Resistance and re-organization of an ecosystem in response to biological invasion: Some hypotheses

Using a dynamic model of Lake Chozas developed by Marchi et al. (2011), we tested three hypotheses about recovery of the indigenous community and water quality after radical changes caused by introduction of an invasive allochthonous crayfish, Procambarus clarkii:

1.

Can the lake resist the pressure of an invasive species, like P. clarkii, by adaptation?

2.

Can the ecosystem recover when all the crayfish are removed and low phosphorus concentrations persist in inflow water?

3.

Does the simulated recovery of submerged vegetation occur at a total phosphorus concentration below 100 mg TP m⁻³, as estimated by Scheffer et al. (1993), Scheffer (1997), Jeppesen et al. (1998) and Zhang et al. (2003)?

We obtained the following answers:

1.

Lake Chozas can at least partly resist by adaptation. A combination of possible parameter changes could lead to a significant increase in eco-exergy.

2.

Removal of the phosphorus represented by crayfish (by harvesting) implies complete recovery of the lake and its eco-exergy, albeit not necessarily with the same organisms having the same properties.

3.

The expected hysteresis created by introduction and harvesting of crayfish is observed under the following conditions: phytoplankton dominance at total phosphorus ≥ about 200-250 mg TP m⁻³ and submerged vegetation returns at total phosphorus < 100 mg TP m⁻³.

     

Modeling the role of macroalgae in a shallow sub-estuary of Narragansett Bay, RI (USA)

Proliferation of macroalgal mats is a frequent consequence of nutrient-driven eutrophication in shallow, photic coastal marine ecosystems. These macroalgae have the potential to significantly modify water quality, plankton productivity, nutrient cycling, and dissolved oxygen dynamics. We developed a model for Ulva lactuca and Gracilaria tikvahiae in Greenwich Bay, RI (USA), a shallow sub-estuary of Narragansett Bay, as part of a larger estuarine ecosystem model. The model predicts the biomass of both species in units of carbon, nitrogen, and phosphorus as a function of primary production, respiration, grazing, decay, and physical exchange, with particular attention to the effects of biomass layering on light attenuation and suppression of metabolic rates. The model successfully reproduced the magnitude and seasonal cycle of area-weighted and peak biomass in Greenwich Bay along with tissue C:N ratios, and highlighted the importance of grazing and inclusion of self-limitation primarily in the form of self-shading to overcome an order of magnitude difference in rates of production and respiration. Inclusion of luxury nutrient uptake demonstrated the importance of internal nutrient storage in fueling production when nutrients are limiting. Macroalgae were predicted to contribute a small fraction of total system primary production and their removal had little effect on predicted water quality. Despite a lack of data for calibration and a fair amount of sensitivity to individual parameter values, which highlights the need for further autecological studies to constrain formulations, the model successfully predicted macroalgal biomass dynamics and their role in ecosystem functioning. Our formulations should be exportable to other temperate systems where macroalgae occur in abundance.     

Modeling seasonal course of carbon fluxes and evapotranspiration in response to low temperature and moisture in a boreal Scots pine ecosystem

Environmental conditions act above and below ground, and regulate carbon fluxes and evapotranspiration. The productivity of boreal forest ecosystems is strongly governed by low temperature and moisture conditions, but the understanding of various feedbacks between vegetation and environmental conditions is still unclear. In order to quantify the seasonal responses of vegetation to environmental factors, the seasonality of carbon and heat fluxes and the corresponding responses for temperature and moisture in air and soil were simulated by merging a process-based model (CoupModel) with detailed measurements representing various components of a forest ecosystem in Hyytiälä, southern Finland. The uncertainties in parameters, model assumptions, and measurements were identified by generalized likelihood uncertainty estimation (GLUE). Seasonal and diurnal courses of sensible and latent heat fluxes and net ecosystem exchange (NEE) of CO₂ were successfully simulated for two contrasting years. Moreover, systematic increases in efficiency of photosynthesis, water uptake, and decomposition occurred from spring to summer, demonstrating the strong coupling between processes. Evapotranspiration and NEE flux both showed a strong response to soil temperature conditions via different direct and indirect ecosystem mechanisms. The rate of photosynthesis was strongly correlated with the corresponding water uptake response and the light use efficiency. With the present data and model assumptions, it was not possible to precisely distinguish the various regulating ecosystem mechanisms. Our approach proved robust for modeling the seasonal course of carbon fluxes and evapotranspiration by combining different independent measurements. It will be highly interesting to continue using long-term series data and to make additional tests of optional stomatal conductance models in order to improve our understanding of the boreal forest ecosystem in response to climate variability and environmental conditions.     

