Water quality changes of the river Meuse assessed by chironomid pupal exuviae

The chironomid fauna of the river Meuse in Belgium and The Netherlands was studied in order to determine spatial water quality variations. Because the river Meuse serves as a source for drinking water production for about five million people in Belgium and The Netherlands, water quality is an essential factor to be studied. From 1984 till 1989 pupal exuviae were collected. Water quality was assessed by several indices and autecological information of the various taxa. With regard to pollution the general trend through the years was the same. The most upstream sampling site, in the Belgian Ardennes, was relatively unpolluted. As a result of untreated industrial and municipal wastewater discharges in the Liège region, water quality deteriorated dramatically at the Dutch border. At the most downstream sampling site, near the river mouth, a slight recovery was observed. Apart from organic and inorganic pollution other environmental factors such as flow velocity and the nature of the riverbed, also play a significant role in the occurrence of the organisms. It is concluded that protective measures need to be taken to restore the affected river ecosystem.The editors deeply regret to have to announce that Nicole Frantzen died in a train accident on 30 November 1992.     

Phases in the development of riverine plankton: Examples from the rivers Rhine and Meuse

Observations on phyto and zooplankton in two hydrographically different rivers were compared in order to discriminate phases in plankton development. Along the longitudinal axis of the River Rhine a gradual increase in the development of phytoplankton was observed, which reached its maximumca. 100 km before the river flows into its artificial sedimentation area. The development of rotifer populations was slightly retarded as compared with that of phytoplankton and highest population densities were only reached in the sedimentation area. Crustaceans developed in significant numbers, not until the river water had entered the sedimentation area. Development of zooplankton coincided here with a strong decrease in the density of phytoplankton. A similar trend in plankton development was observed in the River Meuse, although in this river the highest densities of phyto and zooplankton already occurred in its middle reaches. The differences in the timing of plankton growth in the two rivers are probably caused by differences in flow regime between both rivers. The River Rhine, which is fed by rainwater and melting of glaciers in the Alps, has a relatively constant discharge. On the other hand, the low discharge of the rain-fed River Meuse combined with an increased residence time of the water as a consequence of large numbers of weirs, allows a full cycle of plankton development long before its discharge into the sea. This phenomenon was also reflected in the silicate cycle in the Meuse, where the consumption by planktonic diatoms and the regeneration of silicate of deposits seem to be important. In contrast, in the main branches of the River Rhine only the effects of silicate consumption were detectable.     

One decade of benthic macroinvertebrate biomonitoring in the River Meuse

The benthic macroinvertebrate community of the River Meuse was studied at four locations in Belgium and The Netherlands from 1983–1993. Macroinvertebrates were collected from artificial substrates, placed near the river bottom from June–August. The results were analyzed using autecological information. At an upstream sampling site in the Belgian Ardennes (Hastière) the most diverse fauna was found. One quarter of the taxa found at this site are known to be confined to running waters and none of the dominant or subdominant taxa could be characterized as a specialized riverine taxon. These findings indicate an impoverished riverine fauna. This was especially the case in years with low discharge, indicating a delicately balanced ecosystem, easily susceptible to disturbances in the environment. Further downstream, in the heavily industrialized region of Liège (Hermalle) and just across the Belgian-Dutch border (Borgharen) the macroinvertebrate fauna consisted of taxa tolerating severe organic pollution. The ecosystem at these sites is on the brink of collapse. Recent results indicate an improvement of this situation at Borgharen. The macroinvertebrate fauna in the lowland reach (Keizersveer) indicates ecological recovery. It is concluded that abatement of the severe pollution must be the first step for an ecological rehabilitation of the degraded ecosystem and for the safe drinking water supply for millions of people in Belgium and The Netherlands.This article is dedicated to my friend and colleague Nicole Frantzen, who died in 1992 at the age of 27.     

