The quantification of local substrate heterogeneity in streams and its significance for macroinvertebrate assemblages

The effect of substrate heterogeneity on the structure of stream macroinvertebrate assemblages (total abundance, taxon richness, and evenness) is still not clear, but this could be due to the lack of standard methods for quantifying substrate heterogeneity. An accurate quantification of substrate heterogeneity was obtained from photographs of sampled areas (each 225 cm²), which were used to create maps that were subsequently digitized and analyzed using image analysis software. These maps allowed the calculation of multiple metrics quantifying two aspects of substrate heterogeneity: composition and spatial configuration of substrate patches. The diversity of substrate types (calculated as the Shannon diversity index), and the heterogeneity of patch compactness (calculated as the coefficient of variation of the relationship between patch dimensions) were the metrics explaining more biotic variance at the sample scale, but at higher scales there were no relationships between assemblage structure and substrate heterogeneity. Most variation in substrate heterogeneity occurred at the sample scale, while some metrics varied significantly at riffle or segment scales; these patterns of variation match those of macroinvertebrate assemblages, which had been previously studied. The importance of quantifying substrate heterogeneity and considering the spatial scales of its study are discussed.     

A Mathematical Model for Invasion Range of Population Dispersion Through a Patchy Environment

We focus on the question of how the dispersion of an invading population is affected by the spatial distribution of patches that have resource available for the population’s settlement and reproduction. We have developed and analyzed a mathematical model with a simple stochastic process. The patches are grouped into three classes – free, occupied and abandoned – depending on the state of the patch used by the population. We especially consider the range expanded by invaded patches, the invaded range R, assuming a certain generalized relation between R and the total number of invaded patches k, making use of an index, a sort of fractal dimension, to characterize the spatial distribution of invaded patches. We show that the expected velocity is significantly affected by the nature of spatial distribution of resource patches, and is temporally variable. When the invading population finally becomes extinct at a certain moment, the terminal size of the invaded range at that the moment is closely related to the nature of the spatial distribution of resource patches, which is explicitly demonstrated by our analysis.     

沙冬青灌丛地的土壤颗粒大小分形维数空间变异性分析

通过土壤颗粒大小分形维数对土壤质地定量分析表明,土壤颗粒大小分形维数与沙粒含量（＞0.1mm）呈显著的线性负相关,与粘粉粒含量（＜0.05mm）呈显著的线性正相关关系（p＜0.0001）,沙粒和粘粉粒含量每增加1﹪,分形维数则降低或升高0.022个单位,沙粒增加导致分形维数的降低和粘粉粒增加导致分形维数升高幅度一致,所以土壤颗粒分形维数可以作为评价土地沙质荒漠化程度的定量指标之一.荒漠地带以沙冬青为优势种的沙生植被地段,地表土壤颗粒粗粒化和50～70cm土层土壤颗粒的细粒化,为该区有限降水的深层渗漏提供了基质,为沙冬青的存活提供了保障,一定程度上支持了一些专家提出的概念模型,验证了粗质土壤质地支持以灌木为主的群落的假说.但从空间变异性分析来看,荒漠区沙冬青群落因地表物质的吹蚀和堆积过程频繁以及灌丛沙堆发育程度引起的分枝形态差异,导致了土壤颗粒含量空间变异尺度并未集中在冠幅范围,并不支持灌丛在沙漠生态系统对降尘等细粒物质拦截所形成的＂沃岛＂作用.正是因为以沙冬青为优势种的沙生植被地段土壤颗粒组成的高度空间异质性,并且空间变异的范围并不局限于灌木冠幅范围和冠幅间的裸地,没有为其它植物种的入侵创造土壤基质条件,才使得沙冬青群落在该区稳定存在,为荒漠残遗植物种--沙冬青的迁地保护和干旱沙区植被恢复过程中合理地利用土壤资源,以避免营林失败等问题提供了理论依据.     

