Evaluating the environmental flows of China's Wolonghu wetland and land use changes using a hydrological model, a water balance model, and remote sensing

Environmental flows are critical to sustaining a variety of plant and animal communities in wetlands. However, evaluation of environmental flows is hampered by the problem of hydrological and ecological data shortage, especially in many developing countries such as China. Based on a hydrological model, a water balance model and remote sensing data, we assessed the environmental flows of China's Wolonghu wetland with limited data. The hydrological model provides input data for the water balance model of the wetland, and the remote sensing data can be used to assess land use changes. Integration of these two models with the remote sensing data revealed both the environmental flows of the Wolonghu wetland and the relationships between these environmental flows and land use changes. The results demonstrate that environmental flows have direct and indirect influences on the wetland ecosystem and should be linked to sustainable wetland management.     

Bottom-up effects of plant genotype on aphids, ants, and predators

Theory predicts that bottom-up ecological forces can affect community dynamics, but whether this extends to the effects of heritable plant variation on tritrophic communities is poorly understood. In a field experiment, I contrasted the effects of plant genotype (28 genotypes; 1064 plants), aphid density, and the presence/absence of mutualistic ants in affecting the per capita population growth of a specialist aphid herbivore, as well as the effects of plant genotype on the third trophic level. Plant genotype strongly affected aphid population growth rate, explaining 29% of the total variation in growth rate, whereas aphid density and ant-aphid interactions explained substantially less variation (< 2%) in aphid population growth rate. Plant genotype also had direct and indirect effects on the third trophic level, affecting the abundance of aphid-tending ants and the richness of predators. Multiple regression identified several heritable plant traits that explained 49% of the variation in aphid growth rate and 30% of the variation in ant abundance among plant genotypes. These bottom-up effects of plant genotype on tritrophic interactions were independent of the effects of either initial aphid density or the presence/absence of mutualistic ants. This study shows that plant genotype can be one of the most important ecological factors shaping tritrophic communities.     

Application of a temperature-dependent von Bertalanffy growth model to bullhead (Cottus gobio)

The most studied and commonly applied model of fish growth is the von Bertalanffy model. However, this model does not take water temperature into account, which is one of the most important environmental factors affecting the life cycle of fish, as many physiological processes that determine growth, e.g. metabolic rate and oxygen supply, are directly influenced by temperature. In the present study we propose a version of the von Bertalanffy growth model that includes mean annual water temperatures by correlating the growth coefficient, k, explicitly and the asymptotic length, L_∞, implicitly to water temperature. All relationships include parameters with an obvious biological relevance that makes them easier to identify. The model is used to fit growth data of bullhead (Cottus gobio) at different locations in the Bez River network (Drme, France). We show that temperature explains much of the growth variability at the different sampling sites of the network.     

Herbivores and edaphic factors constrain the realized niche of a native plant

Biotic interactions, such as competition and herbivory, can limit plant species ranges to a subset of edaphically suitable habitats, termed the realized niche. Here we explored the role that herbivores play in restricting the niche of serpentine ecotypes of the native California annual Collinsia sparsiflora. We planted seeds from four populations into a range of natural field environments that varied in the presence/absence of naturally occurring C. sparsiflora and in predicted suitability for growth and survival of the serpentine ecotype of C. sparsiflora. Path analysis was then used to model the direct and herbivore-mediated indirect effects of environmental variables on the survival of C. sparsiflora serpentine ecotypes. We found that C. sparsiflora received more herbivory when planted into areas where serpentine ecotypes of C. sparsiflora were not predicted to persist, and that increased herbivory was associated with decreased survival, suggesting that herbivores may limit the distribution of C. sparsiflora serpentine ecotypes. Additionally, we demonstrated that edaphic environmental variables impacted the survival of C. sparsiflora serpentine ecotypes both directly and indirectly, by altering interactions with herbivores. These indirect effects were probably trait-mediated and probably occurred because edaphic factors may influence plant traits that, in turn, alter attractiveness to herbivores. Although the magnitude of direct effects exceeded the magnitude of indirect effects, many strong herbivore-mediated indirect effects were detected. Thus, interactions between the abiotic environment and insect herbivory contributed to restricting the niche of C. sparsiflora serpentine ecotypes to a subset of available habitat.     

