Habitat specialization and the exploitation of allochthonous carbon by zooplankton

The significance of spatial subsidies depends on consumer resource interactions in the recipient habitat. Lakes are subsidized by terrestrial carbon sources, but the pathways of allochthonous carbon through lake food webs are complex and not well understood. Zooplankton vertically partition resources within stratified lakes in response to life history trade-offs that are governed by predators, the quantity and quality of food, and abiotic conditions (e.g., UV, temperature, and viscosity). We measured habitat specialization of zooplankton in an oligotrophic lake where allochthonous and autochthonous resources varied with depth. During stratification, the quantity and quality of zooplankton food was highest in the hypolimnion. We used a yearlong time series of the delta13C of zooplankton and particulate organic matter (POM) to determine which zooplankton species exploited hypolimnetic rather than epilimnetic resources. Because the delta13C of POM decreased with depth, we used the delta13C of zooplankton to detect inter- and intraspecific variation in habitat selection. We incubated Daphnia pulex at discrete depths in the water column to confirm that the delta13C of zooplankton can indicate habitat specialization. Zooplankton that specialized in the epilimnion relied more on allochthonous carbon sources than those that specialized in the hypolimnion. Therefore, the fate of allochthonous carbon subsidies to lakes depends on spatially explicit consumer-resource interactions.     

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.     

Reciprocal subsidies between freshwater and terrestrial ecosystems structure consumer resource dynamics

Cross-ecosystem movements of material and energy, particularly reciprocal resource fluxes across the freshwater-land interface, have received major attention. Freshwater ecosystems may receive higher amounts of subsidies (i.e., resources produced outside the focal ecosystem) than terrestrial ecosystems, potentially leading to increased secondary production in freshwaters. Here we used a meta-analytic approach to quantify the magnitude and direction of subsidy inputs across the freshwater-land interface and to determine subsequent responses in recipient animals. Terrestrial and freshwater ecosystems differed in the magnitude of subsidies they received, with aquatic ecosystems generally receiving higher subsidies than terrestrial ecosystems. Surprisingly, and despite the large discrepancy in magnitude, the contribution of these subsidies to animal carbon inferred from stable isotope composition did not differ between freshwater and terrestrial ecosystems, likely due to the differences in subsidy quality. The contribution of allochthonous subsidies was highest to primary consumers and predators, suggesting that bottom-up and top-down effects may be affected considerably by the input of allochthonous resources. Future work on subsidies will profit from a food web dynamic approach including indirect trophic interactions and propagating effects.     

A model of macroinvertebrate trophic structure and oxygen demand in freshwater wetlands

We present a model of macroinvertebrate trophic structure, detrital cycling, and dissolved oxygen (DO) dynamics in shallow freshwater wetlands with varying allochthonous subsidies. The model is based on field data of primary and secondary production in municipal wastewater-fed and river-fed constructed wetlands in central Ohio, USA. State variables for primary production include macrophyte, periphyton, and metaphyton biomass. Macroinvertebrate biomass is segregated by functional feeding group and includes collectors, scrapers, shredders, and predators. Model simulations demonstrate the association of water column dissolved oxygen, primary production, allochthonous organic matter, and the structure of the macroinvertebrate community. The quality and quantity of allochthonous carbon is shown to have considerable importance, not only as a food source but also as an oxygen sink. Allochthonous carbon equivalent to 5% of autochthonous production increases the macroinvertebrate standing crop by 4–17%, depending on particle size. A large allochthonous subsidy also reduces the simulated average diel dissolved oxygen and increases the percentage of hypoxia-tolerant macroinvertebrates. Simulations show both the heterotrophic response and the changes in community structure brought about by an allochthonous subsidy.     

