Microenvironmental and vegetational heterogeneity induced by phytogenic nebkhas in an arid coastal ecosystem

The nebkhas of woody plants represent distinct habitats in arid and semiarid ecosystems. Nebkhas are mounds composed of wind-borne sediment within or around shrub canopies. We studied the effects of widely spaced nebkhas of Retama raetam shrub on their microenvironment and associated herbaceous vegetation in the Mediterranean coast of Sinai Peninsula. Our measurements included nebkha size (height and width) and shrub size (canopy height and diameter). We identified four distinct microsites at each nebkha: crest, mid-slope, edge, and internebkha space. We measured soil temperature and moisture, photosynthetically active radiation (PAR), and soil properties. The plant species grown at each microsite were identified and their densities were measured. Average soil temperature and PAR were highest at internebkha space and lowest at nebkha crest. The maximum diurnal temperature and PAR of internebkhas exceeded that of nebkhas. Soil moisture and nutrient concentrations showed a gradient of spatial heterogeneity and were highest at the nebkha edge. Regression analysis indicated that total herbaceous plant density was significantly related to nebkha size, and to shrub canopy diameter and area. Detrended correspondence analysis indicated that patterns of species composition were correlated with the spatial variability in soil moisture and nutrient content along the gradient of increasing distance from the nebkha crest. It is assumed that shrub canopy and its nebkha interact in governing ecosystem functioning in this environment.     

藏锦鸡儿群落土壤资源空间异质性

在内蒙古高原, 藏锦鸡儿(Caragana tibetica)群落分布在草原向荒漠的过渡带上, 在植被区划中常作为由草原进入荒漠的指示物种。藏锦鸡儿是一种旱生垫状矮灌木, 由于其垫状生物学特性, 在地表覆沙的生境中常形成较均匀的、非常醒目的灌丛沙堆。不同大小的灌丛沙堆可对应其不同的发育阶段, 选取不同大小的藏锦鸡儿灌丛沙堆, 划分成小、中、大3组, 小、中两组灌丛沙堆对应其发育阶段, 大沙堆组对应于稳定阶段, 通过测定沙堆间、沙堆内(顶部、中部)及沙堆下的土壤有机质(Soil organic matters, SOM)、全磷(Total phosphorus, TP)和土壤质量含水量(Mass water content of soil, W_m), 分析了藏锦鸡儿群落土壤资源的特点。结果显示, 随着沙堆的发育, 沙堆内和堆下0~20 cm处的土壤有机质、全磷和土壤质量含水量的平均含量呈递增趋势, 大沙堆、中沙堆及堆下0~20 cm处的土壤有机质、全磷和土壤质量含水量的平均含量均大于沙堆间, 小沙堆的土壤有机质、堆下0~20 cm的土壤全磷和质量含水量的平均含量大于沙堆间。不同大小的灌丛沙堆, 在垂直方向上, 土壤有机质、全磷含量基本均呈先增加后减少的趋势, 土壤质量含水量呈递减趋势; 在水平方向, 随着灌丛沙堆的发育, 土壤有机质、全磷和土壤质量含水量在同一部位的同一土层基本呈递增趋势; 不同大小的沙堆表层0~20 cm 处的土壤有机质由顶部、中部、沙堆间依次呈先增加后减小的趋势; 大沙堆和中沙堆的全磷由沙堆顶部、中部、沙堆间依次呈递减趋势, 而小沙堆土壤全磷呈先增加后减少的趋势。随着灌丛沙堆的发育, 藏锦鸡儿灌丛截获的植物残体量呈增加趋势。研究显示, 藏锦鸡儿在发育过程中形成“沃岛效应”, 这种效应不仅表现在沙堆内, 也表现在沙堆下, 同时也引起土壤有机质、全磷和土壤水分的空间异质性分布。     

Role of host identity in effects of phytogenic mounds on plant assemblages and species richness on coastal arid dunes