Species, functional groups, and thresholds in ecological resilience

The cross-scale resilience model states that ecological resilience is generated in part from the distribution of functions within and across scales in a system. Resilience is a measure of a system's ability to remain organized around a particular set of mutually reinforcing processes and structures, known as a regime. We define scale as the geographic extent over which a process operates and the frequency with which a process occurs. Species can be categorized into functional groups that are a link between ecosystem processes and structures and ecological resilience. We applied the cross-scale resilience model to avian species in a grassland ecosystem. A species' morphology is shaped in part by its interaction with ecological structure and pattern, so animal body mass reflects the spatial and temporal distribution of resources. We used the log-transformed rank-ordered body masses of breeding birds associated with grasslands to identify aggregations and discontinuities in the distribution of those body masses. We assessed cross-scale resilience on the basis of 3 metrics: overall number of functional groups, number of functional groups within an aggregation, and the redundancy of functional groups across aggregations. We assessed how the loss of threatened species would affect cross-scale resilience by removing threatened species from the data set and recalculating values of the 3 metrics. We also determined whether more function was retained than expected after the loss of threatened species by comparing observed loss with simulated random loss in a Monte Carlo process. The observed distribution of function compared with the random simulated loss of function indicated that more functionality in the observed data set was retained than expected. On the basis of our results, we believe an ecosystem with a full complement of species can sustain considerable species losses without affecting the distribution of functions within and across aggregations, although ecological resilience is reduced. We propose that the mechanisms responsible for shaping discontinuous distributions of body mass and the nonrandom distribution of functions may also shape species losses such that local extinctions will be nonrandom with respect to the retention and distribution of functions and that the distribution of function within and across aggregations will be conserved despite extinctions.     

Scales of integration for sustainable development governance

Efforts at integration are central in the Sustainable Development Goals and the wider Post-2015 Sustainable Development Agenda. While integration forms a major focus of this agenda, there are fundamental limitations on pursuing this focus. The global governance discussions on integration lack a clear conceptualization of what it is, do not have any tools for measurement or assessment, and are narrowly focused around policy integration between global institutions. This article develops a framework for conceptualizing cross-level and cross-scale integration which addresses these problems and introduces the opportunities and challenges of pursuing different types of integration in global governance. Cross-level integration involves governance efforts which seek to bridge similar institutions with one another (for example, bringing together efforts by environmental ministries and development ministries). Cross-scale integration, in contrast, deals with governance efforts that bring together institutions which have distinct scopes, jurisdictions, and epistemologies. With this important distinction, the article explores the opportunities for cross-scale integration as part of the global sustainable development efforts and particularly expanding the policy integration focus with spatial and temporal integration as well. The conclusion uses the Earth System Governance perspective to focus on the trade-offs that are likely as wider forms of institutional integration are pursued beyond environmental mainstreaming and policy integration.     

Environ indicator sensitivity to flux uncertainty in a phosphorus model of Lake Sidney Lanier, USA

Effective environmental impact assessment and management requires improved understanding of the organization and transformation of ecosystems in which independent agents are linked through an intricate network of energy, matter, and informational interactions. While advances have been made, we still lack a complete understanding of the processes that create, constrain, and sustain ecosystems. Network environ analysis (NEA) provides one approach for building novel ecosystem insights, but it is model dependent. As ecological modeling is an imprecise art, often complicated by inadequate empirical data, the utility of NEA may be limited by model uncertainty. Here, we investigate the sensitivity of NEA indicators of ecosystem growth and development to flow and storage uncertainty in a phosphorus model of Lake Sidney Lanier, USA. The indicators are total system throughflow (TST), total system storage (TSS), total boundary input (Boundary), Finn cycling index (FCI), ratio of indirect-to-direct flows (Indirect/Direct), indirect flow index (IFI), network aggradation (AGG), network homogenization (HMG), and network amplification (AMP). Our results make two primary contributions. First, they demonstrate that five of the indicators – FCI, Indirect/Direct, IFI, AGG and HMG – are relatively robust to the flow and storage uncertainty in the Lake Lanier model. This stability lets us draw robust conclusions about the Lake Lanier ecosystem organization (e.g., phosphorus flux in the lake is dominated by internal processes) in spite of uncertainties in the model. Second, we show that the majority of the indicators co-vary and that most of their common variation could be mapped onto two latent factors, which we interpret as (1) system integration and (2) boundary influences.     

