Light climate and phytoplankton photosynthesis in maritime Antarctic lakes

The responses of phytoplankton populations to seasonal changes in radiation flux in two Antarctic lakes with extensive winter ice-cover are described. A phytoplankton capable of photosynthesis was found throughout the year in both systems. During winter, low incident radiation combined with thick layers of snow and ice prevented in situ photosynthesis becoming detectable. The beginning of spring was marked by a reduction in snow cover which resulted in a considerable increase in surface penetrating radiation. Planktonic algae rapidly adapted to utilise these increased levels efficiently, though they still showed characteristics of strong shade adaptation.Loss of ice cover at the start of the short open water period further increased the radiation levels and a summer population developed which was much less shade adapted. Saturation and photoinhibition effects were widespread during this period as the algae proved unable to utilise high radiation levels efficiently. They were however effective at the radiation fluxes prevalent in the lower part of the rapidly circulating water columns.     

Convergent succession of phytoplankton in microcosms with different inoculum species composition

Summary Different initial mixtures of phyto-and zooplankton from different lakes were grown under identical chemical and physical conditions in medium size (8-and 12–1) laboratory microcosm cultures until convergence of phytoplankton species composition was attained. Five such experiments with four (four experiments) or three (one experiment) microcosm cultures were run. Three experiments were performed with weak stirring which permitted sedimentary elimination of the diatoms. Two experiments were conducted with stronger stirring to prevent sedimentation. In the three sedimentation intensive experiments, the final phytoplankton community was composed of the filamentous chlorophyte Mougeotia thylespora together with a smaller biomass of nanoplanktic algae. In the two sedimentation free experiments the final phytoplankton community consisted of pennate diatoms. Both dissolved nutrient concentrations and the chemical composition of biomass suggested strong nutrient limitation of algal growth rates in the final phase of the experiments. The zooplankton communities at the end of the experiments were composed of species that were apparently unable to ingest the large, dominant algae and that presumably fed on the nanoplanktic undergrowth and the bacteria. There was a distinct sequence of events in all experiments: first, the large zooplankton species (Daphnia and Copepoda) were replaced by smaller ones (Chydorus, Bosmina, rotifers); second, all cultures within one experiment developed the same nutritional status (limitation by the same nutrient); and third, the taxonomic composition of phytoplankton of the different cultures within one experiment converged. The last took 7–9 weeks, with is about 2–3 times as long as the time needed in a phytoplankton competition experiment to reach the final outcome.     

Leading indicators of phytoplankton transitions caused by resource competition

Many important transitions in phytoplankton composition of lakes and oceans are related to shifts in nutrient supply ratios. Some phytoplankton transitions, such as cyanobacteria blooms in freshwater supplies and red tides in coastal oceans, are important for aquatic resource management. Therefore, it would be useful to have leading indicators which precede phytoplankton shifts and could be readily monitored in the field. We investigated potential indicators using a well-understood model of phytoplankton dynamics parameterized to mimic the transition toward cyanobacteria blooms in freshwater lakes. In stationary distributions, performance of the indicators depends on whether the species are capable of stable coexistence over a certain range of nutrient inputs. In transient simulations, however, indicators show consistent responses regardless of the possibility of stable coexistence. Leading indicators occurring 10 to 40 days prior to species shift include shift of lag-1 autoregression coefficient toward 0, low standard deviation, fluctuating skewness, and high kurtosis. These responses are different from those reported for critical transitions such as fold bifurcations. Thus, the indicators reveal clues to the mechanisms of important ecosystem transitions. In practice, indicators should be measured for multiple ecosystem variables, and interpretation of the indicators should be guided by experiments and mechanistic site-specific models to help resolve potential ambiguities.     