Question: Phytogenic mounds (nebkhas) ‐ the natural accumulation of wind‐blown sediments within or around the canopies of plants ‐ have been proposed as important structures for locally maintaining high species richness in coastal and arid ecosystems. Nebkhas are assumed to increase habitat heterogeneity, but what is the importance of the nebkha host species relative to other nebkha characteristics in determining the associated plant assemblages? Are some host species more effective in creating diversity hotspots, or does a single species‐area relationship apply to all nebkhas, regardless of host species? Can the influence of the host be ascribed to its indirect effects on abiotic attributes of the nebkha complex? Methods and location: We investigated plant species richness and composition on nebkhas around six psammophytic species on Mediterranean coastal dunes of the Sinai Peninsula. Results: Plant species richness was significantly related to nebkha size by the single power function according to the general prediction of island biogeography theory, but this relationship was modified ‐ though to a limited degree ‐ by nebkha host species identity. Canonical Correspondence Analysis revealed that nebkha host species identity and nebkha environmental and non‐environmental factors significantly explained species composition on the nebkhas, but host species identity did so to a greater extent. The latter might reflect differences in seed trapping ability or free space for colonization between host species. Conclusion: Differences in community composition and richness among nebkhas formed by different host species represent a key factor in the maintenance of plant diversity on arid coastal dunes.     

Nebkhas of Salvadora persica and their effect on the growth and survival of Prosopis cineraria,Tamarix aphylla,and Capparis decidua trees and shrubs

Few studies were published on the effect of nebkhas (phytogenic mounds) on species diversity and soil resources, but no detailed study has been conducted yet on possible specific influence of nebkhas on growth and survival of the plants associated with them. We studied the nebkhas of Salvadora persica and their effect on growth and survival of three woody species (Prosopis cineraria, Tamarix aphylla, and Capparis decidua) in the Ommanian coast of Hormozgan Province in the south of Iran. The results showed that mean height and mean canopy diameter of P. cineraria and T. aphylla trees and shrubs inhabiting nebkhas of Salvadora persica were considerably higher than those of plants of these species growing outside nebkhas. The reverse occurred in the case of C. decidua. Generally, the percentages of stems with dead parts were significantly lower in plants inhabiting the nebkha sites in comparison to comparable ones growing outside the nebkhas. Salvadora persica nebkhas are enriched with more soil nutrients in comparison to inter-nebkha sites. Soil accumulated per each hectare in the nebkhas of the study area dominated by trees of Salvadora persica amounted to 237.6 m³. This indicates the great importance of nebkhas in the protection of soil and the associating species.     

Changes in spatial point patterns of pioneer woody plants across a large tropical landslide

We assessed whether the relative importance of positive and negative interactions in early successional communities varied across a large landslide on Casita Volcano (Nicaragua). We tested several hypotheses concerning the signatures of these processes in the spatial patterns of woody pioneer plants, as well as those of mortality and recruitment events, in several zones of the landslide differing in substrate stability and fertility, over a period of two years (2001 and 2002). We identified all woody individuals with a diameter >1 cm and mapped them in 28 plots measuring 10 × 10-m. On these maps, we performed a spatial point pattern analysis using univariate and bivariate pair-correlation functions; g (r) and g₁₂ (r), and pairwise differences of univariate and bivariate functions. Spatial signatures of positive and negative interactions among woody plants were more prevalent in the most and least stressful zones of the landslide, respectively. Natural and human-induced disturbances such as the occurrence of fire, removal of newly colonizing plants through erosion and clearcutting of pioneer trees were also identified as potentially important pattern-creating processes. These results are in agreement with the stress-gradient hypothesis, which states that the relative importance of facilitation and competition varies inversely across gradients of abiotic stress. Our findings also indicate that the assembly of early successional plant communities in large heterogeneous landslides might be driven by a much larger array of processes than previously thought.     