环境因素对密云水库上游流域土壤有机碳和全氮含量影响的通径分析

土壤是全球重要的碳库和氮库,在全球碳氮循环中具有重要地位。密云水库是华北地区最大的水库和北京市最重要的地表水水源地,其上游流域山地广布,地形复杂,气候变化明显,土壤类型和植被类型多样,影响土壤碳氮库的环境因素具有较强的变异性。为揭示环境因素对密云水库上游流域土壤有机碳（soil organic carbon,SOC）和全氮（total nitrogen,TN）含量的作用效应,采用野外采样、实验室分析与逐步回归分析和通径分析相结合的方法,研究了气候（温度和降水）、地形（海拔和坡度）、土壤理化性质（土壤容重、含水量、pH值和粘粒含量）等环境因素对流域SOC和TN含量的影响。结果表明：温度、土壤容重、含水量、pH值和粘粒含量对SOC含量的影响显著（P＆lt;0.001）,其中各因子的直接通径系数依次为：土壤含水量（0.439）＞土壤容重（-0.324）＞pH值（-0.238）＞温度（-0.209）＞土壤粘粒含量（0.092）,间接通径系数依次为：土壤容重（-0.425）＞土壤粘粒含量（0.305）＞土壤含水量（0.287）＞pH值（-0.179）＞温度（-0.043）,因此,土壤含水量、pH值和温度主要通过直接作用影响SOC含量,而土壤容重和粘粒含量则主要通过其它因子的作用间接影响SOC含量。海拔、土壤容重、含水量、pH 值和粘粒含量对 TN 含量的影响显著（P＆lt;0.001）,其中各因子的直接通径系数依次为：土壤含水量（0.456）＞海拔（0.234）＞土壤容重（-0.228）＞pH 值（-0.190）＞土壤粘粒含量（0.094）,间接通径系数依次为：土壤容重（-0.484）＞土壤粘粒含量（0.301）＞海拔（0.247）＞土壤含水量（0.257）＞pH 值（-0.202）,因此,土壤含水量主要通过直接作用影响TN含量,而土壤容重和粘粒含量主要通过土壤含水量的作用间接影响TN含量,海拔和土壤pH值的直接作用与间接作用     

Direct and indirect effects of climate and habitat factors on butterfly diversity

Many factors, including climate, resource availability, and habitat diversity, have been proposed as determinants of global diversity, but the links among them have rarely been studied. Using structural equation modeling (SEM), we investigated direct and indirect effects of climate variables, host-plant richness, and habitat diversity on butterfly species richness across Britain, at 20-km grid resolution. These factors were all important determinants of butterfly diversity, but their relative contributions differed between habitat generalists and specialists, and whether the effects were direct or indirect. Climate variables had strong effects on habitat generalists, whereas host-plant richness and habitat diversity contributed relatively more for habitat specialists. Considering total effects (direct and indirect together), climate variables had the strongest link to butterfly species richness for all groups of species. The results suggest that different mechanistic hypotheses to explain species richness may be more appropriate for habitat generalists and specialists, with generalists hypothesized to show direct physiological limitations and specialists additionally being constrained by trophic interactions (climate affecting host-plant richness).     

A model for measuring leaf growth rate of vegetative shoot in the seagrass, Zostera marina

A new model for determining leaf growth in vegetative shoots of the seagrass Zostera marina (eelgrass) is described. This model requires the weights of individual mature and immature whole leaves and leaf plastochrone interval (P_L) as parameters, differing from the conventional leaf marking technique (CLM) that requires cutting and separation between new and old tissue of leaves. The techniques required for the model are the same as for the plastochrone method, but the parameters differ between both methods in use of the weight of individual immature leaves. In a mesocosm study, eelgrass growth was examined, and parameters for the new model and plastochrone method (the weights of individual mature and immature leaves and P_L) were measured. Leaf growth rate was measured using the CLM and determined by the new method and the plastochrone method. The results were then compared between the CLM, the new model, and the plastochrone method. The results obtained with the new model were similar to those obtained with the CLM. However, the results of the plastochrone method differed from those of the CLM, while the weight of immature leaves varied seasonally. The new model was also used to determine leaf growth in a natural eelgrass bed in Mikawa Bay, Japan, and revealed the growth rates in all shoots and those of different ages. This method would be advantageous as an accurate means of direct measurement in fieldwork, and should therefore be a useful tool for monitoring seagrass growth.     