RETRACTED ARTICLE: Removal of lead and cadmium ions from aqueous solutions using the macroalga Caulerpa racemosa

Sediment bulk parameters and fatty acid biomarkers were used to investigate the relative contribution of major sources of particulate organic matter during upwelling and non-upwelling conditions, at organic enriched sites on the south-eastern Brazilian coast. The degradation state of the organic material and its implications for benthic food webs were also evaluated. Temperature and salinity values indicate the intrusion of the South Atlantic Central Water in spring with a strong influence at station 4S. C:N ratios and fatty acid biomarkers suggest that sedimentary particulate organic matter is mainly autochthonous and originated from phytoplankton, zooplankton and bacteria, with a minor terrestrial contribution. Short-chain saturated fatty acids predominated, indicating the prevalence of partially degraded detritus. An exception was station 4S, where polyunsaturated fatty acids dominated, indicating the input of labile organic matter linked to upwelling of the South Atlantic Central Water. Fatty acid concentrations and the organic carbon content of the sediments suggest that food quantity is not a limiting factor for benthic fauna. Despite the high organic background of the sediments, the sporadic inputs of labile organic matter associated with South Atlantic Central Water intrusions may produce an important impact on benthic fauna and on the biogeochemistry of the sediments.     

Seasonal Fluctuations in the Nutritional Value of Particulate Organic Matter in A Lagoon

Water samples were collected on a fortnightly basis in the lagoon of S. Gilla (Sardinia, Mediterranean Sea) in order to study seasonal nutritional fluctuations of particulate organic matter. the lagoon is characterized by high quantities of suspended matter throughout the year. Thermohaline conditions had no effect on particulate matter quantity and composition, but the quantity as well as quality of suspended particles was drastically affected by the wind, the major effecter of sediment resuspension. As a result of sediment resuspension, seston was always richer in inorganic fraction. However, throughout the year of investigation, most particulate organic carbon was quite appealing for filter feeding communities, although the best POM quality was available during phytoplankton blooming. the phytoplankton pool of suspended matter was just a small fraction of the bulk, accounting for only 13% on average of particulate organic carbon. in terms of energy available in the seston, the highest amount was stored in organic matter heterotrophic fraction, whilst the smallest was to be found in living phytoplankton.     

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

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

Predictive Model of the Effects on Lake Metabolism of Decreased Airborne Litterfall through Riparian Deforestation

The importance of airborne allochthonous litter to the carbon and nutrient budgets of lakes has been seldom studied. We complied data on the input of terrestrial litter to develop a simple and speculative model to predict the potential consequences of riparian deforestation on one aspect of lake metabolism, specifically the balance between phytoplankton production and plankton respiration. During the autumn of 1992, 56 litter traps were deployed around the littoral zones of four oligotrophic lakes in a densely forested region of northwestern Ontario, Canada. The airborne litter input was estimated to be 32 g dry weight per meter of forested shoreline per year. Allochthonous litter input per unit offshore distance was related to the size of riparian trees, their proximity to the shoreline, and the elevation of their canopy. Combining our data with those from other studies suggests that terrestrial litter can contribute up to 15% of the total carbon supply to oligotrophic lakes and up to 10% of the total phosphorus supply to lakes with a large surface area relative to that of their drainage basin. These results were incorporated into a simple model that predicts that removal of shoreline trees could increase the ratio of plankton production to respiration in oligotrophic lakes situated within small drainage basins. Such lakes may therefore shift from allotrophy to increasing autotropy (energy self-sustenance) following riparian deforestation.     

Terrestrial, benthic, and pelagic resource use in lakes: results from a three-isotope Bayesian mixing model

Fluxes of organic matter across habitat boundaries are common in food webs. These fluxes may strongly influence community dynamics, depending on the extent to which they are used by consumers. Yet understanding of basal resource use by consumers is limited, because describing trophic pathways in complex food webs is difficult. We quantified resource use for zooplankton, zoobenthos, and fishes in four low-productivity lakes, using a Bayesian mixing model and measurements of hydrogen, carbon, and nitrogen stable isotope ratios. Multiple sources of uncertainty were explicitly incorporated into the model. As a result, posterior estimates of resource use were often broad distributions; nevertheless, clear patterns were evident. Zooplankton relied on terrestrial and pelagic primary production, while zoobenthos and fishes relied on terrestrial and benthic primary production. Across all consumer groups terrestrial reliance tended to be higher, and benthic reliance lower, in lakes where light penetration was low due to inputs of terrestrial dissolved organic carbon. These results support and refine an emerging consensus that terrestrial and benthic support of lake food webs can be substantial, and they imply that changes in the relative availability of basal resources drive the strength of cross-habitat trophic connections.     