Production of Perennial Vegetation in an Oasis-desert Transition Zone in NW China - Allometric Estimation,and Assessment of Flooding and Use Effects

River oases at the southern fringe of the Taklamakan desert in NW China are surrounded by belts of spontaneous vegetation that protect the oases from sand drift. As an important source of forage, fuel and construction wood, this foreland vegetation is also a component part of the agricultural system of the oases but has been, and still is, destroyed through overuse. Within a broader study that aimed to provide a basis for a sustainable management of this foreland vegetation, biomass and production were studied in four vegetation types dominated either by Alhagi sparsifolia, Calligonum caput-medusae, Populus euphratica, or Tamarix ramosissima that were thought to occur under different regimes of natural flooding in the foreland of Qira (Cele) oasis, Xinjiang, NW China. Shoot biomass components were closely correlated to basal area (Calligonum, Populus, Tamarix) or shrub volume and projection area (Alhagi), enabling non-destructive estimation of stand biomass from shoot diameters or shrub dimensions with sufficient precision using allometric regression equations. Relationships between shoot basal area and biomass of the woody species (Calligonum, Populus and Tamarix) agreed with predictions by a theoretical model of plant vascular systems, suggesting that they are determined by hydraulic and mechanical requirements for shoot architecture. Average aboveground biomass densities of typical stands in late summer were 2.97 Mg/ha in Alhagi, 3.6 Mg/ha in a row plantation and 10.9 Mg/ha in homogenous stands of Calligonum, 22–29 Mg/ha in 22 year-old Populus forests and 1.9–3.1 Mg/ha in Tamarix-dominated vegetation. Annual aboveground production including wood and assimilation organs ranged from 2.11 to 11.3 Mg/ha in plantations of Calligonum, 3.17 to 6.12 Mg/ha in Populus, and 1.55 to 1.74 Mg/ha (based on total ground area) or 3.10 to 7.15 Mg/ha (in homogenous stands) in Tamarix. Production of Alhagi is equal to peak biomass. A thinning treatment simulating use by the local population enhanced productivity of Calligonum, Populus and Tamarix. A complete harvest of Alhagi in late August decreased production in the following year. An artificial flood irrigation treatment did not sufficiently increase soil water content except in the uppermost layer and had no clear beneficial effect on growth of the four species and even a negative effect on Alhagi, which was due to increased competition from annual species. As biomass and production with or without artificial irrigation were much higher than values expected for rain-fed desert vegetation at a mean annual precipitation of 35 mm, it is concluded that the existence of all vegetation types studied is probably based on permanent access to groundwater and that natural floods or precipitation do not contribute to their water supply. The effects of agricultural groundwater use in the oasis on groundwater in the foreland of the oasis need further study. Sustainable use of this productive vegetation is possible but requires proper management.     

The Importance of Phytogenic Mounds (Nebkhas) for Restoration of Arid Degraded Rangelands in Northern Sinai

Abstract Natural accumulation of wind‐borne sediments within or around the canopies of plants plays an important role in the ecological and evolutionary dynamics of many coastal and desert ecosystems. The formation of such phytogenic mounds (nebkhas) creates patches that can strongly influence the spatial distribution of plant and soil resources. In land restoration of arid and semiarid environments it is important to study the potential role of such biological patchiness that may provide sites for coexistence of species with different life and growth forms. Our main objective was to test whether the nebkhas of a leguminous shrub, Retama raetam (white broom), promote restoration of herbaceous vegetation and soil in the degraded rangelands of northern Sinai. Vegetation and microclimatic and edaphic characteristics within the nebkhas, as well as within internebkha spaces, were compared for ungrazed and grazed sites. Abundance and richness of herbaceous plants were positively related to nebkha area, which explained more of the variance of abundance and richness in the grazed site than in the ungrazed one. Protection from grazing, especially on nebkhas, was associated with an increase in abundance and richness of herbaceous plants, improved soil microclimate, and increased soil fine particles and nutrient concentrations. The results suggest that management (in casu protection from grazing) of nebkhas of woody perennial shrubs changes rangeland conditions and improves the resource regulatory processes. Furthermore, nebkhas of unpalatable plants have the potential to preserve plant diversity in overgrazed plant communities, because they are effective in capturing and retaining water, soil materials, and propagules within and from nearby areas, resources that would otherwise be lost.     