The long-term effect of artificial destratification on phytoplankton species composition in a subtropical reservoir

1. The response of phytoplankton to the installation of an artificial destratification system in North Pine Dam, Brisbane (Australia) was investigated over an 18 year period (1984–2002); 11 years before and 7 years after installation. 2. An overall increase in phytoplankton abundance was revealed for some groups (in particular, diatoms, cyanobacteria and chlorophytes), but not for others (chlorophytes). Changes in the abundance of chlorophyte functional groups was attributed to eutrophication. 3. A strong spatial gradient in phytoplankton abundance and chlorophyll a was observed, with low abundance in the downstream regions affected by the destratification system which was likely because of light limitation induced by vertical mixing. The upstream region acted as a surrogate for the unaltered state of the reservoir, particularly as an indicator of eutrophication without direct influence from the destratification system. Despite the continuous trend in eutrophication of the reservoir, there has been a definite decrease in the rate of eutrophication (approximately 30%) since the installation of the destratification system at the downstream locations. 4. Correlations of the dominant cyanobacteria Cylindrospermopsis raciborskii with other genera changed after destratification, indicating that prior to destratification the dominance of Cylindrospermopsis was because of its ability to compete for phosphorus, whereas after destratification its dominance was because of its ability to compete for light.     

Competition and resource availability in an annual plant community dominated by an invasive species,Carrichtera annua (L. Aschers.), in South Australia

The last decade has seen spirited debates about how resource availability affect the intensity of competition. This paper examines the effect that a dominant introduced species, Carrichtera annua, has upon the winter annual community in the arid chenopod shrublands of South Australia. Manipulative field experiments were conducted to assess plant community response to changing below-ground resource levels and to the manipulation of the density of C. annua. Changes in the density of C. annua had little effect on the abundance of all other species in the guild. Nutrient addition produced an increase in the biomass of the most abundant native species, Crassula colorata. An analysis of the root distribution of the main species suggested that the areas of soil resource capture of C. annua and C. colorata are largely segregated. Our results suggest that intraspecific competition may be stronger than interspecific competition, controlling the species responses to increased resource availability. The results are consistent with a two-phase resource dynamics systems, with pulses of high resource availability triggering growth, followed by pulses of stress. Smaller plants were nutrient limited under natural field conditions, suggesting that stress experienced during long interpulse phases may override competitive effects after short pulse phases. The observed differences in root system structure will determine when plants of a different species are experiencing a pulse or an interpulse phase. We suggest that the limitations to plant recruitment and growth are the product of a complex interplay between the length and intensity of the pulse of resource availability, the duration and severity of the interpulse periods, and biological characters of the species.     

The phytoplankton species composition and the seasonal periodicity in Lake Vechten from 1956 to 1979

The species composition and seasonal periodicity of the phytoplankton in Lake Vechten (The Netherlands) have been studied over a 20 year period. The results show a more or less regular pattern of algal succession during the year. Phytoplankton growth starts early in the year leading to a maximum in early spring, dominated by Bacillariophyceae and some Chlorophyceae. In summer a second maximum develops mainly consisting of a chrysophycean and a dinophycean species, accompanied by several Chlorophyceae. In September the summer species are gradually succeeded by diatoms. By the end of the summer stratification in October algal numbers decrease rapidly. The winter community consists of several species of Cryptophyceae and Chrysophyceae.When the data of 1975–1979 are compared with those from earlier records, distinct changes can be observed. These changes are discussed with reference to the trophic status of the lake. The vertical distribution of some species is described.     

