Evidence for the dominance of indirect effects in 50 trophic ecosystem networks

Indirect effects are powerful influences in ecosystems that may maintain species diversity and alter apparent relationships between species in surprising ways. Here, we applied network environ analysis to 50 empirically-based trophic ecosystem models to test the hypothesis that indirect flows dominate direct flows in ecosystem networks. Further, we used Monte Carlo based perturbations to investigate the robustness of these results to potential error in the underlying data. To explain our findings, we further investigated the importance of the microbial food web in recycling energy-matter using components of the Finn Cycling Index and analysis of environ centrality. We found that indirect flows dominate direct flows in 37/50 (74.0%) models. This increases to 31/35 (88.5%) models when we consider only models that have cycling structure and a representation of the microbial food web. The uncertainty analysis reveals that there is less error in the I/D values than the ±5% error introduced into the models, suggesting the results are robust to uncertainty. Our results show that the microbial food web mediates a substantial percentage of cycling in some systems (median = 30.2%), but its role is highly variable in these models, in agreement with the literature. Our results, combined with previous work, strongly suggest that indirect effects are dominant components of activity in ecosystems.     

Food web structure and trophic interactions of the tropical highland lake Hayq, Ethiopia

We generated a mass-balance model to figure out the food web structure and trophic interactions of the major functional groups of the Ethiopian highland Lake Hayq. Moreover, the study lay down a baseline data for future ecosystem-based investigations and management activities. Extensive data collection has been taken place between October 2007 and May 2009. Ecotrophic efficiency (EE) of several functional groups including phytoplankton (0.8) and detritus (0.85) was high indicating the utilization of the groups within the system. However, the EE of Mesocyclops (0.03) and Thermocyclops (0.30) was very low implying these resources were rather a ‘sink’ in the trophic hierarchy. Flows based on aggregated trophic level sensu Lindeman revealed the importance of both phytoplankton and detritus to higher trophic levels. The computed average transfer efficiency of 11.5% for the first four trophic levels was within the range for highly efficient African lakes. The primary production to respiration (P/R) ratio (1.05) of Lake Hayq indicates the maturity of the ecosystem. We also modeled the food-web by excluding Tilapia and reduced phytoplankton biomass to get insight into the mass balance before Tilapia was introduced. The analysis resulted in a lower system omnivory index (SOI = 0.016) and a reduced P/R ratio (0.13) that described the lake as immature ecosystem, suggesting the introduction of Tilapia might have contributed to the maturity of the lake. Tilapia in Lake Hayq filled an ecological empty niche of pelagic planktivores, and contributed for the better transfer efficiency observed from primary production to fish yield.     

Ecological role of stomatopods (mantis shrimps) and potential impacts of trawling in a marine ecosystem of the southeast coast of India

Ecosystems are balanced by nature and each component in the system has a role in the sustenance of other components. A change in one component would invariably have an effect on others. Stomatopods (mantis shrimps) are common and ecologically important predatory crustaceans in tropical marine waters. The ecological role of mantis shrimps and potential impacts of trawling in a marine ecosystem were estimated using Ecopath with Ecosim (EwE) Version 5.0 software, by constructing a mass balanced Ecopath model of Parangipettai (Porto Novo) ecosystem. Based on fisheries information from the region, 17 ecological groups were defined including stomatopods. Both primary and secondary data on biomass, P/B, Q/B and diet composition were used as basic inputs. The mass balanced model gave a total system throughput of 14,756 t km⁻² year⁻¹. The gross efficiency of 0.000942 indicated higher contribution of lower food chain groups in the fishery though the mean trophic level was 3.08. The immature and developing stage of the ecosystem was indicated by the ratio of total primary production and total respiration (1.832) and the net system production (2643.30 t km⁻² year⁻¹). Key indices (flow to detritus, net efficiency and omnivory index), split mortality rates and mixed trophic impact of different ecological groups were obtained from the model. A flow diagram was constructed to illustrate the trophic interactions, which explained the biomass flows in the ecosystem with reference to stomatopods. Two temporal simulations were made, with 10 year durations in the mass balanced Ecopath model by using ecosim routine incorporated in EwE software. The effect of decrease in biomass of stomatopods in the ecosystem was well defined, in the first run with increase in stomatopod fishing mortality, and the group showed a high positive impact on benthopelagic fish biomass increase (129%). The simulation with increase in trawling efforts resulted in the biomass decline of different ecological groups as elasmobranchs to 1%, stomatopods to 2%, crabs and lobsters to 36%, cephalopods to 63%, mackerel to 78%, and shrimps to 89%. Present study warns stomatopod discards and further increase in trawling efforts in the region and it explained the need for ecosystem based fisheries management practices for the sustainability of marine fisheries.     

