Habitat connectivity and ecosystem productivity: implications from a simple model

The import of resources (food, nutrients) sustains biological production and food webs in resource-limited habitats. Resource export from donor habitats subsidizes production in recipient habitats, but the ecosystem-scale consequences of resource translocation are generally unknown. Here, I use a nutrient-phytoplankton-zooplankton model to show how dispersive connectivity between a shallow autotrophic habitat and a deep heterotrophic pelagic habitat can amplify overall system production in metazoan food webs. This result derives from the finite capacity of suspension feeders to capture and assimilate food particles: excess primary production in closed autotrophic habitats cannot be assimilated by consumers; however, if excess phytoplankton production is exported to food-limited heterotrophic habitats, it can be assimilated by zooplankton to support additional secondary production. Transport of regenerated nutrients from heterotrophic to autotrophic habitats sustains higher system primary production. These simulation results imply that the ecosystem-scale efficiency of nutrient transformation into metazoan biomass can be constrained by the rate of resource exchange across habitats and that it is optimized when the transport rate matches the growth rate of primary producers. Slower transport (i.e., reduced connectivity) leads to nutrient limitation of primary production in autotrophic habitats and food limitation of secondary production in heterotrophic habitats. Habitat fragmentation can therefore impose energetic constraints on the carrying capacity of aquatic ecosystems. The outcomes of ecosystem restoration through habitat creation will be determined by both functions provided by newly created aquatic habitats and the rates of hydraulic connectivity between them.     

Vertical ecology of the pelagic ocean: classical patterns and new perspectives

Applications of acoustic and optical sensing and intensive, discrete‐depth sampling, in concert with collaborative international research programmes, have substantially advanced knowledge of pelagic ecosystems in the 17 years since the 1996 Deepwater Fishes Symposium of the Fisheries Society of the British Isles. Although the epipelagic habitat is the best‐known, and remote sensing and high‐resolution modelling allow near‐synoptic investigation of upper layer biophysical dynamics, ecological studies within the mesopelagic and deep‐demersal habitats have begun to link lower and upper trophic level processes. Bathypelagic taxonomic inventories are far from complete, but recent projects (e.g. MAR‐ECO and CMarZ, supported by the Census of Marine Life programme) have quantitatively strengthened distribution patterns previously described for fishes and have provided new perspectives. Synthesis of net and acoustic studies suggests that the biomass of deep‐pelagic fishes may be two to three orders of magnitude greater than the total global commercial fisheries landings. Discrete‐depth net sampling has revealed relatively high pelagic fish biomass below 1000 m in some regions, and that gelatinous zooplankton may be key energy vectors for deep‐pelagic fish production. Lastly, perhaps, the most substantive paradigm shift is that vertical connectivity among fishes across classical depth zones is prevalent– suggesting that a whole‐water column approach is warranted for deep ocean conservation and management.     

Ecosystem response to earlier ice break‐up date: Climate‐driven changes to water temperature,lake‐habitat‐specific production,and trout habitat and resource use

Climate warming has yielded earlier ice break‐up dates in recent decades for lakes leading to water temperature increases, altered habitat, and both increases and decreases to ecosystem productivity. Within lakes, the effect of climate warming on secondary production in littoral and pelagic habitats remains unclear. The intersection of changing habitat productivity and warming water temperatures on salmonids is important for understanding how climate warming will impact mountain ecosystems. We develop and test a conceptual model that expresses how earlier ice break‐up dates influence within lake habitat production, water temperatures and the habitat utilized by, resources obtained and behavior of salmonids in a mountain lake. We measured zoobenthic and zooplankton production from the littoral and pelagic habitats, thermal conditions, and the habitat use, resource use, and fitness of Brook Trout (Salvelinus fontinalis). We show that earlier ice break‐up conditions created a "resource‐rich" littoral–benthic habitat with increases in zoobenthic production compared to the pelagic habitat which decreased in zooplankton production. Despite the increases in littoral–benthic food resources, trout did not utilize littoral habitat or zoobenthic resources due to longer durations of warm water temperatures in the littoral zone. In addition, 87% of their resources were supported by the pelagic habitat during periods with earlier ice break‐up when pelagic resources were least abundant. The decreased reliance on littoral–benthic resources during earlier ice break‐up caused reduced fitness (mean reduction of 12 g) to trout. Our data show that changes to ice break‐up drive multi‐directional results for resource production within lake habitats and increase the duration of warmer water temperatures in food‐rich littoral habitats. The increased duration of warmer littoral water temperatures reduces the use of energetically efficient habitats culminating in decreased trout fitness.     