Factors regulating summer phytoplankton in a highly eutrophic Antarctic lake

Lakes from Maritime Antarctica are regarded as systems generally inhabited by metazoan plankton capable of imposing a top-down control on the phytoplankton during short periods, while lakes from Continental Antarctica lacking these communities would be typically controlled by scarcity of nutrients, following a bottom-up model. Otero Lake is a highly eutrophic small lake located on the NW of the Antarctic Peninsula, which has no metazoan plankton. During summer 1996, we studied the density, composition and vertical distribution of the phytoplankton community of this lake with respect to various abiotic variables, yet our results demonstrated neither light nor nutrient limitation of the phytoplankton biomass. Densities of heterotrophic nanoflagellates (HNAN) and ciliates from three different size categories were also studied. Extremely low densities of HNAN (0–155 ind. ml^–1) could be due to feeding competition by bacterivore nanociliates and/or predation by large ciliates. A summer bloom of the phytoflagellate Chlamydomonas aff. celerrima Pascher reached densities tenfold those of previous years (158.10³ ind. ml^–1), though apparently curtailed by a strong peak of large ciliates (107 ind. ml^–1) which would heavily graze on PNAN (phototrophic nanoflagellates). Top-down control can thus occur in this lake during short periods of long hydrologic residence time.     

Succession of phytoplankton species in indoor reservoir models varying in sediment composition and water inlet site

Using four continuous-flow indoor reservoir models, the combined effects of varying water inlet depth and sediment composition on phytoplankton species composition and its succession pattern were studied. Species which are known to bloom in eutrophicated reservoirs were dominant in the species succession in each tank. At any one time, a step-like pattern was observed in the composition of the dominant species of each tank, wherein all the species which appeared in a tank with a lower phosphorus concentration were included in the species composition of the tank with a higher phosphorus concentration in an additive relationship.     

Importance of intermediate disturbances for the species composition and diversity of phytoplankton in two very different Berlin lakes

Species composition and diversity of phytoplankton were studied for several years in two lakes which differ with respect to mixing conditions and nutrient limitation: Schlachtensee regularly stratifies very stably. In contrast, size and wind-exposure predispose Lake Tegel to deeper mixing; additionally, stratification is artificially destabilized by aeration. As the duration of aeration was varied, the study period includes interannual changes in mixing conditions. For both lakes, it also covers trophic change due to restoration; this was especially pronounced in Schlachtensee.Results show that mixing conditions affect species composition on two levels: on a superordinate level, lake morphology or hydrology govern stability of stratification and susceptibility to perturbation, and hence the extent to which motile species can develop. In Schlachtensee, species with some means of actively seeking preferred depths usually dominated during summer stratification: Planktothrix agardhii during the hypertrophic phase, and flagellates since restoration. In contrast, in Lake Tegel deeper mixing as a generally prevailing condition favored non-motile species. Their seasonal pattern was remarkably constant from year to year. Although changes in the extent of mixing were pronounced during the four years studied (1987–1990), these were within a range that affected species composition only slightly: in summer, cyanobacteria and diatoms represented climax species whose dominance was not offset by additional, weather-induced increases of turbulence.On a subordinate level, and within the constraints set by nutrient limitation as well as by grazing pressure, small-scale changes in mixing conditions caused by meteorological cycles were shown to strongly affect species composition and in consequence diversity: Results for the fouryear post-restoration study period at Schlachtensee show that considerable interannual variations of species composition and diversity can be attributed to variations in the frequency of meteorological changes. In accordance with the intermediate disturbance hypothesis (IDH), diversity was lowest during 1989, the year with the longest and most pronounced cycles of fair weather (14 to 27 fair days on end). However, the mechanism for this was rarely a decline of diversity caused by competitive exclusion within single long phases of stable conditions, as conceived by the intermediate disturbance hypothesis. Instead, diversity responded to changes in mixing conditions with a variety of patterns — often with low values during phases of increased mixing and with high values under quiescent conditions, especially during the first calm days just after increased mixing. Thus, not disturbance as such, but rather the rate of change between phases of disturbance and quiescence appears to determine the frequency of high diversity indices.In Lake Tegel, high diversity indices were somewhat more frequent in 1989, the year during which thermal stratification was most stable. For species adapted to frequent or continuous mixing, interjected calm phases with unusually high stability of thermal stratification may represent a disturbance of accustomed conditions. Thus, in turbulent Lake Tegel, meteorological cycles appear to act in reversal to the IDH, in a sense which may be termed intermediate quiescence hypothesis.Phosphorus limitation due to successful restoration was found to decrease winter and vernal diversity in Schlachtensee. Presumably, without nutrient constraints, new populations could grow more rapidly in response to the rapid changes of physical parameters during this season. In contrast, restoration has increased summer diversity, as phosphorus concentrations no longer allow the virtual monocultures of Planktothrix agardhii which prevailed previously. In Lake Tegel, the reduction of phosphorus concentration down to 60 µg/l P in 1989 limited biomass, but this level was still too high to significantly alter species composition or diversity.     

