Phenology and fecundity in 11 sympatric pioneer species of Macaranga (Euphorbiaceae)in Borneo

Reproductive traits of tropical tree species vary predictably in relation to successional stage, but this variation may be due to the species' phylogenetic histories rather than selective pressures imposed by regeneration requirements. Reproductive phenology, tree size at the onset of reproduction, and fecundity of 11 sympatric, closely related Macaranga species were studied to investigate within-species variation in reproductive traits in relation to resource availability, and among-species variation in relation to other life-history traits (shade tolerance, seed size and maximum tree size, H(max)) and consequently the requirements for forest-gap colonization. Nine species reproduced in synchronous episodes, and two species reproduced continuously over 32 mo. Episodic reproduction was most intense in 1992 following a severe drought. For several species, reproductive trees had greater light availability, lower fecundity in lower light levels, and lower growth rates than nonreproductive trees, reflecting resource-limited reproduction. Among species, H(max) was negatively correlated with shade tolerance and seed size. Tree size at the onset of reproduction and fecundity was strongly linked to this axis of life-history variation, but phenological pattern was not. Absolute tree size at the onset of reproduction was positively correlated with H(max) and negatively correlated with shade tolerance. Relative size at reproductive onset was not correlated with shade tolerance or H(max). Fecundity ranged four orders of magnitude among species and was correlated positively with H(max) and negatively with seed size and shade tolerance. The interrelationships among these reproductive and other life-history traits are strongly correlated with the species' requirements for gap colonization.     

Thematic and Spatial Resolutions Affect Model-Based Predictions of Tree Species Distribution

Subjective decisions of thematic and spatial resolutions in characterizing environmental heterogeneity may affect the characterizations of spatial pattern and the simulation of occurrence and rate of ecological processes, and in turn, model-based tree species distribution. Thus, this study quantified the importance of thematic and spatial resolutions, and their interaction in predictions of tree species distribution (quantified by species abundance). We investigated how model-predicted species abundances changed and whether tree species with different ecological traits (e.g., seed dispersal distance, competitive capacity) had different responses to varying thematic and spatial resolutions. We used the LANDIS forest landscape model to predict tree species distribution at the landscape scale and designed a series of scenarios with different thematic (different numbers of land types) and spatial resolutions combinations, and then statistically examined the differences of species abundance among these scenarios. Results showed that both thematic and spatial resolutions affected model-based predictions of species distribution, but thematic resolution had a greater effect. Species ecological traits affected the predictions. For species with moderate dispersal distance and relatively abundant seed sources, predicted abundance increased as thematic resolution increased. However, for species with long seeding distance or high shade tolerance, thematic resolution had an inverse effect on predicted abundance. When seed sources and dispersal distance were not limiting, the predicted species abundance increased with spatial resolution and vice versa. Results from this study may provide insights into the choice of thematic and spatial resolutions for model-based predictions of tree species distribution.     

The direct regeneration hypothesis in northern forests

Question: Can the direct regeneration hypothesis (DRH) be used to predict post‐disturbance regeneration after fire, wind disturbance, and clearcutting in northern forests? Do life‐history traits such as regeneration strategy and shade tolerance influence post‐disturbance regeneration success of tree species? Location: Northern forests in North America. Methods: A meta‐analysis was conducted by collecting published data on pre‐ and post‐disturbance stand compositional characteristics in the northern forests. For each tree species, compositional difference (CD) was calculated as the difference between basal area proportions of the post‐ and pre‐disturbance stands, but for post‐disturbance stands <25 years of age, post‐disturbance proportions were calculated based on relative stem density. Results: Species response to disturbances was best explained by regeneration strategy, while disturbance type had no effect on CD. The proportion of broadleaf trees with either strong or weak vegetative reproduction ability increased after all disturbances. Serotinous species had CD values not significantly different from zero after fire, while CD for semi‐serotinous species was negative. The post‐disturbance proportions of non‐serotinous conifers decreased after all forms of disturbance. Conclusions: All disturbances promote broadleaf trees, regardless of regeneration strategy (suckering, sprouting, or seeding). The DRH is supported for conifers with serotinous cones after fire. Fire causes local extinction of non‐serotinous conifers, while wind and clearcutting only decrease the proportion of non‐serotinous conifers because of partial survival of seed sources and advanced regeneration. This study suggests that increasing stand‐replacing disturbances associated with global climate change will promote broadleaf trees in northern forests.     