Effect of Caragana tibetica nebkhas on sand entrapment and fertile islands in steppe�Cdesert ecotones on the Inner Mongolia Plateau, China

Phytogenic mounds (nebkhas) formed by shrubs are a common phenomenon in arid and semi-arid areas and play important roles in preventing soil erosion and nutrient loss. One feature of nebkhas is the development of fertile islands. We investigated whether fertile islands were present inside and underneath Caragana tibetica-formed nebkhas in the northwest portion of the Ordos on the Inner Mongolia Plateau, China, and if such a fertile island effect increased with the age of the nebkhas. We also analyzed the spatial heterogeneity of the soil properties and quantified the amount of sand trapped by C. tibetica. The morphometric characteristics of C. tibetica nebkhas were investigated in a 4.5 ha area. Soil samples were collected inside, underneath and outside of the nebkhas and the soil organic matter (SOM), total phosphorus (TP), soil moisture (SM) and soil texture were determined. The SOM, TP and SM inside and underneath nebkhas at later stages of growth were higher than those at smaller, younger growth stages, which in turn were higher than those outside the nebkhhas. Inside the nebkhas at the establishment and early growth stages, SOM and TP first increased and then decreased with increasing soil depth, but SM steadily increased. The amount of sand trapped by C. tibetica per unit area was 0.0313 m³m^?2. Nebkhas of C. tibetica primarily accumulated fine sand, which accounted for 74% of the soil, and is significantly higher than that outside the nebkhas. Caragana tibetica significantly increased sand entrapment, and fertile islands are formed inside and underneath C. tibetica nebkhas.     

Patterns of adult abundance in Chthamalus stellatus (Poli) and C. montagui Southward (Crustacea: Cirripedia) emerge during late recruitment

The aim of this study was to investigate when adult distribution patterns are established in the barnacles Chthamalus stellatus and C. montagui. Adult ‘zones’ were identified by analysing field counts of both species at mid and upper shore heights. Monthly collections of cyprids, < 1 month old metamorphs and recruits (all metamorphosed individuals older than approximately 1 month) were made for C. stellatus and C. montagui in natural barnacle beds at six shores in SW Ireland. This was carried out over one year in 1996/1997, using a hierarchical sampling design. Abundance of total recruits (0-3 months old) was compared between adult zones after the main settlement season had ended. In addition, scales of variability in 0-3 month recruitment into adult zones were compared between the species at two scales: shores (1000s of metres) and sites within shores (10s of metres). Older recruits of each species, up to 11 months of age, were also compared between adult zones.The majority of settlement (measured as attached cyprids) occurred between August and October 1996. In October, there was no effect of adult zone on the abundance of total (0-3 month) recruitment up to that point in either species. Despite this homogeneity in recruitment between adult zones, significant spatial variation was found in 0-3 month recruits of both species at both of the scales examined. In C. stellatus the amount of variation associated with the larger scale (shore) was more than twice that of sites or of the residual variation (replicates within sites). 0-3 month recruitment in C. montagui was also most variable at the scale of shores but the residual variability (between replicates within site) was of similar magnitude to that of shores. Variability in 0-3 month C. montagui recruitment was relatively low at the scale of sites.There was a small but consistent input of recruits to adult zones over 9 months of the year, complicating the assessment of when adult patterns were set-up in these species. By June 1997, characteristic patterns of adult dominance had been established at all shores. Settlement had completely ceased by this time and individual barnacles were potentially 11 months old. Neither settlement nor early recruitment are significant in determining adult zonation patterns in these species. Instead, differential mortality patterns in individuals up to the age of 11 months are implicated in determining patterns of distribution of both species.     

