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.     

The independent and interactive effects of snail grazing and nutrient enrichment on structuring periphyton communities

We investigated the independent and interactive effects of nutrient enrichment and snail grazing on structuring periphyton communities in a northern temperate lake. Nutrient releasing substrates and grazer enclosures were used to simultaneously manipulate nutrient availability and herbivory. Periphyton was allowed 18 days to accrue before grazers (Elimia livescens = Goniobasis livescens) were introduced.Addition of nitrogen and phosphorus caused a significant increase in biovolume (p < 0.001), whereas grazing had no significant effect on biovolume but resulted in a shift in species composition. Four taxa were largely responsible for the increase in biovolume on the nutrient enriched substrates: Oedogonium sp, Stigeoclonium tenue, Navicula radiosa var. radiosa and Navicula radiosa var. tenella. By the 28th day, nutrient enrichment caused a shift from a community dominated by diatoms (Bacillariophyceae) to a community dominated by green algae (Chlorophyceae). Blue green algae (Myxophyceae) maintained an equal proportion in high and low-nutrient regimes.Grazing had a more pronounced effect on altering community composition on the nutrient enriched substrates than on the unenriched substrates. Grazing caused a decrease in diversity and an increase in dominance by green algae on the nutrient enriched substrates. The relative biovolume of green algae increased from 64% to 93% on grazed substrates, due to the significant increase in relative abundance of Stigeoclonium tenue. This taxon has both prostate basal cells and erect filamentous cells. The ratio of basal: filamentous cells increased from 4.7 to 5.2 with grazing, suggesting that the heretotrichous growth form of Stigeoclonium tenue is adapted to grazing by virtue of the basal cells which are able to adhere to the substratum and resist being grazed.     

The diatoms ingested by freshwater snails: temporal,spatial, and interspecific variation

Seventeen species of diatoms, representing a broad range of sizes, shapes, and growth habits, were collected from rocks in rapidly-flowing sections of the Mitchell River, North Carolina. The diatoms ingested by adult Goniobasis proxima, juvenile Leptoxis carinata, and adult Physa sp. co-occurring in this habitat were indistinguishable from one another, in spite of great differences in radular morphology. All snails sampled the diatom flora almost randomly, with only one or two of the larger diatom species under-represented in the gut contents. Some snails also seemed to selectively ingest the smaller individuals of the larger diatom taxa, and larger individuals of the smaller diatom taxa. The diatoms identifiable in juvenile Goniobasis guts were somewhat more distinctive, although this seemed to be due at least partly to more mechanical breakage. The diatom flora of quiet, muddy pools was much different from that of shallow, rocky areas, but once again, Goniobasis seemed to sample the available flora randomly. Seasonal variation was also apparent in the diatom diet of Goniobasis. We suggest that in some cases, it may be reasonable to use snails to sample the diatom assemblage present in a particular habitat, if more direct methods are impractical.     

The impact of herbivore type and grazing pressure on benthic microalgal diversity

Benthic microalgae and the herbivores Idothea chelipes (Isopoda) and Littorina littorea (Gastropoda) were used as an experimental model system to study the effect of herbivore type on primary producer diversity. Both herbivores enhanced diversity at intermediate grazing pressures, the enhancing effect being stronger with Littorina . This difference is explained by differences in the spatial heterogeneity of the grazed microalgal biofilm caused by behavioural differences in spatial grazing patterns. Idothea spreads its grazing pressure rather evenly in space ("lawn-mower" type), whereas Littorina creates pronounced spatial heterogeneity by moving slowly over the biofilm and decimating it within the grazing tracks ("bulldozer" type). Idothea grazing did not enhance biomass patchiness beyond the level of ungrazed controls (maximum/minimum from 1.7 to 5.5), whereas Littorina strongly increased patchiness (maximum/minimum from 45 to 850).     

