Small-scale spatial variability in intertidal and subtidal turfing algal assemblages and the temporal generality of these patterns

Spatial and temporal variation in patterns of distribution and abundance of algal assemblages is large and often occurs at extremely small spatial and temporal scales. Despite this, few studies investigate interactions between these scales, that is, how patterns of spatial variation change through time. This study investigated a number of scales of spatial variation (from tens of centimetres to kilometres) in assemblages of intertidal and subtidal turfing algae. Significant differences were found in the composition and abundances of species in assemblages of turf at all spatial scales tested. Much of the variation among assemblages could, however, be explained at the scale of quadrats (tens of centimetres apart) (27±1.4 (SE)% of dissimilarity) with an additional 7±1.2% explained at the scale of sites (tens of metres apart) and 10±1.5% at the scale of locations (kilometres apart). Although the greatest dissimilarity in assemblages occurred at the scale of habitats, this accounted for a relatively small proportion of the overall variation in assemblages. These patterns were consistent through time, that is, at each sampling time the spatial scale explaining the greatest proportion of variation in assemblages was replicate quadrats separated by tens of centimetres. These patterns appear to be due to small-scale variation in patterns of distribution and abundances of the individual species that comprise turfing algal assemblages. The results of this experiment suggest that large scale processes have less effect on patterns of variability of algal assemblages than those occurring on relatively smaller spatial scales and that small-scale spatial variation should not be considered as simply “noise”.     

Variation in Community Structure across Vertical Intertidal Stress Gradients: How Does It Compare with Horizontal Variation at Different Scales?

In rocky intertidal habitats, the pronounced increase in environmental stress from low to high elevations greatly affects community structure, that is, the combined measure of species identity and their relative abundance. Recent studies have shown that ecological variation also occurs along the coastline at a variety of spatial scales. Little is known, however, on how vertical variation compares with horizontal variation measured at increasing spatial scales (in terms of sampling interval). Because broad-scale processes can generate geographical patterns in community structure, we tested the hypothesis that vertical ecological variation is higher than fine-scale horizontal variation but lower than broad-scale horizontal variation. To test this prediction, we compared the variation in community structure across intertidal elevations on rocky shores of Helgoland Island with independent estimates of horizontal variation measured at the scale of patches (quadrats separated by 10s of cm), sites (quadrats separated by a few m), and shores (quadrats separated by 100s to 1000s of m). The multivariate analyses done on community structure supported our prediction. Specifically, vertical variation was significantly higher than patch- and site-scale horizontal variation but lower than shore-scale horizontal variation. Similar patterns were found for the variation in abundance of foundation taxa such as Fucus spp. and Mastocarpus stellatus, suggesting that the effects of these canopy-forming algae, known to function as ecosystem engineers, may explain part of the observed variability in community structure. Our findings suggest that broad-scale processes affecting species performance increase ecological variability relative to the pervasive fine-scale patchiness already described for marine coasts and the well known variation caused by vertical stress gradients. Our results also indicate that experimental research aiming to understand community structure on marine shores should benefit from applying a multi-scale approach.     

Scales of spatial patterns of distribution of intertidal invertebrates

Few comparative studies of spatial patterns at different scales have examined several species in the same habitat or the same species over a range of habitats. Therefore, variability in patterns among species or among habitats has seldom been documented. This study quantifies spatial patterns of a suite of intertidal snails and a species of barnacle using a range of statistical techniques. Variability in densities was quantified from the scale of adjacent quadrats (over a distance of centimeters) to tens of kilometers. Significant differences in abundances occurred primarily at two spatial scales. Small-scale differences were found at the scales of centimeters or 1–2 m and, for many species on many shores, these accounted for most of the variability in abundances from place to place. These are likely to be determined by behavioural responses to small-scale patches of microhabitat. Large-scale differences in abundance were also found in most species at the scale of hundreds of meters alongshore. These are likely to be due to variation in recruitment (and/or mortality) because of limited dispersal by adults of these species. There was little or no additional variation among shores, separated by tens of kilometers, than was shown among patches of shore separated by hundreds of meters. Identification of the scale(s) at which significant differences in abundance are found focus attention on the processes (and the scales at which these processes operate) that influence patterns of distribution and abundance. Some of the advantages and disadvantages of various procedures are discussed.     