The influence of resource subsidies on cave invertebrates: results from an ecosystem-level manipulation experiment

Spatial resource subsidies can greatly affect the composition and dynamics of recipient communities. Caves are especially tractable for studying spatial subsidies because primary productivity is absent. Here, we performed an ecosystem-level manipulation experiment to test the direct influence of detrital subsidies on community structure in terrestrial cave ecosystems. After performing baseline censuses of invertebrates, we removed all organic material from 12 caves and constructed exclusion boxes to prevent natural resource inputs. Next, we stocked each cave with standardized quantities of two major natural subsidies to caves: leaves (leaf packs) and carcasses (commercially supplied rodents), and measured the invertebrate colonization and utilization of these resources for 23 months. Over the course of the experiment, 102 morphospecies were observed. Diplopods and collembolans were most abundant on leaf packs, and dipteran larvae and collembolans were most abundant on the rats. On average, caves receiving either treatment did not differ in species richness, but abundance was significantly higher in rat caves over both the duration of the experiment and the temporal "life" of the individual resources, which were restocked upon exhaustion. Post-manipulation invertebrate communities differed predictably depending on the type of subsidy introduced. Over the course of the experiment, caves that received the same subsidy clustered together based on community composition. In addition, the invertebrate community utilizing the resource changed over the duration of the two-year experiment, and evidence of succession (i.e., directional change) was observed. Results from this study demonstrate how allochthonous resources can drive the community dynamics of terrestrial invertebrates in cave ecosystems and highlight the need for consideration of the surface environment when managing and protecting these unique habitats.     

Exponential Collapse of Benthic Depurative Capacity in A Eutrophic Tropical Marine Ecosystem: A Threshold Response to Organic Nitrogen Loading

Microbial decomposition of particulate organic matter in sediments can substantially modulate eutrophication of nearshore marine environments. Flux budgets for nitrogen compounds and quantification of rates of microbial transformation can provide important information on the process of eutrophication. This survey documents sediment nitrogen budgets for a eutrophic nearshore marine environment at La Parguera on southwest coast of Puerto Rico, including consideration of the organic fraction in addition to microbial transformations.

Sediments of the inshore channel at La Parguera denote the input of heavy organic loads with low redox potentials and high ammonium contents. Benthic fluxes of dissolved organic nitrogen are below those recorded for ammonium. These observations suggest that nearshore tropical sediments subject to heavy nitrogen loads act as a sink for organic nitrogen and that ammonification is a key process in the release of nitrogen from sediments to the water column. Nitrification and denitrification rates are low overall and inversely related to the redox potential. Depuration of excess nitrogen through denitrification is ineffective in these sediments. in contrast to more robust temperate environments, our work demonstrates that tropical marine systems are particularly susceptible to eutrophication given their limited capacity for depuration of excess nitrogen.     

Exponential Collapse of Benthic Depurative Capacity in A Eutrophic Tropical Marine Ecosystem: A Threshold Response to Organic Nitrogen Loading

Microbial decomposition of particulate organic matter in sediments can substantially modulate eutrophication of nearshore marine environments. Flux budgets for nitrogen compounds and quantification of rates of microbial transformation can provide important information on the process of eutrophication. This survey documents sediment nitrogen budgets for a eutrophic nearshore marine environment at La Parguera on southwest coast of Puerto Rico, including consideration of the organic fraction in addition to microbial transformations.