Temporal plankton dynamics in an oligotrophic maritime Antarctic lake

• 1 The population density, diversity and productivity of the microbial plankton in an oligotrophic maritime Antarctic lake were studied for a 15‐month period between December 1994 and February 1996.
• 2 In the lake, concentrations of nutrients and dissolved organic carbon were uniformly low, temperature varied over a small annual range of 0.1–3 °C, and the surface was ice‐covered except during a period of approximately 6 weeks in summer.
• 3 The total of 57 morphotypes of protozoa observed during the study is a higher taxonomic diversity than previously reported from continental Antarctic lakes, but lower than that found in more eutrophic maritime Antarctic lakes. Likewise, planktonic abundance and productivity were lower than has been reported in other lakes on Signy Island, but generally higher than those of lakes on the Antarctic continent.
• 4 There were marked seasonal and interannual variations in planktonic population density.
• 5 Chlorophyll a concentrations ranged from undetectable to 4.2 µg L^‐1 and the greatest rate of primary productivity measured was 4.5 mg C m^‐3 h^‐1. The phytoplankton was dominated by small chlorophytes and chrysophytes, with phototrophic nanoflagellate abundance ranging from 1.1 × 10³ to 1.2 × 10⁷ L^‐1.
• 6 Bacterial densities of 3.6 × 10⁸ to 1.9 × 10¹⁰ L^‐1 were recorded and bacterial productivity reached a peak of 0.36 µg C L^‐1 h^‐1. Numbers of heterotrophic nanoflagellates between 5.0 × 10⁴ and 1.8 × 10⁷ L^‐1, and of ciliates from undetectable to 1.1 × 10⁴ L^‐1 were observed. Naked amoebae were usually rare, but occasionally reached peaks of up to 1.5 × 10³ L^‐1.

     

Resource partitioning and competition for shells between intertidal hermit crabs on the outer coast of Washington

Summary Resource partitioning was quantified for 6 species of intertidal hermit crabs in the genus Pagurus, that occur on the outer coast of Washington. This, together with field evidence of shell shortage and with laboratory experiments to quantify the mechanism of interactions for shells, allowed estimation of the relative intensities of inter-and intraspecific competition between these species. The findings were that: (1) the magnitude of intraspecific competition was greater than any single interspecific competitive effect for all of the species; and (2) the relative proportion of intraspecific competition was greater for the middle and upper intertidal species than for the lower intertidal species. Studies at several outer coast sites supported these generalizations. Both of these findings are consistent with the hypothesis that competitive divergence has occurred in the past. The structure of the outer coast hermit crab assemblage is compared with that of the San Juan Archipelago hermit crab assemblage. Differences between the two do not seem to be the result of adaptive responses to the presence of more competing species in the former group.     

Two species competition in a periodic environment

The classical Lotka-Volterra equations for two competing species have constant coefficients. In this paper these equations are studied under the assumption that the coefficients are periodic functions of a common period. As a generalization of the existence theory for equilibria in the constant coefficient case, it is shown that there exists a branch of positive periodic solutions which connects (i.e. bifurcates from) the two nontrivial periodic solutions lying on the coordinate axes. This branch exists for a finite interval or spectrum of bifurcation parameter values (the bifurcation parameter being the average of the net inherent growth rate of one species). The stability of these periodic solutions is studied and is related to the theory of competitive exclusion. A specific example of independent ecological interest is examined by means of which it is shown under what circumstances two species, which could not coexist in a constant environment, can coexist in a limit cycle fashion when subjected to suitable periodic harvesting or removal rates.Research supported by National Science Foundation Grant No. MCS-7901307     