EFFECTS OF IRRADIANCE AND GRAZING ON LOTIC ALGAL ASSEMBLAGES1

A laboratory experiment was conducted for 75 days to examine how irradiance levels and grazing influence algal biomass and community structure. Twelve laboratory streams were used for experimental analyses, with four channels exposed to one of three irradiance levels (15, 100, or 400 μE·m^?2·s^?1). Three of the four stream at each light level were stocked with the snail Juga silicula (250·m^?2), leaving one stream at each light level without snails. Grazed stream exposed to low light levels developed low amounts of algal biomass (<2 g AFDW·m^?2) and were dominated by adnately attached diatoms. Mean algal biomass increased over time in the grazed streams exposed to intermediate light; by day 75, these streams were characterized by moderate algal biomasses (30-40 g AFDW·m^?2) and filamentous chlorophytes. Algal assemblages in high light, grazed channels had high levels of biomass at day 43 (70 g AFDW·m^?2) that declined to 30 g AFDW·m^?2at day 75 and were dominated by chlorophytes. Among ungrazed streams, algal biomass at day 75 was relatively low in the low light streams (<7g AFDW·m^?2) and was dominated by adnately attached diatoms. Ungrazed streams exposed to intermediate and high light levels had moderate biomasses (23 and 19 g AFDW·m^?2, respectively) and were dominated by chlorophytes and large diatoms. Grazing appeared both to delay and alter successional trajectories of algal assemblages, with alterations most noticeable during early seral stages at intermediate and high light levels. Grazing had the least effect on successional trajectories at low light.     

生态学系统的空间异质性

空间异质性是生态学系统的一个普遍牧场生,生态学家对它在生态学中的重要性已取得了比以往更深刻的认识。试图从空间异质性的含义,空间异质性与尺度和等级的关系,空间异质性的定量描述,空间异质性对生物和非生物过程的影响,以及空间异质性的动态等5个方面综述了有关空间异质性的生态学研究的新进展。     

Grazer species effects on epilithon nutrient composition

1. Field and laboratory experiments were conducted to investigate the excretion stoichiometry of nitrogen (N) and phosphorus (P) of two benthic macroinvertebrate grazers, the crayfish Orconectes propinquus and the snail Elimia livescens, that differ in body stoichiometry (mean body molar N : P 18 and 28, respectively). Crayfish excretion had a significantly higher ammonium : soluble reactive phosphorus (SRP) ratio in the laboratory and in three natural streams than did snails, as predicted by ecological stoichiometry theory. 2. In greenhouse recirculating artificial streams, treatments consisting of crayfish, snails, or no grazers were used to examine responses in dissolved nutrient concentrations and epilithon nutrient composition and limitation. SRP concentrations depended upon the grazer species, with the snail treatment having a higher SRP concentration than other treatments (P < 0.05). Dissolved inorganic N was not affected by grazers, but appeared to be rapidly incorporated in epilithon. 3. Epilithon N content was dependent upon the grazer species present, with the crayfish treatment having a significantly higher N content than other treatments (P = 0.001). No grazer species effects on epilithon P content were found. However, both grazer treatments had significantly lower epilithon P content than the no‐grazer treatment. 4. Traditionally, studies have focused on how grazer‐induced structural changes to epilithon can alter epilithon nutrient dynamics, but this structural mechanism could not solely explain differences in epilithon nutrient contents and ratios in the present study. Our results rather suggest that benthic grazers can alter epilithon nutrient composition and limitation via nutrient excretion. Consequently, macroinvertebrate grazers may serve as ‘nutrient pumps’ that partly regulate the availability of nutrients to algae in stream ecosystems.     

大城市边缘区景观破碎化空间异质性——以北京市顺义区为例

探索大城市边缘区景观破碎化格局及其空间异质性特征,有助于把握景观动态的变化机制及其生态问题.以北京市顺义区这一典型大城市边缘区为例,基于土地利用分类数据,采用有效粒度尺寸方法,对1992年、1999年和2009年景观破碎化空间变异进行了地统计学分析,主要包括尺度依赖性、空间异质性、空间格局及其生态过程的分析.研究结果表明:1)景观破碎化空间变异具有明显的尺度依赖性特征,由空间自相关部分引起的空间异质性较高;1992年和1999年景观破碎化空间总变异相对缓和,2009年空间总变异强烈;通过不同尺度上的异质性对比,选取1000m幅度作为研究区景观破碎化的空间异质性分析较为适宜.2)景观破碎化空间异质性表现出明显的上升趋势,土地利用斑块破碎化不断加剧,其中1999-2009年间的表现尤为强烈.3)景观破碎化高值区呈现出由东部向西部转移的态势,并且其空间布局具有集聚性与分散性的显著特征,空间上相似属性聚集与相异属性聚集的空间格局十分突出.4)景观破碎化空间自相关作用主要来自高强度社会经济活动的影响,并表现出明显的区位特征.该研究结果可为土地、城镇规划以及景观生态建设提供参考.     

