistic dispersal moved seeds a median distance of 7.5 cm. Our simulation model indicated that the combination of ballistic and ant seed dispersal is expected to result in seeds being transported a median net radial dispersal distance of 1.05 m from the canopy edge.Thus in this system, an important function of diplochory may simply be to move a higher proportion of seeds from under the canopy of parent plants than is possible by ballistic dispersal alone. This ‘dispersal-for-distance’ may result in reduced parent–offspring competition or may increase the probability that seeds reach rare safe sites for germination and recruitment.     

Impacts of logging on density-dependent predation of dipterocarp seeds in a South East Asian rainforest

Much of the forest remaining in South East Asia has been selectively logged. The processes promoting species coexistence may be the key to the recovery and maintenance of diversity in these forests. One such process is the Janzen-Connell mechanism, where specialized natural enemies such as seed predators maintain diversity by inhibiting regeneration near conspecifics. In Neotropical forests, anthropogenic disturbance can disrupt the Janzen-Connell mechanism, but similar data are unavailable for South East Asia. We investigated the effects of conspecific density (two spatial scales) and distance from fruiting trees on seed and seedling survival of the canopy tree Parashorea malaanonan in unlogged and logged forests in Sabah, Malaysia. The production of mature seeds was higher in unlogged forest, perhaps because high adult densities facilitate pollination or satiate pre-dispersal predators. In both forest types, post-dispersal survival was reduced by small-scale (1 m(2)) conspecific density, but not by proximity to the nearest fruiting tree. Large-scale conspecific density (seeds per fruiting tree) reduced predation, probably by satiating predators. Higher seed production in unlogged forest, in combination with slightly higher survival, meant that recruitment was almost entirely limited to unlogged forest. Thus, while logging might not affect the Janzen-Connell mechanism at this site, it may influence the recruitment of particular species.     

Co-evolution of seed size and seed predation

Using the evolutionarily stable strategy (ESS) approach in a model for the co-evolution of seed size and seed predation, I show that seed size variation within individual plants is favoured if there is a trade-off in the predator's attack rate for different seed sizes. A single seed size is not evolutionarily stable because a predator that is optimally adapted to one particular seed size cannot prevent invasion by plants with a different seed size. The model generates the following predictions. The ESS consists of a continuous range of seed sizes. Small seeds tend to be attacked more frequently than big seeds. Plants with many resources and plants with low (frequency-independent) juvenile mortality have more variable seeds than plants with few resources and a high juvenile mortality. Seed size variation is higher in fluctuating populations regulated by seed predation alone than in stable populations (partially) regulated by seedling competition. Predator searching behaviour does not directly affect the ESS seed size range, but may have an indirect effect by affecting population stability or the significance of seedling competition as a population regulating mechanism. Moreover, seed size distributions are found to be more skewed in favour of small seeds if predation is spatially non-uniform than if predation is more even. Application of the model to systems of several co-evolving plant and predator species is discussed.     

Contrasting effects of grazing on the early stages of woody encroachment in a Neotropical savanna

Woody encroachment in savannas represents an ecological process of current global interest given its negative impact on ecosystem functioning, particularly on forage production. Traditional savanna models propose competition and niche differentiation as the main mechanisms allowing tree-grass coexistence. Demographic models, instead, propose abiotic and biotic factors as bottlenecks controlling vital rates and transitions from seeds to adult trees. The role played by domestic grazing on woody encroachment is yet controversial. Here, using a multistage tree life approach, we combine both models and evaluate the role of grazing and herbaceous vegetation on woody recruitment in a Neotropical savanna dominated by Vachellia caven, a successful and widely spread encroacher tree species. We performed three experiments to evaluate seed predation, seedling emergence and survival of V. caven by manipulating cattle grazing (grazed and ungrazed areas) and herbaceous vegetation presence (vegetated and unvegetated). Finally, we combined the results of the three experiments to estimate the probability of plant recruitment across these experimental factors. Grazing decreased seed predation by half, did not modify seedling emergence and decreased seedling survival. Herbaceous vegetation did not affect seed predation nor seedling emergence rate, but increased seedling survival. Overall, the net effect of grazing on V. caven recruitment was neutral since the increase in seed availability due to the reduction in seed predation rate was compensated by the negative effect of grazing on seedling survival. Our analysis revealed that cattle grazing and herbaceous vegetation had contrasting effects on the seed and seedling life stages. We propose that in order to restrain the early stages of encroachment, cattle grazing pressure could be managed following the seasonality of demographic tree transitions. Through rotational grazing amongst paddocks, stocking rates could be relaxed during the primary dispersal stage to maximize granivory, and then increased to enhance the chance of seedling consumption and trampling.     

