Large-scale spatial distribution of virioplankton in the Adriatic Sea: testing the trophic state control hypothesis

Little is known concerning environmental factors that may control the distribution of virioplankton on large spatial scales. In previous studies workers reported high viral levels in eutrophic systems and suggested that the trophic state is a possible driving force controlling the spatial distribution of viruses. In order to test this hypothesis, we determined the distribution of viral abundance and bacterial abundance and the virus-to-bacterium ratio in a wide area covering the entire Adriatic basin (Mediterranean Sea). To gather additional information on factors controlling viral distribution on a large scale, functional microbial parameters (exoenzymatic activities, bacterial production and turnover) were related to trophic gradients. At large spatial scales, viral distribution was independent of autotrophic biomass and all other environmental parameters. We concluded that in contrast to what was previously hypothesized, changing trophic conditions do not directly affect virioplankton distribution. Since virus distribution was coupled with bacterial turnover times, our results suggest that viral abundance depends on bacterial activity and on host cell abundance.     

Dynamic of Virioplankton Abundance and Its Environmental Control in the Charente Estuary (France)

The Charente River provides nutrient- and virus-rich freshwater input to the Marennes Oléron Basin, the largest oyster-producing region in Europe. To evaluate virioplankton distribution in the Charente Estuary and identify which environmental variables control dynamic of virioplankton abundance, five stations defined by a salinity gradient (0–0.5, 0.6–5, 13–17, 20–24, and higher than 30 PSU) were surveyed over a year. Viral abundance was related to bacterioplankton abundance and activities, photosynthetic pigments, nutrient concentration, and physical parameters (temperature and salinity). On a spatial scale, virus displayed a decreasing pattern seaward with abundance ranging over the sampling period from 1.4 × 10⁷ to 20.8 × 10⁷ viruses mL⁻¹ making virioplankton the most abundant component of planktonic microorganisms in the Charente Estuary. A good correlation was found between viral and bacterial abundance (r_s = 0.85). Furthermore, bacterial abundance was the most important predictor of viral abundance explaining alone between 66% (winter) and 76% (summer) of viral variability. However, no relation existed between viral abundance and chlorophyll a. Temporal variations in viral distributions were mainly controlled by temperature through the control of bacterial dynamics. Spatial variations of viral abundance were influenced by hydrodynamic conditions especially during the winter season where virioplankton distribution was entirely driven by mixing processes.     

流式细胞仪检测纳帕海高原湿地浮游病毒和浮游细菌丰度

【背景】浮游病毒是水体微生物群落中重要的组成成分,深入研究浮游病毒的时空分布有助于更好地保护和开发当地的微生物资源。【目的】对采集到的纳帕海高原湿地水样中的浮游病毒和浮游细菌进行计数,揭示纳帕海高原湿地浮游病毒的分布规律。【方法】采用流式细胞仪检测2013年12月和2014年9月纳帕海高原湿地7个水样的浮游病毒与浮游细菌丰度,并对影响浮游病毒丰度的因素,如细菌丰度、叶绿素a含量以及其他环境因子进行了相关性分析。【结果】季节分布上,雨季浮游病毒和浮游细菌丰度高于旱季;水平分布上,原水样品的浮游病毒高于湿地水和淤泥水。旱季水样的浮游病毒丰度受到细菌丰度及叶绿素a浓度的影响较大;雨季水样的浮游病毒丰度受到水体的p H值和温度的影响较大。【结论】纳帕海高原湿地的浮游病毒和浮游细菌是比较活跃的。浮游病毒丰度在不同季节、不同采样点受到细菌丰度和叶绿素a浓度等因素的不同影响。在旱季噬菌体而非噬藻体或浮游植物病毒是纳帕海高原湿地中浮游病毒的优势种群。     

Viral abundance and a high proportion of lysogens suggest that viruses are important members of the microbial community in the gulf of trieste

Epifluorescence microscopy and transmission electron microscopy were applied to study virioplankton community in the Gulf of Trieste (northern Adriatic Sea). The total viral abundance was in a range between 2.5 x 10(9)/L and 2.9 x 10(10)/L and was positively correlated with trophic status of the environment. Viruslike particles were significantly correlated with bacterial abundance in all samples studied. Correlations with other physicochemical or biological parameters were not significant. The data suggest that, because of the substantial fraction of tailed viruses present (26%), bacteriophages are an important component of the virioplankton community in the Gulf of Trieste. The abundance of viruslike particles in the seawater changed at hour intervals in a range from 1.3 x 10(9)/L to 5.1 x 10(9)/L. A significant fraction (71%) of the bacterial isolates was inducible in vitro by mitomycin C, and a high occurrence (51%) of lysogenic isolates with more than one phage morphotype present in the lysate was detected. The presence of lysogenic bacteria in the seawater was confirmed in situ with a mitomycin C induction experiment on the natural bacterial population. Results suggest that virioplankton is an abundant component of the microbial community in the Gulf of Trieste.     