Sediments of the inshore channel at La Parguera denote the input of heavy organic loads with low redox potentials and high ammonium contents. Benthic fluxes of dissolved organic nitrogen are below those recorded for ammonium. These observations suggest that nearshore tropical sediments subject to heavy nitrogen loads act as a sink for organic nitrogen and that ammonification is a key process in the release of nitrogen from sediments to the water column. Nitrification and denitrification rates are low overall and inversely related to the redox potential. Depuration of excess nitrogen through denitrification is ineffective in these sediments. in contrast to more robust temperate environments, our work demonstrates that tropical marine systems are particularly susceptible to eutrophication given their limited capacity for depuration of excess nitrogen.     

植物多样性对土壤微生物的影响

生物多样性强烈地影响生态系统的过程.生态系统过程的变化可导致生物多样性衰减并因此导致生态系统功能衰退.植物种丰度和植物功能多样性对土壤细菌群落的代谢活性和代谢多样性有成正比的影响.土壤细菌的代谢活性和代谢多样性随植物种数量的对数和植物功能组的数量而直线上升.其原因可能是由植被流入土壤的物质和能量的多样性和数量的增加,也可能是由土壤动物区系起作用的土壤微生境的多样性的增加造成的.由于植物多样性的丧失所引起的植物生物量的减少对分解者群落有强烈的影响微生物生物量将可能减少,因为在大多数陆地生态系统中,有机碳源限制着土壤微生物的活性.     

Inventories and Distribution of Radiocaesium in Arctic Marine Sediments: Influence of Biological and Physical Processes

Extensive surveys of sediment burdens of radiocaesium, specifically ¹³⁷Cs, and other radioactive contaminants in the Arctic during the 1990's, indicate that almost all anthropogenic radionuclides buried on continental shelves adjacent to Alaska are derived from global bomb fallout. the ¹³⁷Cs (half-life: 30.2y) activities observed in surface (0-4 cm) marine sediments however, vary widely, albeit much less than the expected current inventory resulting from bomb fallout at this latitude (∼100mBq cm^-2). This observed geographical variation provided the opportunity to evaluate physical and biological mechanisms that may affect caesium biogeochemistry on Arctic continental shelves. We investigated whether high biological productivity in portions of the Bering and Chukchi Seas is effective in removing dissolved radiocaesium from the water column, and whether biological production in overlying water affects total radiocaesium inventories in sediments. Based upon C/N ratios in the organic fraction of shallow sediments, we found no evidence that higher inventories or surface activities of radiocaesium are present in areas with higher deposition of particulate organic matter. Based upon stable carbon isotope ratios of organic matter in sediments, we found no evidence that terrestrial runoff contributes proportionally to higher surface activities, although terrestrial runoff may affect total inventories of the radionuclide. Radiocaesium content of surface sediments was significantly correlated with total organic carbon content of sediments and the proportion of sediments in the finest sediment fractions. Because high current flow can also be expected to influence distributions of those sedimentary parameters, we conclude that re-distribution of     

Measuring terrestrial subsidies to aquatic food webs using stable isotopes of hydrogen

Understanding river food webs requires distinguishing energy derived from primary production in the river itself (autochthonous) from that produced externally (allochthonous), yet there are no universally applicable and reliable techniques for doing so. We compared the natural abundance stable isotope ratios of hydrogen (deltaD) of allochthonous and autochthonous energy sources in four different aquatic ecosystems. We found that autochthonous organic matter is uniformly far more depleted in deuterium (lower deltaD values) than allochthonous: an average difference of approximately 100% per hundred. We also found that organisms at higher trophic levels, including both aquatic invertebrates and fish, have deltaD values intermediate between aquatic algae and terrestrial plants. The consistent differences between leaves and algae in deltaD among these four watersheds, along with the intermediate values in higher trophic levels, indicate that natural abundance hydrogen isotope signatures are a powerful tool for partitioning energy flow in aquatic ecosystems.     