Post‐dispersal seed predation and its relations with seed traits: a thirty‐species‐comparative study

Post‐dispersal seed predation is a key process determining the variability in seed survival in forests, where most seeds are handled by rodents. Seed predation is thought to affect seedling regeneration, colonization ability and spatial distribution of plants. Basic seed traits are the essential factors affecting rodent foraging preferences and thus seed survival and seedling recruitment. Many studies have discussed several seed traits and their effects upon seed predation by rodents. However, the results of those previous studies are usually equivocal, likely because few seed traits and/or plant species tend to be incorporated into these studies. In order to elucidate the relationships between seed predation and seed traits, we surveyed the predation of 48 600 seeds in a natural pine forest, belonging to 30 species, for three consecutive years. The results demonstrated that: (i) seed size and seed coat hardness did not significantly affect seed predation; (ii) total phenolics had a negative effect upon seed predation; (iii) positive effects of nitrogen content upon seed predation were found. From our study, it seems that the better strategy to prevent heavy predation is for plants to produce seeds with higher total phenolics content rather than physical defenses (i.e. hard seed coat) or larger seeds. Additionally, rodent foraging preference may depend more on Nitrogen content than other nutrient content of seeds.     

Viability of ecological processes in small Afromontane forest patches in South Africa

Abstract The conservation of biodiversity is dependent on protecting ecosystem‐level processes. We investigated the effects of fragment size and habitat edge on the relative functioning of three ecological processes – decomposition, predation and regeneration of trees – in small Afromontane forests in KwaZulu‐Natal, South Africa. Ten sampling stations were placed in each of four forest categories: the interior of three large indigenous forest fragments (100 m from the edge), the edges of these large fragments, 10 small indigenous fragments (<1 ha) and 10 small exotic woodlands (<0.5 ha). Fragment size and edge effects did not affect the abundance of the amphipod Talitriator africana, a litter decomposer, and overall dung beetle abundance and species richness significantly. Bird egg predation was marginally greater at large patch edges compared with the other forest categories, while seed predation did not differ among forest categories. Tree seedling assemblage composition did not differ significantly among large patch interiors and edges, and small indigenous fragments. Sapling and canopy assemblage composition each differed significantly among these three indigenous forest categories. Thus, while tree recruitment was not negatively affected by patch size or distance from the edge, conditions in small fragments and at edges appear to affect the composition of advanced tree regeneration. These ecological processes in Afromontane forests appear to be resilient to fragmentation effects. We speculate that this is because the organisms in these forests have evolved under fragmented conditions. Repeated extreme changes in climate and vegetation over the Pleistocene have acted as significant distribution and ecological extinction filters on these southern hemisphere forest biota, resulting in fauna and flora that are potentially resilient to contemporary fragmentation effects. We argue that because small patches and habitat edges appear to be ecologically viable they should be included in future conservation decisions.     

Scatter-and clump-dispersal and seedling demography: hypothesis and implications

Summary Fruit-eating animals deposit viable seeds in patterns that determine the conditions under which seeds and seedlings live or die. Many tree species are scatter-dispersed by birds, bats, or other small frugivores that regurgitate, defecate, or drop seeds singly or in pairs. These scatterdispersed plant species normally recruit as isolated individuals, and are unlikely to evolve exceptional resistance to herbivores, pathogens, or to other sources of density-dependent seed or seedling mortality. Other tree species are clump-dispersed by larger terrestrial or arboreal frugivores that defecate seeds in masses which produce bouquets of seedlings. Because their seeds invariably germinate in close proximity to other seedlings, clump-dispersed species necessarily evolve chemical or mechanical defenses against seed predators, pathogens, and herbivores that act in a densitydependent manner.Population and genetic attributes should reflect this basic dichotomy in the conditions of seedling recruitment. I predict that seedlings of scatter-dispersed species rarely survive near parents or in dense aggregations under frugivore roosts. Seed dispersal should be mandatory, often to light gaps or other special habitats. Outbred adults and juveniles are expected to exist at low densities in loose aggregations or random distributions. Seedlings of clump-dispersed trees are pre-adapted for survival in dense aggregations near parents, as well as in fecal clumps. Substantial recruitment of juveniles and young adults should occur from undispersed seeds under and near parent trees. Such species should be common, highly aggregated, and show strong genetic family structure. Because recruitment requires dispersal, scatter-dispersed plant species should be especially vulnerable to loss of dispersal agents. Because offspring consistently recruit near parents, clump-dispersed plants should be less vulnerable to temporary loss of dispersal agents.     

Linking seed dispersal,germination and seedling recruitment in the invasive species <Emphasis Type="Italic">Berberis darwinii</Emphasis> (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.