植物生活史繁殖对策与干扰关系的研究

植物生活史繁殖对策研究是涉及植物的适应或进化、生态系统退化与恢复过程、生物多样性保护等多方面理论生态学和应用生态学研究内容的生态学研究领域。按Grime的植物生活史繁殖对策分类、植物营养繁殖与干扰适应、种子形态学与干扰适应、土壤种子库与干扰适应、植物繁殖体传播和萌发与干扰适应论述了当今极受关注的植物生活史繁殖对策与干扰关系,简述了我国干旱区干扰与植物生活史繁殖对策关系研究。     

Defining species guilds in the Central Hardwood Forest,USA

Tree regeneration outcomes are challenging to generalize and difficult to predict. Many tree species can establish new propagules in a variety of post-disturbance environments and many different reproductive mechanisms may be used. In order to develop conceptual models that accurately reflect reproductive potential, we need a better understanding of the similarities in regeneration ecology among species. We used information from the forest ecology literature to evaluate the reproductive attributes of sixty-two tree species in the central hardwood region of the eastern United States. Each species was classified categorically for features such as flowering, seed production and dispersal, seed dormancy, germination requirements, seedling characteristics, and vegetative reproduction. Cluster analysis (Jaccard's similarity coefficient, complete linkage method) and ordination (homogeneity analysis) were used to separate nine groups (guilds) of species that had similar reproductive attributes. Individual attributes that had high variance in the first and second dimensions included: seed banking, seed dispersal, seedling shade tolerance, and seedbed requirements. Members of each guild had similar levels of reproductive specialization and guilds were either pioneer-like, opportunistic, or persistent. Pioneer guilds included: short-lived or fugitive species that colonize sites rapidly and are too shade intolerant to replace themselves; shade-tolerant species that colonize frequently disturbed sites; and stress-tolerant pioneers that survive on dry or nutrient-poor sites. Opportunistic guilds contained species that are remarkably versatile in their reproductive effort. The most flexible opportunists can colonize new sites, maintain seed in a seed bank, sprout from existing stems and persist as a seedling or sapling bank. Persistent guilds contain species that develop and maintain advance regeneration. These include: species with moderate understory tolerance that regenerate via cycles of dieback and resprouting; and more tolerant species that maintain seedling or sapling banks. Our regeneration guilds may provide a useful approach for more realistically representing large and diverse sets of tree species in forest ecosystem models.     

Interactions between frequency and size of disturbance affect competitive outcomes

Disturbance has many effects on ecological communities, and it is often suggested that disturbance can affect species diversity by altering competitive outcomes. However, disturbance regimes have many distinct aspects that may act, and interact, to influence species diversity. While there are many theoretical models of disturbance-prone communities, few have specifically documented how interactions between different aspects of a disturbance regime change competitive outcomes. Here, we present a model of two plant species subject to disturbance which we then use to examine species coexistence over varying levels of two aspects of disturbance: frequency, and spatial extent (i.e., area disturbed). We show that the competitive outcome is affected differently by changes in each aspect and that the effect of disturbance frequency on species coexistence depends strongly on the spatial extent of the disturbance, and vice versa. We classify the nature of these interactions between disturbance frequency and extent on the basis of the shape of the resulting coexistence regions in a frequency?Cextent parameter plane. Our results illustrate that different types of interaction can result from differences in life-history traits that control species-specific sensitivity to frequency and extent of disturbance. Thus, our analysis shows that the various aspects of disturbance must be carefully considered in concert with the life-history traits of the community members in order to assess the consequences of disturbance.     

Spatially structured intraspecific trait variation can foster biodiversity in disturbed,heterogeneous environments