Simulation modeling to understand how selective foraging by beaver can drive the structure and function of a willow community

Beaver–willow (Castor-Salix) communities are a unique and vital component of healthy wetlands throughout the Holarctic region. Beaver selectively forage willow to provide fresh food, stored winter food, and construction material. The effects of this complex foraging behavior on the structure and function of willow communities is poorly understood. Simulation modeling may help ecologists understand these complex interactions. In this study, a modified version of the SAVANNA ecosystem model was developed to better understand how beaver foraging affects the structure and function of a willow community in a simulated riparian ecosystem in Rocky Mountain National Park, Colorado (RMNP). The model represents willow in terms of plant and stem dynamics and beaver foraging in terms of the quantity and quality of stems cut to meet the energetic and life history requirements of beaver. Given a site where all stems were equally available, the model suggested a simulated beaver family of 2 adults, 2 yearlings, and 2 kits required a minimum of 4 ha of willow (containing about10 stems m⁻²) to persist in a steady-state condition. Beaver created a willow community where the annual net primary productivity (ANPP) was 2 times higher and plant architecture was more diverse than the willow community without beaver. Beaver foraging created a plant architecture dominated by medium size willow plants, which likely explains how beaver can increase ANPP. Long-term simulations suggested that woody biomass stabilized at similar values even though availability differed greatly at initial condition. Simulations also suggested that willow ANPP increased across a range of beaver densities until beaver became food limited. Thus, selective foraging by beaver increased productivity, decreased biomass, and increased structural heterogeneity in a simulated willow community.     

An invasive riparian tree reduces stream ecosystem efficiency via a recalcitrant organic matter subsidy

A disturbance, such as species invasion, can alter the exchange of materials and organisms between ecosystems, with potential consequences for the function of both ecosystems. Russian olive (Elaeagnus angustifolia) is an exotic tree invading riparian corridors in the western United States, and may alter stream organic matter budgets by increasing allochthonous litter and by reducing light via shading, in turn decreasing in-stream primary production. We used a before-after invasion comparison spanning 35 years to show that Russian olive invasion increased allochthonous litter nearly 25-fold to an invaded vs. a control reach of a stream, and we found that this litter decayed more slowly than native willow. Despite a mean 50% increase in canopy cover by Russian olive and associated shading, there were no significant changes in gross primary production. Benthic organic matter storage increased fourfold after Russian olive invasion compared to pre-invasion conditions, but there were no associated changes in stream ecosystem respiration or organic matter export. Thus, estimated stream ecosystem efficiency (ratio of ecosystem respiration to organic matter input) decreased 14%. These findings show that invasions of nonnative plant species in terrestrial habitats can alter resource fluxes to streams with consequences for whole-ecosystem functions.     

Influences of canopy structure and physiological traits on flux partitioning between understory and overstory in an eastern Siberian boreal larch forest

Boreal forests play an important role in the global balance of energy and CO₂. Our previous study of elaborate eddy covariance observations in a Siberian boreal larch forest, conducted both above the forest canopy and at the forest floor, revealed a significant contribution of latent heat flux (LE) from the cowberry understory to the whole ecosystem LE. Thus, in the present study, we examined what factors control the partitioning of whole ecosystem LE and CO₂ flux into the understory and overstory vegetation, using detailed leaf-level physiology (for both understory and overstory vegetation) and soil respiration property measurements as well as a multilayer soil-vegetation-atmosphere transfer (SVAT) model. The modeling results showed that the larch overstory's leaf area index (LAI) and vertical profile of leaf photosynthetic capacity were major factors determining the flux partitioning in this boreal forest ecosystem. This is unlike other forest ecosystems that tend to have dense LAI. We concluded that control of the larch overstory's LAI had a relationship with both the coexistence of the larch with the cowberry understory and with the water resources available to the total forest ecosystem.     

Intertidal invertebrates locally enhance primary production

The contribution of autochthonous vs. allochthonous inputs to productivity is an important determinant of ecosystem function across multiple habitats. In coastal marine systems, nutrients are thought to come primarily from the upwelling of deep, nutrient-rich water. Using experimental manipulations of a dominant tide pool animal, the mussel Mytilus californianus, I show that the presence of mussels greatly increases the supply of inorganic nitrogen and phosphorus. Mussels further had a direct effect on productivity: benthic microalgal abundance increased by a factor of 4-8, while the growth of a red alga was four times greater in the presence of mussels. The increase in nitrite and nitrate associated with mussels further suggests nitrifying activity by microbes. These findings have broad implications for coastal marine systems, including that regenerated nutrients may contribute more to productivity than previously recognized and that the presence of animal-generated nutrients sets the stage for numerous positive interactions.     