Development and ecological importance of phytoplankton in a large lowland river (River Meuse,Belgium)

The ecological importance of the River Meuse phytoplankton with regard to carbon and nutrient transport has been examined in two reaches of the Belgian course of the river.Field measurements of total particulate organic carbon (POC), particulate organic nitrogen (PON) and particulate phosphorus (PP) show that the large autochtonous production of organic matter strongly affects the carbon and nutrient budget of the aquatic system. During the growing season, phytoplankton accounts for nearly 60% of the POC and dominates the PON. Calculations of the carbon and oxygen budget in the upper reach of the Belgian Meuse demonstrates that the ecosystem is autotrophic, i.e. that autochtonous FPOM (fine particulate organic matter) production is the major carbon input. This suggests that in large lowland rivers, primary production (P) may exceed community respiration (R), i.e. P:R>1, whereas they are assumed to be heterotrophic (P:R<1) in the River Continuum concept.The question of maintenance of phytoplankton in turbid mixed water columns is also addressed, and the case of the River Meuse is treated on the basis of studies of photosynthesis and respiration (ETS measurements). The results suggest that the potamoplankton may show some low-light acclimation, through an increase of chlorophyll a relative to biomass, when it comes to deep downstream reaches, and that algal respiration rate may be reduced. A simulation of the longitudinal development of the algal biomass shows the different phases of algal growth and decline along the river and brings support to the importation hypothesis for explaining maintenance of potamoplankton in the downstream reaches.     

Are zebra mussels restricted by toxicant levels in the River Meuse? a review

This paper questions if chemical barriers prevent the occurrence of benthic invertebrates in the river Meuse. To this purpose an ecotoxicological analysis is presented, using published observations on the zebra musselDreissena polymorpha, a tolerant species that maintains populations in the river. Zebra mussels collected or exposed at the Belgian-Dutch border contained high levels of several groups of toxicants, and mixture toxicity is likely to occur. A recently developed bio-assay, using the filtration rate of the zebra mussel, demonstrated strong inhibitory effects of water from the river Meuse. To determine which (combination of) toxicants cause such effects, laboratory experiments with toxicant mixtures were carried out. It was demonstrated that in a mixture of five metals (Cu, Zn, Ni, Cd, Pb) the metals contributed to the toxicity of the mixture below the No Observed Effect Concentrations (NOEC) for these metals when tested individually. The average concentration of Cu, Zn and possibly Pb in Meuse water exceed the NOEC values for filtration rate. Thus, it seems likely that joint effects of different (groups of) toxicants in the river Meuse cause the overall toxicity of the water for the zebra mussel, explaining the marginal populations at the test site, while other river species are absent there. It is suggested that toxicants in the river Meuse restrict the recolonisation of the river by invertebrate species more sensitive than the zebra mussel. This will be studied using invertebrates of different sensitivities (molluscs, arthropods), enabling the assessment of toxicity at different stages of water quality improvement of the river Meuse.     

松花江哈尔滨段浮游植物群落格局及其与环境因子的相关性

松花江是黑龙江在我国境内的最大支流,流经吉林、黑龙江两省。松花江哈尔滨段是哈尔滨市工农业生产与生活用水的重要水资源,近年来由于人类活动的频繁影响,水体质量有所下降。鉴于此,于2018年春、夏、秋三季(4月、8月和10月),在松花江哈尔滨段设置14个采样点,对电导率、总氮和总磷等理化指标进行测定,同时对其浮游植物群落结构及环境因子相关性进行研究。利用群落相似性分析(ANOSIM)探讨浮游植物群落时空分布差异;通过SIMPER分析筛选出影响群落时空格局的关键物种。基于优势种和10项理化指标的冗余分析(RDA)揭示了浮游植物群落生态分布特征并对关键影响因子进行识别。研究结果表明,调查期间共鉴定浮游植物136种,其中优势种15种,群落结构主要以硅藻门和绿藻门的物种构成。松花江哈尔滨段浮游植物群落结构与过去10余年间相比较为稳定。ANOSIM和SIMPER分析表明,季节变化和人为活动干扰对浮游植物群落时空格局具有一定驱动作用,蓝藻门物种的空间分布活动干扰影响明显。RDA分析表明,浮游植物分布特征与水环境时空异质性关系密切,驱动松花江哈尔滨段浮游植物群落时空分布的主要水环境因子为电导率、总磷、浊度和pH。     

Plankton dynamics along a 180 km reach of the Saint Lawrence River from its headwaters in Lake Ontario