Trait variation within populations is an important area of research for empirical and theoretical ecologists. While differences between individuals are doubtlessly ubiquitous, their role for species coexistence is much less clear and highly debated. Both unstructured (random) and structured (linked to space, time or inheritance) intraspecific trait variation (ITV) may modify species interactions with nontrivial consequences for emerging community compositions. In many ecosystems, these compositions are further driven by prevalent disturbance regimes. I therefore explored the effects of unstructured as well as spatially structured ITV under disturbances in a generic ecological model of competing sessile species. Using spatially explicit, individual‐based simulations, I studied how intraspecific variation in life history traits together with interspecific tradeoffs and disturbance regimes shape long‐term community composition. I found that 1) unstructured ITV does not affect species coexistence in the given context, 2) spatially structured ITV may considerably increase coexistence, but 3) spatially clumped disturbances reduce this effect of spatially structured ITV, especially if interspecific tradeoffs involve dispersal distance. The findings suggest that spatially structured ITV with individual trait responses to local habitat conditions differing among species may create or expand humps in disturbance–diversity relationships. Hence, if present, these forms of spatially structured ITV should be included in ecological models and will be important for reliably assessing community responses to environmental heterogeneity and change.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

MORPHOLOGY AND DISPERSAL POTENTIAL OF WIND-DISPERSED DIASPORES OF NEOTROPICAL TREES

Morphological and aerodynamic traits affecting mean potential dispersal distance are quantified for wind-dispersed diaspores of tree species on Barro Colorado Island, Panama. The sample includes 34 species in 16 families and represents six aerodynamic groups. Mass and area (maximum cross section) each vary over six orders of magnitude among the species. In contrast, wing-loading, defined as weight divided by area, varies over only one order of magnitude, as does the rate of descent. While the regression of rate of descent on the square root of wing-loading is significant overall, the slopes vary significantly among five aerodynamic groups. At comparable wing-loading values, diaspores of fluffy kapok fall faster than four other aerodynamic groups and rolling autogyros fall faster than non-rolling autogyros. Assuming the diaspores are released from their typical tree height and experience a mean windspeed of 1.75 m sec^−-1, the expected mean dispersal distance varies among the 34 species from 22 to 194 m. Rate of descent is weakly correlated with shade tolerance of seedlings for a subset of 18 species; rate of descent is more strongly correlated with the log of dry mass of seed for all 34 species. Given these wide differences in dispersal potential, any generalizations about tropical trees that use wind dispersal are of dubious value.     

Co-dominance and succession in forest dynamics: the role of interspecific differences in crown transmissivity

Forests that are composed of two or more tree species with similar ecological strategies appear to contradict the competitive exclusion principle. Beech-maple communities are a well-known example of such a system. On a local scale, a number of mechanisms have been proposed to explain the coexistence of these two species. These are reciprocal replacement, external factors that favour alternatively one or the other species and demographic stochasticity. This paper presents and analyses a simple mathematical model that shows that external factors are not an essential requirement for coexistence. Rather, coexistence requires interspecific differences in light transmissivity through the crowns of adult trees. However, all the three mechanisms mentioned above can be interpreted within the framework of the model. Furthermore, many models of forest dynamics make use of shade tolerance as a key feature in describing successional dynamics. Despite its importance, however, shade tolerance does not have a commonly accepted quantitative definition. Here, a simple scheme is proposed where the relationship between shade tolerance, individual traits (growth and survival) and successional status is defined. This might have important implications in understanding the overall dynamics. Theoretical results have been compared with a number of studies carried out in North American forests. In particular, coexistence in beech-maple communities and the relation between shade tolerance and successional status in a beech-hemlock-birch community have been discussed.     

Two fitness measures for clonal plants and the importance of spatial aspects

The capability of clonal plants to reproduce both sexually and vegetatively and their resulting hierarchical organisation into ramets and genets pose problems for the determination of fitness. We develop ramet-based measures of clonal plant fitness including both modes of reproduction. To this end we use simple population-dynamic equations accounting for limited space, limited dispersal, and disturbance. Neglecting interactions among ramets (r-selection regime) we derive an expression for initial growth rate r as a fitness measure. At higher densities interactions among ramets lead to density control (K-selection regime) and competitive exclusion of genotypes or species may occur. In this case we apply an invasibility criterion to derive an abundance fitness measure: competitiveness C. The optimum of C corresponds to an evolutionary stable strategy. C increases with the proportion of reproductive adults, with inverse module mortality, and with the sum of sexual and vegetative reproduction. The latter are defined as the product of the rate of module production and two correction factors accounting for juvenile mortality before establishment and for the efficiency of space exploitation by propagules. Thus C is equivalent to potential life-time offspring production, in contrast to realized life-time offspring production R₀. Because the correction factor for space exploitation cannot be expressed analytically we obtain it from complementary spatially-explicit individual-based simulations. To illustrate an application of the fitness measure C we predict optimal allocation to vegetative and sexual reproduction. In a homogeneous habitat an intermediate allocation may maximize fitness only if there is a non-linear trade-off between the modes of reproduction. However even if this trade-off is linear, spatially heterogenous disturbances can lead to an intermediate optimum of allocation because seed dispersal with subsequent vegetative spread improves the utilization of available space for recruitment if the spatial extent of single disturbances is much larger than the distance of vegetative dispersal. Thus, our study underlines the importance of spatial features for fitness measures especially of clonal plants.     