Scales of mussel bed complexity: structure, associated biota and recruitment

Hierarchically scaled surveys were carried out on beds of the brown mussel Perna perna (Linnaeus) on the South coast of South Africa. The object was to assess spatial and temporal variations in the complexity of mussel beds and to investigate relationships between mussel bed complexity and mussel recruitment. Complexity was divided into three components: physical complexity; demographic complexity; associated biota. A series of variables within each component were recorded at two different scales (10 and 50 cm) within nested quadrats on three separate occasions. The nested ANOVA design explicitly incorporated spatial scale as levels of the ANOVA. These scales were: shores (areas 1 km in length separated by 25 km); transects (areas 20 m in length separated by 100s of meters); 50x50-cm quadrats separated by meters and 10x10-cm quadrats separated by cm) This approach was intended to generate hypotheses concerning direct associations between recruitment and complexity versus co-variation due external processes. Three main questions were addressed: (1) At what scale does each variable of complexity exhibit greatest significant variation? (2) At these scales is there similar ranking of variables of complexity and recruitment? (3) Within this/these scales, is there any significant relationship between the variables measured and mussel recruitment? On two occasions (Nov. 97 and Mar. 98) the majority of variables showed greatest significant variation at the transect-scale. On a third occasion (Oct. 97) most variables showed greatest significant variation at the quadrat-scale and the site-scale. On all occasions a markedly high percentage of the variation encountered also occurred at the smallest scale of the study, i.e., the residual scale of the ANOVA analyses. Some similarity in the ranking of variables occurred at the transect scale. Within the transect-scale, there was little indication of any relationship between variables of complexity and recruitment. Relationships were inconsistent either among transects or among sampling occasions. Overall, the results suggest that a high degree of variation in mussel bed complexity consistently occurs at very small scales. High components of variance generally also occur at one or more larger scales; however, these scales vary with season. Mussel recruitment does not appear to be directly affected by complexity of mussel beds. Instead it appears external factors may influence both complexity and recruitment independently. In addition recruitment may influence complexity rather than vice versa.     

Spatial and temporal patterns in the recruitment of the intertidal barnacle Chthamalus malayensis Pilsbry (Crustacea: Cirripedia) on the equatorial shores of Peninsular Malaysia and Singapore

Spatial and temporal variation in the recruitment of the intertidal barnacle Chthamalus malayensis was examined over one year (September 2003-August 2004) on the equatorial shores of southern Peninsular Malaysia and Singapore. A nested sampling design was applied for the first time on these shores, over three spatial scales - hundreds of kilometres, kilometres to tens of kilometres and tens of metres - and temporal variation was determined through monthly sampling of recruits. Shores within 2° north of the equator on the East and West coasts of southern Peninsular Malaysia and the southern coast of Singapore were selected as study sites. Generally, all three coasts recruited throughout the year, with varying intensities. There was a clear regional pattern, where the largest number of recruits was found on the East Coast of Malaysia, and the least in Singapore. Analyses of variance (ANOVA) showed that variability occurred on the smallest scale, but only in nine of the twelve months examined, consequently resulting in significant temporal and spatial interaction. Calculated variance components indicated that small-scale variation accounted for most of the overall variability. The potential causes of the spatial and temporal patterns of C. malayensis recruitment, and implications on tropical-temperate comparisons will be discussed.     

Plant coexistence in coastal heaths: Floristic patterns and species attributes

Abstract An investigation was made of plant coexistence in coastal heaths. A 4 km stretch of Pleistocene beach sand was selected, and patterns of variation in heath vegetation on ridges and slopes in the beach system were studied. The sampling design covered variation with geographic distance among transects, plots within transects and quadrats within plots which were orthogonal to the alignment of ridges and slopes. Cover of vascular plant species was scored within each quadrat and both multivariate and univariate analyses tested the hypothesis that variation in the data was associated with environmental difference and geographic distance. There was a strong flohstic variation between the ridge and slope habitats. However, within these habitats there was no significant variation with distance at scales of kilometres, nor at hundreds of metres within transects nor at tens of metres within plots in floristic composition, in numbers of species, and in cover. Though differing floristically, the vegetation of the slopes and ridges had similar curves of importance values and fairly similar spectra of life forms and forms of post-fire regeneration in their constituent species, although the ridges had slightly more obligate-seeding species and fewer species of sedges and herbs than the slopes. The two habitats shared several genera, however, within such genera, species with similar attributes mostly occurred in different habitats. Hypotheses were suggested to explain segregation of species between the two habitats and the coexistence of species within habitats. Since patterns of variation among habitats were consistent over both smaller and larger scales of distance examined, experiments to test hypotheses may be designed so that their findings might be applied generally to the whole system studied.     