The development of a multi-species algal ecodynamic model for urban surface water systems and its application

An ecodynamic model that can simulate four phytoplankton species has been developed to deal with the unique characteristics of urban river systems which has manmade river profile, flow controlled by gates, severe eutrophication status, and fragile aquatic ecosystem. The ecodynamic model was developed referencing two typical models: the water quality simulation model WASP and ecological model CAEDYM. The model can simulate 11 state variables: dissolved oxygen, carbonaceous biochemical oxygen demand, ammonia nitrogen, nitrate nitrogen, organic nitrogen, inorganic phosphorus, organic phosphorus and four phytoplankton species with zooplankton as a boundary condition. The ecodynamic model was applied to Sihai section of the Beijing urban river system, where serious algal blooms broke out in recent years. The dominant phytoplankton species are Cyanophyta, Chlorophyta, Bacillariophyta, and Cryptophyta. Site-specific data on geometry, meteorology, pollution sources, and existing ecosystem parameters were collected and used for model calibration and verification The model results mimic observed trends of water quality and phytoplankton species succession and can be used for forecasting algal blooms as well as assessment of river management measures.     

Predation affects the susceptibility of hard clam Meretrix lusoria to Hg-stressed birnavirus

Predator–prey interaction in aquatic ecosystem is one of the simplest drivers affecting the species population dynamics. Predation controls are recognized as important aspects of ecosystem husbandry and management. In this paper we investigated how predation control cause an increase in host growth in the abundance of hard clam (Meretrix lusoria) populations subject to mercury (Hg)-stressed birnavirus. Here we linked predator–prey relationships with a bioenergetic matrix population model (MPM) associated with a susceptible–infectious–mortality (SIM) model based on a host–pathogen–predator framework to quantify the predator effects on population dynamics of disease in hard clam populations. Our results indicated that relative high predation rates could promote the hard clam abundances in relation to predators that selectively captured the infected hard clam, by which the disease transmission was suppressed. The results also demonstrated that predator-induced modifications in host behavior could have potential negative or positive effects on host growth depending on relative species density and resource dynamics. The most immediate implication of this study for the management of aquatic ecosystem is that, beyond the potential for causing a growth in abundance, predation might provoke greater predictability in aquatic ecosystem species populations and thereby increase the safety of ecosystem production from stochastic environmental events.     

Assessing effects of non-native crayfish on mosquito survival

Introductions of non-native predators often reduce biodiversity and affect natural predator–prey relationships and may increase the abundance of potential disease vectors (e.g., mosquitoes) indirectly through competition or predation cascades. The Santa Monica Mountains (California, U.S.A.), situated in a global biodiversity hotspot, is an area of conservation concern due to climate change, urbanization, and the introduction of non-native species. We examined the effect of non-native crayfish (Procambarus clarkii) on an existing native predator, dragonfly nymphs (Aeshna sp.), and their mosquito larvae (Anopheles sp.) prey. We used laboratory experiments to compare the predation efficiency of both predators, separately and together, and field data on counts of dragonfly nymphs and mosquito larvae sampled from 13 local streams. We predicted a lower predation efficiency of crayfish compared with native dragonfly nymphs and a reduced predation efficiency of dragonfly nymphs in the presence of crayfish. Dragonfly nymphs were an order of magnitude more efficient predators than crayfish, and dragonfly nymph predation efficiency was reduced in the presence of crayfish. Field count data showed that populations of dragonfly nymphs and mosquito larvae were strongly correlated with crayfish presence in streams, such that sites with crayfish tended to have fewer dragonfly nymphs and more mosquito larvae. Under natural conditions, it is likely that crayfish reduce the abundance of dragonfly nymphs and their predation efficiency and thereby, directly and indirectly, lead to higher mosquito populations and a loss of ecosystem services related to disease vector control.     

A system-dynamic model on the competitive growth between Potamogeton malaianus Miq. and Spirogyra sp.