不同尺度苔藓结皮土壤性状的空间分布特征

苔藓结皮是生物结皮的高级阶段,它在防风固沙、改善土壤水分、养分等方面具有十分重要的生态作用,但前期研究多是对结皮区和无结皮覆盖区土壤性状的比较,对结皮斑块内土壤理化因子的空间分布情况尚不明确。运用经典统计学,地统计学以及克里格插值法探讨了土壤水分、有机质、全氮、全磷、总盐在样方和斑块两个尺度的空间变异特征,旨在阐明不同尺度藓类结皮土壤空间异质性的强度,明确不同尺度土壤各性状合适的取样距离。研究结果表明藓类结皮土壤性状在两种尺度均表现出中等程度的变异,随着取样尺度减小,自相关性在空间异质性中占的比重增加,结构因素的影响增强,随机因素的影响减弱。样方尺度的空间异质性大于斑块尺度异质性,且土壤性状的变程较大,采样时可适当增大取样间距;两种尺度下5个土壤性状中,全磷的变程最小,实际取样时应适当缩小取样距离,总盐变程大,自相关程度低,因而取样间距可适当放大。斑块尺度,藓类结皮土壤性状由边缘向中心递增(磷递减)的现象与斑块的边缘效应有关。     

华南海岸英罗港红树植物木榄种群结构的空间异质性

采用位置指数(CE)、分异指数(TC和TH)、Shannon-Wiener多样性指数(D)以及Ripley的K-方程,探讨了华南海岸英罗港树植物木榄种群的分布格局、胸转和树高分异以及冠层结构方面的空间异质性.多数种群呈现随机分布,其个体胸转和树高的分异程度较低;页少数种群呈现集群分布,其个体胸围和树高的分异程度明显.采用地理信息(GIS)对冠层和空隙斑块进行了多种水平垂直尺度的分析,冠层与空隙斑块之间的镶嵌格局因种群而异,这种可基于树冠抽影用Shannon-wiener多样性指数时行定量描述.冠层结构的窨异质性阻碍空间尺度而变化,但这种变化在一定尺度范围内保持相对的稳定.迷一尺度范围可作为木榄红树森更新或生态管理单位的参考尺度.     

华南海岸英罗港红树植物木榄种群结构的空间异质性(英文)

The spatial heterogeneity, including distribution pattern, tree perimeter and height differentiation, and canopy structure heterogeneity, of Bruguiera gymnorrhiza (L.) Lamk populations at Yingtuo Bay, South-China Coast was investigated using the positioning index (CE), differentiation index (TC and TH), Shannon-Wiener diversity index (D), and Ripley‘s K-functions. Most populations showed random distribution and low differentiation in perimeters and heights of individuals, while a few showed clumped distribution and clear differentiation. Canopy and gap patches were analyzed at multiple horizontal and vertical scales using geographic information system (GIS). The mosaic patterns of canopy and gap patches are different among populations, and could be quantitatively described with the Shannon-Wiener diversity index based on crown projection. The spatial heterogeneity of the canopy structure changed with spatial scales, but this kind of change would remain relatively stable over a range of scales. This scale range could be regarded as the referenced scale for a regeneration or ecological management unit for the forest.     

Environmental variability and the ecological effects of spawning Pacific salmon on stream biofilm

1. Variation in resource subsidies can create or reinforce heterogeneity in recipient ecosystems. Related activities of organisms delivering resource subsidies, such as ecosystem engineering by Pacific salmon spawners (Oncorhynchus spp.), also alter heterogeneity. We studied whether heterogeneity in stream environmental conditions and spawner abundances were reflected in the net ecological effects of salmon (i.e. enrichment by resource subsidies and disturbance by ecosystem engineering) on benthic biofilm. 2. We sampled seven Southeast Alaska streams over 3 years, both before and during the salmon run. In each stream and year, stream environmental characteristics and their influence on responses of benthic biofilm [mean and coefficient of variation of chlorophyll a (chl a), ash‐free dry mass (AFDM) and autotrophic index (AFDM:chl a)] to spawners were assessed. 3. Streams and periods before and during the salmon run were distinct based on their environmental characteristics. The responses of most biofilm metrics to spawners were stream‐ and year‐specific, suggesting that the ecological effect of spawners ranged from net enrichment to net disturbance depending on the stream or year studied. The environmental context, especially temperature, large wood, and sediment size, explained >50% of biofilm variability during the run, but <30% over the entire study, suggesting that salmon can alter environmental constraints. 4. Precision of biofilm estimates improved by increasing either the number of streams or the number of years sampled (i.e. spatial or temporal replication). However, combining data from different North Pacific Rim ecoregions inflated the confidence interval as compared with a single ecoregion, indicating the importance of regional environmental contexts for net salmon effects. 5. Our results suggest that biofilm responses to salmon can vary greatly, even within a single ecoregion, and that environmental conditions can modify net salmon effects. Consequently, generalisations about biofilm responses across the native range of salmon may be challenging.     