Cross-habitat effects shape the ecosystem consequences of co-invasion by a pelagic and a benthic consumer

Invasive species can have major impacts on ecosystems, yet little work has addressed the combined effects of multiple invaders that exploit different habitats. Two common invaders in aquatic systems are pelagic fishes and crayfishes. Pelagic-oriented fish effects are typically strong on the pelagic food web, whereas crayfish effects are strong on the benthic food web. Thus, co-invasion may generate strong ecological responses in both habitats. We tested the effects of co-invasion on experimental pond ecosystems using two widespread invasive species, one pelagic (western mosquitofish) and one benthic (red swamp crayfish). As expected, mosquitofish had strong effects on the pelagic food web, reducing the abundance of Daphnia and causing a strong trophic cascade (increase in phytoplankton). Crayfish had strong effects on the benthic food web, reducing the abundance of benthic filamentous algae. Yet, we also found evidence for important cross-habitat effects. Mosquitofish treatments reduced the biomass of benthic filamentous algae, and crayfish treatments increased Daphnia and phytoplankton abundance. Combined effects of mosquitofish and crayfish were primarily positively or negatively additive, and completely offsetting for some responses, including gross primary production (GPP). Though co-invasion did not affect GPP, it strongly shifted primary production from the benthos into the water column. Effects on snail abundance revealed an interaction; snail abundance decreased only in the presence of both invaders. These results suggest that cross-habitat effects of co-invaders may lead to a variety of ecological outcomes; some of which may be unpredictable based on an understanding of each invader alone.     

Carbon and Nitrogen Cycling in a Shallow Productive Sub-Tropical Coastal Embayment (Western Moreton Bay, Australia): The Importance of Pelagic–Benthic Coupling

Climatic variables, water quality, benthic fluxes, sediment properties, and infauna were measured six times over an annual cycle in a shallow sub-tropical embayment to characterize carbon and nutrient cycling, and elucidate the role of pelagic–benthic coupling. Organic carbon (OC) inputs to the bay are dominated by phytoplankton (mean 74%), followed by catchment inputs (15%), and benthic microalgae (BMA; 9%). The importance of catchment inputs was highly variable and dependent on antecedent rainfall, with significant storage of allochthonous OC in sediments following high flow events and remineralization of this material supporting productivity during the subsequent period. Outputs were dominated by benthic mineralization (mean 59% of total inputs), followed by pelagic mineralization (16%), burial (1%), and assimilation in macrofaunal biomass (2%). The net ecosystem metabolism (NEM = production minus respiration) varied between ?4 and 33% (mean 9%) of total primary production, whereas the productivity/respiration (p/r) ranged between 0.96 and 1.5 (mean 1.13). Up to 100% of the NEM is potentially removed via the demersal detritivore pathway. Dissolved inorganic nitrogen (DIN) inputs from the catchment contributed less than 1% of the total phytoplankton demand, implicating internal DIN recycling (pelagic 23% and benthic 19%) and potentially benthic dissolved organic nitrogen (DON) fluxes (27%) or N fixation (up to 47%) as important processes sustaining productivity. Although phytoplankton dominated OC inputs in this system, BMA exerted strong seasonal controls over benthic DIN fluxes, limiting pelagic productivity when mixing/photic depth approached 1.3. The results of this study suggest low DIN:TOC and net autotrophic NEM may be a significant feature of shallow sub-tropical systems where the mixing/photic depth is consistently less than 4.     

罗非鱼对附着藻类和浮游植物的影响

鱼类通过牧食和营养盐排泄可以对水体生态系统产生影响,杂食性鱼类由于可摄食不同生境中的食物,可使生境之间的耦合作用发生变化。罗非鱼是我国南方很多水体的优势种,食物包括敞水生境的浮游植物和基质表层生境的附着藻类等。为了解罗非鱼对浮游植物和附着藻类的影响,实验在室外模拟条件下,分别设置罗非鱼组和无鱼对照组的两组处理,分析了罗非鱼对附着藻类及浮游植物生物量(叶绿素a)等的影响。结果表明:(1)罗非鱼显著地降低了附着藻类生物量,罗非鱼组中的附着藻类叶绿素a的平均值为0.15 mg·cm^-2,显著低于对照组中的1.26mg·cm^-2;(2)罗非鱼显著地增加了浮游植物的生物量,罗非鱼组中的浮游植物叶绿素a平均值为31.99μg·L^-1,显著高于对照组中的14.99μg·L^-1。研究结果显示,杂食性的罗非鱼可以促进系统的附着藻类向浮游植物转化。从控制浮游植物生物量的角度看,湖泊等水体的管理应该对罗非鱼密度加以有效控制。     

