On the dynamics of grazing systems in the semi-arid succulent Karoo: The relevance of equilibrium and non-equilibrium concepts to the sustainability of semi-arid pastoral systems

Two long-term mechanistic models of grazing systems in the semi-arid succulent Karoo have been used to study factors that influence vegetation changes, livestock productivity and sustainability of the ecosystem. In this region of low and highly variable rainfall, goats and sheep feed on vegetation comprising perennial shrubs and annuals. A previously published model of the Namaqualand system (the “standard” model) explicitly simulates three guilds of perennial shrubs, a guild of annuals, forage consumption, growth of goats and goat reproductive and survival rates. The model also simulates variable rainfall and predicts that, if no steps are taken to control the goat population, stock numbers will vary widely between years and the population of the different plant guilds will fluctuate. Plots of model output indicate that the system is driven by rainfall. Temporal changes in the relative abundance of each guild vary with different sequences of rainfall having similar long-term mean and variability. A single run of the model may display equilibrial, disequilibrial and threshold behaviour. Thus, the system exhibits complex dynamics. If animal numbers are held constant at the long-term average of variable stock or at the recommended stocking rate then the cover of palatable shrubs decreases and that of toxic plants increases substantially. A “simplified” model based on an aggregated forage variable and equilibrium dynamics is inadequate to describe the behaviour of this system.     

Emergent properties modelled with the functional structural tree growth model ALMIS: Computer experiments on resource gain and use

The functional structural tree growth model ALMIS uses the individual based modelling approach and is implemented in the object-oriented programming language SIMULA. All features (state variables) and functions (processes) are specified locally, on the level of the single plant organs. Increasing numbers of “copies” (objects) of these elementary units, Internodes, Leaves, Meristems, Roots, and Root tips, form the growing tree. Various procedures (e.g. Photosynthesis, Nutrient_uptake, Transport, Storage, Mobilisation, Respiration, Growth) are employed to describe carbon and nutrient uptake, and matter fluxes between the different plant organs. Combining plant physiology and architecture, ALMIS allows studying the effect of single ecophysiological and structural processes on whole tree growth and in the tree–environment system. Some of these effects, driven by microclimate, self-shading, variable nutrient availability, variable transport dynamics, and branching patterns are exemplified. From the interactions at the organ and sub-organ levels new features emerge at higher levels of plant organisation. These so-called emergent properties are, for example, lifetime spectrum of single organs, space filling (“architecture”) and self-thinning of the crown. The most prominent emergent properties are the different growth forms of trees resulting from simulations under various conditions. Their causal interrelations are discussed in detail.     

Relationships between nutrients and macroalgal biomass in a brazilian coastal lagoon: the impact of a lock construction

The Piratininga Lagoon is a coastal, choked, and brackish ecosystem in SE Brazil, where uncontrolled discharge of domestic sewage led to eutrophication, increasing massive proliferation of benthic macroalgae, and decrease of the lagoon mean depth. In 1992, a dam was constructed by the local Municipality at its tidal channel aiming at stabilising its water level. Main physico-chemical parameters, together with macroalgae biomass, nutrient concentrations in the water column, particulate organic carbon, and chlorophyll a were recorded monthly at 4 sampling stations within the Piratininga lagoon from April 1994 to April 1995. The data, compared with “before-lock” existing studies, show that nutrients and chlorophyll a concentrations significantly increased after the lock construction. Based on the functioning of the ecosystem, we propose to harvest the algal mats before their decomposition period in order to partly remove the nutrient stocks from the lagoon and the future ecosystem modelling to predict the impact of natural and anthropogenic eutrophication.     