Spatial heterogeneity in soil particle size: does it affect the yield of plant communities with different species richness?

Aims Soil heterogeneity is ubiquitous in many ecosystems. We hypothesized that plant communities with higher species richness might be better adapted to soil heterogeneity and produce more biomass than those with lower richness. This is because there is niche differentiation among species and different species can complement each other and occupy a broader range of niches when plant species richness is high. However, no study has tested how soil particle heterogeneity affects the yield of plant communities, and whether such effects depend on the spatial scale of the heterogeneity and the species richness within the communities.Methods In a greenhouse experiment, we sowed seeds of four-species or eight-species mixtures in three heterogeneous treatments consisting of 32, 8 or 2 patches of both small (1.5mm) and large quartz (3.0mm) particles arranged in a chessboard manner and one homogeneous treatment with an even mixture of small and large quartz particles.Important findings Biomass production was significantly greater in the communities with high species richness than those with low species richness. However, soil particle heterogeneity or its interactions with patch scale or species richness did not significantly affect biomass production of the experimental communities. This work indicates that plant species richness may have a bigger impact on plant productivity than soil particle heterogeneity. Further studies should consider multiple sets of plant species during longer time periods to unravel the potential mechanisms of soil heterogeneity and its interactions with the impacts of species richness on community yield and species coexistence.     

Hydrological disturbance overrides the effect of substratum roughness on the resistance and resilience of stream benthic algae

1. Habitat heterogeneity in lotic systems is usually associated with the availability of refuges. Heterogeneous habitats (here, rough substrata) should mediate the effect of high‐flow disturbances by protecting benthic algae, thus increasing the resistance and resilience of the system. Additionally, the ability of algae to resist a disturbance and recover after it should be dependent on biological traits that confer resistance and resilience. 2. We designed a field experiment, simulating a high‐flow event with bed movement, to test the effect of substratum roughness on the resistance (assessed as the similarity between samples collected before and immediately after disturbance) and resilience (the similarity between samples collected before and 7 and 15 days after disturbance) of five algal life forms. We evaluated whether algal resistance and resilience were higher on rough than on smooth substrata, and whether the life forms differed in their ability to resist and recover from a disturbance. 3. Rough substrata had higher species richness than smooth substrata at all sampling periods, even immediately after the disturbance. There was no significant effect of substratum roughness on algal resistance and resilience, for both species richness and density of the total assemblage. Neither did roughness affect the resistance and resilience of the total algal assemblage or of the algal life forms separately, when evaluated using multivariate data sets (presence‐absence and quantitative). 4. Algal life forms differed in resistance and resilience; adnate/prostrate and erect/stalked species were more resistant and resilient than the other life forms (filamentous, motile and metaphytic). Additionally, motile species resisted and recovered better than did species that are only loosely associated with the substratum (metaphytic species). 5. Substratum roughness had no pronounced effect on benthic algal resistance and resilience. The results of this and some other studies suggest that the intensity of disturbance determines the importance of habitat heterogeneity and flow refuges for benthic algae in streams.     

Influence of habitat heterogeneity on the distribution of larval Pacific lamprey (Lampetra tridentata) at two spatial scales

1. Spatial patterns in channel morphology and substratum composition at small (1–10 metres) and large scales (1–10 kilometres) were analysed to determine the influence of habitat heterogeneity on the distribution and abundance of larval lamprey. 2. We used a nested sampling design and multiple logistic regression to evaluate spatial heterogeneity in the abundance of larval Pacific lamprey, Lampetra tridentata, and habitat in 30 sites (each composed of twelve 1‐m² quadrat samples) distributed throughout a 55‐km section of the Middle Fork John Day River, OR, U.SA. Statistical models predicting the relative abundance of larvae both among sites (large scale) and among samples (small scale) were ranked using Akaike's Information Criterion (AIC) to identify the ‘best approximating’ models from a set of a priori candidate models determined from the literature on larval lamprey habitat associations. 3. Stream habitat variables predicted patterns in larval abundance but played different roles at different spatial scales. The abundance of larvae at large scales was positively associated with water depth and open riparian canopy, whereas patchiness in larval occurrence at small scales was associated with low water velocity, channel‐unit morphology (pool habitats), and the availability of habitat suitable for burrowing. 4. Habitat variables explained variation in larval abundance at large and small scales, but locational factors, such as longitudinal position (river km) and sample location within the channel unit, explained additional variation in the logistic regression model. The results emphasise the need for spatially explicit analysis, both in examining fish habitat relationships and in developing conservation plans for declining fish populations.