Patterns of gammaridean amphipod abundance and species composition associated with dominant subtidal macroalgae from the western Antarctic Peninsula

The communities of gammaridean amphipods associated with eight dominant macroalgal species were examined near Palmer Station, Western Antarctic Peninsula. A total of 78,415 individuals belonging to 32 amphipod taxa were identified with mean densities ranging up to 20 individuals/g algal wet wt. The most abundant amphipod taxon, Metaleptamphopus pectinatus, was found to associate predominately with the brown alga Desmarestia menziesii, while the second most common taxon, Jassa spp. occurred primarily on the red alga Gigartina skottsbergii. Non-metric multidimensional scaling analysis demonstrated that the population densities of each amphipod species and amphipod species composition were similar on the same algal species but dissimilar on different species of algae. Comparisons of amphipod communities associated with a given algal species but from different sampling sites indicated that although the structure of species-specific macroalgal-associated amphipod communities can vary across spatial scales of 3 km, 50% of the macroalgal species examined showed no significant inter-site differences in associated amphipod community structure. Spearman rank correlation analyses showed that higher abundances of amphipods occurred on the macroalgae with the highest number of branches. As many Antarctic amphipods are known consumers of macroalgae, their remarkable abundances are likely to play a significant role in mediating energy and nutrient transfer in nearshore Antarctic Peninsular macroalgal communities.     

An analysis of competitive interactions between 3 hermit crab species

Summary Competition for empty gastropod shells in a group of three sympatric hermit crabs (Pagurus hirsutiusculus, Pagurus granosimanus, and Pagurus beringanus) was studied in the San Juan Archipelago, Washington State. Estimates of the competitive effects of each species on the others' shell supplies were derived using field data on shell utilization and the results of laboratory experiments to determine rates of acquisition and exchange of shells and preferences for different shell species. Each species experienced approximately an order of magnitude more intraspecific competition than interspecific competition for empty shells. This resulted from differences in preference for shell shapes, shell size use, and habitat use between P. hirsutiusculus and P. granosimanus, and largely from differences in habitat use between P. beringanus and the other two species. Experiments involving the release and recensusing of marked empty shells were used to estimate competitive effects more directly for the interaction between P. hirsutiusculus and P. granosimanus. Results were consistent with the estimates derived from data on resource partitioning. Possible causes of the low levels of interspecific competition are discussed, and results are compared with studies of other organisms that estimated both inter- and intra-specific competition.     

Temperature-influenced species competition in mass cultures of marine phytoplankton

Five marine phytoplankton species (Phaeodactylum tricornutum, Thalassiosira pseudonana, Skeletonema costatum, Monochrysis lutheri, and Dunaliella tertiolecta) were grown in enriched laboratory continuous cultures and natural populations were mass cultured outdoors for 16 months. Competition among the species was shown to be highly dependent on temperature, although the actual production of plant organic matter at the low growth rates used was relatively independent of this variable. Control of marine species in mass cultures does not appear economically feasible, but this drawback may be overcome by selecting herbivorous shellfish that are capable of assimilating those temperature-dependent phytoplankton species dominating in a particular locale.     