Impacts of an invasive snail (Tarebia granifera) on nutrient cycling in tropical streams: the role of riparian deforestation in Trinidad, West Indies

Non-native species and habitat degradation are two major catalysts of environmental change and often occur simultaneously. In freshwater systems, degradation of adjacent terrestrial vegetation may facilitate introduced species by altering resource availability. Here we examine how the presence of intact riparian cover influences the impact of an invasive herbivorous snail, Tarebia granifera, on nitrogen (N) cycling in aquatic systems on the island of Trinidad. We quantified snail biomass, growth, and N excretion in locations where riparian vegetation was present or removed to determine how snail demographics and excretion were related to the condition of the riparian zone. In three Neotropical streams, we measured snail biomass and N excretion in open and closed canopy habitats to generate estimates of mass- and area-specific N excretion rates. Snail biomass was 2 to 8 times greater and areal N excretion rates ranged from 3 to 9 times greater in open canopy habitats. Snails foraging in open canopy habitat also had access to more abundant food resources and exhibited greater growth and mass-specific N excretion rates. Estimates of ecosystem N demand indicated that snail N excretion in fully closed, partially closed, and open canopy habitats supplied 2%, 11%, and 16% of integrated ecosystem N demand, respectively. We conclude that human-mediated riparian canopy loss can generate hotspots of snail biomass, growth, and N excretion along tropical stream networks, altering the impacts of an invasive snail on the biogeochemical cycling of N.     

Development of indicators of ecosystem functioning in a temperate shelf sea: a combined fieldwork and modelling approach

A conceptual model of the main carbon and nitrogen flows through pelagic and benthic food webs was used to identify the key biogeochemical processes representing ecosystem functioning, and to select indicators of each of these processes. A combined fieldwork and modelling approach was used to provide the data required to evaluate the indicators in terms of their suitability for assessing and managing the impacts of climate change and demersal trawling. Four of our 16 proposed indicators (phytoplankton production and productivity, near-bed oxygen concentrations and oxygen penetration of the seabed) met the majority of criteria we used for evaluating indicators. Five indicators (depth of anoxic sediment, zoobenthos biomass, production, productivity and bioturbation potential) did not comply with sufficient criteria to be considered as good indicators. Six of our proposed indicators (zooplankton biomass, size structure, production and productivity; ecosystem productivity; ecosystem balance) could not be assessed for sensitivity and specificity using our models, and therefore need to be addressed in future work aimed at improving both the models and the fieldwork. Our results indicate that evaluation of indicators is difficult, because of the number and variety of human pressures which need to be considered in reality, and the interactions between these pressures and the ecosystem components which they affect. The challenge will be to establish if there are indeed any indicators which are able to meet the majority of criteria for good indicators in holistic ecosystem-based assessments.     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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Integrative ecosystem analyses of phytoplankton dynamics in the York River estuary (USA)

The relative importance of biotic (top-down) vs. abiotic (bottom-up) controls on phytoplankton dynamics was investigated in the York River estuary, Virginia (USA) by a combination of extensive analyses of long-term data sets collected by the U.S. Environmental Protection Agency (EPA) monitoring program over 17 years (1984–2001), field studies (1996–1997) and ecosystem modeling analyses. Results from the analysis of long-term data records collected at three stations along the salinity gradient suggested that phytoplankton are more likely controlled by abiotic mechanisms such as resource limitation than biotic mechanisms such as grazing since annual cycles of primary production and phytoplankton biomass were similar and no grazing effects were evident based on the observed relationship between phytoplankton and zooplankton biomass (R ² < 0.2, p > 0.1). This scenario was supported by short-term field observations made over an annual cycle at three stations in the mid-channel of the estuary where both chlorophyll a and primary production demonstrated similar patterns of seasonal variation. Ratios of fluorescence before and after acidification at all study sites were relatively high suggesting low grazing pressure in the estuary. A tidally-averaged, size-structured plankton ecosystem model was previously developed and verified for the lower York River estuary. The validated ecosystem model was also used to examine this issue and simulation results supported the importance of bottom-up control in the York River estuary.     