Viral Abundance and a High Proportion of Lysogens Suggest That Viruses Are Important Members of the Microbial Community in the Gulf of Trieste

Epifluorescence microscopy and transmission electron microscopy were applied to study virioplankton community in the Gulf of Trieste (northern Adriatic Sea). The total viral abundance was in a range between 2.5 × 10⁹/L and 2.9 × 10¹⁰/L and was positively correlated with trophic status of the environment. Viruslike particles were significantly correlated with bacterial abundance in all samples studied. Correlations with other physicochemical or biological parameters were not significant. The data suggest that, because of the substantial fraction of tailed viruses present (26%), bacteriophages are an important component of the virioplankton community in the Gulf of Trieste. The abundance of viruslike particles in the seawater changed at hour intervals in a range from 1.3 × 10⁹/L to 5.1 × 10⁹/L. A significant fraction (71%) of the bacterial isolates was inducible in vitro by mitomycin C, and a high occurrence (51%) of lysogenic isolates with more than one phage morphotype present in the lysate was detected. The presence of lysogenic bacteria in the seawater was confirmed in situ with a mitomycin C induction experiment on the natural bacterial population. Results suggest that virioplankton is an abundant component of the microbial community in the Gulf of Trieste.     

纳帕海高原湿地浮游病毒丰度及与环境因子相关性研究

目的利用EFM技术对纳帕海湿地不同季节7个水样样品中的浮游病毒丰度开展了调查,并对影响浮游病毒丰度的环境因子进行了相关性分析。方法元素检测,叶绿素a检测,荧光显微镜检测浮游细菌及浮游病毒的丰度。结果从生态分布上来看旱季和雨季浮游病毒丰度的平均值分别为3.63×10~6/mL和3.71×10~7/mL,浮游病毒丰度季节性变化不显著。从对影响浮游病毒丰度的环境因子相关性分析来看,旱季浮游病毒与Chl-a具有弱负相关性(r=0.49,P0.05);而雨季浮游病毒丰度与Chl-a具有显著正相关性(r=0.37,P0.05),表明雨季纳帕海湿地浮游病毒丰度受Chl-a含量的影响较大。旱季浮游病毒丰度与温度显著负相关,虽然在雨季浮游病毒丰度与温度也呈负相关,但并不显著;在雨季,水体pH变化较明显,而浮游病毒丰度与水体pH呈显著负相关,表明湿地的来水主要在雨季,而离子组成变化大。虽然在雨季和旱季,从多点采样的平均值来看浮游病毒丰度并无明显差异,但从一些局部的采样点来看,雨季的浮游病毒丰度显著高于旱季,说明在纳帕海区域浮游病毒的分布并不平均,因采样点环境的不同会产生明显的区别。结论对纳帕海高原湿地浮游病毒生态分布的调查,是对该地域生态学研究的补充,使得这一独特地理环境的微生物研究更加系统化。     

Communities of bacteria and viruses in waters of the Gulf of Ob and Taz Estuary

The study of the most abundant components in freshwater plankton in the Gulf of Ob and Taz Estuary in the summer–autumn season has demonstrated that the abundance and biomass of bacteria are stable and typical for mesotrophic waters during active microalgae vegetation. The abundance of viral particles varies in the range which is reported for unproductive or medium-productive water bodies. The environmental factors affecting affect the development and patterns of bacterio- and virioplankton distribution are considered.     