A system-dynamic model has been built to evaluate the competition between submerged macrophytes Potamogeton malaianus Miq. (PM) and filamentous green algae Spirogyra sp. (SP). The data background is based on a spring–summer and an autumn–winter experiment carried out in artificial field ponds. The experiments had the aim to acquire a knowledge base necessary to a successful restoration of submerged macrophyte vegetation in Lake Taihu, China by use of P. malaianus Miq. The model mainly focuses on variations in water volume; biomass dynamics of P. malaianus Miq., Spirogyra sp. and zoobenthos; nutrients cycling between water column, P. malaianus Miq., Spirogyra sp., zoobenthos, detritus and sediment. Sixteen state variables are included in the model: biomass of P. malaianus Miq., Spirogyra sp. and zoobenthos; nitrogen in sediments, detritus, in P. malaianus Miq., in Spirogyra sp. and zoobenthos; total dissolved nitrogen; phosphorus in sediments, detritus, in P. malaianus Miq., in Spirogyra sp. and in zoobenthos; total dissolved phosphorus, and water volume of the experiment pond. The calibration and validation of the model show a good accordance with the results of the spring–summer experiment and the autumn–winter experiment.     

Non-linear ecological processes, fires, environmental heterogeneity and shrub invasion in northwestern Patagonia

The emergent behaviors of nature are not only the sum of interactions among ecosystem parts but also depend on the organization of these interactions. Fire, climate and vegetation patterns produce non-linear fire propagation across the landscape. Environmental heterogeneity, like outcrop presence and hare density, increases landscape patchiness and makes possible the occupation of fire refuges by plants, like Fabiana imbricata shrubs. We monitored shrub recruitment and cover during nine postfire years in northwestern Patagonia grasslands and we studied the long-term population dynamics under different environmental conditions through a matrix model, exploring different fire frequencies and spring precipitation regimes. Both, the field monitoring and the model seem to confirm the relationships between shrub invasion and fire. The climate change forecast predicts an increase in the frequency of El Niño Southern Oscillation phenomena that could causes more coupled fires—rainy springs in northwestern Patagonia, and consequently, more recruitment windows for shrubs, like F. imbricata. The matrix model also indicates that this scenario would be the most favourable for shrub invasion. Our results contribute to the knowledge of the ecosystem properties and processes, providing useful information to improve the grasslands sustainable use.     

有机磷细菌的分离鉴定及其对MC-RR的响应(The Isolation and Identification of Organic Phosphorus Bacteria and Its Response to MC-RR)

从云南滇池水样中分离出具有解磷能力的有机磷细菌P-2,并利用现代分子生物学技术进行了初步鉴定.用0.01、5mg·L-1微囊藻毒素(MC-RR)处理有机磷细菌P-2,研究了MC-RR对其生长、细胞内酸碱磷酸酶活性(ACP和AKP)以及培养液中可溶性磷酸盐含量的影响.结果表明,高浓度MC-RR能显著抑制有机磷细菌的生长,延缓其细胞增殖,抑制细胞内酸碱磷酸酶活性以及培养液中可溶性磷酸盐含量的升高,因而可能改变或减缓生态系统中磷循环的进程,这表明微囊藻毒素在一定程度上可能调节水体细菌功能群落.     

Estimating the critical phosphorus loading of shallow lakes with the ecosystem model PCLake: Sensitivity, calibration and uncertainty

There is a vast body of knowledge that eutrophication of lakes may cause algal blooms. Among lakes, shallow lakes are peculiar systems in that they typically can be in one of two contrasting (equilibrium) states that are self-stabilizing: a ‘clear’ state with submerged macrophytes or a ‘turbid’ state dominated by phytoplankton. Eutrophication may cause a switch from the clear to the turbid state, if the P loading exceeds a critical value. The ecological processes governing this switch are covered by the ecosystem model PCLake, a dynamic model of nutrient cycling and the biota in shallow lakes. Here we present an extensive analysis of the model, using a three-step procedure. (1) A sensitivity analysis revealed the key parameters for the model output. (2) These parameters were calibrated on the combined data on total phosphorus, chlorophyll-a, macrophytes cover and Secchi depth in over 40 lakes. This was done by a Bayesian procedure, giving a weight to each parameter setting based on its likelihood. (3) These weights were used for an uncertainty analysis, applied to the switchpoints (critical phosphorus loading levels) calculated by the model. The model was most sensitive to changes in water depth, P and N loading, retention time and lake size as external input factors, and to zooplankton growth rate, settling rates and maximum growth rates of phytoplankton and macrophytes as process parameters. The results for the ‘best run’ showed an acceptable agreement between model and data and classified nearly all lakes to which the model was applied correctly as either ‘clear’ (macrophyte-dominated) or ‘turbid’ (phytoplankton-dominated). The critical loading levels for a standard lake showed about a factor two uncertainty due to the variation in the posterior parameter distribution. This study calculates in one coherent analysis uncertainties in critical phosphorus loading, a parameter that is of great importance to water quality managers.     