Herbivory reduces plant interactions with above- and belowground antagonists and mutualists

Herbivores affect plants through direct effects, such as tissue damage, and through indirect effects that alter species interactions. Interactions may be positive or negative, so indirect effects have the potential to enhance or lessen the net impacts of herbivores. Despite the ubiquity of these interactions, the indirect pathways are considerably less understood than the direct effects of herbivores, and multiple indirect pathways are rarely studied simultaneously. We placed herbivore effects in a comprehensive community context by studying how herbivory influences plant interactions with antagonists and mutualists both aboveground and belowground. We manipulated early-season aboveground herbivore damage to Cucumis sativus (cucumber, Cucurbitaceae) and measured interactions with subsequent aboveground herbivores, root-feeding herbivores, pollinators, and arbuscular mycorrhizal fungi (AMF). We quantified plant growth and reproduction and used an enhanced pollination treatment to determine if plants were pollen limited. Increased herbivory reduced interactions with both antagonists and mutualists. Plants with high levels of early herbivory were significantly less likely to suffer leaf damage later in the summer and tended to be less attacked by root herbivores. Herbivory also reduced pollinator visitation, likely due to fewer and smaller flowers, and reduced AMF colonization. The net effect of herbivory on plant growth and reproduction was strongly negative, but lower fruit and seed production were not due to reduced pollinator visits, because reproduction was not pollen limited. Although herbivores influenced interactions between plants and other organisms, these effects appear to be weaker than the direct negative effects of early-season tissue loss.     

Sensitivity of system stability to model structure

A community is stable, and resilient, if the levels of all community variables can return to the original steady state following a perturbation. The stability properties of a community depend on its structure, which is the network of direct effects (interactions) among the variables within the community. These direct effects form feedback cycles (loops) that determine community stability. Although feedback cycles have an intuitive interpretation, identifying how they form the feedback properties of a particular community can be intractable. Furthermore, determining the role that any specific direct effect plays in the stability of a system is even more daunting. Such information, however, would identify important direct effects for targeted experimental and management manipulation even in complex communities for which quantitative information is lacking. We therefore provide a method that determines the sensitivity of community stability to model structure, and identifies the relative role of particular direct effects, indirect effects, and feedback cycles in determining stability. Structural sensitivities summarize the degree to which each direct effect contributes to stabilizing feedback or destabilizing feedback or both. Structural sensitivities prove useful in identifying ecologically important feedback cycles within the community structure and for detecting direct effects that have strong, or weak, influences on community stability. The approach may guide the development of management intervention and research design. We demonstrate its value with two theoretical models and two empirical examples of different levels of complexity.     

Bird metacommunities in temperate South American forest: vegetation structure, area, and climate effects

Spatial structure in metacommunities and their relationships to environmental gradients have been linked to opposing theories of community assembly. In particular, while the species sorting hypothesis predicts strong environmental influences, the neutral theory, the mass effect, and the patch dynamics frameworks all predict differing degrees of spatial structure resulting from dispersal and competition limitations. Here we study the relative influence of environmental gradients and spatial structure in bird assemblages of the Chilean temperate forest. We carried out bird and vegetation surveys in South American temperate forests at 147 points located in nine different protected areas in central Chile, and collected meteorological and productivity data for these localities. Species composition dissimilarities between sites were calculated, as well as three indices of bird local diversity: observed species richness, Chao estimate of richness, and Shannon diversity. A stepwise multiple regression and partial regression analyses were used to select a small number of environmental factors that predicted bird species diversity. Although diversity indices were spatially autocorrelated, environmental factors were sufficient to account for this autocorrelation. Moreover, community dissimilarities were not significantly related to distance between sites. We then tested a multivariate hypothesis about climate, vegetation, and avian diversity interactions using a structural equation modeling (SEM) approach. The SEM showed that climate and area of fragments have important indirect effects on avian diversity, mediated through changes in vegetation structure. Given the scale of this study, the metacommunity framework provides useful insights into the mechanisms driving bird assemblages in this region. Taken together, the weak spatial structure of community composition and diversity, as well as the strong environmental effects on bird diversity, support the interpretation that species sorting has a predominant role in structuring avian assemblages in the region.     