Spatial distribution and morphologic diversity of virioplankton in Lake Donghu, China

The spatial distribution and morphological diversity of virioplankton were determined in Lake Donghu which contains three trophic regions: hypertrophic, eutrophic and mesotrophic region. Virioplankton abundance measured by transmission electron microscope (TEM) ranged from 7.7 × 10⁸ to 3.0 × 10⁹ ml^–1, being among the highest observed in any natural aquatic system examined so far. The spatial distribution of virioplankton was correlated significantly with chlorophyll a concentration (r = 0.847; P < 0.01) at the sampling sites in Lake Donghu. 76 morphotypes were observed. Most morphotypes have tails, belonging to Siphoviridae, Myoviridae and Podoviridae. The majority of tailed phages in the lake were Myoviridae. Morphotypes which were rarely reported, such as prolate-headed virus-like particles, lemon-shaped virus-like particle, and viruses resembling Tectiviridae and Corticoviridae were all observed in the lake. It is concluded that the high viral abundance might be associated with high density of phytoplankton including algae and cyanobacteria. There was high viral diversity in this eutrophic shallow lake. In addition, cyanophage represented an important fraction of the virioplankton community in Lake Donghu.     

Analytical evidence for scale-invariance in the shape of species abundance distributions

The distribution of species abundances within an ecological community provides a window into ecological processes and has important applications in conservation biology as an indicator of disturbance. Previous work indicates that species abundance distributions might be independent of the scales at which they are measured which has implications for data interpretation. Here we formulate an analytically tractable model for the species abundance distribution at different scales and discuss the biological relevance of its assumptions. Our model shows that as scale increases, the shape of the species abundance distribution converges to a particular shape given uniquely by the Jaccard index of spatial species turnover and by a parameter for the spatial correlation of abundances. Our model indicates that the shape of the species abundance distribution is taxon specific but does not depend on sample area, provided this area is large. We conclude that the species abundance distribution may indeed serve as an indicator of disturbances affecting species spatial turnover and that the assumption of conservation of energy in ecosystems, which is part of the Maximum Entropy approach, should be re-evaluated.     

Relative importance of spatial processes and environmental factors in shaping alpine meadow communities

Aims Spatial processes and environmental control are the two distinct, yet not mutually exclusive forces of community structuring, but the relative importance of these factors is controversial due to the species-specific dispersal ability, sensitivity towards environmental variables, organism's abundance and the effect of spatial scale. In the present paper, we explored spatial versus environmental control in shaping community composition (i.e. β-diversity) and species turnover (i.e. change of β-diversity) at an alpine meadow along a slope aspect gradient on the Qinghai–Tibetan Plateau at different spatial scales of sampling (quadrats and plots), by taking account of seed dispersal mode and abundance.Methods We examined the relative importance of spatial processes and environmental factors using all species and four additional subsets of selected species. Moreover, we attempted to explore the effect of scale (quadrat refers to scale of ~0.3 m and plot of ~8 m) on their counter balance. The data were analyzed both by variation partitioning and multiple regressions on distance matrices. The spatial structure was modelled using Moran's eigenvector maps (MEM).Important findings Both spatial processes and environmental factors were important determinants of the community composition and species turnover. The community composition in the alpine meadow was controlled by spatially structured environment (17.6%), space independent of environment (18.0%) and a negligible effect of environment independent of space (4.4%) at the scale of quadrats. These three components contributed 21.8, 9.9 and 13.9%, respectively, at the scale of plots. The balance between the forces at different spatial scales drove community structures along the slope aspect gradient. The importance of environmental factors on β-diversity at alpine meadow increased with scale while that of spatial processes decreased or kept steady, depending on dispersal mode and abundance of species comprising the subset. But the 'pure' effect of spatial processes on species turnover increased with scale while that of environmental factors decreased. This discrepancy highlights that β-diversity and species turnover were determined jointly by spatial processes and environmental factors. We also found that the relative roles of these processes vary with spatial scale. These results underline the importance of considering species-specific dispersal ability and abundance of species comprising the communities and the appropriate spatial scale in understanding the mechanisms of community assembly.     