We test the hypothesis that phytoplankton biomass decrease upon entry into the St. Lawrence River from headwaters in Lake Ontario is attributable to a range of causal factors including, decreased photosynthetic fitness due to turbulence, cell loss due to increased flocculation and subsequent sedimentation, decreases in nutrients, and loss due to grazing. In order to test this, changes in phytoplankton and crustacean zooplankton concentrations were examined during four transects along the river, from 8 km offshore in Lake Ontario to the hydroelectric power dam 180 km downstream. Both phytoplankton biomass, measured as chlorophyll-a, and zooplankton decreased markedly upon entry of lake water into the river. Phytoplankton community composition and size changed little over the river reach and tended to reflect that in Lake Ontario. Total phosphorus increased with transit of river water downstream despite low tributary inputs of water into this reach of river. Light availability was high, photosynthetic efficiency suggested that phytoplankton were not physiologically stressed during transit in turbulent waters, and there was no direct evidence of flocculation causing sedimentation of phytoplankton. Grazing by the benthic community (filtering insect larvae and dreissenid mussels) is inferred to be a dominant biological factor as is the geomorphology in this reach of the river, which includes large littoral areas, shoals, and reaches of high water velocity that can increase particle contact in the water column with benthic grazers. The findings of this study have a bearing on understanding how changing water levels in a regulated river might alter areas of benthic grazing.     

The zooplankton of the lower river Meuse, Belgium: seasonal changes and impact of industrial and municipal discharges

Between 1991 and 1993, samples were collected upstream and downstream of the industrial basin and urban centre of Liège. Rotifers and crustaceans (cladocerans and copepods) were identified and counted. Their population dynamics were related to physical and chemical factors (temperature, oxygen, ammonium, nitrates, nitrites, phosphates) and to phytoplankton biomass. The zooplankton was dominated by rotifers; crustaceans (cladocerans and copepods) were less abundant. There was a succession of groups and species, some thriving in the spring and others in summer or autumn. The dominant rotifer species were Brachionus calyciflorus Pallas, Brachionus angularis Gosse, Keratella cochlearis (Gosse) and Synchaeta spp.; B. calyciflorus and B. angularis are spring species. K. cochlearis was present between May and November. Crustacean biomass was important in summer and autumn, but the faunal spectrum and biomass also varied with sampling location. Low spring and summer discharges allowed the phytoplankton to develop significantly. The zooplankton development followed a similar pattern. During low flow, when plankton populations become established, some declines in phytoplankton could only be explained by sedimentation and grazing pressure by zooplankton. Although these factors provided a good explanation of the longitudinal variation, some local conditions (e.g. oxygen deficit, high level of phosphate) also induced changes (e.g. industrial and municipal waste water discharge).     

Using XAD resins to study the effects of reduced organic micropollutant concentrations in River Rhine and Meuse water on phytoplankton growth

Laboratory incubation experiments were used to study the effect of reduced concentrations of organic micropollutants in water from the rivers Rhine and Meuse on the specific growth rate of the river phytoplankton community. Before incubation, part of the water sampled was treated with XAD-4 and XAD-8 resins to absorb dissolved organic compounds. Four dilutions were made by mixing untreated water with XAD-treated water in the ratios 100:0 (control), 70:30, 40:60 and 0:100. The phytoplankton specific growth rate increased significantly with the increased fraction treated with XAD in all but one incubation experiment. In these experiments, the specific growth rate was on average 9% higher in the fraction in which 100% was treated with XAD than in the controls. In the Rhine and Meuse river water, phytoplankton growth seemed to be inhibited by organic compounds. This inhibition was ascribed to the presence of dissolved organic micropollutants. Removing organic micropollutants using XAD resins to study the toxic effects of these compounds on field phytoplankton communities can be concluded to be a promising tool for risk assessment of micropollutants but needs to be supported by additional methodological research.     

Phytoplankton community response to hydrological variations in oxbow lakes with different levels of connection to a tropical river

Changes to the structure of the phytoplankton community and to the physical and chemical variables of the water were investigated in oxbow lakes with different levels of connection to a tropical river and subject to annual hydrological pulse variations. The selected lentic environments are located at the mouth region of the main tributary in a reservoir built for water storage and electric power generation. The temporal variation of phytoplankton in the studied lentic environments can be attributed mainly to the hydrological level of the river. A similar variation pattern of the ecological attributes was observed in the structure of the phytoplankton community in the connected lakes and Paranapanema River, evidencing the high degree of association that the lacustrine systems maintain with the river. The highest values of richness and diversity for connected environments were observed at the end of the emptying period and in the drought. However, considering the isolated lake, the highest values of these attributes were recorded during the flooding period.     