Steady states and sensitivities of commonly used pelagic ecosystem model components

Pelagic, coupled ocean circulation-ecosystem models, are widely used in climate research. These tools aim to quantify fluxes of nutrients and carbon in the ocean and are, increasingly, the base of future projections. For this purpose it is crucial to quantify and identify the sources of uncertainties. In contrast to physical models, the underlying equations for ecosystem models are derived from empirical relationships rather than based on first principles. This resulted in the development of a multitude of different ecosystem models - different in respect to both, underlying principles and complexity. Clearly, the question arises, to what extent the sensitivities of these models are comparable.This study focuses on the intrinsic dynamics of some widely used, simple (containing 2-3 prognostic variables) ecosystem models in a 0-D framework (i.e., comprising only the well-mixed oceanic surface layer). A suite of differing model approaches is tuned such that their behavior is similar. The setup resembles the well-mixed oceanic surface layer in the Baltic proper. It is illustrated that strong differences between the model approaches appear due to exemplary, anticipated changes in the external nutrient and light conditions. Herewith, we demonstrate the well-known, but rarely demonstrated fact that, apparent consistency between modeled prognostic variables with today's data bases is not necessarily a good measure of forecast skill. The causes which lead to the different sensitivities are illustrated by considering the steady state solutions. It is pointed out, that apparently small changes in the model formulations can result in very different dynamical behavior and an enormous spread between the model approaches, despite the feasibility to tune a common behavior in a limited range of light and nutrient supply. In our examples, the sensitivity is mainly a function of the formulation of the loss rate of phytoplankton. It is thus, in particular, the formulation of highly unknown heteorotrophic processes that determines the model sensitivity.     

Estimation of algal growth parameters from vertical primary production profiles

Phytoplankton maximum growth rate and the saturation light intensity, I_s, can be estimated from vertical profiles of primary production by non-linear least-squares estimation. Solution through the normal equations leads to formulae which are relatively simple and easy to implement. The computation of confidence contours is demonstrated, and the results are contrasted to the confidence limits on the parameters individually. These can be quite misleading due to model non-linearity and correlation between parameter estimation.The procedure has been applied to primary production data from Lake Balaton, a shallow lake in Hungary. The growth rate-temperature relation is analysed by separating the parameters into two groups characteristic for “warm” and “cold” water phytoplankton, respectively. A bell-shaped curve is found for “cold” water communities, with an optimum at about 7–9°C, whereas the “warm” water phytoplankton exhibits a strong exponential dependency in the temperature range of interest (up to 25°C). I_s also appears to be related to temperature except for the “cold” water group, where I_s is essentially constant. However, a roughly linear relation with considerably less scatter is obtained when I_s is plotted directly versus day-averaged solar radiation. This apparent fast adaptation is attributed to the extremely short turnover time in Lake Balaton. Maximum growth rates of 10–20 d^?1 have been found for temperatures between 20 and 25°C. These results and a critical appraisal of available literature suggest that the common notion of maximum growth rates being in the order of 1–3 d^?1 needs revision, at least for lakes with relatively high summer temperatures.     

Aspects of a zooplankton,phytoplankton, phosphorus system

In this paper a three-dimensional dynamical system which models the three-species system made up of phytoplankton, zooplankton and organic phosphorus nutrient in a lake environment is studied. The system is part of a more general limnological model for eutrophic lakes and impoundments which has been developed by Battelle Northwest Laboratories. It is shown that this system, henceforth referred to as Z-P-P, has a phase portrait comprised of a plane portrait embedded in the three-dimensional space R³ as an “attractor”.¹ Under a small perturbation of the nutrient equation it is shown that the system is essentially a classical Volterra-Lotka system embedded in a three-dimensional phase space R³.The system derived from Z-P-P by the addition of a term to the nutrient equation which represents the organic phosphorus contribution of dying phytoplankton is also considered. The equilibria of this system are studied and what can be deduced of the phase portrait is compared with that of the above systems. It is found that these phase portraits are qualitatively indifferent to the form of the growth rate functions for zooplankton and phytoplankton provided they are monotone increasing. Some discussion about the stability of these systems is included. Throughout this paper results are interpreted in limnological terms.     

Relating effect and response traits in submersed aquatic macrophytes.

