Spatio-temporal scales of dispersal limitation in the recolonization of a semi-arid mediterranean old-field

Dispersal and establishment limitations have been proposed as important mechanisms that control vegetation colonization processes. While many studies have emphasized the role of dispersal limitation in the dynamics of plant communities, little is known about the spatial and temporal scales at which dispersal limitation affects plant population dynamics. In this study we investigate spatial and temporal scales of dispersal limitation in a system of semi-arid Mediterranean old-fields recolonized by the dominating dwarf shrub Sarcopoterium spinosum (Rosaceae). Our analysis was based on a combination of two complementary approaches: 1) age analysis of S. spinosum shrubs growing at different distances from source populations, and 2) a corresponding analysis of changes in vegetation cover since recolonization onset (ca 55 yr ago), based on information obtained from historical aerial-photographs. Shrubs sampled in plots near the source populations were similar in their age distribution to the source populations, but significantly different from shrubs sampled far from the source populations. Increasing distance from source populations was associated with a decrease in median age of the shrubs, and in the density and relative frequency of adult shrubs (>15 yr). These patterns of changes in age structure occurred over distances of several tenths of meters. Young age groups (<5 yr) were scarcely represented in all plots, suggesting that establishment limitation was also important in determining recolonization rates. Analysis of the aerial photographs supported the results of the age analysis, showing evidence for both distance effect and establishment limitation. We conclude that dispersal limitation may have a long-term impact on vegetation patterns, even at small spatial scales, and that dispersal limitation interacts with establishment limitation in determining recolonization processes.     

Variable Gene Dispersal Conditions and Spatial Deforestation Patterns Can Interact to Affect Tropical Tree Conservation Outcomes

Tropical lowland rain forest (TLRF) biodiversity is under threat from anthropogenic factors including deforestation which creates forest fragments of different sizes that can further undergo various internal patterns of logging. Such interventions can modify previous equilibrium abundance and spatial distribution patterns of offspring recruitment and/or pollen dispersal. Little is known about how these aspects of deforestation and fragmentation might synergistically affect TLRF tree recovery demographics and population genetics in newly formed forest fragments. To investigate these TLRF anthropogenic disturbance processes we used the computer program NEWGARDEN (NG), which models spatially-explicit, individual-based plant populations, to simulate 10% deforestation in six different spatial logging patterns for the plant functional type of a long-lived TLRF canopy tree species. Further, each logging pattern was analyzed under nine varying patterns of offspring versus pollen dispersal distances that could have arisen post-fragmentation. Results indicated that gene dispersal condition (especially via offspring) had a greater effect on population growth and genetic diversity retention (explaining 98.5% and 88.8% of the variance respectively) than spatial logging pattern (0.2% and 4.7% respectively), with ‘Near’ distance dispersal maximizing population growth and genetic diversity relative to distant dispersal. Within logged regions of the fragment, deforestation patterns closer to fragment borders more often exhibited lower population recovery rates and founding genetic diversity retention relative to more centrally located logging. These results suggest newly isolated fragments have populations that are more sensitive to the way in which their offspring and pollen dispersers are affected than the spatial pattern in which subsequent logging occurs, and that large variation in the recovery rates of different TLRF tree species attributable to altered gene dispersal regimens will be a likely outcome of fragmentation. Conservation implications include possible manual interventions (manual manipulations of offspring dispersers and/or pollinators) in forest fragments to increase population recovery and genetic diversity retention.     

A spatially explicit analysis of seedling recruitment in the terrestrial orchid Orchis purpurea

Seed dispersal and the subsequent recruitment of new individuals into a population are important processes affecting the population dynamics, genetic diversity and spatial genetic structure of plant populations. Spatial patterns of seedling recruitment were investigated in two populations of the terrestrial orchid Orchis purpurea using both univariate and bivariate point pattern analysis, parentage analysis and seed germination experiments. Both adults and recruits showed a clustered spatial distribution with cluster radii of c. 4-5 m. The parentage analysis resulted in offspring-dispersal distances that were slightly larger than distances obtained from the point pattern analyses. The suitability of microsites for germination differed among sites, with strong constraints in one site and almost no constraints in the other. These results provide a clear and coherent picture of recruitment patterns in a tuberous, perennial orchid. Seed dispersal is limited to a few metres from the mother plant, whereas the availability of suitable germination conditions may vary strongly from one site to the next. Because of a time lag of 3-4 yr between seed dispersal and actual recruitment, and irregular flowering and fruiting patterns of adult plants, interpretation of recruitment patterns using point patterns analyses ideally should take into account the demographic properties of orchid populations.     