鄂尔多斯草地退化过程中植被地上生物量空间分布的异质性

应用地统计学方法分析研究了鄂尔多斯草地荒漠化过程中植被地上生物量空间分布的异质性。发现从本氏针茅(Stipa bungeana)、本氏针茅 油蒿(Artemisia ordosicn)、油蒿、油蒿 牛心朴子(cynanchum komarovii)、牛心朴子群落的5个退化系列群落中,植被地上生物量空间格局异质化程度及其相关范围变化非常明显。变异函数的基台值(C0 C)在5个退化群落中的变化表明,在本氏针茅群落中,植被地上生物量空间分布异质性较低(0．48),本氏针茅 油蒿群落增强(19．47),油蒿群落显著增强(1522),油蒿 牛心朴子群落减弱(171．6),牛心朴子群落显著减弱(4．164)。变异函数的相关范围(A0)表明,在本氏针茅群落空间相关范围较小(53．9cm),本氏针茅 油蒿群落增加(77．8cm),油蒿群落减小(51．7cm),油蒿 牛心朴子群落增加(62．9cm),牛心朴子群落增加(94．6cm)。空间变异比分析表明,由空间自相关因素引起的空间异质性占主要部分(74．2％～80．7％)。空间分布格局图(Kriging map)分析进一步显示,退化群落中植被地上生物量格局变化显著。从本氏针茅群落退化到油蒿群落的中度荒漠化过程中,植被地上生物量空间分布的异质性的增强。由油蒿群落退化到牛心朴子群落的严重荒漠化过程中,植被地上生物量空间分布的异质性减弱。     

干旱地区山地荒漠草原阴坡植物群落空间异质性

在样线调查基础上 ,以 Shannon- Wiener指数 ,群落盖度 ,DCA排序轴为区域化随机变量 ,应用半方差、分形分析等方法 ,对干旱地区山坡植物群落空间异质性进行了研究。结果表明 ,在整个山坡尺度上植被格局的空间异质性不大 ,小尺度上植被格局的空间异质性较大 ,尺度依赖性较强 ,不同群落类型的空间格局不同 ,随尺度变化的规律也不一样。严重放牧干扰强烈影响干旱山地植物群落的空间格局 ,生境中的牧道格局 ,斑块格局使山坡植物群落多样性空间异质性更加复杂 ,牧道效应是导致群落空间格局周期性振荡的重要因子。去势对应分析 (DCA)排序第一轴特征值体现了综合生态因子对群落格局作用的结果 ,DCA第二轴半方差变化包含了大量信息 ,其变化具有周期性。群落多样性空间格局强烈影响着干旱山地生态系统的各种生态学过程 ,这些作用机制有待进一步研究     

Multi-scale effects of resource patchiness on foraging behaviour and habitat use by longnose dace, Rhinichthys cataractae

1. We examined the response of a predatory benthic fish, the longnose dace ( Rhinichthys cataractae ), to patchiness in the distribution of benthic macroinvertebrates on cobbles at three hierarchical spatial scales during summer and autumn 1996, and spring 1997 in a southern Appalachian stream. 2. At the primary scale (four to five individual cobbles separated by <1 m), the intensity of foraging was not correlated with the biomass of benthic macroinvertebrates/cobble, regardless of season. 3. At the secondary scale (i.e. foraging patches <5 m in diameter) we found that benthic macroinvertebrates were patchily distributed in summer, but not in autumn or spring. Concomitantly, in summer, longnose dace foraged on cobbles with a significantly higher biomass of benthic macronvertebrates than nearby, randomly selected cobbles with similar physical conditions (i.e. longnose dace tended to avoid low-prey foraging patches). In contrast, when benthic macroinvertebrates were distributed homogeneously (spring and autumn), dace did not select patches with a significantly higher biomass of benthic macroinvertebrates than that available on randomly selected cobbles. 4. At the tertiary scale (i.e. stream reaches 11–19 m long), the biomass of benthic macroinvertebrates (per cobble per reach) was patchily distributed (i.e. differed significantly among reaches) in all seasons. Among reaches with physical characteristics preferred by longnose dace, (i.e. erosional reaches dominated by cobble/boulder substratum and high current velocity), we detected a significant, positive correlation between the biomass of benthic macroinvertebrates/cobble and longnose dace density in all seasons. 5. Our results demonstrated that both spatial and temporal patchiness in resource availability influenced significantly the use of both foraging patches and stream reaches by longnose dace.     

黄土高原南麓县域耕地土壤速效养分时空变异

在生态脆弱的黄土高原南麓合阳县,以1983、2006年耕地土壤速效氮、磷、钾含量为对象,利用空间自相关、变异函数及分形维数等方法,研究县域土壤速效养分时空变异特征及主要影响因素.结果表明,从1983到2006年,除土壤速效钾外,耕地速效氮、磷的平均含量分别提高73.98％和92.69％;土壤速效养分全局Moran's I指数和空间相关距均呈递减变化、分维数增加,预示其空间结构减弱,随机变异性增强.土壤速效氮、磷含量在绝大部分研究区域内呈现不同程度累积,而速效钾含量则在59.65％的耕地上减少;导致其不同变化的因素有耕地利用方式、施肥管理、灌溉、土壤类型等,其中以土地利用方式和施肥管理措施的影响最大.     