Spatial analyses of intertidal assemblages on sheltered rocky shores

Abstract Understanding processes in complex assemblages depends on good understanding of spatial and temporal patterns of structure at various spatial scales. There has been little quantitative information about spatial patterns and natural temporal changes in intertidal assemblages on sheltered rocky shores in temperate Australia. Natural changes and responses to anthropogenic disturbances in these habitats cannot be accurately measured and assessed without quantitative data on patterns of natural variability in space and through time. This paper describes some suitable quantitative methods for examining spatial and temporal patterns of diversity and abundances of highshore, midshore and lowshore intertidal assemblages and the important component species for a number of shores in a bay that has not been severely altered by human disturbance. Despite a diverse flora and fauna on these shores, the midshore and lowshore assemblages on sheltered shores were characterized by a few species which were also the most important in discriminating among assemblages on a shore and, for each assemblage, among different shores. The same set of species was also important for measuring small-scale patchiness within each assemblage (i.e. between replicate sites on a shore). Therefore, these data provide a rationale for selecting species that are useful for measuring differences and changes in abundance among places and times at different scales and, hence, can be used in the more complex sampling designs necessary to detect environmental impacts. There was considerable spatial variability in all assemblages and all species (or taxa) examined at scales of metres, tens of metres and kilometres. There were no clear seasonal trends for most measures, with as much or more variability at intervals of 3 months as from year to year. Most interactions between spatial and temporal measures were at the smallest scale, with different sites on the same shore generally showing different changes from time to time. The cause(s) of this apparently idiosyncratic variability1 were not examined, but some potential causes are discussed. These data are appropriate for testing hypotheses about the applicability of these findings to other relatively undisturbed sheltered shores, about effects of different anthropogenic disturbances on sheltered intertidal assemblages and to test hypotheses about differences in intertidal assemblages on sheltered versus wave-exposed shores.     

Contrasting spatial heterogeneity of sessile organisms within mussel (Perna perna L.) beds in relation to topographic variability

We examined the spatial heterogeneity in three sessile rocky shore organisms, the mussel Perna perna, the barnacle Octomeris angulosa (Sowerby) and the red alga Gelidium pristoides (Turn.) at a range of continuous local scales along horizontal transects within mid- and upper mussel beds of South African shores. We also examined the relationships between variability of organisms and topographic features (rock depressions, slope, aspect), and between mussel, barnacle and algal variability over the same scales. To estimate spatial heterogeneity, we analyzed scaling properties of semivariograms using a fractal approach. Relationships between different variables at the different scales were examined by cross-semivariograms. Spatial dependence of P. perna variability increased with spatial dependence of topographic variability, so that scaling regions of mussel and topographic distributions corresponded well. This relationship often improved with larger local scales (mussel cover increased with depressions, steeper slope and aspect towards waves), while at smaller spatial scales, variability in mussel cover was less well explained by variability in topography. The variability of the barnacle O. angulosa exhibited spatial dependence, even on topographically unstructured shores. In contrast, the distribution of the alga G. pristoides revealed high fractal dimensions, showing spatial independence on topographically unstructured shores. Algae also showed a very strong negative relationship with mussels at most local scales, and a negative relationship with barnacles in upper zones, especially at larger local scales. Barnacles may show clear spatial dependence because of hydrodynamics (at larger local scales) and the need to find a future mate in close proximity (at smaller local scales), while algae may show a strong negative relationship with mussels because of competition for space.     

Similarity of rocky intertidal assemblages along the Pacific coast of Japan: effects of spatial scales and geographic distance

Factors and processes affecting community structures operate at various spatial and temporal scales. We analyzed how similarities of rocky intertidal assemblages vary at different spatial scales using a nested, hierarchical sampling design. Intertidal assemblages consisting of algae, sessile animals, and mobile animals were censused on five rock walls at each of five shores chosen within each of six regions along the Pacific coast of Japan, encompassing 1,800 km of coastlines. Based on this sampling design, similarities in assemblages were calculated using both qualitative (presence/absence) and quantitative (percent cover and density) data, and compared at three spatial levels: (1) rock level (the finest spatial scale, encompassing several to hundreds of meters), (2) shore level (the intermediate spatial scale, encompassing several to tens of kilometers), and (3) region level (the broadest spatial scale, encompassing hundreds to thousands of kilometers). Cluster analysis showed that assemblages were categorized into distinct regional groups except for the two southern regions, but they did not separate clearly from each shore. A nested analysis of similarities revealed significant variation in similarities among regions and among shores within each region, with the former showing greater variation. Similarity was negatively correlated with geographic distance at the regional level but not at the shore or the rock levels. At the regional level, similarity decreased more rapidly with distance for mobile animals than sessile organisms. The analyses highlighted the importance of broad-scale abiotic/biotic factors such as oceanic current conditions and biogeographic factors in determining observed patterns in similarity of rocky intertidal assemblages.     