Scale‐specific processes shape plant community patterns in subtropical coastal grasslands

Processes responsible for shaping community patterns act at specific spatial scales. In this study, we aimed at disentangling the effects of climate, soil and space as drivers of variation in a coastal grassland plant community. We were specifically interested in evaluating the relative influence of those processes at broad and fine spatial scales as well as when considering species groups with good and poor long‐distance dispersal capacity. We sampled grassland vegetation at 16 sites distributed along a latitudinal gradient of more than 500 km in subtropical southern Brazil and used variation partitioning procedures to ascertain the relative influence of climatic, edaphic and spatial processes on variation in species composition at different spatial scales, considering the entire community and subsets with only species from the Asteraceae family (good long‐distance dispersal) and Poaceae (poor long‐distance dispersal). Climatic filters were the most responsible for shaping grassland community composition at the broad scale, while edaphic filters showed higher importance at the fine scale. When not considering the influence of spatial scale, we observed higher influence of climate structured in space. Composition patterns of species with poor long‐distance dispersal (Poaceae) were more closely related to spatial variables than those of species with effective dispersal (Asteraceae). Our results stressed the importance of addressing different spatial scales to rightly ascertain the magnitude that different drivers exert on plant community assembly. Dividing the community into groups with different dispersal abilities proved useful for a more detailed understanding of the community assembly processes.     

New interpretations of fine-scale spatial genetic structure

Recent methodological advances permit refined inferences of evolutionary processes from the fine-scale spatial genetic structure of plant populations. In this issue of Molecular Ecology, Born et al. (2008) exploit the full power of these methods by examining effects of ancient and recent landscape histories in an African rainforest tree species. The authors first detected admixture of distinct gene pools that may have formed in Pleistocene forest refuges. Then, comparing across six study populations in Gabon, the authors found similar patterns of fine-scale spatial genetic structure despite natural and anthropogenic variation in population density. The latter results suggest that enhanced gene dispersal may compensate for low population densities in fragmented landscapes.     

<Emphasis Type="Italic">PRUNUS</Emphasis>: a spatially explicit demographic model to study plant invasions in stochastic,heterogeneous environments

To model the invasion of Prunus serotina invasion within a real forest landscape we built a spatially explicit, non-linear Markov chain which incorporated a stage-structured population matrix and dispersal functions. Sensitivity analyses were subsequently conducted to identify key processes controlling the spatial spread of the invader, testing the hypothesis that the landscape invasion patterns are driven in the most part by disturbance patterns, local demographical processes controlling propagule pressure, habitat suitability, and long-distance dispersal. When offspring emigration was considered as a density-dependent phenomenon, local demographic factors generated invasion patterns at larger spatial scales through three factors: adult longevity; adult fecundity; and the intensity of self-thinning during stand development. Three other factors acted at the landscape scale: habitat quality, which determined the proportion of the landscape mosaic which was potentially invasible; disturbances, which determined when suitable habitats became temporarily invasible; and the existence of long distance dispersal events, which determined how far from the existing source populations new founder populations could be created. As a flexible “all-in-one” model, PRUNUS offers perspectives for generalization to other plant invasions, and the study of interactions between key processes at multiple spatial scales.     

DECOUPLING OF SHORT‐ AND LONG‐DISTANCE DISPERSAL PATHWAYS IN THE ENDEMIC NEW ZEALAND SEAWEED CARPOPHYLLUM MASCHALOCARPUM (PHAEOPHYCEAE,FUCALES)1