Ecosystem engineering by invasive exotic beavers reduces in-stream diversity and enhances ecosystem function in Cape Horn,Chile

Species invasions are of global significance, but predicting their impacts can be difficult. Introduced ecosystem engineers, however, provide an opportunity to test the underlying mechanisms that may be common to all invasive engineers and link relationships between changes in diversity and ecosystem function, thereby providing explanatory power for observed ecological patterns. Here we test specific predictions for an invasive ecosystem engineer by quantifying the impacts of habitat and resource modifications caused by North American beavers (Castor canadensis) on aquatic macroinvertebrate community structure and stream ecosystem function in the Cape Horn Biosphere Reserve, Chile. We compared responses to beavers in three habitat types: (1) forested (unimpacted) stream reaches, (2) beaver ponds, and (3) sites immediately downstream of beaver dams in four streams. We found that beaver engineering in ponds created taxonomically simplified, but more productive, benthic macroinvertebrate assemblages. Specifically, macroinvertebrate richness, diversity and number of functional feeding groups were reduced by half, while abundance, biomass and secondary production increased three- to fivefold in beaver ponds compared to forested sites. Reaches downstream of beaver ponds were very similar to natural forested sections. Beaver invasion effects on both community and ecosystem parameters occurred predominantly via increased retention of fine particulate organic matter, which was associated with reduced macroinvertebrate richness and diversity (via homogenization of benthic microhabitat) and increased macroinvertebrate biomass and production (via greater food availability). Beaver modifications to macroinvertebrate community structure were largely confined to ponds, but increased benthic production in beaver-modified habitats adds to energy retention and flow for the entire stream ecosystem. Furthermore, the effects of beavers on taxa richness (negative) and measures of macroinvertebrate biomass (positive) were inversely related. Thus, while a generally positive relationship between diversity and ecosystem function has been found in a variety of systems, this work shows how they can be decoupled by responding to alterative mechanisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Lake morphometry predicts the degree of habitat coupling by a mobile predator

Habitat coupling is an ecosystem process whereby semi-discontinuous habitats are connected through the movement of energy and nutrients by chemical, physical or biological processes. One oft-cited example is that of littoral–pelagic coupling in lakes. Theory has argued that such habitat coupling may be critical to food web dynamics, yet there have been few empirical studies that have quantified ecological factors that affect the degree of habitat coupling in ecosystems. Specifically, the degree to which habitat coupling occurs across important physical gradients has largely been ignored. To address this, we investigate the degree of littoral habitat coupling (i.e. the degree to which a top predator lake trout, Salvelinus namaycush, derives energy from the littoral zone) along a gradient of lake shape, where lake shape modifies the relative quantity of coupled epilimnetic benthic and pelagic habitats within each lake. Herein we demonstrate that littoral habitat coupling is intensified in simple circular lakes compared to their reticulate counterparts in seven Canadian Shield lakes. Although the more reticulate lakes had larger areas of epilimnetic benthic habitat, littoral food sources comprised 11% compared to 24% of lake trout diet in reticulate and circular lakes, respectively. This heightened interaction in circular lakes also appears to translate into increased omnivory in more circular lakes compared to reticulate lakes such that lake trout of circular lakes have a significantly lower trophic position than lake trout of reticulate lakes (F_1,5=6.71 p=0.05). These results suggest that it is the accessibility of littoral production via thermal refugia, and not the amount of littoral production, that determines the degree to which lake trout couple littoral and pelagic habitats in lakes.     

Sea ice phenology and timing of primary production pulses in the Arctic Ocean

Arctic organisms are adapted to the strong seasonality of environmental forcing. A small timing mismatch between biological processes and the environment could potentially have significant consequences for the entire food web. Climate warming causes shrinking ice coverage and earlier ice retreat in the Arctic, which is likely to change the timing of primary production. In this study, we test predictions on the interactions among sea ice phenology and production timing of ice algae and pelagic phytoplankton. We do so using the following (1) a synthesis of available satellite observation data; and (2) the application of a coupled ice‐ocean ecosystem model. The data and model results suggest that, over a large portion of the Arctic marginal seas, the timing variability in ice retreat at a specific location has a strong impact on the timing variability in pelagic phytoplankton peaks, but weak or no impact on the timing of ice‐algae peaks in those regions. The model predicts latitudinal and regional differences in the timing of ice algae biomass peak (varying from April to May) and the time lags between ice algae and pelagic phytoplankton peaks (varying from 45 to 90 days). The correlation between the time lag and ice retreat is significant in areas where ice retreat has no significant impact on ice‐algae peak timing, suggesting that changes in pelagic phytoplankton peak timing control the variability in time lags. Phenological variability in primary production is likely to have consequences for higher trophic levels, particularly for the zooplankton grazers, whose main food source is composed of the dually pulsed algae production of the Arctic.     