Foraging for nutrients, responses to changes in light, and competition in tropical deciduous tree seedlings

We evaluated (1) the responses of two co-occurring tropical tree species, Heliocarpuspallidus and Caesalpiniaeriostachys, to changes in light, (2) the ability of these species to search for and exploit a fertilized soil patch, (3) the relationship between the capacity to forage for a fertilized patch and the capacity to respond to changes in light availability and (4) how the relationship between light and nutrient acquisition influenced the competitive interactions between these species. Plants of the two species were exposed to a factorial combination of high (H) and low (L) light intensity and fertilized (+Fp) and unfertilized (−Fp) nutrient patches for 50 days. Half of the plants from H were then transferred to L (HL treatment), and half of the plants from L were transferred to H (LH). The remaining plants were kept in their original light condition and grown for another 50 days. Plants were grown in these light and patch treatments alone (one plant per pot) and in interspecific competition (one plant per species resulting in two plants per pot). Both species exploited fertilized patches by increasing root biomass and length in the patch. This enhanced plant productivity and growth rate mainly under LH and HH conditions for Heliocarpus and the HH condition for Caesalpinia). When plants in the HH light environment were grown with an unfertilized patch, plant biomass and relative growth rates (RGRs) were even lower than␣under the LL light environment [(HH–Fp)<LL]. However, the combined activity of shoot and roots when above- and below-ground resources were temporally and spatially heterogeneous influenced plant productivity and growth rate. The benefit from light increase (LH) was reduced when grown with an unfertilized patch. Larger reductions in root biomass, length and density in the patch, and in plant biomass and RGR, were exhibited by Heliocarpus than by Caesalpinia. These results suggest a close relationship between root foraging and light capture, where the benefit of the exploitation of the patch will be reflected in whole-plant benefit, if enough light is captured above-ground. In addition, the results suggest a change in the expected plant responses to light due to heterogeneity in soil nutrients, even though the fertilized patch was only a small proportion of the total soil volume. Leaf characteristics such as specific leaf area responded only to light conditions and not to patchily distributed nutrients. Root characteristics responded more strongly to nutrient heterogeneity. Competition modified the pattern of foraging under both high- and low-light conditions in Heliocarpus by 50 days, and the ability to forage for a fertilized patch under LL after 100 days of growth for Caesalpinia. Even though plant growth and productivity are greatly reduced under low-light conditions (HL and LL), competition modifies the ability of species to forage for a rich patch (especially for the fast-growing species Heliocarpus). Received: 24 November 1997 / Accepted: 15 June 1998     

Potential phytoplankton indicator species for monitoring Baltic coastal waters in the summer period

There are very few time series documenting clear trends of change in the biomass of total phytoplankton or single taxa that coincide with trends of increasing nutrient concentrations. Weekly or biweekly monitoring since 1997 on a cross section of the central Gulf of Finland (NE Baltic Sea) with similar climatic and hydrographic conditions, but different nutrient levels, provided a uniform dataset. In order to evaluate seasonal (June–September) patterns of phytoplankton succession, more than 1,200 samples were statistically analyzed by selecting 12 dominant taxa using wet weight biomass values. In addition, the continuously measured hydrographic parameters on board the ships of opportunity, and simultaneous nutrient analyses gave high frequency information on the water masses. The objective of this study was to identify the taxa that may prove indicative in the assessment of eutrophication in the appropriate monitoring time periods. None of the most common bloom-forming species (Aphanizomenon sp., Nodularia spumigena, and Heterocapsa triquetra) showed reliable correlations with enhanced nutrient concentrations. The species we suggest as reliable eutrophication indicators—oscillatorialean cyanobacteria and the diatoms Cyclotella choctawhatcheeana and Cylindrotheca closterium—showed the best relationships with total phosphorus concentrations. Their maxima appear toward the end of July or in August–September when phytoplankton community structure is more stable, and less frequent observations may give adequate results. Another diatom, Skeletonema costatum, exhibited stronger correlations with dissolved inorganic and total nitrogen in June, during the period of the summer phytoplankton minimum. Guest editors: J. H. Andersen & D. J. Conley Eutrophication in Coastal Ecosystems: Selected papers from the Second International Symposium on Research and Management of Eutrophication in Coastal Ecosystems, 20–23 June 2006, Nyborg, Denmark     

Aggregation pattern of individuals and the outcomes of competition within and between species: Differential equation models