The processes that produce and maintain genetic structure in organisms operate at different timescales and on different life‐history stages. In marine macroalgae, gene flow occurs through gamete/zygote dispersal and rafting by adult thalli. Population genetic patterns arise from this contemporary gene flow interacting with historical processes. We analyzed spatial patterns of mitochondrial DNA variation to investigate contemporary and historical dispersal patterns in the New Zealand endemic fucalean brown alga Carpophyllum maschalocarpum (Turner) Grev. Populations bounded by habitat discontinuities were often strongly differentiated from adjoining populations over scales of tens of kilometers and intrapopulation diversity was generally low, except for one region of northeast New Zealand (the Bay of Plenty). There was evidence of strong connectivity between the northern and eastern regions of New Zealand’s North Island and between the North and South Islands of New Zealand and the Chatham Islands (separated by 650 km of open ocean). Moderate haplotypic diversity was found in Chatham Islands populations, while other southern populations showed low diversity consistent with Last Glacial Maximum (LGM) retreat and subsequent recolonization. We suggest that ocean current patterns and prevailing westerly winds facilitate long‐distance dispersal by floating adult thalli, decoupling genetic differentiation of Chatham Island populations from dispersal potential at the gamete/zygote stage. This study highlights the importance of encompassing the entire range of a species when inferring dispersal patterns from genetic differentiation, as realized dispersal distances can be contingent on local or regional oceanographic and historical processes.     

Does spatial heterogeneity blur the signature of dispersal syndromes on spatial patterns of woody species? A test in a tropical dry forest

Spatial patterns of adult plants are a consequence of several ecological processes related to seed dispersal and recruitment. Dispersal limitation, mediated by dispersal syndrome, is considered a key factor in the formation of adult plant spatial patterns. Although this initial pattern determined by dispersal has been thoroughly studied, the subsequently modification by the effect of additional ecological factors, such as habitat heterogeneity is less understood. We explored the relative importance of dispersal syndrome and spatial heterogeneity on the realization of spatial patterns of adult trees in an Ecuadorian tropical dry forest. The spatial distribution of 28 species was modeled with four different spatial point processes each: homogeneous Poisson (HPP), inhomogeneous Poisson (IPP), homogeneous Poisson cluster (HPCP), and inhomogeneous Poisson cluster process (IPCP). These models allowed us to discern between effects of random processes, habitat heterogeneity, limited dispersal, and joint effects of habitat heterogeneity and limited dispersal. We employed Akaike's information criterion (AIC) to select the model which best fit the spatial pattern of each species. The best model of each species was used to analyze differences in cluster size and degree of aggregation, between dispersal syndromes. Seventy‐five percent of the species showed inhomogeneous patterns. IPCP yielded the best fit for the spatial distribution of 50% of species in the studied forest and was the prevalent model for the three dispersal syndromes. Thus, the effect of spatial heterogeneity was prevalent in the distribution of most species in this dry tropical forest. Only 21% of species had spatial patterns compatible with random mechanisms associated to limited dispersal around parent sources. Clearly, ignoring habitat heterogeneity could bias the analysis of relationships between dispersal syndrome and species patterns.     

Generation of Spatial Patterns in Boreal Forest Landscapes

Boreal forests are composed of a few plant species with contrasting traits with respect to ecosystem functioning and spatial patterning. Early successional deciduous species, such as birch and aspen, disperse seeds widely, do not tolerate low light and nitrogen availabilities, have rapidly decaying litter, and are highly preferred by herbivores. These later succeed to conifers, such as spruce and fir, which disperse seeds locally, tolerate low light levels and low nitrogen availability, have litter that decays slowly, and are unpalatable to most mammalian herbivores. Although there are also early successional conifers, such as jack pine and Scots pine, the aspen-birch-spruce-fir successional sequence is the most common over much of North America, and (without fir) in Fennoscandia and Siberia. The course of succession in these forests is controlled partly by seed dispersal and selective foraging by mammalian herbivores. Both of these processes are spatially dynamic, but little is known about how their spatial dynamics may affect ecosystem processes, such as nitrogen cycling or productivity. We present spatially explicit models that demonstrate the following: (a) Spatially explicit seed dispersal results in more clumped distribution of tree species and persistence of greater paper birch biomass than uniform seed rain across the landscape. Such results are consistent with current spatially explicit population models of dispersal and coexistence. (b) With localized seed dispersal, the concentrations of available soil nitrogen are distributed in larger patches with sharp transitions from low to high nitrogen availability near patch edges. In contrast, with a uniform seed rain, the distribution of soil nitrogen availability was more uniform and “hotspots” were more localized. Thus, the spatial pattern of an ecosystem process (nitrogen cycling) is determined by seed dispersal and competition for light among competing populations. (c) A dispersing herbivore, such as moose, that selectively forages on early successional deciduous species with high quality litter, such as aspen or birch, and discriminates against late successional conifers, such as spruce or fir, imposes higher-order repeated patterns of plant species and biomass distribution on the landscape. Thus, seed dispersal and herbivore foraging correlate properties in adjacent patches but in different ways, and different spatial patterns emerge. Other processes, such as insect outbreaks, fire, and water flow, also may correlate properties between adjacent patches and result in additional patterns. Received 8 February 1999; accepted 28 May 1999.     