Habitat loss, trophic collapse, and the decline of ecosystem services

The provisioning of sustaining goods and services that we obtain from natural ecosystems is a strong economic justification for the conservation of biological diversity. Understanding the relationship between these goods and services and changes in the size, arrangement, and quality of natural habitats is a fundamental challenge of natural resource management. In this paper, we describe a new approach to assessing the implications of habitat loss for loss of ecosystem services by examining how the provision of different ecosystem services is dominated by species from different trophic levels. We then develop a mathematical model that illustrates how declines in habitat quality and quantity lead to sequential losses of trophic diversity. The model suggests that declines in the provisioning of services will initially be slow but will then accelerate as species from higher trophic levels are lost at faster rates. Comparison of these patterns with empirical examples of ecosystem collapse (and assembly) suggest similar patterns occur in natural systems impacted by anthropogenic change. In general, ecosystem goods and services provided by species in the upper trophic levels will be lost before those provided by species lower in the food chain. The decrease in terrestrial food chain length predicted by the model parallels that observed in the oceans following overexploitation. The large area requirements of higher trophic levels make them as susceptible to extinction as they are in marine systems where they are systematically exploited. Whereas the traditional species-area curve suggests that 50% of species are driven extinct by an order-of-magnitude decline in habitat abundance, this magnitude of loss may represent the loss of an entire trophic level and all the ecosystem services performed by the species on this trophic level.     

Trophic relationships in the recently impounded Bagré reservoir in Burkina Faso

The trophic dynamics of Bagré reservoir which has been recently impounded in Burkina Faso was based on the data collected during 1997–1998 period using the Ecopath model and software. Total fish biomass is 22.63 t km⁻² and mainly represents trophic levels (TLs) 2 and 3. The trophic food chain is relatively long and the overall transfer efficiency is quite low. Grazing foodweb based on primary producers is prominent in the reservoir ecosystem and detritus plays a less significant role. Seasonal and long-term variations in water quality have significant influences on the lower TLs clearly showing a bottom-up functioning of the ecosystem. Environmental degradations, such as siltation occurring in the lake, suggest possible risks in limiting ecosystem productivity.     

What is hidden behind the concept of ecosystem efficiency in energy transformation?

The number of energy transformation levels in trophic webs is usually below five, but can be extended up to ten when parasites and hyper-parasites are included. Research on the structure and function of food webs is relevant to the complexity–stability–productivity debate. The aim of this theoretical analysis is to link energetic and connectional aspects of ecosystems with information theory. Updating an energetic model reported by Ricklefs [Ecologia, Zanichelli Editore S.p.A., Bologna, Italy, 1993, p. 896], our approach is integrated with a static analysis of food webs. The length of food webs is theoretically associated with the average ecological efficiency which can be empirically correlated with the effective connectance between species. Furthermore, the advantage of greater complexity when applied to a signalling network is qualitatively addressed.The overall efficiency of energy transformation into biomass throughout a trophic web, in an ecosystem with a given number of species, is the resultant of the various ecological efficiencies, η, at the transitions between the trophic levels. However, we propose that an increment in effective connectance and interspecies connectivity based on a superimposed signalling web may increase the η values, despite the fact that signalling per se has an energetic cost. According to this hypothesis, ecosystem stability would not be necessarily reduced by increasing the number of trophic levels, N, whenever stability in terms of persistence is improved by a cost-efficient regulatory network.     