上海市内不同水质的河道春季浮游细菌群落结构分析

2008年4月采集上海市城郊结合地区的4条河道午潮港、横港、朝阳河和曹杨环浜的水样, 测定水样的相关理化数据, 计算水环境质量综合指数; 提取水样中浮游细菌的总DNA, 进行变性梯度凝胶电泳(DGGE)分析; 并对水体中的可培养浮游细菌的数量进行测定。结果表明, 污染严重的河道浮游细菌的生物量较高, 而多样性明显偏低, 河道不同采样点的微环境影响浮游细菌的组成, 河道中浮游细菌的组成与生物量的变化与水环境质量密切相关。     

The structure of phytoplankton and physicochemical characteristics of the Kolyma River (Northeastern Siberia) in summer

The first results of a study of the phytoplankton and chemical composition of water of the entire Kolyma River are reported. The study revealed the spatial structure of the phytoplankton communities of river sections with various hydrologic conditions. The high diversity of the Kolyma River phytoplankton was established. It was acknowledged that the elemental composition and physical characteristics of the water are determined by environmental factors, mainly permafrost soils. Water quality was estimated by the physicochemical characteristics, sabrobic algae, and phytoplankton biomass of the water.     

安康水库表层浮游藻类群落结构及其与环境因子的关系

为了解安康水库表层水质与浮游藻类群落现状以及浮游藻类群落结构与环境因子之间的关系,于2012年1—12月对水库表层水进行每月一次的采样,分别对浮游藻类分布状况和理化因子进行分析.利用Shannon指数(H)和Pielou均匀度指数(J)分析浮游藻类群落特征,根据理化指标和综合营养状态指数(TSI)评价水体营养状态.共检测到浮游藻类7门110属,丰度为0.11×10⁴～2.08×10⁴ cells·L^-1.藻类组成、污染指示种分布情况、多样性分析和TSI显示安康水库表层属于中污染水质,处于中营养状态.此外,高密度投饵水产养殖及生活污水直接排放入库对水质产生负面影响,支流岚河的水质状况也较差.典范对应分析(CCA)结果显示,测定的8个环境因子中,不同季节影响浮游藻类群落组成及分布的环境因子不同,且氮是影响浮游藻类组成的主要营养盐因子.理化指标分析显示,安康水库表层水大体满足Ⅱ类水标准,水质良好.但部分采样点的总氮出现差于Ⅱ类水标准的现象,表明安康水库水质有变差的趋势.     

Impact of hydrology on the chemistry and phytoplankton development in floodplain lakes along the Lower Rhine and Meuse

The impact of hydrology (floods, seepage) on the chemistry of water and sediment in floodplain lakes was studied by a multivariate analysis (PCA) of physico-chemical parameters in 100 lakes within the floodplains in the lower reaches of the rivers Rhine and Meuse. In addition, seasonal fluctuations in water chemistry and chlorophyll-a development in the main channel of the Lower Rhine and five floodplain lakes along a flooding gradient were monitored. The species composition of the summer phytoplankton in these lakes was studied as well.At present very high levels of chloride, sodium, sulphate, phosphate and nitrate are found in the main channels of the rivers Rhine and Meuse, resulting from industrial, agricultural and domestic sewage. Together with the actual concentrations of major ions and nutrients in the main channel, the annual flood duration determines the physico-chemistry of the floodplain lakes. The river water influences the water chemistry of these lakes not only via inundations, but also via seepage. A comparison of recent and historical chemical data shows an increase over the years in the levels of chloride both in the main channel of the Lower Rhine and in seepage lakes along this river. Levels of alkalinity in floodplain lakes showed an inverse relationship with annual flood duration, because sulphur retention and alkalinization occurred in seepage waters and rarely-flooded lakes. The input of large quantities of nutrients (N, P) from the main channel has resulted, especially in frequently flooded lakes, in an increase in algal biomass and a shift in phytoplankton composition from a diatom dominated community towards a community dominated by chlorophytes and cyanobacteria.     