模拟水流扰动对香溪河库湾浮游病毒-宿主动态的影响

【背景】浮游病毒是淡水生态系统的重要组成部分,在调节浮游细菌和藻类群落结构及调控系统物质循环过程中起着重要的作用。水库具有不同于湖泊的水动力过程,产生的扰动可能影响浮游病毒的调控功能。【目的】研究水力扰动对浮游病毒-宿主动态的影响,为阐释水库生境下浮游病毒生态功能提供理论依据。【方法】以香溪河库湾原水为材料,模拟不同流速扰动对病毒-宿主动态的影响;通过病毒丰度、宿主丰度、宿主裂解率、宿主溶源诱导率等参数的变化反映这种动态变化过程,并分析其与环境因子间的关系。【结果】0.05 m/s和0.10 m/s的流速扰动强度对浮游植物和浮游细菌生长有显著促进作用,但扰动对浮游病毒丰度的影响不显著;扰动能促进病毒介导的浮游植物和细菌裂解率上升,而且0.05 m/s扰动强度的促进作用大于0.10 m/s;同时,扰动显著降低了浮游植物溶源诱导率,但引起浮游细菌溶源诱导率的显著上升(P<0.05)。【结论】模拟扰动对浮游病毒-宿主动态过程产生了显著的影响,表明水库浮游病毒维持种群延续的生态策略可能与湖泊浮游病毒存在差异。     

南海北部海域春季浮游细菌和病毒空间分布及其影响因素

应用流式细胞检测技术测定了2014年春季南海北部海域浮游细菌和病毒丰度,研究了其水平和垂直分布特征并对其与环境因子的相关性进行了分析。结果表明,调查海区浮游细菌和病毒丰度分别介于1.28×10~4—9.96×10~5个/m L和4.69×10~5—5.39×10~7个/m L之间,二者丰度随水深的增加基本呈现逐渐下降的趋势,而水平分布趋势不明显。浮游细菌和病毒丰度与温度、p H和溶解氧显著正相关,与水深、盐度、活性磷酸盐、硅酸盐、硝酸盐和总氮则呈显著负相关关系(P0.01),说明该海域细菌和病毒数量受到上述环境因子的共同调控。分析浮游细菌和病毒的相互关系发现,VBR(Virus to bacteria ratio)平均32.23,最小值位于S11站位25m层,最大值则位于S7站位75m层,分别为4.80和264.63,VBR值小于100的站位占到调查站位总数的95.6%。VBR值除与细菌呈显著负相关关系外(P0.01),与其它环境因子相关性不明显(P0.05),说明该海区细菌是病毒的主要寄主,病毒可能主要是以噬菌体的状态存在。     

Loss of Estuarine Bacteria by Viral Infection and Predation in Microcosm Conditions

The bacterioplankton density in Ria de Aveiro, a shallow estuarine ecosystem, varied in the broad range of 1.9-10.6 109 cells L-1. The range of values was about 2 times higher in brackish water than in marine water. At high tide bacterial abundance was 2-3 times lower than at low tide. The overall variation in virioplankton was in the range of 2.4-25.0 1010 particles L-1. Brackish water was about 2 times richer in viral particles than the marine water. Near low tide the virioplankton was 2-3 times higher that at high tide. Viral density followed the pattern of bacterial abundance (it explained 40% of virioplankton variation). The viruses to bacterium ratio varied, throughout tidal cycles, by a factor of about 10 establishing the range 4.7-55.6 (average 17.6). This ratio was rather similar in the two estuarine zones. We compared the effects of infection and predation on the control of bacterioplankton size in the two zones of the estuary. The approach to this question was conducted in experimental microcosms, set up in six combinations of plankton variables affecting the presence/absence of predators, virus-to-bacterium ratio (10-fold increase), virus-to-bacterium distance (2.2-fold increase), and bacterial growth rate. The results showed that predation was similar, in a percent basis, in marine (69%) and brackish water (73%). Viral infection was, however, higher in brackish water (59%) than in the marine water (36%). We conclude that the bacterioplankton along the salinity gradient evolves under biological pressures that are in different balance in the marine and brackish water zones. The effect of viral lysis on bacterial communities with enhanced growth (after yeast extract addition) was masked even when the initial ratio was 10-fold greater than in the natural samples. The high density of the virioplankton did not preclude the large and rapid increase in bacterial density. We suggest that the dynamics of the equilibrium between bacteria and viruses in the environment is driven to higher numerical levels during periods of intensive bacterial growth. On the contrary, at low bacterial growth rates the temporarily increased virus-to-bacterium ratio may drive the equilibrium to its lowest levels.     