EcoTroph: Modelling marine ecosystem functioning and impact of fishing

EcoTroph (ET) is a model articulated around the idea that the functioning of aquatic ecosystems may be viewed as a biomass flow moving from lower to higher trophic levels, due to predation and ontogenetic processes. Thus, we show that the ecosystem biomass present at a given trophic level may be estimated from two simple equations, one describing biomass flow, the other their kinetics (which quantifies the velocity of biomass transfers towards top predators). The flow kinetic of prey partly depends on the abundance of their predators, and a top-down equation expressing this is included in the model. Based on these relationships, we simulated the impact on a virtual ecosystem of various exploitation patterns. Specifically, we show that the EcoTroph approach is able to mimic the effects of increased fishing effort on ecosystem biomass expected from theory. Particularly, the model exhibits complex patterns observed in field data, notably cascading effects and ‘fishing down the food web’. EcoTroph also provides diagnostic tools for examining the relationships between catch and fishing effort at the ecosystem scale and the effects of strong top-down controls and fast-flow kinetics on ecosystems resilience. Finally, a dynamic version of the model is derived from the steady-state version, thus allowing simulations of time series of ecosystem biomass and catches. Using this dynamic model, we explore the propagation of environmental variability in the food web, and illustrated how exploitation can induce a decrease of ecosystem stability. The potential for applying EcoTroph to specific ecosystems, based on field data, and similarities between EcoTroph and Ecopath with Ecosim (EwE) are finally discussed.     

Spatial and mass balanced trophic models of La Rinconada Marine Reserve (SE Pacific coast), a protected benthic ecosystem: Management strategy assessment

Steady-state, dynamic, and spatial models were constructed for the benthic system of La Rinconada Marine Reserve off northern Chile (SE Pacific coast). We examined data on biomass, P/B ratios, catches, food spectrum, consumption, and the dynamics of commercial and non-commercial populations using three theoretical frameworks: Ecopath, Ecosim, and Ecospace. The biomass of the scallop Argopecten purpuratus and the clam Tagelus dombeii were the most relevant compartments of the studied ecosystem. Among the carnivores, the functional crab group Cancer spp. was the most relevant. The Rhodophyta was the dominant macroalga compartment of the system. The results obtained using mixed trophic impacts (MTI) showed that the predatory snail Thais chocolata propagated higher magnitudes of direct and indirect effects on the other species or functional groups. The sea star Luidia magallanica and Rhodophyta had the least effects on the remaining compartments. According to the Ecosim estimates (increasing mortality by fishing), the scallop A. purpuratus had the highest impact on the other compartments. The Ecospace model showed similar qualitative and quantitative effects for changes in biomass under three different exploitation scenarios (by subsystems and globally). Nevertheless, the greatest changes were provoked by using the top-down control and the vulnerabilities estimated by Ecosim. System recovery times were highest with increased mortality of the asteroid L. magallanica and the carnivorous snail T. chocolata, suggesting that the sea star could be considered to be a top predator with a top-down control. The F_MSY estimated for the scallop A. purpuratus was close to the F_i originally entered in Ecopath, limiting the design and execution of an exploitation plan within ecologically sustainable boundaries. The situation was different (F_MSY ? F_i) for the other commercial species, making possible multi-species exploitation programs. The Ecospace trophic-spatially explicit model shows a similar pattern of direct and indirect effects generated when exerting exploitation separately by subsystems. Therefore, habitat rotation of fisheries is not justified.     