Plant dispersal in the sub-Antarctic inferred from anisotropic genetic structure

Climatic conditions and landscape features often strongly affect species' local distribution patterns, dispersal, reproduction and survival and may therefore have considerable impacts on species' fine-scale spatial genetic structure (SGS). In this study, we demonstrate the efficacy of combining fine-scale SGS analyses with isotropic and anisotropic spatial autocorrelation techniques to infer the impact of wind patterns on plant dispersal processes. We genotyped 1304 Azorella selago (Apiaceae) specimens, a wind-pollinated and wind-dispersed plant, from four populations distributed across sub-Antarctic Marion Island. SGS was variable with Sp values ranging from 0.001 to 0.014, suggesting notable variability in dispersal distance and wind velocities between sites. Nonetheless, the data supported previous hypotheses of a strong NW-SE gradient in wind strength across the island. Anisotropic autocorrelation analyses further suggested that dispersal is strongly directional, but varying between sites depending on the local prevailing winds. Despite the high frequency of gale-force winds on Marion Island, gene dispersal distance estimates (σ) were surprisingly low (<10 m), most probably because of a low pollen dispersal efficiency. An SGS approach in association with isotropic and anisotropic analyses provides a powerful means to assess the relative influence of abiotic factors on dispersal and allow inferences that would not be possible without this combined approach.     

基于零模型的宁夏荒漠草原优势种群点格局分析

植物种群空间分布格局是多种生态过程综合作用的结果。明确植物优势种群个体的空间分布格局与形成机制有助于认识种群生态适应对策与群落多样性维持机制。以宁夏荒漠草原优势种群蒙古冰草、短花针茅、牛枝子和牛心朴子为研究对象,采用完全空间随机零模型分析其种群空间分布格局特征,并通过异质泊松零模型与泊松聚块零模型探讨生境异质性、扩散限制等因子在其空间分布格局形成过程中的作用。结果显示:(1)完全空间随机零模型下,4个物种在4 m尺度范围内表现为聚集分布,随尺度增大,逐渐过渡到随机分布和均匀分布。(2)在排除生境异质性的异质泊松零模型下,蒙古冰草种群在整个研究尺度上表现为随机分布;牛枝子、短花针茅和牛心朴子种群仅分别在0—0.2、0.1—0.4 m与0—0.2 m尺度范围内发生偏离,表现为均匀分布与聚集分布,其他尺度均为随机分布。(3)在排除扩散限制的泊松聚块零模型下,所研究种群均表现为随机分布。综上,荒漠草原优势种群在小尺度范围内主要表现为聚集分布;生境异质性与扩散限制均是驱动其空间分布格局形成的重要因子,相对而言,小尺度空间范围内扩散限制的作用更为显著。     

不同空间尺度下的肉果植物扩散过程和机理

肉果植物扩散的生态学过程在最近得到生态学者们的广泛关注,其扩散过程包括果实搬运、果实消耗、种子雨、种子取食、种子库动态、萌发和幼苗定居等。许多过程涉及到果食性动物和肉果植物之间的互惠的协同进化关系。对最近15a关于肉果植物扩散的研究论文进行了综述,探讨在生境、微生境、景观和区域等常用的空间尺度上,肉果植物扩散和定居过程的格局与机理。     

Contemporary pollen and seed dispersal in a Prunus mahaleb population: patterns in distance and direction