Reliably predicting the consequences of short- or long-term changes in the environment is important as anthropogenic pressures are increasingly stressing the world's ecosystems. One approach is to examine the manner in which biota respond to changes in the environment ("response traits") and how biota, in turn, affect ecosystem processes ("effect traits"). I compared the response and effect traits of four submersed aquatic macrophytes to understand how water level management may affect wetland plant populations and ecosystem processes. I measured resource properties (nutrients in sediment and water), non-resource properties (pH, alkalinity, sediment temperature, oxygen production), and biotic properties (periphyton biomass) in replicated outdoor monocultures of Stuckenia pectinata, Potamogeton nodosus, P. crispus, and Zannichellia palustris. After seven weeks, three of eight replicates of each species treatment were subjected to a temporary water draw-down that desiccated aboveground plant parts. The four species differed in their effects on ecosystem properties associated with nutrient uptake and photosynthetic activity. Shoot growth rate was negatively correlated with light transmittance to the sediment surface whereas root growth rate and root:shoot ratio were correlated with a species' ability to deplete nutrients in sediment interstitial water. Occupation of space in the water column was correlated with water alkalinity and pH and with sediment temperature. Root growth rate was related simultaneously to species effects on sediment nutrient dynamics and recovery of ecosystem properties after water draw-down. This suggests that this morphological trait may be used to predict the effects of environmental change on ecosystem functioning within the context of water level management. Expanding these analyses to more species, different environmental stressors, and across aquatic and terrestrial ecosystems should enhance predictions of the complex effects of global environmental change on ecosystem functioning.     

岩溶生态系统的土壤

岩溶生态系统的土壤是岩溶地区岩石、大气、水、生物等四大圈层相互作用的产物，成土速率较慢，土壤剖面和空间分布受地貌演化阶段和岩溶双层空间结构的控制。在碳酸盐岩的差异侵蚀和土壤丧失的作用下，岩溶生态系统的土壤逐渐向裂隙和低洼部位退缩，形成景观中的土壤资源斑块，成为岩溶生态系统的养分库、水库和种子库，这种自然演替形成的小尺度上的土壤斑块和生境异质性对于维持岩溶景观的健康状况是非常重要的。今后的研究应着重从岩溶表层生态系统的运行过程中来把握岩溶生态系统土壤的动态特征和相互间的反馈关系。     

An ecosystem model for estimating potential shellfish culture production in sheltered coastal waters

A generic ecosystem model has been developed for estimating the potential production of shellfish culture and the effect of that cultivation on the pelagic ecosystem in sheltered coastal waters. The model describes the dynamics of a simple food web, nutrient cycling and growth of shellfish. The design of the model is closely tied to the temporal and spatial scales that are important in determining the sustainable production level for a particular embayment. The pelagic ecosystem, mussel energetics, population dynamics and hydrodynamics are coupled to allow fully dynamic predictions of the effect of the shellfish density. When applied to Beatrix Bay, an intensive culture embayment in the Pelorus Sound of New Zealand, the model successfully captured main features of the observed system behaviour. The hydrodynamic regime of the bay controls mussel growth and production. Although high fluxes of water into the bay suppress nutrient and carbon cycling signals in the system, the model simulations demonstrated that the mussel cultivation can have considerable effects on the ecosystem of the bay including food depletion and nutrient cycling. One of the most obvious effects is nutrient enhancement through mussel excretion at low cultivation densities, which promotes primary production particularly during the N-limitation period in summer. The sensitivity analysis identified uncertainty in some parameters and indicated areas for which experimental studies could lead to model improvement. The modelling exercise has established a primary predictive tool for managing mussel aquaculture of a coastal embayment to estimate relationships between the stock level and the growth rate of mussels, and the potentially achievable harvest and stocking density.     

