Polymorphism of mannose phosphate isomerase in North Sea and Baltic Sea populations of the amphipods Gammarus zaddachi and G. salinus

Genetic variation at the mannosephosphate isomerase (MPI) locus was examined in the euryhaline sibling species Gammarus zaddachi Sexton and G. salinus Spooner. Both crustacean amphipods share identical enzyme mobilities, following electrophoresis on vertical starch gels. The MPI locus turned out to be highly polymorphic; it is encoded by 6 alleles in G. zaddachi and 7 alleles in G. salinus. Geographic variation of allelic diversity was studied in samples from 9 G. zaddachi and 10 G. salinus populations, primarily obtained from Baltic Sea and North Sea sites. Patterns of inter- and intraspecific heterogeneity are described. Differences in allelic composition exist between Baltic and North Sea samples of G. salinus. In G. zaddachi, levels of polymorphism are higher in North Sea populations than in those from Baltic Sea areas. The significance of these findings is discussed in the light of previous biochemical genetic investigations on the population structure of the two amphipods considered.     

Predator-inducible changes in blue mussels from the predator-free Baltic Sea

Blue mussels, Mytilus spp., have inhabited the brackish Baltic Sea, an environment lacking predatory crabs and starfish, for several thousand years. In this paper we examined whether Baltic Mytilus that were transplanted to the North Sea showed predator-inducible plasticity like their "marine" conspecifics. Our experiments showed that native North Sea Mytilus changed their morphology when exposed to waterborne scents from shore-crabs and starfish. These predators induced different kinds of changes, with emphasis on shell thickness and adductor muscle size, respectively. Baltic Mytilus responded in a similar way to crab scents, whereas starfish scents had a relatively weak effect on the morphology. Crab and starfish scent induced strengthening of the byssal attachment in North Sea Mytilus, with crabs providing more stimulation than starfish. Baltic mussels also improved the byssal attachment when exposed to either of the predators, but the attachment strength, as well as the response to crabs, were relatively weaker than that of North Sea mussels. We conclude that inducible plasticity still is present in Baltic Mytilus, despite their recent evolution in a predator-free environment. There is probably no strong selective pressure against inducible plasticity, but it could also be maintained in the population by gene flow from Mytilus in the adjoining North Sea. The question whether Baltic Mytilus are M. edulis or M. trossulus may also be relevant for the present results.     

Genetic differentiation in Hediste diversicolor (Polychaeta: Nereididae) for the North Sea and the Baltic Sea

Genetic differentiation between North Sea and Baltic Sea Hediste diversicolor (O.F. Müller, 1776) (Polychaeta: Nereididae) populations was studied by allozyme electrophoresis on starch gel. Thirteen loci were analyzed in eight populations. The level of genetic variation was very low (mean H _o = 0.000 to 0.015). Differentiation between H. diversicolor populations is quite high (F _ST = 0.892) and reflected by three enzyme loci (MDH-I*, MDH-II*, IDH-I*). The reduced gene flow (N _m<1) may be explained by the limited dispersal capacity of the species. Regardless of whether found in the North Sea or Baltic Sea, there appear to be two different genetic types which are parapatric or sympatric in some places. The two types hybridize at three localities, but no signs of hybridization have been found at one (Tallinn). Received: 26 June 1997 / Accepted: 10 September 1997     

Icy heritage: ecological evolution of the postglacial Baltic Sea reflected in the allozymes of a living fossil,the priapulid Halicryptus spinulosus