The influence of spatial distribution pattern on the outcomes of intra- and interspecific competition is studied theoretically. The models developed are the generalized logistic andVolterra equations, whereLloyd 's indices of intra- and interspecies mean crowding were incorporated with their assumed linear relationship to mean density in order to express the intensity of crowding which is really effective to the existing individuals. It is shown that while the increasing patchiness of distribution has a pronounced effect of promoting the intraspecific competition and lowering the equilibrium density for individual populations, it generally relaxes the interspecific competition, making it easy for different species sharing the same niche, which would otherwise be incompatible, to coexist stably. These models thus provide a simplest theoretical basis to explain why many insect populations in nature are kept relatively rare in number and why a number of allied species often coexist freely sharing the same resource, against the “competitive exclusion principle” deduced from the originalVolterra equations.     

Effects of light and nutrient availability on dry matter and N allocation in six successional grassland species

Summary Recent discussions on determinants of competitive success during succession require the study of the combined effect of light and nutrient availability on growth and allocation. These effects can be used to predict the outcome of competition at changing resource availabilities. This work is part of a study on the successional sequence in permanent grassland starting after fertilizer application is stopped, but with continued mowing, in order to restore former species-rich communities. This yields a successional sequence which proceeds from grasslands with a high nutrient availability and a closed canopy, to grasslands with a low nutrient availability and an open canopy. If allocation is related to competitive ability, species from the productive stages would be expected to allocate more biomass and nitrogen to leaves, which could make them better competitors for light, while species from the unproductive stages would allocate more biomass to roots, which could make them better nutrient competitors. This study reports on growth, specific leaf area (SLA), vertical display of leaves, and allocation of biomass and nitrogen of six grassland species from this successional sequence at 16 combinations of light and nutrient supply. Species from the poorer successional stages reached a lower final dry weight than species from the richer stages, over all treatment combinations. The experimental design made it possible to test for unique effects of the resource ratio effect of light and nutrients on allocation characteristics. This resource-ratio effect was defined as the ratio light intensity/(light intensity + nutrient supply rate), using standardized levels for the treatments. The within-species variation (plasticity) in both allocation of dry matter and nitrogen was linearly related to this resource-ratio effect. Some interspecific differences in this relationship were found which could be related to the position of the species along the successional gradient. However, the range of plasticity in allocation pattern expressed within each species was much larger than the differences between species. It was concluded that allocation differences between these grassland species are relatively unimportant, given the large amount of plasticity in these traits. Interspecific differences in SLA and vertical stature seemed to be more important in explaining the position of species along the successional gradient.     

The vertical distribution of phytoplankton in stratified water columns

What determines the vertical distribution of phytoplankton in different aquatic environments remains an open question. To address this question, we develop a model to explore how phytoplankton respond through growth and movement to opposing resource gradients and different mixing conditions. We assume stratification creates a well-mixed surface layer on top of a poorly mixed deep layer and nutrients are supplied from multiple depth-dependent sources. Intraspecific competition leads to a unique strategic equilibrium for phytoplankton, which allows us to classify the distinct vertical distributions that can exist. Biomass can occur as a benthic layer (BL), a deep chlorophyll maximum (DCM), or in the mixed layer (ML), or as a combination of BL+ML or DCM+ML. The ML biomass can be limited by nutrients, light, or both. We predict how the vertical distribution, relative resource limitation, and biomass of phytoplankton will change across environmental gradients. We parameterized our model to represent potentially light and phosphorus limited freshwater lakes, but the model is applicable to a broad range of vertically stratified systems. Increasing nutrient input from the sediments or to the mixed layer increases light limitation, shifts phytoplankton towards the surface, and increases total biomass. Increasing background light attenuation increases light limitation, shifts the phytoplankton towards the surface, and generally decreases total biomass. Increasing mixed layer depth increases, decreases, or has no effect on light limitation and total biomass. Our model is able to replicate the diverse vertical distributions observed in nature and explain what underlying mechanisms drive these distributions.