Ecological perspectives for the restoration of plant communities in European temperate forests

Simultaneously with increasing afforestation efforts in western Europe, among conservationists the consciousness is growing that protecting areas to conserve biodiversity will not be sufficient in the long term, and that also the ecological restoration of more or less severely altered areas will be necessary. The probability that recently established forest stands develop towards their ecological reference (i.e. ancient forest) depends largely on the possibility of the target species to colonize them. We focused on the colonization ability of forest plant species and particularly on so-called ancient forest plant species. Major constraints for ecological forest restoration are the spatial characteristics of the target site (isolation, shape and area), imposing dispersal limitations, and in the duration and intensity of the historical land use, leading to changes in habitat characteristics influencing recruitment probability. We reviewed the ecological literature with respect to these constraints and conclude that it takes at least a century to restore the understorey layer of recent forests, even when the target stand is adjacent to a well-developed ancient forest. Both recruitment and dispersal limitation of the target species are responsible for this. Newly established forests should therefore be situated at a minimal distance of the ancient forest source. In other cases, forest plant species will not be able to colonize the newly established forest on a measurable time scale and artificial introduction of forest plant species can be taken into consideration. The negative effects of habitat characteristics, and mainly high soil nutrient values in the recent forest stand can be mitigated by soil nutrient lowering measures. Disturbances in the recent forest should be minimized to maintain a high canopy closure level, preventing light demanding, high competitive species from establishing a stable population. An additional negative consequence of soil disturbances is that it stimulates germination of species from the soil seed bank, which is mainly composed of highly competitive or ruderal species.     

Seed dispersal and seedling recruitment of trees at different successional stages in a temperate forest in northeastern China

Aims Spatial distribution of adult trees in a forest community is determined by patterns of both seed dispersal and seedling recruitment. The objectives of our study were to understand the processes of seed dispersal and seedling recruitment of dominant tree species in a temperate forest of northeastern China and to identify the factors constraining seed dispersal and seedling establishment at different stages of forest succession.Methods During three summer and autumn sessions between 2006 and 2008, altogether 113080 seeds from 22 different tree species were collected in three large field plots representing different forest types in the Changbai Mountain region of northeastern China. The spatial distribution of seed abundance was analyzed using a Syrjala test. Regeneration success of nine major tree species was assessed using variables defining 'limitations' in 'seeds' and 'seedling establishment'.Important findings We found that seed production fluctuated between years and varied greatly with forest types. Four tree species, Acer spp., Fraxinus mandshurica, Tilia amurensis and Betula spp., had the greatest seed production and the widest range of seed dispersal, whereas Quercus mongolica showed the most sustained seed production pattern. The spatial patterns of seed abundance differed significantly among forest types and years. The tree species investigated in this study differed in the degree of seed limitation, as well as in limitation of seedling establishment. There were both negative and positive correlations between seed density and seedling density, depending on site and parental tree density. Seeds of 16 tree species were found in the Populus davidiana–Betula platyphylla forest (PBF) plot, 11 in the conifer and broad-leaved mixed forest (CBF) plot but only 8 in the broad-leaved-Korean pine mixed forest (BKF) plot. The number of seed-contributing species was not only greater in the secondary forests (CBF and PBF plots) than in the primary forest (BKF plot) but was also more variable during the 3 years of assessment. Results from the correlations between seed density and seedling occurrence and that between parental tree density or seed weight and dispersal limitation confirm our intuitive expectations, i.e. heavy seeds had greater dispersal limitation but higher establishment success than light seeds.     

The effect of landscape fragmentation and forest continuity on forest floor species in two regions of Denmark

Abstract. Forest species composition was recorded in 82 forests in the Himmerland and Hornsherred regions in Denmark and analysed with respect to isolation (distance to other forests and areas of forest), forest continuity (older or younger than 200 yr), soil pH, tree species composition and seed dispersal groups. Continuity and isolation measures were correlated with forest species richness in Hornsherred. Myrmecochorous, autochorous, anemoballistic and endozoochorous species were markedly fewer in recent than in ancient forests. In Himmerland, patterns were much weaker and few significant correlations were found between forest species richness or different seed dispersal groups and continuity or isolation of the forests. Differences between the two regions may result from less intensive land use, a more humid climate and a smaller species pool with less species with short distance dispersal in Himmerland. Landscape fragmentation therefore appears to limit forest species’recolonization more in Hornsherred than in Himmerland.     