A process-based model designed for filling of large data gaps in tower-based measurements of net ecosystem productivity

In this paper we present a simple hybrid gap-filling model (GFM) designed with a minimum number of parameters necessary to capture the ecological processes important for filling medium-to-large gaps in Flux data. As the model is process-based, the model has potential to be used in filling large gaps exhibiting a broad range of micro-meteorological and site conditions. The GFM performance was evaluated using “Punch hole” and extrapolation experiments based on data collected in west-central New Brunswick. These experiments indicated that the GFM is able to provide acceptable results (r² > 0.80) when >500 data points are used in model parameterization. The GFM was shown to address daytime evolution of NEP reasonably well for a wide range of weather and site conditions. An analysis of residuals indicated that for the most part no obvious trends were evident; although a slight bias was detected in NEP with soil temperature. To explore the portability of the GFM across ecosystem types, a transcontinental validation was conducted using NEP and ancillary data from seven ecosystems along a north-south transect (i.e., temperature–moisture gradient) from northern Europe (Finland) to the Middle East (Israel). The GFM was shown to explain over 75% of the variability in NEP measured at most ecosystems, which strongly suggests that the GFM maybe successfully applied to forest ecosystems outside Canada.     

From individuals to ecosystem function: toward an integration of evolutionary and ecosystem ecology

An important goal in ecology is developing general theory on how the species composition of ecosystems is related to ecosystem properties and functions. Progress on this front is limited partly because of the need to identify mechanisms controlling functions that are common to a wide range of ecosystem types. We propose that one general mechanism, rooted in the evolutionary ecology of all species, is adaptive foraging behavior in response to predation risk. To support our claim, we present two kinds of empirical evidence from plant-based and detritus-based food chains of terrestrial and aquatic ecosystems. The first kind comes from experiments that explicitly trace how adaptive foraging influences ecosystem properties and functions. The second kind comes from a synthesis of studies that individually examine complementary components of particular ecosystems that together provide an integrated perspective on the link between adaptive foraging and ecosystem function. We show that the indirect effects of predators on plant diversity, plant productivity, nutrient cycling, trophic transfer efficiencies, and energy flux caused by consumer foraging shifts in response to risk are qualitatively different from effects caused by reductions in prey density due to direct predation. We argue that a perspective of ecosystem function that considers effects of consumer behavior in response to predation risk will broaden our capacity to explain the range of outcomes and contingencies in trophic control of ecosystems. This perspective also provides an operational way to integrate evolutionary and ecosystem ecology, which is an important challenge in ecology.     

Simulation of thermodynamic transmission in green roof ecosystem

Green roofs entail the creation of vegetated space on the top of artificial structures. They can modify the thermal properties of buildings to bring cooling energy conservation and improve human comfort. This study evaluates the thermodynamic transmission in the green roof ecosystem under different vegetation treatments. Our model simulation is based on the traditional Bowen ratio energy balance model (BREBM) and a proposed solar radiation shield effectiveness model (SEM). The BREBM investigates energy absorption of different components of radiation, and the SEM evaluates the radiation shield effects. The proposed model is tested and validated to be efficient to simulate solar energy transmission in green roofs, with some major findings. Firstly, the solar radiation transmission processes might be considered as free vibration motion. Daytime positive heat storage of the green roof is 350-520 W·m⁻² on an hourly basis. Nighttime or afternoon negative value registers a rather constant magnitude of −60 W·m⁻². Daily net average is positive around 155-210 W·m⁻². Secondly, solar radiation vibration is highly correlated with plant structure. The canopy reflectance and transmittance are strongly correlated (R² = 0.87). The multi-layer shrub treatment has the highest shield effectiveness (0.34), followed by two-layer groundcover (0.27), and single-layer grass (0.16). Green roof vegetation absorbs and stores large amounts of heat to form an effective thermal buffer against daily temperature fluctuation. Vegetated roofs drastically depress air temperature in comparison with bare ground (control treatment). Finally, the thermodynamic model is relatively simple and efficient for investigating thermodynamic transmission in green roof ecosystem, and it could be developed into a broad solar radiant land cover model.     