Distribution of microbial populations and their relationship with environmental parameters in the coastal waters of Qingdao,China

In order to understand the large‐scale distribution of microbial populations simultaneously and their relationship with environmental parameters, flow cytometry was used to analyse samples collected from 46 stations in the coastal waters of Qingdao in spring, 2007. The distribution of virus was significantly and positively correlated with heterotrophic bacteria. Two groups of picophytoplankton (Synechococcus and picoeukaryotes) were detected; however, Prochlorococcus was not found. Picoeukaryotes and nanophytoplankton were abundant in the near‐shore waters, whereas Synechococcus was abundant in the off‐shore areas. No variation was found in vertical distribution of virus, heterotrophic bacteria, Synechococcus and nanophytoplankton abundances, except picoeukaryotes abundance in the bottom layer was dramatically lower than that in the upper layers. Correlation analyses indicated that the relationship between abiotic variables and heterotrophic bacteria, pico‐ and nanophytoplankton was closer than that between abiotic variables and virioplankton. Temperature and nutrients were the synchronous factors controlling the growth of heterotrophic bacteria, pico‐ and nanophytoplankton in the coastal waters of Qingdao in spring. The results suggested that synergistic and antagonistic effects existed among microbial groups.     

仿刺参养殖池塘浮游病毒丰度与环境因子的相关性分析

为了揭示仿刺参养殖池塘生态系统中浮游病毒与环境因子的关系,于2008年3—11月对大连市谢屯地区的仿刺参养殖池塘中的浮游病毒丰度进行了定期检测,同时对水温、pH、溶解氧、盐度、叶绿素a含量、化学需氧量、无机氮、活性磷酸盐、异养细菌等因子进行了监测,对浮游病毒丰度与这些环境因子之间的相关性进行了分析。结果表明:仿刺参养殖池塘中浮游病毒的平均丰度为8.32×1010VLPs.L-1(最高值为4月的18.2×1010VLPs.L-1,最低值为11月的1.31×1010VLPs.L-1),外海水中浮游病毒平均丰度为6.45×1010VLPs.L-1(最高值为4月的12.6×1010VLPs.L-1,最低值为6月的2.02×1010VLPs.L-1),仿刺参养殖池塘中营养盐、水温、pH及盐度对浮游病毒丰度的影响较大,而外海水中叶绿素a和异养细菌对浮游病毒丰度的影响较大。     

桂西南喀斯特季节性雨林枯立木的空间格局及生境关联性分析

枯立木是森林生态系统物质循环和能量流动的重要组成部分,对森林群落动态和演替具有重要作用。该研究以位于桂西南的弄岗北热带喀斯特季节性雨林15 hm~2森林动态监测样地为平台,以样地中2011年第一次植被调查数据胸径(diameter at breast height,DBH)≥1 cm的枯立木为研究对象,在充分考虑空间自相关的基础上分析其不同径级(小径级,1 cm≤DBH5 cm;中径级,5 cm≤DBH15 cm;大径级,15 cm≤DBH)的空间分布格局、枯立木多度与环境因子的相关性及生境关联性等。结果表明:枯立木所有个体和小径级分别在0~4 m和0~5 m尺度上表现为聚集分布,其它尺度为随机分布;中径级和大径级在0~30 m尺度上均为随机分布。大径级与凹凸度表现为显著负相关,所有个体和小径级与干旱度指数和样方内所有植株个体数为显著正相关,所有个体和小径级与样方内最大胸径和平均胸径呈显著负相关。所有个体和小径级与3种生境类型间的关联性并不显著;中径级与山坡呈正关联性、与山脊呈负关联性、与山谷无显著关联性;大径级与山谷呈正关联性、与山坡无显著关联性、与山脊呈负关联性。在喀斯特季节性雨林中,枯立木空间分布格局总体表现为随机分布,仅小径级在小尺度内聚集分布,而较大径级在样地中表现出一定的生境关联性;树木死亡过程可能是小尺度内密度制约、大尺度内生境过滤和个体自然衰老等生态学过程形成的。     

Viral abundance, decay, and diversity in the meso- and bathypelagic waters of the north atlantic