香溪河水质空间分布特性研究

运用聚类分析和主成分分析对香溪河19个样点水质的理化特性进行研究,聚类分析表明,根据各采样点之间水质组分的相似性可将香溪河大致分为3个河段,分别属于不同的亚流域,各亚流域问的特征差异显著,对各河段水质的主成分分析表明,上述3河段的主要水质信息差异很大,第1河段(在河流上游)水质的信息主要体现为总碱度和硬度,第2河段(河流中游)主要体现为可溶性磷酸盐、总磷和氯离子,第3河段(河流下游)则为pH、亚硝酸盐氮、总氮和COD,文中结合香溪河流域地理环境背景,探讨了香溪河水质空间分布格局的成因,为分析流域水质状况及成因提供了一条简单有效的途径。     

Monitoring of algae in Dutch rivers: does it meet its goals?

Observations on phytoplankton in the lower reaches of the rivers Rhine and Meuse were carried out in the framework of the national monitoring programme of the main water systems in The Netherlands. Seasonal changes in density and species composition were analysed to detect the major variables and to discuss whether this monitoring meets its goals. Phytoplankton reached peak densities of 140 and 65 μg L^-1 Chl a in the Rhine and Meuse, respectively. Overall density was correlated with seasonal variation in water discharge rather than with nutrient concentrations, that are high in both rivers. The position of sampling sites in relation to the downstream development of the plankton was very important, especially in the regulated Meuse. Despite hydrographic and chemical differences between the two rivers, many species, predominantly diatoms and green algae, were shared. The occurrence of the diatom Skeletonema subsalsum in the Rhine and the green algae Neodesmus danubialis, Micractinium pusillum and Pseudotetrastrum punctatum in the Meuse (1992, but not 1996) was interpreted as a feature related, respectively, to the high salinity of the Rhine and specific riverine conditions of the Meuse. In general the potamoplankton was characterised as an opportunistic assemblage exploiting the high nutrient contents and disturbed hydrography of both rivers. Nevertheless, the phytoplankton contributed to the characterisation of the present river communities, ones that differ rather strongly in historic records. Although year to year variation in phytoplankton density is mainly related to variation in water discharge, phytoplankton biomass did increase as a result of eutrophication over a period of decades. The widespread nature of many of today's potamoplankton species in the two rivers render these organisms less useful for indicating short-term changes in water quality. However the importance of phytoplankton in trophic relationships merits its inclusion in the monitoring of riverine ecosystems. This revised version was published online in June 2006 with corrections to the Cover Date.     

英德西南旅游区水源河流水质与浮游生物群落特征

通过野外采样及实验室分析,调查了英德市西南旅游区水源河流上、中、下游的水质状况及浮游生物的多样性。结果表明,(1)英德市西南旅游区上、中、下游河段亚硝酸盐、氨、硝酸盐、总氮、镉、铅、铬、镍、铜、锌的含量低于国家标准,铝、钙、钴、铁、钼、硒的含量超过国家标准,而总磷、砷、镁、锰的含量在部份区段已超过国家标准;(2)浮游动物种类的个体数在整条河中以桡足类最多,上游通天岩中浮游动物个体数最多,下游碧落湖浮游动物多样性最低;此次调查总共发现68种浮游植物,其中主要集中在下游碧落湖,在中游碧落洞绿藻、蓝藻、硅藻总个体数最低,上游通天岩绿藻及甲藻总个体数最多,而下游碧落湖的蓝藻和硅藻的总个体数最多;(3)总氮、总磷、铅、钙、浮游动物Shannon-Weaver指数与浮游植物Shannon-Weaver指数从上游向下游增加,而锰与镍含量则成相反趋势。文中探讨了水中元素之间、元素与生物之间、浮游动植物之间的相互关系。本文对风景区水源的生态环境的研究风景区旅游资源开发与管理提供参考。     

Order and disorder in the river continuum: the contribution of continuity and connectivity to floodplain meadow biodiversity