Inter-annual variability of ecosystem production in boreal jack pine forests (1975-2004) estimated from tree-ring data using CBM-CFS3

We describe and apply a method of using tree-ring data and an ecosystem model to reconstruct past annual rates of ecosystem production. Annual data on merchantable wood volume increment and mortality obtained by dendrochronological stand reconstruction were used as input to the Carbon Budget Model of the Canadian Forest Sector (CBM-CFS3) to estimate net ecosystem production (NEP), net primary production (NPP), and heterotrophic respiration (Rh) annually from 1975 to 2004 at 10 boreal jack pine (Pinus banksiana Lamb.) stands in Saskatchewan and Manitoba, Canada. From 1975 (when sites aged 41-60 years) to 2004 (when they aged 70-89 years), all sites were moderate C sinks except during some warmer than average years where estimated Rh increased. Across all sites and years, estimated annual NEP averaged 57 g Cm⁻² yr⁻¹ (range −31 to 176 g Cm⁻² yr⁻¹), NPP 244 g Cm⁻² yr⁻¹ (147-376 g Cm⁻² yr⁻¹), and Rh 187 g Cm⁻² yr⁻¹ (124-270 g Cm⁻² yr⁻¹). Across all sites, NPP was related to stand age and density, which are proxies for successional changes in leaf area. Regionally, warm spring temperature increased NPP and defoliation by jack pine budworm 1 year previously reduced NPP. Our estimates of NPP, Rh, and NEP were plausible when compared to regional eddy covariance and carbon stock measurements. Inter-annual variability in ecosystem productivity contributes uncertainty to inventory-based assessments of regional forest C budgets that use yield curves predicting averaged growth over time. Our method could expand the spatial and temporal coverage of annual forest productivity estimates, providing additional data for the development of empirical models accounting for factors not presently considered by these models.     

Integrated model projections of climate change impacts on a North American lake

Climate change is likely to impact terrestrial and aquatic ecosystems via numerous physical and biological mechanisms. This study outlines a framework for projecting potential impacts of climate change on lakes using linked environmental models. Impacts of climate drivers on catchment hydrology and thermal balance in Onondaga Lake (New York State) are simulated using mechanistic models HSPF and UFILS4. Outputs from these models are fed into a lake ecosystem model, developed in AQUATOX. Watershed simulations project increases in the magnitude of peak flows and consequent increases in catchment nutrient export as the magnitude of extreme precipitation events increases. This occurs concurrently with a decrease in annual stream discharge as a result of increased evapotranspiration. Simulated lake water temperatures increase by as much as 5 °C during the 2040-2069 time period, accompanied by a prolonging of the duration of summer stratification. Projected changes include shifts in the timing of nutrient cycling between lake sediments and water column. Plankton taxa projected to thrive under climate change include green algae and Bosmina longirostris. Responses for species at higher trophic levels are mixed. Benthic macroinvertebrates may either prosper (zebra mussels) or decline (chironomids), while fish (e.g., gizzard shad) exhibit high seasonal variability without any clear trend.     

Calculating ecological carrying capacity of shellfish aquaculture using mass-balance modeling: Narragansett Bay, Rhode Island

Increasing growth in the aquaculture industry demands ecosystem-based techniques for management if that growth is to be ecologically sustainable and promote equity among users of the ecosystems in which it occurs. Models of carrying capacity can be used to responsibly limit the growth of aquaculture in increasingly crowded coastal areas. Narragansett Bay, Rhode Island, USA is one such crowded coastal region experiencing a rapid increase in bivalve aquaculture. An ecosystem mass-balance model was used to calculate the ecological carrying capacity of bivalve aquaculture. Cultured oyster biomass is currently at 0.47 t km⁻² and could be increased 625 times without exceeding the ecological carrying capacity of 297 t km⁻². This translates to approximately 38,950 t of harvested cultured oysters annually which is 4 times the total estimated annual harvest of finfish. This potential for growth is due to the high primary productivity and large energy throughput to detritus of this ecosystem. Shellfish aquaculture has potential for continued growth and is unlikely to become food limited due, in part, to the large detritus pool.     