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.     

No effect of naturally-occurring rock types on settlement or survival in the intertidal barnacle, Tesseropora rosea (Krauss)

The settlement and early survival of young barnacles, Tesseropora rosea (Krauss), on four different, naturally-occurring rock substrata were investigated in a four-factor experiment. Three factors assessed variation on three spatial scales: among shores on the New South Wales coast in eastern Australia i.e., 10 to 600 km apart; among sites within these shores i.e., 20 to 50 m apart; within these sites, i.e., < 3 m². Variation on these scales was usually significant for settlement density and proportional survival 30–60 days post-settlement. The fourth factor, different rock types, showed no consistent trends for settlement density or proportional survival. Previously observed differences in these phenomena among shores cannot, therefore, be ascribed to differences in rock substrata.     

SPATIAL SCALE OF GENETIC STRUCTURE AND AN INDIRECT ESTIMATE OF GENE FLOW IN EELGRASS,ZOSTERA MARINA

In this study, the first investigation of population structure in an aquatic angiosperm, I show that populations of a marine angiosperm (eelgrass, Zostera marina) are genetically differentiated at a number of spatial scales. I find also that there is no correspondence between geographic and genetic distances separating subpopulations, an increasingly common result in spatially stratified studies of genetic structure in marine invertebrates. F-statistics, calculated for two years from electrophoretic variation at five polymorphic allozyme loci, indicate significant genetic differentiation among sampling quadrats within each of two bays (θ = 0.064-0.208), between tide zones within a bay (θ = 0.025-0.157) and between bays (θ = 0.079). Spatial autocorrelation analysis was used to explore genetic differentiation at smaller spatial scales; estimated patch sizes (within which genetic individuals are randomly associated) indicated no appeciable genetic structure at scales less than 20 m × 20 m. Calculated values of F-statistics were a function of the spatial scale from which samples were drawn: increasing the size of the “subpopulation” included in calculation of fixation indices for the same “total” sample resulted in an increase in the magnitude of f (e.g., from 0.092 to 0.181) and a decrease in θ (e.g., from 0.186 to 0.025). On the basis of the best estimate of the spatial scale of subpopulations, the effective number of migrants per generation (N_em) ranges from 1.1 to 2.8. Genetic consequences of the disturbance regime in the eelgrass habitat sampled were extreme variation between years in the allele richness and proportion of heterozygotes in a sample and a positive relationship between the extinction probability of patches and the genetic variance among them. The changes in F-statistics as a function of sampling scale and the observation that θ among sampled quadrats was positively associated with the probability of extinction among quadrats indicated that indirect estimates of gene flow (N_em) calculated from θ should be cautiously interpreted in populations that may not yet be in drift-migration equilibrium.     

SPATIAL VARIATION PATTERNS IN THE MARINE MACROALGA XIPHOPHORA GLADIATA SSP. GLADIATA (PHAEOPHYTA). I. PARTITIONING OF MORPHOLOGICAL CARIATION INTO SPATIAL SCALES1

The geographic expression of morphological variation in xiphophora gladiata ssp. gladiata was examined throughout the range of the taxon. The spatial scales on which this variation occurred were identified using nested analyses of variance (ANOVA) on 14 morphometric characters from 1224 plants collected at 12 sites. These characters were remarkably homogeneous in the scales at which their variation was expressed. The greater part of the morphological variation in X. gladiata ssp. gladiata (over 90% for most characters was expressed between adjacent primary clumps; that is, over scales of about 100 mm. Since it was previously shown that the clumps of plants are themselves spatially discrete, the spatial expression of morphology in this taxon forms a true mosaic with abrupt between-cells boundaries. Much of the remaining variation in most characters was expressed at a scale of about 300 km. At intermediate scales, that is, over a few meters and (in most characters) a few kilometers, there appears to be negative additional variance. In effect, the mixture of clumps at these scales tends to be a uniform rather than a random selection of the available morphological variety of clumps within each region. This observed pattern is compared and contrasted with those previously found in the confamilial species Fucus distichus and F. evanescens.     