空间分析方法在微生物生态学研究中的应用

微生物生态正在受到越来越多的关注,对其研究也渐趋深入。然而由于微生物个体微小的特点及研究手段的限制,多数研究还停留在探索阶段,研究方法也在不断完善当中。近年来,较多的研究开始探讨空间因素在微生物多样性和分布中的影响,对空间分布的探讨有助于更好地认识生态过程,是一种有力的研究手段。微生物空间分析方法已经成为微生物生态学领域中重要的研究方向之一,我国空间方法在微生物生态研究中的应用还没有得到普遍的重视。从不同研究角度出发,结合空间统计的作用,对空间统计方法在微生物生态研究中的应用的必要性及现状做了评述。介绍了空间自相关性的检验,方差图,Mantel检验,Kriging插值等方法在微生物生态研究中的应用,并论述了微生物研究中的尺度问题。这一梳理,对丰富微生物生态学研究中的新方法、新手段具有一定价值。     

Tufa deposition and periphyton overgrowth as factors affecting the ciliate community on travertine barriers in different current velocity conditions

A characteristic of aquatic systems in karstic region is the formation of tufa – the product of calcium carbonate precipitation. Artificial substrata (glass slides) were used to investigate the influence of tufa deposits at two different current velocities (5 cm s^–1 and 50 cm s^–1) on the ciliate assemblages in periphyton. After two-month exposure periods, periphyton biomass and tufa deposit were c. three times greater at 50 cm s^–1 than at 5 cm s^–1. Ciliate population density was also higher on artificial substrata exposed in a lotic than in a lentic microhabitat (the overall mean number of ciliates at 5 cm s^–1 was 122 ind. cm^–2, and at 50 cm s^–1 497 ind. cm^–2). At each of the two observed current velocities, a Principal Components (PCA) ordination of the colonized ciliate associations showed a spatial separation of the associations that developed under two different conditions of tufa deposition. During the period of greater tufa deposit, associated with greater periphyton overgrowth rate, the ciliate communities had higher species diversity (a higher number of species and a lower number of individuals). Species diversity of ciliates had a positive nonlinear relation to tufa deposition rate. These results suggest that artificial surfaces covered by a rough tufa layer associated with greater periphyton biomass offer diverse conditions for colonization by ciliates.     

Inconsistency in the experimentally derived relationship between epilithon abundance and the micro-distribution of Agapetus monticolus (Trichoptera)

Abstract Studies of the micro-distribution of herbivorous lotic macroinvertebrates have tended to rely on short-term experiments. While these experiments have provided information on the factors that determine the distributions of herbivorous invertebrates, they ignore the complexity of the epilithon as a food source, any interaction between physical and biological factors and the possibility that the factors that determine an invertebrate species’ distribution may vary through time. Five colonization experiments were performed in an Australian upland stream to determine the role of epilithon abundance in the distribution of an abundant invertebrate (Agapetus monticolus: Trichoptera). Artificial cobbles with different types and amounts of epilithon were placed in the Taggerty River at two sites and the response of A. monticolus recorded. The experiments revealed that, although A. monticolus expressed a preference for patches of epilithon with abundant periphyton, the response varied among experiments performed at different times and was always secondary to the flow preferences of A. monticolus. Studies that fail to acknowledge the presence of temporal variation in the response of animals to their environment run the risk that they will over- or underestimate the importance of particular factors because of the timing of their experiment.     

科尔沁沙地景观格局特征分析

用修改分维数,分维数和景观多样性指数方法,对科尔沁沙地没沙漠化土地景观空间格局２进行了研究。结果表明,在所划分的人工固沙区,围封保护区和流动沙丘区三种沙地景观类型中,人工固沙区景观空间格局最复杂,修改分维数Ｄｍ为１．５４７６;其次为围封保护区Ｄｍ为１．４９７５;流动沙丘景观格局最简单Ｄｍ为１．４２０５。     

High assemblage persistence in heterogeneous habitats: an experimental test with stream benthic algae

1. The persistence of biological assemblages is positively affected by spatial heterogeneity. This influence may be indirect, through increased species richness. Another possibility is the increased availability of refuges from disturbances, which would prevent local loss. 2. We conducted a field experiment to test the hypothesis that greater roughness (a form of spatial heterogeneity) on the surface of substrata allows higher persistence of assemblages of stream benthic algae and that this relationship does not depend on species richness. Samples were taken on six occasions from smooth and rough artificial substrata used for algal colonisation. We calculated the persistence of assemblages using two analytical approaches: the mean distance to group centroid and the sum of the Euclidean distances between consecutive sampling occasions, both in a multivariate space. We also subsampled the data to take into account differences in species richness between treatments and thus to evaluate the effect of species richness on persistence. 3. Assemblages on rough substrata were more persistent than assemblages on smooth substrata. The effects detected were not due to the greater species richness on rough substrata, since a higher persistence of the assemblages on rough substrata remained after the subsampling procedures. 4. Our results indicate a strong positive relationship between substratum roughness and the persistence of stream benthic algal assemblages. We suggest that this is due to the presence of physical refuges in heterogeneous habitats.     