Influence of land use history on seed banks in European temperate forest ecosystems: a review

This study summarises European research on seed banks in temperate forest systems and analyses for differences in seed bank composition between geographically scattered forests with a different land use history. Special attention is given to seed bank characteristics of ancient forest species. Results of Detrended Correspondence Analysis suggest that historical land use is a key factor in determining the seed bank composition. Particularly seed banks of forests on former heathland sites differ from seed banks of ancient forest due to a high contribution of early successional species. The effect of former land use decreases after 50 yr, due to seed senescence. Total seed density decreases with recent forest age. Seed bank composition of eastern European forests is different from northern or western European forests, a difference which is mainly caused by species with a higher Ellenberg indicator value for continentality, temperature and reaction. In general, ancient forest species are poorly represented. Only a limited number is mentioned to have a persistent seed bank, and their densities are relatively low, which means that restoration of typical ancient forest vegetation can not rely on the seed bank. However, there is still a considerable lack of knowledge concerning seed bank and germination characteristics of forest species.     

Spatio‐temporal colonization patterns of forest plant species in a mixed deciduous forest

Abstract. An integrated analysis of the colonization patterns of forest plant species was carried out in a 34‐ha, mixed deciduous forest in northern Belgium. First, we sought to describe the relationships between land use history and environmental conditions. Land use history and soil type were related and negative correlations between pH and secondary forest age were found. The density of the shrub layer increases with secondary forest age. Litter quantity and cover of Urtica dioica were mainly indirectly influenced by land use history. Litter starts accumulating at low pH values and high shrub density and Urtica dioica grows vigorously on nutrient enriched soils where much light can reach the ground. Next, the importance of these human‐altered environmental conditions for the colonization of forest plant species was assessed relative to the importance of dispersal limitation. Therefore, the distribution of 16 forest species was mapped and species‐specific spatio‐temporal isolation measures were calculated. The analysis revealed that the colonization patterns of the slowly colonizing species (i.e. ‘ancient forest plant species’) are best explained by a combination of spatio‐temporal isolation, soil type, pH and the (non‐)cover of Urtica dioica. By contrast, spatio‐temporal isolation was never a limiting factor for good colonizing forest species. Our results suggest that colonization of ‘ancient forest plant species’ is hampered by a combination of dispersal‐ and recruitment limitation and that the relative importance of both factors is species‐specific.     

Seed-deposition and recruitment patterns of Clusia species in a disturbed tropical montane forest in Bolivia

Spatial patterns of seed dispersal and recruitment of fleshy-fruited plants in tropical forests are supposed to be driven by the activity of animal seed dispersers, but the spatial patterns of seed dispersal, seedlings and saplings have rarely been analyzed simultaneously. We studied seed deposition and recruitment patterns of three Clusia species in a tropical montane forest of the Bolivian Andes and tested whether these patterns changed between habitat types (forest edge vs. forest interior), distance to the fruiting tree and consecutive recruitment stages of the seedlings. We recorded the number of seeds deposited in seed traps to assess the local seed-deposition pattern and the abundance and distribution of seedlings and saplings to evaluate the spatial pattern of recruitment. More seeds were removed and deposited at the forest edge than in the interior. The number of deposited seeds decreased with distance from the fruiting tree and was spatially clustered in both habitat types. The density of 1-yr-old seedlings and saplings was higher at forest edges, whereas the density of 2-yr-old seedlings was similar in both habitat types. While seedlings were almost randomly distributed, seeds and saplings were spatially clustered in both habitat types. Our findings demonstrate systematic changes in spatial patterns of recruits across the plant regeneration cycle and suggest that the differential effects of biotic and abiotic factors determine plant recruitment at the edges and in the interior of tropical montane forests. These differences in the spatial distribution of individuals across recruitment stages may have strong effects on plant community dynamics and influence plant species coexistence in disturbed tropical forests.     