The stability of trophic mass-balance models of marine ecosystems: a comparative analysis

Dynamic simulations of 18

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mass-balance marine trophic models are used to explore the stability of systems when briefly impacted by a fishery on the key ‘wasp–waist' populations occurring at intermediate trophic levels. The results are related to different ecosystem goal functions previously identified as representative of three attributes of ecosystems development: community complexity, homeostasis and energetics. System recovery time, the time required for all functional groups to returns to baseline level, and here used as a measure of model stability, was inversely correlated to Finn's Cycling Index, i.e. to the fraction of ecosystem throughput that is recycled, and to the mean length of trophic pathways in the systems. Systems with higher capacity to recycle detritus are systems with a higher ability to recover from perturbations. The results are in agreement with the E.P. Odum's theory of ecosystem development, where recycling is interpreted as a chief positive feedback mechanism that contributes to stability in the mature systems by preventing overshoots and destructive oscillations due to external impacts.     

Trophic diversity in amphipods within a temperate eelgrass ecosystem as determined by gut contents and C and N isotope analysis

A conjoint analysis of gut contents and stable C and N isotopes was applied to determine the main food sources and feeding habits of dominant amphipods in an eelgrass bed (Zostera marina) in Gwangyang Bay, Korea. Gut content observations demonstrated that, while Gammaropsis japonicus and Jassa slatteryi are herbivorous, feeding on epiphytes and detritus, Pontogeneia rostrata and Monocorophium acherusicum are omnivorous, feeding on mesozooplankton fragments and detritus. Stable isotope data confirmed that epiphytes, detritus, and mesozooplankton fragments were major food sources for amphipods in the eelgrass bed. Isotopic mixing model calculations clearly showed an interspecific difference in diet composition. A high isotopic dissimilarity between amphipod taxa demonstrated interspecific trophic diversity, reflecting their herbivorous (G. japonicus and J. slatteryi) and omnivorous (P. rostrata and M. acherusicum) feeding habits and confirmed the detrivorous feeding habits of caprellids. Such trophic diversity at interspecific level of the amphipod species indicates that they use different food resources within their microhabitats and play species-specific functional roles as mediators in trophic pathways from producers to higher-level consumers of the eelgrass ecosystem. Finally, our findings suggest that information on the species-specific trophic ecology of amphipods is needed to better understand their potential role in the trophic dynamics and carbon flow of seagrass bed ecosystems.     

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.     

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

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

1.

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

2.

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

3.

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

We obtained the following answers:

1.

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

2.

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

3.

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

     

The swimming crab Charybdis smithii: distribution, biology and trophic role in the pelagic ecosystem of the western Indian Ocean

Surface “swarms” of the swimming crabs Charybdis smithii are still considered as an unusual phenomenon in the open Indian Ocean, although their dense pelagic aggregations were already reported in waters off the Indian coast and in the northern Arabian Sea. Based on an extensive large-scale data series taken over 45 years, we demonstrate that C. smithii is common in the pelagic provinces of the western Indian Ocean driven by the wind monsoon regime. Swimming crabs are dispersed by the monsoon currents throughout the equatorial Indian Ocean. They aggregate at night in the upper 150-m layer, where their estimated biomass derived from pelagic trawling data can exceed 130 kg km⁻². Abundance of C. smithii can reach >15,000 ind. km⁻² in July (i.e. the peak of the south-west monsoon), declines by 50-fold in March and is negligible in May. C. smithii is an important prey for more than 30 species of abundant epipelagic top predators. In turn, it feeds on mesopelagic species. This swimming crab is a major species of the intermediate trophic levels and represents a crucial seasonal trophic link in the open ocean ecosystem of the western Indian Ocean. Outbursts in pelagic waters of huge biomasses of ordinarily benthic crustaceans (C. smithii and Natosquilla investigatoris) are a remarkable feature of the Indian Ocean, although similar, but smaller, events are reported in the Pacific and Atlantic Oceans.

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