Multivariate analysis of the mud crab Scylla serrata (Brachyura: Portunidae) from four locations in Southeast Asia

The large, edible mud crab Scylla serrata (Forskål) exhibits different phenotypes which are recognised by the fishermen of Southeast Asia and are given local names such as “white”, “green”, “red” or “black” crabs on the basis of their colouration. A preliminary study using multivariate techniques was undertaken in order to examine the degree of dissimilarity between the different morphs of commercially fished stocks of S. serrata. Twenty-two morphometric and 20 meristic characters were measured on male crabs from four locations in three countries in Southeast Asia, including Surat Thani in Thailand where “black” and “white” morphs exist sympatrically. Canonical variate analysis (CVA) on the morphometric data discriminated three groups with no chain-linking. Surat Thani “white” crabs and those from south Vietnam were phenotypically similar and formed one distinct group; a second group contained crabs from Ranong (Thailand) and Sarawak which were phenotypically “black”; the third group contained “black” crabs from Surat Thani. CVA on meristic data confirmed two groupings, but implied that the Surat Thani “black” morph may only be a variant of the Surat Thani/Vietnam “white” form, rather than a third species. These findings are discussed in relation to the ecology, fishery management and aquaculture potenial of S. serrata.     

蒙古栎林生态系统的水分循环与养分循环关系

本文从群体水平上研究了蒙古栎林生态系统中的水分循环与养分循环的关系。结果表明,降雨是系统养分的重要输入。降雨对林冠的淋洗淋溶作用有助于养分的归还及缩短养分循环的途径。蒙古栎树干茎流量大,且其中含有的养分元素较多,从而使蒙古栎的根系能获得更多的水分与养分,有助于适应干旱、瘠薄的立地环境。径流是系统的水分输出,也是养分输出。蒙古栎林的下渗流约占整个径流的90％,且养分元素含量较低,从而有助于养分的积累。降雨在蒙古栎林内的分配格局不一,导致系统养分积累的明显差异。本文还探讨了干扰对养分循环与水分循环的影响。结果表明,采伐森林导致水分分配的变化,也使系统的养分发生亏损。只有把养分循环与水分循环结合起来,才能较完整地评价森林的生态功能。     

Environmental periodicity and ecosystem stability

Simple ecosystem models involving a primary producer, nutrients, and a herbivore are developed with metabolic dependence on periodic functions representing solar radiation and temperature. Simulation results suggests that the models yield generally stable output provided that equilibrium values for the state variables are positive in the nonperiodic analog. Linear dependence of plant growth on nutrient supply leads to a larger stability domain than that produced by hyperbolic dependence.     

Sparse regression for analyzing the development of foliar nutrient concentrations in coniferous trees

Analyzing and predicting the development of foliar nutrient concentrations are important and challenging tasks in environmental monitoring. This article presents how linear sparse regression models can be used to represent the relations between different foliar nutrient concentration measurements of coniferous trees in consecutive years. In the experiments the models proved to be capable of providing relatively good and reliable predictions of the development of foliage with a considerably small number of regressors. Two methods for estimating sparse models were compared to more conventional linear regression models. Differences in the prediction accuracies between the sparse and full models were minor, but the sparse models were found to highlight important dependencies between the nutrient measurements better than the other regression models. The use of sparse models is, therefore, advantageous in the analysis and interpretation of the development of foliar nutrient concentrations.     

Biological interactions and environmental effects in the economics of pest control

The problem of pest control is tackled in a context of an ecosystem that consists of prey-predator populations with human interaction through pesticide application. The control, aimed at reducing pest damage, results in two undesirable external effects: reduction of beneficial predator population and environmental contamination. The untapped natural equilibrium is compared with equilibrium resulting from decentralized and centralized economic decision making. It is shown that, under certain conditions, myopic decision rules increase rather than decrease the pest damage. The “user cost” (or benefit) is shown to be crucial in determining the optimal centralized policy, and its relations to the various components of the system are analyzed. The components of the user cost are analyzed to determine the level of taxes or subsidies that will yield the optimal policy.     