Migration of herbaceous plant species across ancient–recent forest ecotones in central Belgium

1 We studied the migration of forest plant species using their percentage cover and frequency in 197 plots distributed over 26 transects across ecotones between ancient and recent deciduous forests in the Meerdaal forest complex in central Belgium. The recent forest stands varied in age between 36 and 132 years, and all occurred on silty, well-drained soils.
2 The total cover, number and diversity of field layer species did not differ significantly between ancient and recent forest stands.
3 The number and cover of the ancient forest plant species and of ant-dispersed species correlated positively with the age of the recent forest and negatively with both the duration of its former agricultural land use and the distance to the ancient forest. This implies a slow colonization of the recent forest stands by these species; all species were, however, able to migrate across the ecotones.
4 The cover of four species ( Anemone nemorosa , Lamium galeobdolon , Convallaria majalis and Polygonatum multiflorum ) declined along the transect, suggesting that they are limited by seed dispersal. Their colonization rates, calculated from the occurrence of the farthest individual, ranged from < 0.05 to 1.15 m year⁻¹ and for other measures from < 0.05 to 0.65 m year⁻¹. Anemone and Lamium appeared to colonize the recent forest by establishment of isolated individuals, while Polygonatum and Convallaria expanded populations from existing patches on the border between ancient and recent forest.
5 Several forest species were able to colonize the recent forest rapidly, where some of them even reached a higher abundance, due to the increased availability of colonization sites with a higher nutrient content and a thinner organic layer.     

植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

Environmental limitation contributes to the differential colonization capacity of two forest herbs

Question: The recovery of forest plant communities in post‐agricultural landscapes is largely determined by dispersal constraints, but can environmental legacies of former land use additionally limit the recolonization of recent forests by forest herbs? Location: Ancient forest and recent forest on former heavily fertilized agricultural land (Muizenbos, northern Belgium). Methods: Seeds and adults of two forest herbs with similar life‐history traits, but contrasting colonization capacity – the fast‐colonizing Geum urbanum and the slow‐colonizing Primula elatior– were introduced into both ancient and recent forest sites. Soil conditions and plant tissue nutrient concentrations were measured to characterize habitat quality. To determine whether the introduced species could successfully establish and persist, we monitored recruitment, longevity and adult performance during 8 years in permanently marked plots. Results: Phosphorus availability was ten times higher in recent forest soils and was also reflected in the plant tissue samples. Species longevity was clearly lower in recent forest sites indicating higher turnover. The fast‐colonizing G. urbanum counterbalanced this lower longevity by new establishment, while the slow‐colonizing P. elatior dropped below the number of originally introduced individuals. Additionally, G. urbanum performed better in recent forest sites in contrast to P. elatior. Conclusions: Even when dispersal constraints of the slow‐colonizing forest herb P. elatior are eliminated through introduction, environmental conditions in recent forest sites additionally restrict its recruitment, longevity and performance. These experimental results suggest that environmental constraints may strengthen the differences in colonization capacity among forest herbs if slow dispersers also tend to be less likely to establish.     

Dispersal limits natural recruitment of African mahoganies

The abundance of forest tree species may be locally limited by the inability of species to disperse to all sites suitable for germination and establishment. This phenomenon of "dispersal limitation" has been suggested to promote the maintenance of high species diversity in tropical forests by slowing down competitive exclusion. We present the first direct experimental evaluation of dispersal limitation in tropical forests, and of its importance relative to other factors affecting recruitment. Seed addition, litter removal and light availability effects on seedling establishment were evaluated in a full factorial experiment on two African mahogany species, Entandrophragma utile and Khaya anthotheca . Seedling recruitment after 18 months, was ∼22 times greater in seed addition treatments (with addition rates corresponding to the tail of seed shadows of large reproductive trees) compared to controls. This seed addition effect was an order of magnitude greater than either gap or litter removal effects, with similar results observed in both logged and primary forest locations. We conclude that dispersal limitation strongly limits seedling establishment for the two species studied. An important "applied corollary" of this result is that seed supplementation may substantially increase local abundance of valuable or rare species in disturbed or managed tropical forests.     

森林种子雨研究进展与展望

种子雨阶段是植物更新的关键环节,它连接着繁殖生产与植物后续生活史阶段,对群落结构有着重要的影响.虽然早在19世纪中叶达尔文就认识到种子扩散的重要性,然而对种子雨的广泛研究于20世纪80年代初才开始.本文聚焦于森林木本植物种子雨研究,首先介绍了种子雨监测方法,包括收集器的布置,种子雨的收集、分离和鉴定.然后综述了种子雨的4个主要研究方向:种子产量的时间和空间变化格局(包括季节变化、年际变化和空间变化)、增补限制及其在物种多样性维持中的作用、验证负密度效应假说、种子雨与其他生活史阶段(土壤种子库、幼苗、幼树及母树)的比较.未来还需要加强对种子雨的长期监测,开展增补限制的跨纬度比较研究,探讨植物早期更新阶段负密度效应沿纬度梯度的变化规律,加强数学模型以及分子标记和稳定同位素技术等新手段的运用.     