SPATIAL DYNAMICS OF A HIMANTHALIA ELONGATA (FUCALES, PHAEOPHYTA) POPULATION

In dense monospecific stands of plants intraspecific competition usually results in self-thinning, the concurrent increase in biomass and decrease in density over time. Self-thinning may also result in a change in the spatial pattern of individuals, but so far the spatial dynamics of marine plants has not been investigated. The brown alga Himanthalia elongata ( L.) S. F. Gray forms dense monospecific stands on many northern temperate rocky shores, and various attributes (including its simple form) facilitated the study of the spatial dynamics of this species .
The spatial pattern of settling zygotes was examined in the laboratory. In the absence of water movement, substratum heterogeneity, and a point source, zygotes usually settled in clumps rather than randomly. Within the clumps zygotes appeared to be regularly distributed at a scale similar to the size of the zygotes themselves. Furthermore, the clumps themselves seemed to be regularly distributed. On the shore, well-established stands of "button-stage" Himanthalia populations were examined during a period of extensive growth and self-thinning. Individual plants were initially highly regular in spatial pattern but became less so over time. The pattern of plants dying during self-thinning was also highly regular and probably reflected existing spatial regularity. However, using a hypothesis of mortality as a random event, I found that smaller plants had a less than average survival potential, while larger plants had a greater than average chance. A consideration of the spatial pattern of plants alive at the end of the study revealed regularity at a scale of 2–7 mm but a random spatial pattern at larger scales, which might indicate a small sphere of influence of competing individuals. The best predictor of mean nearest neighbor distances in the populations was mean plant diameter .     

Inconsistency in short-term temporal variability of microgastropods within and between two different intertidal habitats

Small-scale temporal variation in abundances of fauna in marine soft sediments has long been recognised. Many studies on rocky intertidal shores have, however, focused on larger fauna in single habitats and have primarily examined relatively long time-scales. The implications of small-scale variability are frequently not adequately addressed in the studies of changes in fauna over longer time-scales. Without knowledge of the magnitude of variation at smaller scales, comparisons across longer time-scales may be confounded. In this study, the temporal variability of a number of co-existing species of microgastropods in patches of two different intertidal habitats (coralline turf and sediment) in Botany Bay, New South Wales, Australia, was measured using a nested, hierarchical sampling design incorporating temporal scales of weeks, 1 and 3 months. In addition to habitats, there were also spatial scales of metres between plots and 100s of metres between the locations. There was generally a lack of consistency in the trends of variance for the three temporal scales at the smallest spatial scale of plots. In addition, the different species, including those that were closely related, showed different patterns of variation, depending on the habitat and site. These data show the importance of incorporating adequate scales of sampling in different habitats when analysing the distribution and abundance of microbenthos in intertidal habitats.     

四川卧龙国家级自然保护区多空间尺度下绿尾虹雉的生境选择

动物的生境选择具有空间尺度依赖性, 在不同空间尺度上影响生境选择的环境因素有所不同。研究不同空间尺度下动物生境选择的关键影响因子及其季节性变化, 对于全面了解物种的生境资源需求和开展生境保护具有重要意义。绿尾虹雉(Lophophorus lhuysii)是中国特有的高山雉类, 国家一级重点保护野生动物, 具有极高的保护价值。然而, 目前尚未对其不同尺度和时期的生境选择进行过探究。本研究于2019年10月至2020年10月, 在四川卧龙国家级自然保护区的羊角湾、魏家沟和文扎都3个区域共布设15条样线、303个样方, 并结合红外相机监测(176个红外相机位点), 对保护区内绿尾虹雉种群的生境利用状况进行了调查, 使用主成分分析和逻辑斯蒂回归模型分别从景观和微生境两个尺度对繁殖期(3‒8月)和非繁殖期(9月至翌年2月)的生境选择模式进行了分析。结果显示, 在景观尺度上, 在繁殖期和非繁殖期都显著偏好海拔较高(3,700‒ 4,300 m)、坡度较小(27°‒33°)、靠近阳坡、草甸和流石滩比例较高而森林和灌丛比例较低的生境。在微生境尺度上, 绿尾虹雉在繁殖期显著偏好岩石盖度较高的生境; 而非繁殖期则显著偏好草本盖度较高、灌木盖度和落叶盖度较低的生境。研究表明, 绿尾虹雉在景观和微生境尺度上均对生境有明显的选择性, 并且其微生境选择还存在季节性变化, 反映了该物种在生活史不同阶段具有不同的资源需求。本研究丰富了绿尾虹雉的基础生态学信息, 为卧龙及其他自然保护区绿尾虹雉的生境管理和种群保护工作提供了参考。     