Modelling above-ground herbage mass for a wide range of grassland community types

Whereas it is recognized that management of plant diversity can be the key to reconciling production and environmental aims, most grassland models are tailored for high-value grass species. We proposed to adapt a mono-specific grass model to take into account specific features of species-rich permanent grasslands, especially over the reproductive phase. To this end, we used the concept of plant functional type (PFT), i.e. the grouping of plant species according to plant traits determined by the response of plant species to different management practices (land use and fertilization) and characterizing of agronomic properties of the corresponding species. In the model, weather and nutrient availability act upon rates of biophysical processes (radiation capture and use, plant senescence). These rates are modified over times due to PFT-specific parameters determined experimentally which represent the different strategies of plant species regarding growth. The integration of these parameters into the model made it possible to predict herbage biomass accumulation rate under different management practices for a wide range of plant communities differing in their PFT composition. The model was evaluated in two steps, first by analyzing separately the effects of PFT and an indicator of nutrient availability on herbage accumulation and then by conducting a sensitivity analysis. It was validated using two independent datasets; a cutting experiment running over the whole growing season to examine the consistency of the model outputs under different cutting regimes, and a monitoring of meadows and pastures in spring over a whole growth cycle to assess the model’s ability to reproduce growth curves. Although a good fit was observed between the simulated and observed data, the few discrepancies noticed between field data and predicted values were attributed mainly to the potential presence of non-grass species. More specifically, we noticed that nutrient (mainly nitrogen) availability is the main driver of plant growth rate, and that PFT determines the times at which this rate changes in relation to the phenological characteristics of species present. We concluded that integration of the PFT concept into the initial mono-specific growth model is especially suited to evaluating the consequences of management practices on species-rich permanent grasslands to meet feed production targets.     

养分管理与农田杂草生物多样性和遗传进化的关系研究进展

施肥被看作是一项潜在降低农田杂草对作物干扰的有效农作管理措施。不同施肥处理可形成具有不同养分状况的土壤条件,而土壤条件的差异是导致杂草生物多样性的重要因素。论证养分管理可显著改变农田杂草发生状况和杂草生物多样性,强调了深入开展不同养分管理模式对农田杂草生物多样性和遗传进化影响研究的重要性,提出了如何利用养分管理措施调节农田杂草,在控制杂草对农业危害的同时,保持一定农田可控杂草生物多样性的科学问题。长期N、P和K平衡配施,可维持杂草群落和杂草种子库的生物多样性与稳定性,兼顾杂草控制和杂草多样性保护,实现平衡施肥的经济效应与生态效应的统一。同时,农业生态系统中不同养分管理措施条件下的地表光辐射变化特征、不同光辐射变化对杂草种群种子萌发及植株建立的影响、优势杂草种群的确定及其生理生态和遗传进化规律是今后研究的重点和难点。     

Influence des conditions hivernales sur les productions phyto- et zooplanctoniques en Méditerranée Nord-Occidentale. II. Biomasse et production phytoplanctonique

Hydrological observations, and measurements of nutrient chemistry, plankton biomass, and production were carried out during the Médiprod I cruise of the R.V. “Jean-Charcot”. The March cruise was characterized by almost winter conditions, exhibiting strong vertical mixing of water masses in the offshore region and nutrient transport up to the photic zone. According to the working hypothesis, the strong vertically mixed area (e.g. Station 15) and the surrounding oligotrophic area are separated by an intermediate zone, where both nutrients and stability have produced phytoplankton-bloom conditions. During the April cruise, highest biomass and production rates were encountered everywhere in the offshore region, and especially in the previously mixed area of central divergence. Production was as high as 2 gC·m^-2 day^-1, and the standing crop of chlorophyll was 3 mg·m^-3; such values are rather important for the so called “poor” Mediterranean Sea. Salinity-phosphate and chlorophyll-phosphate diagrams are presented. Biomass and production rates are in agreement with the potential fertility based on the nutrient content of the waters. The disappearance of 1 μatg P·PO₄ by photosynthetic uptake corresponds to 7.7 mg chlorophyll a, which represents the autotrophic biomass remaining after grazing by the simultaneous zooplankton bloom. Biomass and production features are analyzed in regard to interaction of both nutrient availability and the stability of water masses. Stability conditions can be created either by intrusion of local mixing in a stratified oligotrophic area (“winter bloom”), or by thermal stratification of the upper layer (“spring bloom”). In the latter case, the highest biomasses are present in the zone where the nutrients were previously introduced by mixing. The oligotrophic situation remained constant during the two crunises in the surrounding coastal area, which is characterized by low-salinity water and, therefore, absence of vertical nutrient transport into the photic zone. Chlorophyll pigment concentration and photosynthetic rates in the “Cote d'Azur” region are similar to those in the “Provence” region; this situation may result more from upwelling of nutrient-rich intermediate water than from the mixing process which predominates in the latter region.     