Population structure and adult plant performance of forest herbs in three contrasting habitats

In Flanders, forests have been subjected to subsequent periods of reclamation and reforestation, leading to a mosaic of forest patches of various ages and with different abiotic conditions, depending on the duration and intensity of the past agricultural activities. While extensive research has been done on the relative importance of dispersal limitation and establishment limitation in explaining colonization patterns of so-called ancient forest plant species, little is known about the viability of populations of these species once they have colonized recent forests. Therefore, a comparative study on population structure and plant performance was carried out for three forest plant species with different colonization capacity ( Primula elatior , Geum urbanum and Arum maculatum ) in three different habitat types: ancient forests, recently established forests and small landscape elements (hedgerows). Contrary to the expectation, it was shown that adult plant performance – a combination of vegetative as well as reproductive characteristics – of the three species was generally higher in recent forests and small landscape elements. This also led to higher seedling densities and proportions in these habitat types. Adult densities, however, were extremely low in newly established forests, suggesting reduced seedling and juvenile survival under these circumstances, probably due to higher light availability and hence increased competition from species like Urtica dioica . Although there might be no immediate establishment limitation, installing a shrub layer in recent forests might enhance survival and result in a stable population structure. Hedgerows on the other hand seemed to contain viable populations of certain forest plant species and can therefore function as an important colonization source when incorporated in recently established forests.     

Factors limiting plant recruitment in a tropical Afromontane Forest

Predicting how forest species composition may change in response to global change is essential for meaningful management. Which species are most likely to successfully recruit depends on a multitude of factors, but processes operating at the seed-to-seedling transition being especially important. Here, we explore how insufficient seed dispersal (dispersal limitation) and ecological filtering of seedlings (establishment limitation) influence species recruitment in an Afromontane Forest. We combined census data from seeds in seed traps and seed-seedling transition plots in a Nigerian montane forest to measure dispersal and establishment limitation across species, life history strategies, and seed sizes. From 106 seed traps and 318 seedling plots, we recorded 11,278 seeds from 38 species and 2688 seedlings of 61 species. All species except for one liana showed strong dispersal limitation while 23 out of the 30 species showed establishment limitation. The two lianas sampled had significantly lower dispersal limitation compared to the trees investigated in the study. Our results suggest that both dispersal and establishment limitation are markedly high in this Afromontane forest than would be expected in a lowland counterpart.     

辽东山区次生林木本植物空间分布

森林木本植物的空间格局有助于揭示群落结构的形成机制与潜在的生态学过程,且对林分经营具有一定指导意义。在0—50 m尺度范围内综合分析了辽东山区4 hm2温带次生林样地多度10的树种空间格局。研究发现:(1)在完全随机零模型下,大部分树种呈现聚集格局,聚集格局树种的比例随尺度增加而降低;在32 m的较大尺度下,随尺度增加,随机和规则格局成为树种分布的主要形式;(2)在异质性泊松过程零模型下,55.9%的树种呈现随机格局,其余大部分树种在10 m的尺度下呈现聚集格局,且随尺度增加,规则格局成为主要形式;(3)在完全随机零模型下,树种属性(林层、径级和多度)显著地影响种群聚集度,而在异质性泊松过程零模型下,树种属性对种群聚集度不存在显著影响。综上,生境异质性、扩散限制和树种属性部分解释了辽东山区次生林木本植物空间分布格局,相对而言,生境异质性的效应更为突出。研究结果有助于揭示次生林群落生物多样性的维持机制。     

Epiphytic bromeliad communities in secondary and mature forest in a tropical premontane area