Manipulation of the Pelagic Food Web by Stocking with Predacious Fishes

Two water bodies, which are quite different with respect to nutrient load and hydrophysical conditions, are used to perform long-term experiments in the whole water on the manipulation of the pelagic food web. Experimental water 1: Bautzen Reservoir (Dresden County, GDR); hypereutrophic, mean depth=7.4 m; extremely exposed to wind. Experimental water 2: Small pond in a former quarry (Dresden County); mesotrophic; mean depth=7.0 m; extremely protected against wind. Only the results of Experiment 2 are given in detail. Experiment 1 is not yet finished. Experimental water 2 was investigated in 1979 and 1980 when no predatory fish species were present, and in 1981 after introduction of predators (mainly Salmo gairdneri). The response of the ecosystem can be summarized as follows: (1) The biomass of the zooplankton-eating fish (mainly Leucaspius delineatus) decreases rapidly. (2) The biomass of the herbivorous zooplankton increases to nearly 400%. (3) This finding reveals that the dense population of an invertebrate predator (Chaoborus flavicans) is not able to compensate for the feeding pressure of the small fish. But the intensive feeding activity of the young larvae of Chaoborus leads to a strong increase (200 to 300%) in the mean individual body size of the crustaceans during late summer and autumn, which supports the “balanced predation hypothesis”. (4) The remarkable enhanced grazing pressure of the herbivorous zooplankton on the phytoplankton does not exert any effect on the total phytoplankton biomass. This result is interpreted as a consequence of growth limitation of the algae due to low nutrient (Fe, P) supply in that mesotrophic water body. But the phytoplankton composition does reveal a strong response to the enhanced grazing pressure. The Secchi depth increases as a consequence of this change in the phytoplankton composition. The conclusion is drawn that, when using biomanipulation as a means of water quality management, it is obviously necessary to take into account the complex interrelationships between fish stocks, predacious invertebrates, herbivorous zooplankton, phytoplankton as well as nutrient load and hydrophysical processes in the particular water.     

Morphological differentiation in introduced pumpkinseed Lepomis gibbosus (L.) occupying different habitat zones in Portuguese reservoirs

Introduced pumpkinseed Lepomis gibbosus sampled from four habitat zones (fluvial pelagic, fluvial littoral, lacustrine pelagic and lacustrine littoral) in three Portuguese reservoirs were used to test the hypotheses that habitats with the least similar characteristics will show the most differentiation, and that morphological differences will relate to functional adaptations to flow and trophic habitats. Results from discriminant function analysis and ANCOVA showed that there were significant differences in external morphology in pumpkinseed captured from the four habitat zones in all three reservoirs. Littoral and pelagic differentiation was stronger than fluvial and lacustrine differentiation in all of the reservoirs, and the most significant variable that differentiated pumpkinseed from the littoral and pelagic habitats was body depth. The illustration of external morphological differentiation in pumpkinseed along both habitat dimensions demonstrates the high degree of morphological plasticity of this introduced species.     

Evaluation of the Danish mussel fishery: suggestions for an ecosystem management approach

In Limfjorden, Denmark, an extensive mussel fishery exploits the wild stocks of Mytilus edulis with annual landings of 80,000–100,000 t of mussels. During the last 10 years the impact of mussel dredging on the ecosystem has been studied, including the effect of resuspension of sediment and nutrients and the impoverishment of in- and epi-fauna assemblages. Furthermore, dredging changes the physical structure and complexity of the seabed which affects mussel growth and interactions among zoobenthic species. The blue mussel constitutes the dominant fraction of the zoobenthic suspension feeders, and is important for the transport of material and energy from the pelagic to benthic systems and the control of phytoplankton biomass. In order to evaluate the impact on clearance capacity of a reduction in mussel densities due to mussel dredging, mussel filtration activity measured in situ has been related to the mixing of the water column and the amount of near-bed phytoplankton. Fishery practice for mussel dredging in Limfjorden is discussed in relation to its known impact on the ecosystem and the ecological role of the mussels, and modifications towards an ecosystem management approach and a more sustainable fishery are suggested. The suggested modifications include: a fishery practice where the mussel beds are thinned out when the mussels have attained good quality, and a transplantation practice of mussels from areas with a high mortality to areas with a high growth rate. Both practices intensify the production in a certain area, leaving other areas open for alternative production or for permanent closure for the benefit of the benthic flora and fauna. In addition, other shellfish species represent interesting new resources for fishing or aquaculture. Habitat restoration, such as the relaying of mussel shells from the mussel industry, is another important management tool that should be included in an ecosystem management approach of the mussel fishery. Electronic Publication     

Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

ANTARCTIC AQUATIC ECOSYSTEMS AS HABITATS FOR PHYTOPLANKTON

1. The Southern Ocean is a large-scale, relatively homogeneous upwelling ecosystem whose phytoplankton apparently grows suboptimally over much of its area. By contrast there is a wide variety of freshwater habitats in the Antarctic and in some of these phytoplankton growth efficiency is very high. The two habitats share similar temperature and irradiance regimes, but differ markedly in availability of inorganic nutrients, in grazing pressure and in the time- and space-scales on which various physical processes act. 2. Concentrations of inorganic nutrients in the marine ecosystem have been represented as being in excess of phytoplankton requirements, but the ionic composition of some nutrient pools may not conform to phytoplankton preferences. 3. Nutrient-limitation determines phytoplankton production in Antarctic lakes and gives rise to gross differences between lakes. 4. Irradiance in the water column varies greatly over the year in both marine and freshwater ecosystems. Most algae are shade-adapted, with the ability to utilize low irradiance but with sub-optimal response to high irradiance. However, local phytoplankton maxima may attain very high carbon fixation and growth rates. 5. Consistently low temperatures characterize both systems. Their effects on photo-synthetic carbon uptake mirror shade-adaptation. Division rates of marine phytoplankton may however be very much higher than predicted for ambient temperatures. 6. Vertical mixing is important in both ecosystems and influences the environment experienced by phytoplankton cells. This appears to have little effect on the average performance of phytoplankton in the strongly mixed surface water column of the Southern Ocean, where the mixed depth may exceed 100 m. This can be related partly to the shade-adapted photosynthetic response. Euphotic depths range from 20 to 100 m. 7. Strong vertical mixing under ice-free conditions in lakes may maximize photosynthetic efficiency, whilst distinct vertical stratification in permanently ice-covered lakes gives rise to segregation of nutrient uptake and regeneration. 8. Physical removal of phytoplankton biomass by grazing is locally important in the Southern Ocean, in contrast to the estimated mean mesoscale impact of grazing. Vertical sedimentation losses appear important in the context of mixing depth and generation time, and may be modified by vertical circulation of water. 9. Loss of phytoplankton biomass from lakes during the ice-free period is dominated by physical removal via the lake outflow. Grazing is generally unimportant, except where larvae of otherwise nektobenthic zooplankton hatch in synchrony with a phytoplankton maximum. Sedimentation is important under ice-cover.     

Biomass flow and structure of a tropical upwelling ecosystem in La Guajira, Colombian Caribbean

La Guajira is an exploited tropical upwelling ecosystem in the Colombian Caribbean coast. A trophic model of 27 functional groups was constructed using the ECOPATH 5.0 Beta software to integrate the available information on the ecosystem. The model allowed a comparison with other trophic flow models of upwelling ecosystems. Total system biomass (68 t/km2/year), net system production (1,248.5 t/km2/year), and total system throughput (3,275 t/km2/year) make La Guajira moderate when compared with other systems. The largest amount of energy throughput is achieved from trophic level I to II (68.93 %), although an important proportion of the total flow originates from detritus (32 %). The production/respiration ratio exceeds 1, suggesting that La Guajira is an immature ecosystem and is in development, as determined by its low ascendency (33.7 %) and high development capacity (66.3 %), similar to other upwellings that have values of ascendency between 20 % and 35 %. Although the basic input data were good and covered 1995 to 2000, appropriate information is still not available on some trophic groups such as biomass (for phytoplankton, invertebrates, catfishes and pelagic predator fishes), secondary production data (invertebrates, pelagic predator fishes, and small pelagic fishes), and seabird and mammal populations, which are top trophic levels and an essential part of upwelling ecosystems.