Life history variation across a riverine landscape: intermediate levels of disturbance favor sexual reproduction in the ant‐dispersed herb Ranunculus ficaria

Global climate change can fundamentally alter disturbance regimes across landscapes, but little is known about how species adjust their life histories to shifts in disturbance regimes. In plants, dispersal by seeds may permit rapid re‐colonization under frequent disturbances, but often seed‐dispersing animals may be absent and local dispersal by vegetative diaspores may be a more efficient means of occupying open space in the vicinity of the plant. We tested the effect of disturbances due to inundation on the investment in seed production by the ant‐dispersed plant, Ranunculus ficaria ssp. bulbifer. During seed ripening we collected 392 plants within a landscape mosaic of 39 sites with different levels of inundation. We measured the mass of fruits and other tissues (leafs, roots, bulbils, stalks) and described plant growth form. We found that fruit numbers and masses were more variable among plants than numbers and masses of other tissues. We then standardized fruit mass against the mass of other tissues and other growth‐form parameters. Standardized fruit mass showed a highly significantly hump‐shaped relationship with the level of inundation disturbances. This pattern was consistent across 12 small‐scale transects and thus not confounded by spatial autocorrelation within landscapes. The pattern was also confirmed by analyses that simultaneously accounted for disturbance and morphological co‐variables. We thus conclude that plants invested most heavily into reproduction by seeds under intermediate levels of disturbance. Under intermediate disturbance, seeds are beneficial for rapidly re‐colonizing open space after disturbance while the seed‐dispersers are still available. The life history of mutualists such as ant‐dispersed plants and ants may thus change across a landscape, reflecting small‐scale variation in the disturbance regime.     

Herbaceous plant strategies in disturbed habitats

A systematic theoretical evaluation has been made of three important plant life history traits: adult longevity, seed longevity and seed mass, where seed mass is interpreted as being indicative of dispersal distance and seedling vigour. This model study examined the role of these three traits in relation to environmental disturbance. We chose temperate grasslands, widespread in north Western Europe and northern and eastern America, as our reference system for our simulations. Eight plant strategies were defined by allowing two levels in each of the three and combining them in all eight possible ways. A simple, spatially explicit model was developed to simulate competition among individuals with these eight trait combinations at different levels of disturbance.
Simulation results were compared with the actual occurrence over a disturbance gradient of species with similar plant trait combinations in a large database from the Sheffield area (UK). This showed that with increasing disturbance level, non-dormant perennials, dormant perennials, non-dormant annuals and dormant annuals, respectively, became dominant but only if small-seeded, indicating the relative viability of these particular strategies with respect to disturbance.
A new prediction from the model was that stable coexistence occurs between plant strategies with dormant and with non-dormant seeds over a range of levels of disturbance. Plant strategies with large seeds were inferior to small-seeded ones if competitive ability of seedlings is proportional to seed weight. This difference was highest at low seed densities and low germination probabilities, indicating that large-seeded species secure no advantage from being dormant (i.e. having a low germination probability). Finally, the results indicated that dormancy is superior to dispersal as a method of coping with disturbance.     

Seeds tend to disperse further in the tropics

Despite the importance of seed dispersal in a plant's life cycle, global patterns in seed dispersal distance have seldom been studied. This paper presents the first geographically and taxonomically broad quantification of the latitudinal gradient in seed dispersal distance. Although there is substantial variation in the seed dispersal distances of different species at a given latitude, seeds disperse on average more than an order of magnitude further at the equator than towards the poles. This pattern is partially explained by plant life‐history traits that simultaneously associate with seed dispersal distance and latitude, including dispersal mode and plant height. The extended seed shadow of tropical plants could increase the distance between conspecific individuals. This could facilitate species coexistence and contribute to the maintenance of high plant diversity in tropical communities. The latitudinal gradient in dispersal distance also has implications for species’ persistence in the face of habitat fragmentation and climate change.     