We analyzed the differences in species richness, community composition, population structure and within-tree location of epiphytic bromeliads in contiguous secondary and mature forests in a premontane area in Costa Rica. Diversity in the mature forest was highest, and the communities differed in their composition as well as in the recruitment rates of the dominant species. Guzmania monostachia and Catopsis nutans dominated the secondary forests, whereas Tillandsia fasciculata and T. tricolor were more abundant in the mature forest. The secondary forest species showed high rates of seedling recruitment while the opposite was found for the mature forest species. Species presence and abundance among and within habitats did not correlate with their physiological (i.e. CAM vs. C₃ photosynthesis) or morphological attributes. The spatial distribution patterns were similar among habitats; bromeliads tended to aggregate on a few relatively large phorophytes. The species shared a similar vertical stratification within habitats, except for the two dominant species in the early and mid-successional stages, although its ecological implication is not clear. With some exceptions, conspecifics of different ages were located on similar substrate types (i.e. stems, primary, secondary, or tertiary branches) within the tree-crowns, which suggests limited within-tree dispersion. Differences in species composition and rates of seedling recruitment among secondary and mature forest may arise from ecophysiological differences among species; however, the combined effect of seed availability and dispersal differences may have a larger influence. Thus, epiphyte community assembly can only be understood when the differences in habitat conditions, the availability of propagules, their dispersal characteristics and requirements for seedling establishment are known.     

Former land use affects the nitrogen and phosphorus concentrations and biomass of forest herbs

The colonization rates of understorey plants into forests growing on former agricultural land differ remarkably among species. Different dispersal and recruitment largely account for the contrasting colonization rates, but different effects of the soil legacies of former agricultural land use on plant performance may also play a role. Seven herbaceous forest species were sampled in paired post-agricultural and ancient forest stands to study whether land-use history has an effect on the aboveground nutrient concentrations (N, P and N:P ratios) and biomass of forest herbs and, if so, whether slow and fast colonizing species respond differently. Results showed that P concentrations were significantly affected by former land use with higher concentrations in the post-agricultural stands. N concentrations were unaffected and N:P ratios were significantly higher in the ancient stands. Nutrient concentrations varied considerably among species, but the variation was unrelated to their colonization capacity. Six out of the seven species had higher biomass in the post-agricultural stands relative to the ancient stands, and the degree to which the species increased biomass was positively related to their colonization capacity, i.e., the fast colonizing species showed the strongest increase. Such differential responses to past land use may contribute to the contrasting colonization capacity of forest plants. Land-use history thus affected both the nutrient concentrations and biomass of forest herbs, and only the biomass response was related to colonization capacity.     

Seed- vs. microsite-limited recruitment in a myrmecochorous herb

Dispersal in myrmecochorous herbs is frequently short-distance and this might limit patch occupancy. Here we ask whether recruitment and patch occupancy in the myrmecochere Helleborus foetidus are seed- or microsite-limited and what role dispersal plays in recruitment limitation. In pine forests from two widely separated mountain systems in the Iberian Peninsula, we performed two seed addition experiments to explore the nature of recruitment limitation in this perennial herb. In an introduction experiment, we added seeds in patches far from reproductive individuals, which rarely receive any seed. In an augmentation experiment, we added seeds in patches beneath reproductive plants. Each experiment considered three main factors: seed addition (added or not added), rodent exclusion (excluded or not excluded), and microhabitat (scrub-covered or not scrub-covered). The augmentation experiment tests whether recruitment is limited by seed availability (exploring also the possible role of rodent granivory in such limitation) while the introduction experiment tests whether the empty patches occur either because they are unsuitable for the species (microsite limitation) or due to a low probability of seed arrival (dispersal limitation). Adding seeds almost always increased seedling emergence and recruitment, demonstrating that recruitment in this species is seed-limited and refuting the possibility of microsite limitation. The comparison of the introduction and augmentation experiments indicated that Helleborus recruitment dynamics and patch occupancy are constrained by additive effects of fecundity and dispersal limitation.     