Pollination and seed dispersal determine the spatial pattern of gene flow in plant populations and, for those species relying on pollinators and frugivores as dispersal vectors, animal activity plays a key role in determining this spatial pattern. For these plant species, reported dispersal patterns are dominated by short-distance movements with a significant amount of immigration. However, the contribution of seed and pollen to the overall contemporary gene immigration is still poorly documented for most plant populations. In this study we investigated pollination and seed dispersal at two spatial scales in a local population of Prunus mahaleb (L.), a species pollinated by insects and dispersed by frugivorous vertebrates. First, we dissected the relative contribution of pollen and seed dispersal to gene immigration from other parts of the metapopulation. We found high levels of gene immigration (18.50%), due to frequent long distance seed dispersal events. Second, we assessed the distance and directionality for pollen and seed dispersal events within the local population. Pollen and seed movement patterns were non-random, with skewed distance distributions: pollen tended moved up to 548 m along an axis approaching the N-S direction, and seeds were dispersed up to 990 m, frequently along the SW and SE axes. Animal-mediated dispersal contributed significantly towards gene immigration into the local population and had a markedly nonrandom pattern within the local population. Our data suggest that animals can impose distinct spatial signatures in contemporary gene flow, with the potential to induce significant genetic structure at a local level.     

What causes conspecific plant aggregation? Disentangling the role of dispersal,habitat heterogeneity and plant–plant interactions

Spatial patterns of plant species are determined by an array of ecologica factors including biotic and abiotic environmental constraints and intrinsic species traits. Thus, an observed aggregated pattern may be the result of short‐distance dispersal, the presence of habitat heterogeneity, plant–plant interactions or a combination of the above. Here, we studied the spatial pattern of Mediterranean alpine plant Silene ciliata (Caryophyllaceae) in five populations and assessed the contribution of dispersal, habitat heterogeneity and conspecific plant interactions to observed patterns. For this purpose, we used spatial point pattern analysis combined with specific a priori hypotheses linked to spatial pattern creation. The spatial pattern of S. ciliata recruits was not homogeneous and showed small‐scale aggregation. This is consistent with the species’ short‐distance seed dispersal and the heterogeneous distribution of suitable sites for germination and establishment. Furthermore, the spatial pattern of recruits was independent of the spatial pattern of adults. This suggests a low relevance of adult‐recruits interactions in the spatial pattern creation. The difference in aggregation between recruits and adults suggests that once established, recruits are subjected to self‐thinning. However, seedling mortality did not erase the spatial pattern generated by seed dispersal, as S. ciliata adults were still aggregated. Thus, the spatial aggregation of adults is probably due to seed dispersal limitation and the heterogeneous distribution of suitable sites at seedling establishment rather than the presence of positive plant–plant interactions at the adult stage. In fact, a negative density‐dependent effect of the conspecific neighbourhood was found on adult reproductive performance. Overall, results provide empirical evidence of the lack of a simple and direct relationship between the spatial structure of plant populations and the sign of plant–plant interactions and outline the importance of considering dispersal and habitat heterogeneity when performing spatial analysis assessments.     

Spatial genetic structure of the southeastern North American endemic, Ceratiola ericoides (Empetraceae)

Understanding the spatial distribution of genetic diversity (i.e., spatial genetic structure [SGS]) within plant populations can elucidate mechanisms of seed dispersal and patterns of recruitment that may play an important role in shaping the demography and spatial distribution of individuals in subsequent generations. Here we investigate the SGS of allozyme diversity in 2 populations of the southeastern North American endemic shrub, Ceratiola ericoides. The data suggest that the 2 populations have similar patterns of SGS at distances of 0-45 m that likely reflect the isolation by distance (IBD) model of seed dispersal. However, at distances >or=50 m, the pattern of SGS differs substantially between the 2 populations. Whereas one population continues to reflect the classical IBD pattern, the second population shows a marked increase in autocorrelation coefficient (r) values at 50-75 m. Furthermore, r values at these distances are as much as 33% higher than at 0-5 m where the highest r value would be predicted by IBD. A likely explanation is the differing frequencies of 2 fruit morphologies in these populations and the greater role that birds play in seed dispersal in the second population.     