矸石山不同造林模式对土壤养分及酶活性影响的研究

本文以抚顺矸石山立地条件下树龄为10年的刺槐Robinia pseudoacacia L.、火炬Rhus typhina Nutt、白榆Ulmus pumila L.、沙棘Hippophae rhamnoides L.林以及植被自然恢复裸地（CK）5种植被恢复模式为研究对象,按土壤发生学层次进行采样,对土壤有机质、碱解氮、有效磷、速效钾、pH、过氧化氢酶、蔗糖酶和脲酶等指标进行研究,结果表明：与CK相比,4种造林模式显著提高了Ah层的土壤有机质和养分含量,其中,有机质、有效磷改良效果最好的是白榆模式,分别增加了194.8%、442.9%;速效钾改良效果最好的是刺槐模式,增加了262.0%;碱解氮则是火炬模式改良效果最好,增加了509.2%。不同造林模式与CK相比增加了各层次的过氧化氢酶、脲酶、蔗糖酶活性,其中,蔗糖酶和过氧化氢酶活性Ah层最大的是火炬模式,增加了134.5%和880.9%;脲酶酶活性最大的是白榆模式,增加了119.1%。从垂直角度看,有机质、碱解氮、过氧化氢酶、脲酶随土层加深含量降低,在C层达到最小值,而速效钾、有效磷、蔗糖酶的最小值则出现在AC层;不同模式对土壤pH影响不一,垂直规律不明显。     

Biodiversity and trophic structure of nematode assemblages in seagrass systems: evidence for a coupling with changes in food availability

Nematode assemblage composition, trophic structure and biodiversity were followed over an annual cycle in a Posidonia oceanica bed of the NW Mediterranean to test the response to temporally changing food availability (sediment organic matter, bacterial and microphytobenthic biomass). The sediment-water interface of the seagrass meadow was characterised by high particulate organic matter concentrations. Also, seagrass sediments had high organic matter content, chloropigment concentrations and bacterial biomass. All organic matter components (i.e. carbohydrates, proteins and lipids) changed temporally, with higher concentrations in winter and spring and lower concentrations in summer; however, overall, large amounts of organic compounds were potentially available for (seagrass) benthic consumers throughout the year. Nematode assemblage in the P. oceanica bed maintained a high genus number (88 genera), and a trophic structure tightly coupled with the composition of the potential food sources. In agreement with the relevance of microphytobenthos, epigrowth-feeder nematodes (2A) represented the dominant trophic guild. The biomass of predator nematodes (2B) was significantly correlated with the biomass of other nematodes. Non-selective deposit feeders (1B) displayed a close relationship with the concentration of proteins, carbohydrates and biopolymeric carbon (i.e. labile organic detritus). Conversely to what was expected, epigrowth feeders were not correlated with chloropigment concentrations, but showed a significant relationship with the number of dividing bacteria, possibly suggesting a trophic plasticity of the 2A feeding guild. The coupling between temporal changes in food sources and temporal variability of the nematode trophic structure was highlighted by a CANOCO analysis, which allowed us to identify and associate, at each sampling time, nematode genera and their feeding habits, with specific environmental variables and food indicators. Furthermore, species diversity (H') and evenness (J) calculated on nematodes identified to the genus level displayed evident temporal changes, also reflected by the index of trophic diversity (ITD). Both structural and functional diversity were coupled with high concentration and highly heterogeneous composition of the food sources (including organic detritus, microphytobenthic algae and bacteria) potentially available to nematodes. These data suggest that temporal changes in quantity and quality of food sources do not only influence nematode dynamics and trophic composition, but also influence nematode structural and functional diversity.     