两类植物型沙丘上植物群落的异同及其对沙丘形态的响应

为探究相同环境条件下发育的两类植物型沙丘上植物群落的异同点,以及相异点与沙丘形态的关系,对毛乌素沙地南缘盐碱地上相间分布的抛物线形沙丘和白刺灌丛沙丘分别进行形态学、植物群落学及土壤理化性质的调查分析。结果显示:抛物线形沙丘的水平尺度虽为白刺灌丛沙丘的12-23倍,但两类沙丘上物种数相当,其植物分属12科31属39种和12科30属33种,均以菊科、藜科、禾本科、豆科植物为主(占70%以上),其中共有植物17种,群落相似度0.66;两类沙丘上的群落建群种不同,优势种的重叠度较低,抛物线形沙丘的不同部位共统计到7个植物群丛,可分为沙生植物群落和喜湿耐盐碱群落,白刺灌丛沙丘上均以白刺为建群种,油蒿、冰草、雾冰藜、沙蓬、狗尾草为主要优势种;沙丘形态造成其不同部位风沙活动及土壤水分、PH值和全盐含量的差异是两类沙丘上植物群落相异的重要影响因素。     

Do riparian plant community characteristics differ between <Emphasis Type="Italic">Tamarix</Emphasis> (L.) invaded and non-invaded sites on the upper Verde River,Arizona?

Invasion by Tamarix (L.) can severely alter riparian areas of the western U.S., which are globally rare ecosystems. The upper Verde River, Arizona, is a relatively free-flowing river and has abundant native riparian vegetation. Tamarix is present on the upper Verde but is a minor component of the vegetation (8% of stems). This study sought to determine whether riparian vegetation characteristics differed between sites where Tamarix was present and sites where Tamarix was absent during the invasion of the upper Verde. We hypothesized that herbaceous understory and woody plant communities would differ between Tamarix present and absent sites. Our hypothesis was generally confirmed, the two types of sites were different. Tamarix present sites had greater abundance of all vegetation, native understory species, graminoids, and native trees, and a positive association with perennial native wetland plant species. Tamarix absent sites had greater abundance of exotic plants and upland adapted plants and an association with greater abiotic cover and litter. These results are contrary to other reports of Tamarix association with depauperate riparian plant communities, and suggest that Tamarix invasion of a watershed with a relatively natural flow regime and a robust native plant community follows similar establishment patterns as the native riparian plant community.     

甘肃民勤不同演化阶段白刺灌丛沙堆分布格局及特征

通过对甘肃民勤不同演化阶段灌丛沙堆的调查,研究了其空间格局、形态和表层土壤水分特征及其差异,并对研究区的生态环境现状进行了分析。结果表明,研究区灌丛沙堆以稳定阶段为主,但这种稳定状态可能是暂时的,随着距离沙丘距离的变化,不同演化阶段灌丛沙堆呈现出有规律的分布,植被、表层土壤水分以及风力的相互作用是引起这种分布的主要原因。随着灌丛沙堆的演化,灌丛沙堆的形态参数呈现不同的变化趋势,雏形阶段灌丛沙堆的高度与水平尺度之间无明显相关性,发育阶段灌丛沙堆高度随水平尺度的增加呈现二次函数增长,稳定阶段灌丛沙堆高度随水平尺度的增加呈现对数函数增长,活化阶段灌丛沙堆高度与水平尺度的相关性较差。     

More than just drought: complexity of recruitment patterns in Mediterranean forests

Understanding community dynamics during early life stages of trees is critical for the prediction of future species composition. In Mediterranean forests drought is a major constraint for regeneration, but likely not the only factor determining the observed spatial patterns. We carried out a sowing experiment aimed at identifying main filters during seed-seedling transition. Specifically, we studied seed fate (predation, fungi infection, emergence) and subsequent seedling performance (mortality during the first summer and overall recruitment after 2 years) of four co-occurring Mediterranean tree species (Quercus ilex, Quercus faginea, Juniperus thurifera, Pinus nigra). We related these processes to the dominant species composition, microhabitat heterogeneity, herb cover and seed mass. The identity of the dominant species in the forest canopy was more important for recruitment than the forest canopy being dominated by conspecific vs. heterospecific species. The patterns we found suggest that biotic interactions such as facilitation (lower mortality under the canopies) and herb competition (during emergence of J. thurifera) are relevant during recruitment. Moreover, our results pointed to ontogenetic conflicts regarding the seed mass of Q. faginea and to density-dependent seed mortality for Q. ilex, rarely described in Mediterranean ecosystems. We propose that our study species experience population growth in forests dominated by heterospecifics where the recruitment success depends on habitat heterogeneity and on moderated biotic and abiotic stresses created by each species. Our results reveal patterns and mechanisms involved in recruitment constraints that add complexity to the well-known drought-related processes in Mediterranean ecosystems.     