A heterogeneity test for fine-scale genetic structure

For organisms with limited vagility and/or occupying patchy habitats, we often encounter nonrandom patterns of genetic affinity over relatively small spatial scales, labelled fine-scale genetic structure. Both the extent and decay rate of that pattern can be expected to depend on numerous interesting demographic, ecological, historical, and mating system factors, and it would be useful to be able to compare different situations. There is, however, no heterogeneity test currently available for fine-scale genetic structure that would provide us with any guidance on whether the differences we encounter are statistically credible. Here, we develop a general nonparametric heterogeneity test, elaborating on standard autocorrelation methods for pairs of individuals. We first develop a 'pooled within-population' correlogram, where the distance classes (lags) can be defined as functions of distance. Using that pooled correlogram as our null-hypothesis reference frame, we then develop a heterogeneity test of the autocorrelations among different populations, lag-by-lag. From these single-lag tests, we construct an analogous test of heterogeneity for multilag correlograms. We illustrate with a pair of biological examples, one involving the Australian bush rat, the other involving toadshade trillium. The Australian bush rat has limited vagility, and sometimes occupies patchy habitat. We show that the autocorrelation pattern diverges somewhat between continuous and patchy habitat types. For toadshade trillium, clonal replication in Piedmont populations substantially increases autocorrelation for short lags, but clonal replication is less pronounced in mountain populations. Removal of clonal replicates reduces the autocorrelation for short lags and reverses the sign of the difference between mountain and Piedmont correlograms.     

Defining the limits to local density: alternative views of abundance–environment relationships

1. In many ecosystems, the local abundance of organisms is spatially heterogeneous. Ecologists often seek to explain this variation by modelling the central tendency of abundance as a function of a single dominant factor (central response, CR). An alternative approach is to model maximum and minimum abundance in relation to the dominant factor (limiting response, LR), thereby acknowledging that multiple factors may constrain abundance and create scatter in the relationship. In many ecosystems, including streams, abundance–environment relationships are traditionally expected to be CR models with a single dominant factor determining local abundance, but this hypothesis lacks rigorous test. This omission is of concern because CR modelled relationships form the foundations of many ecological tests, predictive and management tools that, consequently, may provide erroneous interpretations. 2. In a survey designed to minimise variance in the data, we related densities of three taxa of stream invertebrates to near‐bed flow. Data were analysed using both ordinary least squares (OLS) regression and quantile regression, which have different characteristic results depending on whether the relationship is better described as a CR or LR model. For all three taxa, invertebrate responses to flow conformed most closely to LR models and were best described by quantile regression, although OLS regression revealed broadly similar general trends in this case. 3. Based on the statistically modelled relationships, we were able to hypothesise which ecological mechanisms limited maximum abundance at this site, including dislodgement with high flow (Heptageniidae, Leuctridae), reduced oxygen uptake at low flow (Baetidae) and reduced provision of interstitial spaces at high flow (Leuctridae). Scatter in the relationships was attributable to multiple factors that may operate at different scales, including inter‐individual variation, alternative environmental gradients and biotic processes at the patch scale, and potential constraints of dispersal and settlement operating outside the patch. 4. We illustrate how the choice of CR or LR model to describe abundance–environment relationships may affect tests of ecological theory and applied ecology. If our results are typical of other streams and other types of systems, greater use of LR models may transform the way many ecologists view ecosystems, thus having positive consequences for survey and experimental designs and highlighting weaknesses in existing management tools that are underpinned by CR models.     

红树植物木榄幼树斑块形状的分形分析

应用分形理论分析了山口国家级红树林自然保护区木榄幼树斑块形状的分形特征。木榄幼树斑块在整体水平上的分形维数为 1 .1 2 ,而单个斑块的分形维数介于 1 .1 7～ 1 .3 7之间。分形维数的大小与斑块形状的复杂程度密切相关。频谱法和周长 -面积法都适用于分析幼树斑块形状的分形特征。     

The effects of a new reservoir on the attached diatom communities in Huntington Creek,Utah, U.S.A.

A study was made to determine whether a new reservoir being constructed on Huntington Creek, Utah would have significant effects on the attached diatom communities of the stream. Cluster analysis and other statistical methods indicated only small differences in the diatom communities before and after construction of the dam.