Integrating the Study of Non-native Plant Invasions across Spatial Scales

Non-native (alien, exotic) plant invasions are affecting ecological processes and threatening biodiversity worldwide. Patterns of plant invasions, and the ecological processes which generate these patterns, vary across spatial scales. Thus, consideration of spatial scale may help to illuminate the mechanisms driving biological invasions, and offer insight into potential management strategies. We review the processes driving movement of non-native plants to new locations, and the patterns and processes at the new locations, as they are variously affected by spatial scale. Dispersal is greatly influenced by scale, with different mechanisms controlling global, regional and local dispersal. Patterns of invasion are rarely documented across multiple spatial scales, but research using multi-scale approaches has generated interesting new insights into the invasion process. The ecological effects of plant invasions are also scale-dependent, ranging from altered local community diversity and homogenization of the global flora, to modified biogeochemical cycles and disturbance regimes at regional or global scales. Therefore, the study and control of invasions would benefit from documenting invasion processes at multiple scales.     

Implications of recurrent disturbance for genetic diversity

Exploring interactions between ecological disturbance, species’ abundances and community composition provides critical insights for ecological dynamics. While disturbance is also potentially an important driver of landscape genetic patterns, the mechanisms by which these patterns may arise by selective and neutral processes are not well‐understood. We used simulation to evaluate the relative importance of disturbance regime components, and their interaction with demographic and dispersal processes, on the distribution of genetic diversity across landscapes. We investigated genetic impacts of variation in key components of disturbance regimes and spatial patterns that are likely to respond to climate change and land management, including disturbance size, frequency, and severity. The influence of disturbance was mediated by dispersal distance and, to a limited extent, by birth rate. Nevertheless, all three disturbance regime components strongly influenced spatial and temporal patterns of genetic diversity within subpopulations, and were associated with changes in genetic structure. Furthermore, disturbance‐induced changes in temporal population dynamics and the spatial distribution of populations across the landscape resulted in disrupted isolation by distance patterns among populations. Our results show that forecast changes in disturbance regimes have the potential to cause major changes to the distribution of genetic diversity within and among populations. We highlight likely scenarios under which future changes to disturbance size, severity, or frequency will have the strongest impacts on population genetic patterns. In addition, our results have implications for the inference of biological processes from genetic data, because the effects of dispersal on genetic patterns were strongly mediated by disturbance regimes.     

Going against the flow: a case for upstream dispersal and detection of uncommon dispersal events

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GENE DISPERSAL AND SPATIAL GENETIC STRUCTURE

Spatial autocorrelation statistics have been studied in theoretical population genetic models and widely used in experimental studies of spatial structure in many plant and animal populations. However, the statistical properties of spatial autocorrelation statistics have remained uncharacterized. Little is known about how values of spatial autocorrelation statistics in population samples depend on the level of dispersal and scheme of sampling. In this paper, we characterize the statistical properties of join-count spatial autocorrelation statistics for population genetic surveys under various conditions of dispersal and sampling. The results indicate generally high statistical power. These results can provide a method to estimate gene dispersal based on standing spatial patterns of genetic variation observed within populations.     

Assessment of spatial discordance of primary and effective seed dispersal of European beech (Fagus sylvatica L.) by ecological and genetic methods

Spatial discordance between primary and effective dispersal in plant populations indicates that postdispersal processes erase the seed rain signal in recruitment patterns. Five different models were used to test the spatial concordance of the primary and effective dispersal patterns in a European beech (Fagus sylvatica) population from central Spain. An ecological method was based on classical inverse modelling (SSS), using the number of seed/seedlings as input data. Genetic models were based on direct kernel fitting of mother‐to‐offspring distances estimated by a parentage analysis or were spatially explicit models based on the genotype frequencies of offspring (competing sources model and Moran‐Clark's Model). A fully integrated mixed model was based on inverse modelling, but used the number of genotypes as input data (gene shadow model). The potential sources of error and limitations of each seed dispersal estimation method are discussed. The mean dispersal distances for seeds and saplings estimated with these five methods were higher than those obtained by previous estimations for European beech forests. All the methods show strong discordance between primary and effective dispersal kernel parameters, and for dispersal directionality. While seed rain was released mostly under the canopy, saplings were established far from mother trees. This discordant pattern may be the result of the action of secondary dispersal by animals or density‐dependent effects; that is, the Janzen‐Connell effect.