不同沙源供给条件下柽柳灌丛与沙堆形态的互馈关系——以策勒绿洲沙漠过渡带为例

以塔克拉玛干沙漠南缘策勒绿洲—沙漠过渡带为研究区域,根据沙源供给丰富程度的明显差异,在固定沙地、半固定沙地、半流动沙地和流动沙地随机选择100个相对独立的柽柳灌丛沙堆,对每个沙堆形态进行测量,并对沙堆上的柽柳灌丛进行调查,分析研究不同沙源供给条件下柽柳灌丛形态特征和沙堆形态变化及其相互关系。结果表明:(1)在4种沙源供给条件下,柽柳灌丛高度相对稳定,灌丛长度、宽度、树冠投影面积、总枝条数、新枝率、基部新枝率和生物量随着沙源供给丰富度的增加而相应增加。(2)随沙源供给丰富度的增加沙堆尺度显著增大,外形向着沿风向拉长的不对称圆丘状发展,风影沙尾轮廓逐渐清晰。沙堆形态参数间存在良好的相关性,其中长度与宽度协同增长,回归曲线在流动沙地为二次函数关系,在其它3种沙地呈线性相关;底面积与高度表现为良好的二次函数关系;底面积与体积间存在显著的线性关系。(3)柽柳灌丛形态参数与沙堆形态参数间存在良好的相关性。其中树冠面积与沙堆体积间为显著线性关系,而树冠大小又与总枝条数以及新枝率密切相关,柽柳灌丛沙堆的适应机理应归于不断增加的沙源供给而产生的风积沙埋对柽柳灌丛生长的促进作用及灌丛与沙堆间的互馈效应。该区域柽柳沙包的形成以一定灌幅的存在为前提,可以判定是土地退化的产物,其中半流动沙地生境相对于其他沙地正处于急速退化中。     

Effects of Aspect on Clonal Reproduction and Biomass Allocation of Layering Modules of Nitraria tangutorum in Nebkha Dunes

The formation of many nebkha dunes relies on the layering of clonal plants. The microenvironmental conditions of such phytogenic nebkha are heterogeneous depending on the aspect and slope. Exploring the effects of aspect on clonal reproduction and biomass allocation can be useful in understanding the ecological adaptation of species. We hypothesized that on the windward side layering propagation would be promoted, that biomass allocation to leaves and stems of ramets would increase, and that the effects of aspect would be greater in the layering with larger biomass. To test these hypotheses, we surveyed the depth of germination points of axillary buds, the rate of ramet sprouting, the density of adventitious root formation points, and the biomass of modules sprouting from layering located on the NE, SE, SW and NW, aspects of Nitraria tangutorum nebkhas. The windward side was located on the NW and SW aspects. The results indicated that conditions of the NW aspect were more conducive to clonal reproduction and had the highest rate of ramet sprouting and the highest density of adventitious formation points. For the modules sprouting from layering on the SW aspect, biomass allocation to leaves and stems was greatest with biomass allocation to adventitious roots being lowest. This result supported our hypothesis. Contrary to our hypothesis, the effects of aspect were greater in layering of smaller biomass. These results support the hypothesis that aspect does affect layering propagation capacity and biomass allocation in this species. Additionally, clonal reproduction and biomass allocation of modules sprouting from layering with smaller biomass was more affected by aspect. These results suggest that the clonal growth of N. tangutorum responses to the microenvironmental heterogeneity that results from aspect of the nebkha.     