Genetic variation of 16 allozyme loci in 397 Halicryptus spinulosus (Priapulida) revealed overall polymorphism of P=0.438 and Hardy-Weinberg expectations for heterozygosity of H _e=0.060 for Baltic Sea stocks, H _e=0.143 for the White Sea and H _e=0.121 for Iceland. Maximal unbiased standard distances of D=0.0693 separated Baltic and White Sea populations. Nordic and Baltic populations could be distinguished by allozymes, but Baltic subsamples proved cohesive. Gene flow amounted to effective exchange values per generation of N _m=2.94 over 650 km of continuous habitat, N _m=10.65 over 175 km, and N _m=13.85 over 20 km. Gene flow started to decrease with geographic distance beyond a dispersal threshold of 20 km, but hierarchical G _ST-statistics indicated light isolation by distance beyond a minimum of 8 km. Gene flow is high for a benthic worm assumed to lack dispersal by pelagic larva, a paradox which cannot be resolved now. Baltic populations are characterized by lower heterozygosity than Nordic stocks. In the Baltic Basin, temporally continuous brackish-water conditions have only existed for the past 7000 years. The two possible colonization routes of H. spinulosus to the geologically young Baltic Sea imply genetic drift, whether by founder effect (sweepstake colonization from Iceland) or by refugial bottlenecking during the Ancylus phase of the Baltic Basin after a direct connection to the White Sea had been sequestered. Continued genetic drift is emphasized by lower heterozygosity in the ecologically unstable Belt Sea compared to the central Baltic. Allozymes falsify the reduced-mutability hypothesis to explain bradytelic evolution of Priapulida. Regional genetic homogeneity, ample polymorphism, and preference for anoxic black mud qualify H. spinulosus populations as indicators of microevolutionary responses to water circulation regimes or pollution in the Baltic Sea.     

Genetic variability and level of differentiation in North Sea and Baltic Sea populations of the green alga<Emphasis Type="Italic"> Cladophora rupestris</Emphasis>

Cladophora rupestris is a perennial filamentous macroalga belonging to the Chlorophyta. It is widely distributed on both sides of the northern Atlantic Ocean and penetrates into the brackish Baltic Sea down to ca. 4 psu salinity. In this paper we present evidence for genetic differentiation of a Baltic form of this marine alga. We assessed genetic structure within and among 11 populations ranging along a salinity gradient from the Norwegian coast to the northern Baltic Sea proper. Samples of 328 individuals were studied using starch-gel protein electrophoresis to evaluate genetic variability and interpopulation differentiation based on allozymes. Of 11 loci examined, only one was polymorphic. For this locus, encoding superoxide dismutase (SOD-3), a total of seven alleles were distinguished. We found two genetically differentiated groups of populations of C. rupestris, one Baltic Sea group and one North Sea group, with a distinct border in the southern Kattegat near the entrance to the Baltic Sea. The genetic differentiation for SOD-3, expressed as pairwise F_ST values between the populations, was generally higher within the Baltic Sea group (0.10-0.43) than within the North Sea group (0.05-0.10); in the latter group also fewer pairs of populations differed significantly. Pairs of populations from different groups had the highest F_ST values (0.20-0.60). Hierarchical analysis of variance showed that 29.6% of the total variation in the SOD-3 locus was explained by variation between the two groups, while only 4.2% was explained by variation among the populations within the groups. The remaining variation (66.2%) was found within the populations.     

Photosynthetic production and its regulating factors in the Baltic Sea

Measurements of primary production in the Baltic Sea over a period of 2 years (1969/1971) by the ¹⁴C method are reported. The rate of total annual production for the Baltic Sea proper and the Gulf of Finland including coastal areas appears to lie between 35 and 40 gC·m², a rate which can be assumed as characteristic for oligotrophic waters with a low production rate. An apparent increase in productivity between the 2 years could be noted. Calculations for the water mass below the halocline show that between 0.7 and 0.8 ml/l oxygen is used up during a period of 3 months required for the development of anoxic conditions. The major portion of the organic matter in the Baltic Sea undergoes decomposition in the sub-photic zone above the halocline. The ratio of organic material in deep water to the total available oxidizable matter appears to remain constant with time; therefore, the apparent increase in productivity in the photic layer does not affect apprecibly the development of anoxic conditions in deep water. Of the different areas investigated, the Gulf of Finland seems to be the most productive. In the southern Baltic Sea, the rate of production over 5 years between 1966 and 1971 has not changed appreciably. In fact, it shows a trend toward a slight decrease, whereas, in other areas of the Baltic Sea, the rate of production indicates an apparent increase. Comparison with oceanic coastal areas of the west coast of Sweden shows that the rate of production there is higher than in the Baltic Sea. Of the micronutrients controlling productivity in the photic layer of the Baltic Sea, nitrate has a stronger limiting effect on plant growth than phosphate, a fact which is in agreement with existing conceptions on the subject. Dissolved iron does not seem to exert any appreciable influence on the productivity.     