Cavity excavation and enlargement as mechanisms for indirect interactions in an avian community

Direct and indirect species interactions within ecological communities may play a strong role in influencing or maintaining community structure. Complex community interactions pose a major challenge to predicting ecosystem responses to environmental change because predictive frameworks require identification of mechanisms by which community interactions arise. Cavity-nesting communities are well suited for mechanistic studies of species interactions because cavity nesters interact through the creation of and competition for cavity-nest sites. In this study, we use a cavity-nest web as a predictive framework for identifying potential indirect species interactions within a cavity-nesting community. From 2002 to 2005, we monitored abundance and nests of cavity-nesting birds in the longleaf pine (Pinus palustris) ecosystem. Using a nest-web approach, we identified a potential indirect interaction between the Red-cockaded Woodpecker (Picoides borealis) and large secondary cavity nesters, mediated by the Northern Flicker (Colaptes auratus). We used structural equation modeling to test a path model of this interaction, using cavity excavation and enlargement as mechanisms which drive the relationship between these species. Through experimental manipulation of cavity availability, we blocked links described in our model, confirming cavity creation and enlargement as processes that influence community structure. We found that a single-species management technique could potentially disrupt this indirect relationship by affecting Northern Flicker cavity-excavation behavior. This study is the first demonstration of how experimental cavity manipulation can be used to test inferred processes derived from a nest web and highlights the need to understand how mechanisms underlying species interactions can complicate ecosystem responses to environmental change.     

Allowing macroalgae growth forms to emerge: Use of an agent-based model to understand the growth and spread of macroalgae in Florida coral reefs,with emphasis on Halimeda tuna

The growth patterns of macroalgae in three-dimensional space can provide important information regarding the environments in which they live, and insights into changes that may occur when those environments change due to anthropogenic and/or natural causes. To decipher these patterns and their attendant mechanisms and influencing factors, a spatially explicit model has been developed. The model SPREAD (SPatially-explicit Reef Algae Dynamics), which incorporates the key morphogenetic characteristics of clonality and morphological plasticity, is used to investigate the influences of light, temperature, nutrients and disturbance on the growth and spatial occupancy of dominant macroalgae in the Florida Reef Tract. The model species, Halimeda and Dictyota spp., are modular organisms, with an “individual” being made up of repeating structures. These species can also propagate asexually through clonal fragmentation. These traits lead to potentially indefinite growth and plastic morphology that can respond to environmental conditions in various ways. The growth of an individual is modeled as the iteration of discrete macroalgal modules whose dynamics are affected by the light, temperature, and nutrient regimes. Fragmentation is included as a source of asexual reproduction and/or mortality. Model outputs are the same metrics that are obtained in the field, thus allowing for easy comparison. The performance of SPREAD was tested through sensitivity analysis and comparison with independent field data from four study sites in the Florida Reef Tract. Halimeda tuna was selected for initial model comparisons because the relatively untangled growth form permits detailed characterization in the field. Differences in the growth patterns of H. tuna were observed among these reefs. SPREAD was able to closely reproduce these variations, and indicate the potential importance of light and nutrient variations in producing these patterns.     

Impact of cannibalism on predator-prey dynamics: size-structured interactions and apparent mutualism

Direct and indirect interactions between two prey species can strongly alter the dynamics of predator-prey systems. Most predators are cannibalistic, and as a consequence, even systems with only one predator and one prey include two prey types: conspecifics and heterospecifics. The effects of the complex direct and indirect interactions that emerge in such cannibalistic systems are still poorly understood. This study examined how the indirect interaction between conspecific and heterospecific prey affects cannibalism and predation rates and how the direct interactions between both species indirectly alter the effect of the cannibalistic predator. I tested for these effects using larvae of the stream salamanders Eurycea cirrigera (prey) and Pseudotriton ruber (cannibalistic predator) by manipulating the relative densities of the conspecific and heterospecific prey in the presence and absence of the predator in experimental streams. The rates of cannibalism and heterospecific predation were proportional to the respective densities and negatively correlated, indicating a positive indirect interaction between conspecific and heterospecific prey, similar to "apparent mutualism." Direct interactions between prey species did not alter the effect of the predator. Although both types of prey showed a similar 30% reduction in night activity and switch in microhabitat use in response to the presence of the predator, cannibalism rates were three times higher than heterospecific predation rates irrespective of the relative densities of the two types of prey. Cumulative predation risks differed even more due to the 48% lower growth rate of conspecific prey. Detailed laboratory experiments suggest that the 3:1 difference in cannibalism and predation rate was due to the higher efficiency of heterospecific prey in escaping immediate attacks. However, no difference was observed when the predator was a closely related salamander species, Gyrinophilus porphyriticus, indicating that this difference is species specific. This demonstrates that cannibalism can result in the coupling of predator and prey mortality rates that strongly determines the dynamics of predator-prey systems.     