The drivers of avian‐haemosporidian prevalence in tropical lowland forests of New Guinea in three dimensions

Haemosporidians are among the most common parasites of birds and often negatively impact host fitness. A multitude of biotic and abiotic factors influence these associations, but the magnitude of these factors can differ by spatial scales (i.e., local, regional and global). Consequently, to better understand global and regional drivers of avian‐haemosporidian associations, it is key to investigate these associations at smaller (local) spatial scales. Thus, here, we explore the effect of abiotic variables (e.g., temperature, forest structure, and anthropogenic disturbances) on haemosporidian prevalence and host–parasite networks on a horizontal spatial scale, comparing four fragmented forests and five localities within a continuous forest in Papua New Guinea. Additionally, we investigate if prevalence and host–parasite networks differ between the canopy and the understory (vertical stratification) in one forest patch. We found that the majority of Haemosporidian infections were caused by the genus Haemoproteus and that avian‐haemosporidian networks were more specialized in continuous forests. At the community level, only forest greenness was negatively associated with Haemoproteus infections, while the effects of abiotic variables on parasite prevalence differed between bird species. Haemoproteus prevalence levels were significantly higher in the canopy, and an opposite trend was observed for Plasmodium. This implies that birds experience distinct parasite pressures depending on the stratum they inhabit, likely driven by vector community differences. These three‐dimensional spatial analyses of avian‐haemosporidians at horizontal and vertical scales suggest that the effect of abiotic variables on haemosporidian infections are species specific, so that factors influencing community‐level infections are primarily driven by host community composition.     

Regeneration dynamics in arid subtropical thicket,South Africa

In arid environments nurse-plants modify localised habitats and create regeneration opportunities for seedlings vulnerable to hostile conditions created by biotic and abiotic factors. Facilitation is thus recognised as an important process structuring plant communities in harsh environments. Here we use spatial patterns of species association and recruitment to infer species replacement patterns in arid subtropical thicket of South Africa. Although our study site was floristically impoverished, all major plant functional groups that characterise subtropical thicket were present. Portulacaria afra clumps comprised approximately 50% of the study site by area. The mean and median clump size was 16.5 and 6.9 m², respectively, indicating the prevalence of small individuals in the population. Approximately 90% of tree seedlings were recorded under P. afra clumps and 93% of P. afra seedlings were recorded under woody shrubs. P. afra seedlings were recorded more frequently than expected beneath Rhigozum obovatum compared with other woody shrub species. No clear recruitment patterns were recorded for R. obovatum. Lycium cinereum, a woody shrub, and the stem-succulent Psilocaulon absimile were distributed more frequently on nutrient rich patches than expected and both these species are replaced by grass as the nutrient rich patch ages. Mature trees were generally recorded growing to the south of the assumed founding P. afra stem indicating that tree establishment was more frequent on the shaded side of P. afra clumps. However, most trees grew towards the sunny north and east-facing aspects. Plant species replacement patterns are facilitated by nurse-plant effects in arid subtropical thicket. These recruitment patterns together with our inferred species replacement on nutrient rich patches result in a predictable sequence of species replacement that is cyclic in nature.     

Fine-scale spatial genetic dynamics over the life cycle of the tropical tree Prunus africana

Studying fine-scale spatial genetic patterns across life stages is a powerful approach to identify ecological processes acting within tree populations. We investigated spatial genetic dynamics across five life stages in the insect-pollinated and vertebrate-dispersed tropical tree Prunus africana in Kakamega Forest, Kenya. Using six highly polymorphic microsatellite loci, we assessed genetic diversity and spatial genetic structure (SGS) from seed rain and seedlings, and different sapling stages to adult trees. We found significant SGS in all stages, potentially caused by limited seed dispersal and high recruitment rates in areas with high light availability. SGS decreased from seed and early seedling stages to older juvenile stages. Interestingly, SGS was stronger in adults than in late juveniles. The initial decrease in SGS was probably driven by both random and non-random thinning of offspring clusters during recruitment. Intergenerational variation in SGS could have been driven by variation in gene flow processes, overlapping generations in the adult stage or local selection. Our study shows that complex sequential processes during recruitment contribute to SGS of tree populations.