Dispersal and recruitment limitation in native versus exotic tree species: life‐history strategies and Janzen‐Connell effects

Life‐history traits of invasive exotic plants are typically considered to be exceptional vis‐à‐vis native species. In particular, hyper‐fecundity and long range dispersal are regarded as invasive traits, but direct comparisons with native species are needed to identify the life‐history stages behind invasiveness. Until recently, this task was particularly problematic in forests as tree fecundity and dispersal were difficult to characterize in closed stands. We used inverse modelling to parameterize fecundity, seed dispersal and seedling dispersion functions for two exotic and eight native tree species in closed‐canopy forests in Connecticut, USA. Interannual variation in seed production was dramatic for all species, with complete seed crop failures in at least one year for six native species. However, the average per capita seed production of the exotic Ailanthus altissima was extraordinary: > 40 times higher than the next highest species. Seed production of the shade tolerant exotic Acer platanoides was average, but much higher than the native shade tolerant species, and the density of its established seedlings (≥ 3 years) was higher than any other species. Overall, the data supported a model in which adults of native and exotic species must reach a minimum size before seed production occurred. Once reached, the relationship between tree diameter and seed production was fairly flat for seven species, including both exotics. Seed dispersal was highly localized and usually showed a steep decline with increasing distance from parent trees: only Ailanthus altissima and Fraxinus americana had mean dispersal distances > 10 m. Janzen‐Connell patterns were clearly evident for both native and exotic species, as the mode and mean dispersion distance of seedlings were further from potential parent trees than seeds. The comparable intensity of Janzen‐Connell effects between native and exotic species suggests that the enemy escape hypothesis alone cannot explain the invasiveness of these exotics. Our study confirms the general importance of colonization processes in invasions, yet demonstrates how invasiveness can occur via divergent colonization strategies. Dispersal limitation of Acer platanoides and recruitment limitation of Ailanthus altissima will likely constitute some limit on their invasiveness in closed‐canopy forests.     

Dynamics of seedling populations and tree species coexistence in a forest: a simulation study

Theoretically, temporal variation of reproduction promotes species coexistence of sessile and polycarpic organisms when the reproduction is synchronized within species but independent among species. Monopoly of vacant sites and high relative population growth rate of minor species in the absence of propagules of other species is the essence of this mechanism. The mechanism is expected to work in forests, but persistent populations of seedlings may affect the promotion of coexistence. Using a tree-based simulation model of forest dynamics, it was demonstrated that the number of coexisting tree species was sensitively affected by the seedling establishment rate. The coexistence was not enhanced by temporal variation of reproduction when seedling establishment rate was low. This is because the reproducing minor species fail to monopolize vacant sites and allow the establishment of seedlings of other species in later years. High mortality of established seedlings under shade also suppressed coexistence. This is likely to be the result of a reduced storage effect of the population of seedlings. A forest structure and dynamics pattern that appears when tree species coexistence is promoted by fluctuating reproduction was searched for, and the number of coexisting species was varied by manipulating the seedling establishment rate. No distinct difference other than the species number itself was found between species-rich and species-poor forests. For example, the seedling population size varied, reflecting the temporal variation of reproduction, irrespective of the seedling establishment rate. Further strategy development is needed to validate the proposed mechanisms of species coexistence.     

How to survive as a pioneer species in the Antarctic benthos: minimum dispersal distance as a function of lifetime and disturbance

Very few details exists concerning the dispersal traits of Antarctic species and dispersal distances in particular are mostly unknown. Especially the general low number of mesoplanktonic larvae has caught attention, leading to the formulation of Thorson’s rule. From this concept, originally concerning only trophic aspects, sometimes a reduced dispersal distance is deduced. Using a generic simulation model we show that in a benthic habitat exposed to iceberg scouring, even short dispersal phases of few hours are sufficient for a pioneer species to persist. This is very surprising for a pioneer species which should be able to disperse widely and colonise distant disturbed areas which are free of superior competitors. Our model revealed that the reason for this is the non-linear dependence of the dispersal distance on the disturbance regime and on species longevity. Thus, it is the combined effect of life history and disturbance traits which is important here: a sufficiently high disturbance frequency due to iceberg scouring and a long individual lifetime due to the low temperature decrease minimum dispersal distances required for persistence and thus coexistence and present an additional explanation for the relative rarity of planktonic larvae.     