Interactions between cod, herring and sprat in the changing environment of the Baltic Sea: A dynamic model analysis

The interactions between cod (Gadus morhua), herring (Clupea harengus) and sprat (Sprattus sprattus) in the Central Baltic Sea were examined with a simple dynamic model, an alternative to more complicated and data-demanding multispecies and ecosystem models. The main aims of the study were to compare the effect of alternative structures on the model output and examine the control relationships in the fish assemblage under different environmental conditions. The effect of environmental conditions was modelled using a stock-recruitment equation for cod incorporating an environmental index. The model output was especially sensitive to the functional response in predation by cod on herring and sprat. The type II functional response led to a collapse of the clupeid stocks when cod was abundant, while the type III response produced more realistic stock dynamics. According to the simulations, an abundant cod stock was able to keep the sprat stock at a low level, while the herring stock was less affected and benefited from the decreased density of sprat. Simulation of different fishing scenarios indicated that reducing fishing mortality to the level currently advised by ICES would allow the recovery of the cod stock even in unfavourable environmental conditions.     

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.     

Are there two species of the polychaete genus Marenzelleria in Europe?

North Sea and Baltic Sea populations of the introduced polychaete Marenzelleria viridis (Verrill, 1873) reproduce at different times (spring and autumn, respectively). Enzyme separation by starch gel electrophoresis revealed major differences between specimens from the Baltic Sea and those from the North Sea (collected in 1992 and 1993) but a high degree of homogeneity among populations from the same sea. Three enzyme loci, glucose-6-phosphate isomerase (GPI-A, GPI-B) and malate dehydrogenase (MDH), were fixed to 100% by different alleles in the North and Baltic Sea populations, respectively. Different alleles are dominant for mitochondrial aspartate aminotransferase (mAAT) with allele frequencies of ca. 0.97 in all sampled populations from the North Sea and Baltic Sea, respectively, but heterozygotes were found in all populations. These genetic differences could be due to environmentally induced selection or genetically different origins of the populations, suggesting that populations of the genus Marenzelleria in the North and Baltic Seas may be two different species.     

Recruitment and attrition of associated plants under a shading crop canopy: Model selection and calibration

Associated plant and animal diversity provides ecosystem services within crop production systems. The importance of the maintenance or restoration of diversity is therefore increasingly acknowledged. Here we study the population dynamics of associated annual plants (‘weeds’) during the growth of a crop in a season and introduce a minimal model to characterize the recruitment and attrition of the associated plants under the influence of shading by the crop. A mechanistically based, logistic, light interception model was parameterized with light interception measurements in two single crops (barley and rye) and in mixtures of these cereals with peas. Population dynamics data were collected for the annuals Papaver rhoeas, Centaurea cyanus, Chrysanthemum segetum, and Misopates orontium. A minimal population dynamics model was identified for each annual plant species, using system identification techniques as model selection and calibration.     

The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

Carbon, nitrogen, oxygen and sulfide budgets in the Black Sea: A biogeochemical model of the whole water column coupling the oxic and anoxic parts