The impact of detritivorous fishes on a mangrove estuarine system

Sundarban mangrove ecosystem in India is one of the largest detritus based ecosystem of the world and it supplies the detritus and nutrients to the adjacent Hooghly-Matla estuarine complex. In this estuary a group of fish completely detritivorous in nature, belonging to the genus Mugil spp. is present. This group of fish is expected to have important effects on the trophic dynamics of ecosystems, but exact nature of these effects is not known. In order to study the impact that detritivory by fish may have on the estuarine food chain, we developed mathematical formulations. We run two models, one with phytoplankton, zooplankton, carnivorous fish, detritus and nutrient and without this group of fish and a second one after including this fish in the system. In our model this group of fish has no major impact on primary productions of the estuarine system but has extensive role in total fish production. Coexistence of detritivorous fish and carnivorous fish occurs within reasonable parameter range. We have tested different growth rates of phytoplankton, grazing rates and predation rates of zooplankton, detritivorous fish and carnivorous fish for total system equilibrium. Carnivorous fish predation rate on detritivorous fish and detritivorous fish grazing rate on detritus are very important. Different foraging ratios are also tested in this study. Foraging preference of carnivorous fish on detritivorous fish appears significant for the system equilibrium.     

On maturation and spawning in some penaeid prawns of the Arabian Gulf

A mathematical model is presented of the ecosystem in the upper layers of the marine pelagic zone. The model has been constructed on the basis of presumed connections between biotic and abiotic ecosystem parameters typical of the Black Sea. Details on total model relations as well as on model analysis and construction may be obtained after gaining more insight into the behaviour of models constructed for the upper pelagic ecosystem of different marine aquatoria. The model is characterized by a rather complex behaviour. Realization of the model on the electronic computer is made by employing the method of random trajectories. As a result, a qualitative picture of model behaviour under different conditions is revealed and statistical characteristics of the parameters are obtained.     

Temperature effects on vital rates of different life stages and implications for population growth of Baltic sprat

Baltic sprat (Sprattus sprattus balticus S.) is a key species in the pelagic ecosystem of the Baltic Sea. Most stocks of small pelagic species are characterized by natural, fishery-independent fluctuations, which make it difficult to predict stock development. Baltic sprat recruitment is highly variable, which can partly be related to climate-driven variability in hydrographic conditions. Results from experimental studies and field observations demonstrate that a number of important life history traits of sprat are affected by temperature, especially the survival and growth of early life stages. Projected climate-driven warming may impact important processes affecting various life stages of sprat, from survival and development during the egg and larval phases to the reproductive output of adults. This study presents a stage-based matrix model approach to simulate sprat population dynamics in relation to different climate change scenarios. Data obtained from experimental studies and field observations were used to estimate and incorporate stage-specific growth and survival rates into the model. Model-based estimates of population growth rate were affected most by changes in the transition probability of the feeding larval stage at all temperatures (+0, +2, +4, +6?°C). The maximum increase in population growth rate was expected when ambient temperature was elevated by 4?°C. Coupling our stage-based model and more complex, biophysical individual-based models may reveal the processes driving these expected climate-driven changes in Baltic Sea sprat population dynamics.