西双版纳热带季节雨林四种珍稀濒危树种种子萌发对脱水和光的响应

绒毛番龙眼(Pometia tomentosa)、琴叶风吹楠(Horsfieldia pandurifolia)、箭毒木(Antiaris toxicaria)和思茅木姜子(Litseapierrei var. szemois)是西双版纳热带季节雨林乔木层的优势树种,也属于珍稀濒危物种。实验研究了4个树种的种子在经过不同脱水水平、人工遮荫条件下的不同光照水平以及热带季节雨林之林下和林窗的萌发特性。结果显示:脱水对于种子萌发特征具有显著影响,绒毛番龙眼和琴叶风吹楠经48h的连续脱水后丧失了萌发能力;箭毒木和思茅木姜子具较强的脱水耐性,经96h的连续脱水后仍具有较高的种子含水量,且种子萌发率分别为15%和33%。干旱季散布的种子比雨季散布的种子具有更强的脱水耐性。遮荫及森林种子萌发试验中,除绒毛番龙眼外,其它3个树种的种子萌发率以及平均萌发周期在不同光水平或生境下差异显著。琴叶风吹楠和思茅木姜子在30%的光照以及林窗中心的萌发率要显著低于低光处理(10%和3.5%光照)和林下,而箭毒木在30%和10%光水平以及林窗中心获得较高萌发率。雨季传播的具有干燥敏感性的种子,可以减少种子由于干燥而引起的死亡。试验结果可以为4种珍稀濒危植物的保护和人工繁育提供一定理论依据。     

Reproduction of invasive Amur honeysuckle (Lonicera maackii) and the arithmetic of an extermination strategy

Ecological restorations often require removal of invasive species. The abundance of invasives has tended to catalyze research emphasizing removal, not broader understandings, of species mechanisms for persistence in the landscape (e.g. reproductive output and seed dispersal). Asiatic shrub honeysuckles (Lonicera spp.) are pernicious invaders throughout eastern North America. Heavy tree canopy cover apparently reduces growth and reproductive output in Lonicera maackii, which is widespread through the lower Midwestern United States. To help focus control efforts more effectively, we quantified the effect of tree canopy cover on vegetative growth, flowering, and fruit production under three canopy densities. Mean vegetative growth of flowering shoots was not affected by canopy cover. All aspects of sexual reproduction (flower production, fruit set, fruit number, fruit mass, seed number, and seed size) were strongly reduced by moderate shade. Although all individuals modify community and ecosystem properties, a limited number of high light individuals might also provide the greatest proportion of the seeds. Through model simulation of honeysuckle population structure in relation to canopy cover, we argue that it can sometimes be more efficient to initially target reproductive individuals in the high light edge and interior gap environments than to immediately focus on all individuals in the forest interior.     

Spatial genetic structure in Milicia excelsa (Moraceae) indicates extensive gene dispersal in a low-density wind-pollinated tropical tree

In this study, we analysed spatial genetic structure (SGS) patterns and estimated dispersal distances in Milicia excelsa (Welw.) C.C. Berg (Moraceae), a threatened wind-pollinated dioecious African tree, with typically low density (∼10 adults/km²). Eight microsatellite markers were used to type 287 individuals in four Cameroonian populations characterized by different habitats and tree densities. Differentiation among populations was very low. Two populations in more open habitat did not display any correlation between genetic relatedness and spatial distance between individuals, whereas significant SGS was detected in two populations situated under continuous forest cover. SGS was weak with a maximum S _p-statistic of 0.006, a value in the lower quartile of SGS estimates for trees in the literature. Using a stepwise approach with Bayesian clustering methods, we demonstrated that SGS resulted from isolation by distance and not colonization by different gene pools. Indirect estimates of gene dispersal distances ranged from σ _g = 1 to 7.1 km, one order of magnitude higher than most estimates found in the literature for tropical tree species. This result can largely be explained by life-history traits of the species. Milicia excelsa exhibits a potentially wide-ranging wind-mediated pollen dispersal mechanism as well as very efficient seed dispersal mediated by large frugivorous bats. Estimations of gene flow suggested no major risk of inbreeding because of reduction in population density by exploitation. Different strategy of seed collection may be required for reforestation programmes among populations with different extent of SGS.