Carbon, nitrogen, oxygen and sulfide budgets are derived for the Black Sea water column from a coupled physical-biogeochemical model. The model is applied in the deep part of the sea and simulates processes over the whole water column including the anoxic layer that extends from ?115 m to the bottom (?2000 m). The biogeochemical model involves a refined representation of the Black Sea foodweb from bacteria to gelatinous carnivores. It includes notably a series of biogeochemical processes typical for oxygen deficient conditions with, for instance, bacterial respiration using different types of oxidants (i.e denitrification, sulfate reduction), the lower efficiency of detritus degradation, the ANAMMOX (ANaerobic AMMonium OXidation) process and the occurrence of particular redox reactions. The model has been calibrated and validated against all available data gathered in the Black Sea TU Ocean Base and this exercise is described in Gregoire et al. (2008). In the present paper, we focus on the biogeochemical flows produced by the model and we compare model estimations with the measurements performed during the R.V. KNORR expedition conducted in the Black Sea from April to July 1988 (Murray and the Black Sea Knorr Expedition, 1991). Model estimations of hydrogen sulfide oxidation, metal sulfide precipitation, hydrogen sulfide formation in the sediments and water column, export flux to the anoxic layer and to the sediments, denitrification, primary and bacterial production are in the range of field observations.With a simulated Gross Primary Production (GPP) of 7.9 mol C m⁻² year⁻¹ and a Community Respiration (CR) of 6.3 mol C m⁻² year⁻¹, the system is net autotrophic with a Net Community Production (NCP) of 1.6 mol C m⁻² year⁻¹. This NCP corresponds to 20% of the GPP and is exported to the anoxic layer. In order to model Particulate Organic Matter (POM) fluxes to the bottom and hydrogen sulfide profiles in agreement with in situ observations, we have to consider that the degradation of POM in anoxic conditions is less efficient that in oxygenated waters as it has often been observed (see discussion in Hedges et al., 1999). The vertical POM profile produced by the model can be fitted to the classic power function describing the oceanic carbon rate (C_R=Z^−α) using an attenuation coefficient α of 0.36 which is the value proposed for another anoxic environment (i.e. the Mexico Margin) by Devol and Hartnett (2001). Due to the lower efficiency of detritus degradation in anoxic conditions and to the aggregation of particles that enhanced the sinking, an important part of the export to the anoxic layer (i.e. 33%, 0.52 mol C m⁻² year⁻¹) escapes remineralization in the water column and reaches the sediments. Therefore, sediments are active sites of sulfide production contributing to 26% of the total sulfide production.In the upper layer, the oxygen dynamics is mainly governed by photosynthesis and respiration processes as well as by air-sea exchanges. ?71% of the oxygen produced by phytoplankton (photosynthesis+nitrate reduction) is lost through respiration, ?21% by outgasing to the atmosphere, ?5% through nitrification and only ?2% in the oxidation of reduced components (e.g. Mn²⁺, Fe²⁺, H₂S).The model estimates the amount of nitrogen lost through denitrification at 307 mmol N m⁻² year⁻¹ that can be partitioned into a loss of ?55% through the use of nitrate for the oxidation of detritus in low oxygen conditions, ?40% in the ANAMMOX process and the remaining ?5% in the oxidation of reduced substances by nitrate.In agreement with data analysis performed on long time series collected since the 1960s (Konovalov and Murray, 2001), the sulfide and nitrogen budgets established for the anoxic layer are not balanced in response to the enhanced particle fluxes induced by eutrophication: the NH₄ and H₂S concentrations increase.     

Colonization of Europe by two American genetic types or species of the genus Marenzelleria (Polychaeta: Spionidae). An electrophoretic analysis of allozymes

Allozyme electrophoresis was conducted in an attempt to identify the origin ofMarenzelleria sp. found in the North Sea and Baltic Sea. The analysis covered eight enzymes with ten loti from nine populations found on the North American Atlantic toast, these populations in the North Sea and five populations in the Baltic. The North Sea spionids correspond to the Type IMarenzelleria from North American coastal waters between Barnstable Harbor (Massachusetts) and Cape Henlopen (Delaware). Nei's genetic distance between these North American populations and those from the North Sea wasD = 0.010 to 0.020. TheMarenzelleria sp. found in the Baltic Sea very probably stems from North American populations of Type II found from the region of Chesapeake Bay (Trippe Bay) south to the Currituck Sound (North Carolina). The genetic distance between these North American populations and the Baltic populations isD = 0.000 to 0.001. The invaders appear to have lost little of their genetic variation while colonizing the North and Baltic Seas. Probably, both colonizing events tan be attributed to large numbers of individuals reaching Europe simultaneously on one or more occasions. In addition, aMarenzelleria Type III was found by electrophoresis among specimens from Currituck Sound (North Carolina), rohere it is sympatric withMarenzelleria Type II. Salinity is discussed as an important factor for the establishment ofMarenzelleria Type I in the North Sea and Type II in the Baltic Sea.     