Plant recruitment bottlenecks in temperate forest fragments: seed limitation and insect herbivory

Numerous studies have documented declines in plant diversity in response to habitat loss in fragmented landscapes. However, determining the mechanisms that lead to species loss is challenging using solely a correlative approach. Here we link correlative assessments of plant community composition with seed additions for a focal species to test the hypothesis that distributions of forests plants within a fragmented landscape are limited by seed dispersal. Woody plant species richness of fragments declined as fragments (n=26) became more isolated by agricultural fields. We predicted that if these isolation effects were driven by poor dispersal rather than other effects associated with habitat loss, then plants should vary in their response to isolation in relation to their seed size (i.e., stronger effects for plants with larger seeds). As predicted under this dispersal limitation hypothesis, sensitivity of bird-dispersed shrubs to isolation was related to their seed mass, with species with heavy seeds (e.g., Lindera benzoin) exhibiting stronger declines in presence across isolation gradients than species with light seeds. Seed addition experiments were performed for Lindera benzoin in two high isolation forest fragments (nearest neighbor mean distance=803 m) where Lindera was naturally absent, and two low isolation fragments (nearest neighbor mean distance=218 m) with naturally occurring Lindera populations. Seed addition and control plots (n=50 1 m² plots per fragment) were monitored for 13 censuses over 3 years. Across all four fragments, seed additions resulted in significant increases in Lindera seedling recruitment with no differences in final seedling establishment among fragments. However, insect herbivory was higher on Lindera seedlings in high isolation compared to low isolation fragments and was negatively correlated with seedling survival over some years. Consistent with prior work, our results confirm that seed dispersal plays a significant role in affecting plant diversity in fragmented landscapes. However, results also suggest the need for a better understanding of how additional processes, such as herbivory, may be altered as habitat is lost and what effects such changes have for forest plants.     

Effects of habitat alteration on the effectiveness of plant-avian seed dispersal mutualisms: Consequences for plant regeneration

Anthropogenic habitat alteration may affect the dispersal service provided by avian seed dispersers, ultimately causing regeneration collapse, through a decay in both the quantitative (seed removal) and qualitative (seed arrival to safe sites) components of seed dispersal effectiveness. However, despite its implications for management in real-world landscapes, few studies have investigated the shifts in components of seed dispersal effectiveness resulting from habitat alteration. We advocate the use of stage-specific transition probabilities, combined with data on seed shadows and bird abundance and mobility, for a mechanistic inference of the consequences for recruitment of the disruption of plant-frugivore mutualism in altered habitats. Such an approach allows the identification of regeneration bottlenecks, evaluates the differential contribution to recruitment of quantitative and qualitative components of seed dispersal, and provides the means to compare seed dispersal limitation. We exemplify our conceptual approach with studies of seed dispersal and recruitment in the wild olive tree in unaltered and severely altered adjacent sites. We show that simplification of the habitat substantially affected bird abundance, diversity and mobility, which caused a reduction in fruit removal and a concomitant simplification of the seed shadows compared to the unaltered site. Linked to these shifts, postdispersal seed survival and seedling emergence and survival were affected. The final outcome of habitat alteration was the collapse of the regeneration dynamics with very few seeds escaping the influence of maternal plants and reaching the safest sites for recruitment (dispersal limitation). As predicted, the collapse in the regeneration dynamics resulted from severe decays in the quantitative but especially in the qualitative components of seed dispersal effectiveness. Management of fleshy-fruited plant populations in altered habitats should thus pay attention to landscape elements that promote frugivore abundance, diversity and mobility and that alleviate the dispersal limitation.     

Land crabs as key drivers in tropical coastal forest recruitment

Plant populations are regulated by a diverse assortment of abiotic and biotic factors that influence seed dispersal and viability, and seedling establishment and growth at the microsite. Rarely does one animal guild exert as significant an influence on different plant assemblages as land crabs. We review three tropical coastal ecosystems–mangroves, island maritime forests, and mainland coastal terrestrial forests–where land crabs directly influence forest composition by limiting tree establishment and recruitment. Land crabs differentially prey on seeds, propagules and seedlings along nutrient, chemical and physical environmental gradients. In all of these ecosystems, but especially mangroves, abiotic gradients are well studied, strong and influence plant species distributions. However, we suggest that crab predation has primacy over many of these environmental factors by acting as the first limiting factor of tropical tree recruitment to drive the potential structural and compositional organisation of coastal forests. We show that the influence of crabs varies relative to tidal gradient, shoreline distance, canopy position, time, season, tree species and fruiting periodicity. Crabs also facilitate forest growth and development through such activities as excavation of burrows, creation of soil mounds, aeration of soils, removal of leaf litter into burrows and creation of carbon-rich soil microhabitats. For all three systems, land crabs influence the distribution, density and size-class structure of tree populations. Indeed, crabs are among the major drivers of tree recruitment in tropical coastal forest ecosystems, and their conservation should be included in management plans of these forests.