海门市蛎岈山牡蛎礁海洋特别保护区生态系统健康评价

在分析海门市蛎岈山牡蛎礁海洋特别保护区海域生态系统现状及特点基础上,构建了包含环境、结构、稳定性3个方面24个指标的海洋特别保护区生态系统健康评价指标体系,并采用熵权综合指数模型和模糊综合评价模型对保护区的生态系统健康状况进行评价及对比分析。熵权综合指数法评价结果表明,保护区生态系统处于健康趋向亚健康的状态;熵权模糊综合模型评价结果表明,保护区生态系统处于健康状态。由于模糊综合评价模型能反映保护区各生态要素的健康状况,其评价结论更趋近于保护区客观现状,更适用于海洋保护区健康评价。     

On the role of entropy in water quality modelling

The use of the entropy principle in phenomenological water quality models is not only necessary, but also of great advantage. A deterministic ecosystem model must obey the 2nd law of thermodynamics. Gibb's equation is a constraint additional to the balances of mass, energy and momentum. The entropy principle supports the unified treatment of physical, chemical and biological processes in water bodies, offers stability criteria and controls the further development of the aquatic ecosystems. Thermodynamic criteria also allow the determination of the bifurcation points of the model equations. Especially near these points the state and structure of the ecosystem can be strongly changed by fluctuations of the variables and parameters of the ecosystem.Results of the thermodynamic theory of selforganizing systems (Glansdorff and Prigogine, 1971; Nicolis and Prigogine, 1977) are of very great importance for water quality modelling. Furthermore, the entropy principle bridges the phenomenological, stochastic and cybernetic approaches to water quality modelling.While the paper deals with general aspects of the role of entropy in water quality modelling, the basic system of equations, taking the entropy principle into account, can be found in a previous paper (Mauersberger, 1978).     

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.     

Using the functional response of a consumer to predict biotic resistance to invasive prey

Predators sometimes provide biotic resistance against invasions by nonnative prey. Understanding and predicting the strength of biotic resistance remains a key challenge in invasion biology. A predator's functional response to nonnative prey may predict whether a predator can provide biotic resistance against nonnative prey at different prey densities. Surprisingly, functional responses have not been used to make quantitative predictions about biotic resistance. We parameterized the functional response of signal crayfish (Pacifastacus leniusculus) to invasive New Zealand mud snails (Potamopyrgus antipodarum; NZMS) and used this functional response and a simple model of NZMS population growth to predict the probability of biotic resistance at different predator and prey densities. Signal crayfish were effective predators of NZMS, consuming more than 900 NZMS per predator in a 12-h period, and Bayesian model fitting indicated their consumption rate followed a type 3 functional response to NZMS density. Based on this functional response and associated parameter uncertainty, we predict that NZMS will be able to invade new systems at low crayfish densities (< 0.2 crayfish/m2) regardless of NZMS density. At intermediate to high crayfish densities (> 0.2 crayfish/m2), we predict that low densities of NZMS will be able to establish in new communities; however, once NZMS reach a threshold density of -2000 NZMS/m2, predation by crayfish will drive negative NZMS population growth. Further, at very high densities, NZMS overwhelm predation by crayfish and invade. Thus, interacting thresholds of propagule pressure and predator densities define the probability of biotic resistance. Quantifying the shape and uncertainty of predator functional responses to nonnative prey may help predict the outcomes of invasions.     

Development of Lake Ladoga ecosystem models: modeling of the phytoplankton succession in the eutrophication process. I

New models of Lake Ladoga ecosystem and the results of modeling are presented. In the first part the model of phytoplankton succession in the process of anthropogenic eutrophication of the lake is considered under the evolution of the phosphorus loading. The still continued anthropogenic eutrophication of the lake started in 1962 when the phosphorus load began to increase. Since 1962 during the evolution of the lake’s state from oligotrophic to developed mezotrophic one, the structure of phytoplankton community dominating species was significantly changed as well as its total productivity. The system state in the model is described by 14 parameters: nine phytoplankton complexes, zooplankton, dissolved organic matter, detritus, dissolved mineral phosphorus and dissolved oxygen. The number of parameters of this model is noticeably larger than that of previous models created by the authors. The relative dynamics of phytoplankton complexes in the lake’s ecosystem evolution was simulated by the new model. It is shown that the modeling results are adequately corresponding to the observation data. The results of phytoplankton structure modeling allow to estimate the impact of phytoplankton on the water quality as well as give the prediction of the lake’s ecosystem evolution with the changes of the phosphorus loading.