Functional divergence in heat shock response following rapid speciation of Fucus spp. in the Baltic Sea

In the Baltic Sea, the broadly distributed brown alga Fucus vesiculosus coexists in sympatry over part of its range (south west Gulf of Bothnia) with the Baltic endemic F. radicans sp. nov, while further north in colder and lower-salinity areas of the Baltic F. radicans occurs alone (north west Gulf of Bothnia). F. radicans appears to have arisen via rapid speciation from F. vesiculosus within the recent history of the Baltic (ca. 7500 BP). Possible functional divergence between the two species was investigated by comparing stress-responsive gene expression in a common-garden experiment. The experiment used two allopatric populations of Fucus vesiculosus from the Skagerrak (North Sea) and Central Baltic, as well as F. radicans from the same Central Baltic site. The two species in sympatry displayed divergent heat shock responses, while F. vesiculosus populations from allopatric sites did not. F. radicans was more sensitive to heat shock at 25°C, either alone or together with high irradiance and desiccation, than Baltic or Skagerrak F. vesiculosus. The results indicate that rapid functional divergence in the inducible heat shock response has occurred between sympatric species on a timescale of thousands of years.     

Modelling cyanobacteria in shallow coastal seas

There is an increasing need to describe cyanobacteria bloom dynamics using ecosystem models. We consider two fundamentally different ways how cyanobacteria are currently implemented: a simple approach without explicit consideration of the life cycle which assumes that cyanobacteria grow due to nitrogen fixation alone and an advanced approach that computes the succession of four different stages of the cyanobacteria life cycle based on internal quotas of energy and nitrogen. To qualitatively and quantitatively intercompare these different approaches and with observations, we use the Baltic Sea ecosystem model ERGOM coupled to the one-dimensional water column model GOTM. Four experiments are carried out: three, using the simple approach with either (a) a prescribed constant minimum production, (b) no minimum value or (c) a prescribed constant minimum concentration, and one with (d) the full predictive life cycle. The model data of 35 years (1970-2005) are analyzed for the timing of the bloom, the interannual variability, the annual mean nitrogen fixation rates and the effect of cyanobacteria on eukaryotic phytoplankton. The results show significant differences. In the climatological seasonal mean, only the advanced approach which resolves the life cycle produces a realistic bloom onset and duration. The interannual variability of blooms is unrealistically small in the experiments with a prescribed minimum value. Annual mean nitrogen fixation rates diverge by up to 30% between the four model solutions. Finally, the representation of the cyanobacteria also influences the seasonal cycle of eukaryotic phytoplankton, i.e., flagellates. This study demonstrates that the way how cyanobacteria are implemented in coupled biological-physical models strongly determines the fluxes into the system and between the individual compartments.     

Large-scale diversity and biogeography of benthic copepods in European waters

A large-scale database concerning benthic copepods from the Arctic, Baltic Sea, North Sea, British Isles, Adriatic Sea and Crete was compiled to assess species richness, biodiversity, communities, ecological range size and biogeographical patterns. The Adriatic showed the highest evenness and the most species-rich communities. Assemblages from the North Sea, British Isles, Baltic and Crete had a lower evenness. The British Isles were characterised by impoverished communities. The ecological specificity of copepod species showed two diverging trends: higher specificity of species in more diverse assemblages was observed in the Adriatic, North Sea and Baltic. A uniformly high species specificity disregarding sample diversity was found on Crete and in the British Isles. Benthic copepod communities showed distinct patterns that clearly fit the predefined geographical regions. Communities were distinguishable and β-diversity was found to be high around Europe, indicating a high species turnover on the scale of this investigation. The British Isles and the North Sea were found to be faunistic links to the Baltic and the Arctic.     