The effects of direct and indirect constraints on biological communities

Human activities are expected to result in a diversity of directional or stochastic constraints that affect species either directly or by indirectly impacting their resources. However, there is no theoretical framework to predict the complex and various effects of these constraints on ecological communities. We developed a dynamic model that mimics the use of different resource types by a community of competing species. We investigated the effects of different environmental constraints (affecting either directly the growth rate of species or having indirect effects on their resources) on several biodiversity indicators. Our results indicate that (i) in realistic community models (assuming uneven resource requirements among species) the effects of perturbations are strongly buffered compared to neutral models; (ii) the species richness of communities can be maximized for intermediate levels of direct constraints (unimodal response), even in the absence of trade-off between competitive ability and tolerance to constraints; (iii) no such unimodal response occurs with indirect constraints; (iv) an increase in the environmental (e.g., climatic) variance may have different effects on community biomass and species richness.     

Temporal and spatial variation in the growth rates of Baltic herring (<Emphasis Type="Italic">Clupea harengus membras</Emphasis> L.) larvae during summer

It has been suggested that larval survival determines the year-class strength in most marine fish species. During their growth and development, the ability of the larvae to catch prey and avoid predation will increase. However, the factors affecting short-term changes in the growth of Baltic Sea herring have been little studied in the field. We collected Baltic herring (Clupea harengus membras L.) larvae from five different towing areas in the Archipelago Sea (SW Finland) during May and June 1989, right after the main spawning season. Twenty thousand two hundred and ten larvae were analysed and the area-specific growth rate (i.e. increase in standard length) was estimated by tracing the larval cohorts from the length-frequency data. This represents the first Baltic herring study with daily sampling during a long study period. The growth rate was related to environmental factors, such as temperature, number of zooplankters, and wind speed and direction. Large variation in larval growth rate occurred between areas: lowest and highest growth rates were 0.18 and 0.52 mm·day^-1. Temperature was an important variable controlling larval-fish growth rate. An increase of one 1°C in average water temperature corresponded to an increase in growth rate of 0.043 mm·day^-1. This may have been caused either by a direct temperature effect (changes in metabolic rate) or by the indirect effect of changes in food availability. We also found the densest herring populations in the areas with highest average water temperature. However, temperature and larval growth rate both increased towards the inner archipelago.     

区域生态环境服务价值评估 --以延安市宝塔区为例

将小尺度行政区域作为研究对象,建立区域生态环境服务价值评估模型。模型重点考虑了2个方面:第一,主体生态系统对区域生态环境服务价值的贡献大小及主体生态系统之间的相互影响;第二,估算主体生态系统价值时考虑直接使用价值、间接使用价值、非使用价值的重复或遗漏及生态环境服务价值随社会经济的变化。以陕西延安市宝塔区为例进行评估,得到2002年宝塔区生态环境服务价值为118322.214万元。     

Relevant variables to predict macrophyte communities in running waters

In both predictive theoretical and empirical models for aquatic plant communities in running waters, the development and competition are many times explained in terms of nutrients. Minerals necessary for growth are generally not assumed to be limiting, although they influence the important pH-value. At the same time it is known that factors such as oxygen-concentration, solar energy, salinity, dimension of the system and soil characteristics (including river sediments) influence the development of the community, and should be considered in modelling. Effects of water quantity and water quality on macrophytes are reviewed. These conditions are caused by processes in the landscape, characterised by a set of nested variables which explain the distribution of macrophyte species and communities. Relevant variables are described and grouped on three scales: regional, local and site conditions. Case studies with direct and indirect gradient analysis are presented. Statistical tests (stepwise regression with forward selection) reveal that each species distribution is explained by a characteristic set of relevant variables, ranging from soil type and dimension of the system, to nutrient and salinity concentration.