To elucidate the potential importance of deep-water viruses in controlling the meso- and bathypelagic picoplankton community, the abundance, decay rate, and diversity of the virioplankton community were determined in the meso- and bathypelagic water masses of the eastern part of the subtropical North Atlantic. Viral abundance averaged 1.4 x 10(6) ml(-1) at around 100 m of depth and decreased only by a factor of 2 at 3,000 to 4,000 m of depth. In contrast, picoplankton abundance decreased by 1 order of magnitude to the Lower Deep Water (LDW; 3,500- to 5,000-m depth). The virus-to-picoplankton ratio increased from 9 at about 100 m of depth to 110 in the LDW. Mean viral decay rates were 3.5 x 10(-3) h(-1) between 900 m and 2,750 m and 1.1 x 10(-3) h(-1) at 4,000 m of depth, corresponding to viral turnover times of 11 and 39 days, respectively. Pulsed-field gel electrophoresis fingerprints obtained from the viral community between 2,400 m and 4,000 m of depth revealed a maximum of only four bands from 4,000 m of depth. Based on the high viral abundance and the low picoplankton production determined via leucine incorporation, we conclude that the viral production calculated from the viral decay is insufficient to maintain the high viral abundance in the deep North Atlantic. Rather, we propose that substantial allochthonous viral input or lysogenic or pseudolysogenic production is required to maintain the high viral abundance detected in the meso- and bathypelagic North Atlantic. Consequently, deep-water prokaryotes are apparently far less controlled in their abundance and taxon richness by lytic prokaryotic phages than the high viral abundance and the virus-to-picoplankton ratio would suggest.     

A multi-scale comparison of trait linkages to environmental and spatial variables in fish communities across a large freshwater lake

Species present in communities are affected by the prevailing environmental conditions, and the traits that these species display may be sensitive indicators of community responses to environmental change. However, interpretation of community responses may be confounded by environmental variation at different spatial scales. Using a hierarchical approach, we assessed the spatial and temporal variation of traits in coastal fish communities in Lake Huron over a 5-year time period (2001–2005) in response to biotic and abiotic environmental factors. The association of environmental and spatial variables with trophic, life-history, and thermal traits at two spatial scales (regional basin-scale, local site-scale) was quantified using multivariate statistics and variation partitioning. We defined these two scales (regional, local) on which to measure variation and then applied this measurement framework identically in all 5 study years. With this framework, we found that there was no change in the spatial scales of fish community traits over the course of the study, although there were small inter-annual shifts in the importance of regional basin- and local site-scale variables in determining community trait composition (e.g., life-history, trophic, and thermal). The overriding effects of regional-scale variables may be related to inter-annual variation in average summer temperature. Additionally, drivers of fish community traits were highly variable among study years, with some years dominated by environmental variation and others dominated by spatially structured variation. The influence of spatial factors on trait composition was dynamic, which suggests that spatial patterns in fish communities over large landscapes are transient. Air temperature and vegetation were significant variables in most years, underscoring the importance of future climate change and shoreline development as drivers of fish community structure. Overall, a trait-based hierarchical framework may be a useful conservation tool, as it highlights the multi-scaled interactive effect of variables over a large landscape.     

Similar Processes but Different Environmental Filters for Soil Bacterial and Fungal Community Composition Turnover on a Broad Spatial Scale

Spatial scaling of microorganisms has been demonstrated over the last decade. However, the processes and environmental filters shaping soil microbial community structure on a broad spatial scale still need to be refined and ranked. Here, we compared bacterial and fungal community composition turnovers through a biogeographical approach on the same soil sampling design at a broad spatial scale (area range: 13300 to 31000 km²): i) to examine their spatial structuring; ii) to investigate the relative importance of environmental selection and spatial autocorrelation in determining their community composition turnover; and iii) to identify and rank the relevant environmental filters and scales involved in their spatial variations. Molecular fingerprinting of soil bacterial and fungal communities was performed on 413 soils from four French regions of contrasting environmental heterogeneity (Landes<Burgundy≤Brittany<<South-East) using the systematic grid of French Soil Quality Monitoring Network to evaluate the communities’ composition turnovers. The relative importance of processes and filters was assessed by distance-based redundancy analysis. This study demonstrates significant community composition turnover rates for soil bacteria and fungi, which were dependent on the region. Bacterial and fungal community composition turnovers were mainly driven by environmental selection explaining from 10% to 20% of community composition variations, but spatial variables also explained 3% to 9% of total variance. These variables highlighted significant spatial autocorrelation of both communities unexplained by the environmental variables measured and could partly be explained by dispersal limitations. Although the identified filters and their hierarchy were dependent on the region and organism, selection was systematically based on a common group of environmental variables: pH, trophic resources, texture and land use. Spatial autocorrelation was also important at coarse (80 to 120 km radius) and/or medium (40 to 65 km radius) spatial scales, suggesting dispersal limitations at these scales.