Use of seaweeds for monitoring trace elements in coastal waters

Concentrations of a wide range of trace elements: arsenic, cadmium, cobalt, chromium, hafnium, nickel, thorium, uranium, zinc and the rare earth elements, cerium, europium, samarium, terbium and ytterbium were determined by instrumental neutron activation analysis in the brown alga,Fucus vesiculosus from Eckwarder Hörne, North Sea and from Rügen, Baltic Sea. Another brown alga,Sargassum filipendula from Sri Lanka, Indian ocean (representing an unpolluted control station) was similarly investigated. Cobalt, chromium and nickel concentrations were highest inF. vesiculosus from the North Sea while zinc was highest in samples from the Baltic Sea, reflecting high levels of these elements in coastal waters of the North and the Baltic sea. Cadmium, cobalt, nickel and zinc levels were lowest inS. filipendula from Sri Lanka, probably demonstrating lower levels of those elements in coastal waters. Concentration levels of hafnium, thorium, uranium, and the rare earth elements were highest inS. filipendula. Two years later in 1994,S. filipendula along withUlva sp. (green alga) was resampled from the same sampling site, and in addition to the above elements, six other trace elements (Ag, Ba, Br, Rb, Se and Sr) were determined.Sargassium filipendula showed a particular affinity for Ag, As, Br and Sr. For the other elements, marginal concentration differences were observed betweenS. filipendula andUlva sp., probably reflecting the regional background levels. Substantially higher concentrations of Hf, Th, U, and the rare earths were found again in the 1994Sargassum andUlva samples, reflecting the effect of a substrate rich in rare earth elements. The brown algae used in this study may be used to monitor trace elements in coastal waters.     

New indicator species in the Baltic zooplankton in 1972

The occurrence of zooplankton species, indicating a salinewater influx from the North Sea into the Baltic Sea in 1972, is described. Particular attention is paid to 8 species so far unknown in this region (Ectopleura dumortieri, Rosacea plicata, Tomopteris kefersteini, Aetideus armatus, Centropages typicus, Eucalanus elongatus, Metridia lucens, Electra pilosa). The appearance of these zooplankters in the Southern Baltic Sea is discussed in relation to the hhdrographic changes taking place in this area in 1972. Based on these data, assessments are made regarding the intensity of dynamic exchange processes in the Baltic and North Atlantic waters in 1972.     

Ecosystem states: Creating a data-derived, ecosystem-scale ecological response model that is explicit in space and time

Increasing difficulties associated with balancing consumptive demands for water and achieving ecological benefits in aquatic ecosystems provide opportunities for new ecosystem-scale ecological response models to assist managers. Using an Australian estuary as a case study, we developed a novel approach to create a data-derived state-and-transition model. The model identifies suites of co-occurring birds, fish, benthic invertebrates and aquatic macrophytes (as ‘states’) and the changing physico-chemical conditions that are associated with each (‘transitions’). The approach first used cluster analysis to identify sets of co-occurring biota. Differences in the physico-chemical data associated with each state were identified using classification trees, with the biotic distinctness of the resultant statistical model tested using analysis of similarities. The predictive capacity of the model was tested using new cases. Two models were created using different time-steps (annual and quarterly) and then combined to capture both longer-term trends and more-recent declines in ecological condition. We identified eight ecosystem states that were differentiated by a mix of water-quantity and water-quality variables. Each ecosystem state represented a distinct biotic assemblage under well-defined physico-chemical conditions. Two ‘basins of attraction’ were identified, with four tidally-influenced states, and another four independent of tidal influence. Within each basin, states described a continuum of relative health, manifest through declining taxonomic diversity and abundances. The main threshold determining relative health was whether freshwater flows had occurred in the region during the previous 339 days. Canonical analyses of principal coordinates tested the predictive capacity of the model and demonstrated that the variance in the environmental data set was well captured (87%) with 52% of the variance in the biological data set also captured. The latter increased to >80% when long- and short-term biological data were analysed separately, indicating that the model described the available data for the Coorong well. This approach thus created a data-derived, multivariate model, where neither states nor transitions were determined a priori. The approach did not over-fit the data, was robust to patchy or missing data, the choice of initial clustering technique and random errors in the biological data set, and was well-received by local natural resource managers. However, the model did not capture causal relationships and requires additional testing, particularly during future episodes of ecological recovery. The approach shows significant promise for simplifying management definitions of ecological condition and, via scenario analyses, can be used to assist in manager decision-making of large, complex aquatic ecosystems in the future.