Scale specific determinants of tree diversity in an old growth temperate forest in China

The major processes generating pattern in plant community composition depend upon the spatial scale and resolution of observation, therefore understanding the role played by spatial scale on species patterns is of major concern. In this study, we investigate how well environmental (topography and soil variables) and spatial variables explain variation in species composition in a 25-ha temperate forest in northeastern China. We used new variation partitioning approaches to discover the spatial scale (using multi-scale spatial PCNM variables) at which environmental heterogeneity and other spatially structured processes influence tree community composition. We also test the effect of changing grain of the study (i.e. quadrat size) on the variation partitioning results. Our results indicate that (1) species composition in the Changbai mixed broadleaf-conifer forest was controlled mainly by spatially structured soil variation at broad scales, while at finer scales most of the explained variation was of a spatial nature, pointing to the importance of biotic processes. (2) These results held at all sampling grains. However, reducing quadrat size progressively reduced both spatially and environmentally explained variance. This probably partly reflects increasing stochasticity in species abundances, and the smaller proportion of quadrats occupied by each species, when quadrat size is reduced. The results suggest that environmental heterogeneity (i.e. niche processes) and other biotic processes such as dispersal work together, but at different spatial scales, to build up diversity patterns.     

Spatial variation in polyphenolic content of Ascophyllum nodosum (Fucales,Phaeophyta)

Spatial variation in polyphenolic content in annual shoots of the brown alga Ascophyllum nodosum was quantified using a hierarchical sampling design. Three sampling levels, covering distances of 10⁰–10⁶ m, were used. Comparisons were made between two areas, Tjärnö on the Swedish west-coast and the Isle of Man in the Irish Sea, with very different types of environmental conditions. No significant differences in mean polyphenolic levels were found between the two study areas (6.6% of dry mass at Tjärnö and 9.2% at the Isle of Man), whereas significant and substantial differences were found among sites within areas (range 5.7%–11.4%) and among quadrats within sites (range 3.7%–13.1 %). The extensive variation at the smaller spatial scales points out the importance of using thorough sampling procedures at all levels in large-scale studies on algal polyphenolics, e.g. biogeographical comparisons, which have been neglected in several previous studies. Moreover, the results imply that experiments on causal factors of polyphenolic variation should be designed to explain the spatial scales on which the factors are impportant. This study also investigated the relationship between polyphenolic concentration and both plant size and mean area of annual shoots. The mean area was used as an estimate of the mean growth rate of the annual shoots within an individual. No significant relationships were found between shoot growth rate, or plant size, and polyphenolic levels in annual shoots at any of the three spatial scales that were investigated.     

Effects of canopy gaps on the genetic structure of Camellia japonica saplings in a Japanese old-growth evergreen forest

The genetic structure of Camellia japonica saplings was investigated in relation to canopy conditions in an old-growth evergreen forest in Tsushima, Japan. To elucidate effects of canopy gaps on genetic structure, a 1 ha study site was divided into 20 x 20 m quadrats, which were classified into a gap quadrats (GAP), closed canopy quadrats (CLS) and mixed quadrats. Five GAP quadrats and six CLS quadrats were analyzed separately. Isolation-by-distance was tested by examining the correlation between genetic distance and geographic distance. A significant positive correlation was detected for GAP quadrats, whilst that for CLS quadrats was significantly smaller and not significantly different from zero. On the other hand, an analysis using Moran's I spatial autocorrelation coefficients indicates that the genetic structure is weaker in GAP quadrats than in CLS quadrats in short distance classes. The values were significantly positive for both types of quadrat. These results, along with our field observations on flowering, suggest that canopy gaps affect the genetic structure of C. japonica saplings in two distinct ways. First, canopy gaps may promote flowering and mating in an isolation-by-distance manner within canopy gaps. Second, canopy gaps may promote seed production and resulting overlap in seed shadows may weaken fine-scale genetic structures.