Aim Aspects of connectivity and continuity operating in the River Meuse were analysed for their contribution to the biodiversity of the floodplain. From this analysis of the diversity and composition of the meadow communities, we aimed to derive effective biodiversity conservation strategies. Location The River Meuse is one of the larger rivers in the European Western Plains ecoregion. The alluvial plains of the river have a long history of cultivation, and for these plains the floodplain meadow vegetation is a highly appreciated and valuable nature conservation asset. Method We sampled floodplain meadows from 400 km of the six geomorphic reaches of the middle to lower course of the River Meuse. For each, 50 vascular plant relevés were recorded, representing the spectrum of floodplain meadow communities of that reach. Beta diversity was calculated to quantify similarity in species pools between the reaches. A dissimilarity formula was used to determine the turnover between the reaches, and these dissimilarities were compared with a Mantel test to detect whether species composition of the floodplain meadows exhibited connectivity and continuity between the reaches. Species richness for the floodplain vegetation data of the reaches was compared with data for riparian invertebrate communities. The vegetation data sets were ordinated using detrended correspondence analysis (DCA) to reveal patterns in the floodplain meadow species composition, and the DCA axes were related to plant functional groups and population strategies. The axis scores of the species and plots were linked to river and plant species traits. Results We did not observe an overall continuity trend in similarity, nor one in diversity in a downstream direction. Lateral connectivity was highlighted by the dissimilarity between the reaches and in the influxes of species from adjacent ecoregions. The DCA ordination showed statistically significant separations between reaches and between the plant functional groups. The second DCA axis related to the longitudinal gradient of the river, whereas the first axis showed stronger correlations with river and plant species traits. We termed this axis the ‘disorder axis’. Plant invaders and avoiders are located at the left side of the disorder axis, whereas the true river‐adapted categories of resisters and endurers are at the right extremity. Contributions to the disorder were identified in terms of connectivity with adjacent ecoregions and the physical disturbance regime of natural and anthropogenic perturbations, resulting in community changes between the reaches. Main conclusions We concluded that a single overall strategy for biodiversity conservation of the river or its floodplains is not feasible. Strategies can, however, be derived for separate river reaches based on functional groups in the communities, the disorder characteristics of the reach, and the influence of surrounding ecoregions.     

Temporal fluctuations of the phytoplankton community of the Baía River, in the upper Paraná River floodplain, Mato Grosso do Sul, Brazil

Subsurface samples were taken monthly (March/1993 through February/1994) in the Baía River, a tributary of the right bank of the Paraná River (22° 43 S and 53° 17 W). We analyzed temporal changes in the phytoplankton community in relation to density, biomass, richness and species diversity, equitability, size structure, and dominance. We related these to regional climate and hydrology, and to the physical and chemical variables of the water column. We determined 119 taxa, wich were numerically dominated by the class Chlorophyceae with 37 taxa. The classes Cyanophyceae and Bacillariophyceae contributed the most abundant biomass and were responsible for the two peaks that were observed. The high water period was generally characterized by lower phytoplankton biomass, higher richness and species diversity, and higher density of nannoplanktonic species such as Monoraphidium tortile in March, and Cryptomonas brasiliensis in January. During low water, however, the highest values of phytoplankton biomass were recorded, with heterocytous cyanophytes dominating during the phase of greater stability of the water column, and filamentous diatoms during periods of mixing turbulence. The fluctuations observed indicate that the hydrosedimentological regime of the Paraná River, together with the climatological factors, constitute the main forcing functions acting on the Baía River phytoplankton.     

Phytoplankton assemblages across the marine to low-salinity transition zone in a blackwater dominated estuary

Along the salinity gradient between the freshwater reach ofthe Suwannee River and the marine waters of the Gulf of Mexico,the spatial and temporal composition and abundance of phytoplanktonwas examined in relation to physical and chemical water-columnvariables (i.e. salinity, nutrient and light availability andtemperature). The goal of this study was to explore the applicabilityof the concepts of ecotone and ecocline to the distributionof phytoplankton. This was undertaken in a relatively open estuarydominated by a tannin-colored river, the Suwannee River, alongthe central northwest coast of Florida, USA. Hierarchical clusteranalysis and non-metric multi-dimensional scaling (MDS) wasused to define biologically distinct regions within the estuarybased on the composition of phytoplankton. Variations in phytoplanktonassemblages in relation to environmental variables were investigatedusing a canonical correspondence analysis. The results indicatethat in addition to the alignment of phytoplankton compositionalong a salinity gradient, several key environmental factorsinfluence phytoplankton assemblages within hydrologically distinctregions of the estuary supporting an ecocline view. However,the distribution of dinoflagellate assemblages were in contrastto the more temporally fluid spatial distributions of cyanobacteriaand diatoms in the Suwannee River estuary. The spatially definednature of the dinoflagellate assemblages was more characteristicof an ecotone rather than an ecocline.