三江源区不同建植年代人工草地群落演替与土壤养分变化

研究了了三源区不同建植期人工修复草地在不同演替阶段毒杂草[主要是甘肃马先蒿(Pedicularis kansuensis)]的入侵规律、数量特征,植物群落物种组成、生物苗和草地质最以及土壤养分、微生物活性的变化规律.结果表明,不同建植期人工修复草地植物群落的种类组成、植物功能群组成和群落数量特征存在显著差异.随着演替时间的推移,人工草地群落盖度、高度、物种数、生物最和多样性指数均表现出"V"字型变化规律,杂类草--甘肃马先蒿的数量特征变化尤为明显,在4 a的人工草地群落中开始局部入侵,在5～6 a的人工草地群落中大面积入侵,其入侵速度、入侵面积达到高峰期.土壤的含水量、容重、土壤中有机质、氮素和磷素在演替过程(7 a、9 a草地)中逐渐降低,到一定时期又逐步增加;随着演替的进行,不同建植期人工草地的土壤微牛物生物量碳和酶活性均呈"V"字型,变化.对于退化生态系统的恢复首先是植被恢复,其次是土壤肥力的恢复.土壤有机质等养分的积累、微生物活性的改善不仅能使土壤-植物复合系统的功能得以恢复,同时也能促进物种多样性的形成,有利于人工草地群落稳定性的提高.在试验区尽管植被恢复演替进行得比较缓慢,但从土壤发展的角度看,仍属进展演替.所以,在退化高寒草甸的恢复过程中,若降低和有效控制外界的干扰(如围栏封育),可为退化草地恢复提供繁殖体与土壤环境,实现人工草地逐步向恢复(正向)演替进行.图3表6参34     

Inventories and Distribution of Radiocaesium in Arctic Marine Sediments: Influence of Biological and Physical Processes

Abstract

Extensive surveys of sediment burdens of radiocaesium, specifically ¹³⁷Cs, and other radioactive contaminants in the Arctic during the 1990′s, indicate that almost all anthropogenic radionuclides buried on continental shelves adjacent to Alaska are derived from global bomb fallout. the ¹³⁷Cs (half-life: 30.2y) activities observed in surface (0–4 cm) marine sediments however, vary widely, albeit much less than the expected current inventory resulting from bomb fallout at this latitude (～100mBq cm^?2). This observed geographical variation provided the opportunity to evaluate physical and biological mechanisms that may affect caesium biogeochemistry on Arctic continental shelves. We investigated whether high biological productivity in portions of the Bering and Chukchi Seas is effective in removing dissolved radiocaesium from the water column, and whether biological production in overlying water affects total radiocaesium inventories in sediments. Based upon C/N ratios in the organic fraction of shallow sediments, we found no evidence that higher inventories or surface activities of radiocaesium are present in areas with higher deposition of particulate organic matter. Based upon stable carbon isotope ratios of organic matter in sediments, we found no evidence that terrestrial runoff contributes proportionally to higher surface activities, although terrestrial runoff may affect total inventories of the radionuclide. Radiocaesium content of surface sediments was significantly correlated with total organic carbon content of sediments and the proportion of sediments in the finest sediment fractions. Because high current flow can also be expected to influence distributions of those sedimentary parameters, we conclude that re-distribution of     

Suspended particulate organic matter in a Mediterranean submarine cave

Submarine caves display a paucity of benthic density and biomass that may be related to low trophic resources. Analysis of organic carbon, organic nitrogen, carbohydrate, protein and lipid content of suspended particulate matter was made during the period July 1985–July 1987 in a Mediterranean cave (Marseille, France) in order to determine any differences in the particulate organic matter composition along an horizontal transect. Particulate organic matter content clearly declined from the entrance of the cave to the dark inner area. This impoverishment could not be explained by a simple decrease in a few organic compounds, but appeared to be related to the combination of a decrease in both the amount and the composition of the suspended particles. Three progressive levels of impoverishment were identified towards the dark inner area of the cave: (i) decreasing amounts of seston; (ii) decreasing organic content of particles; (iii) increasing proportions of the geopolymeric (i.e., humic) components in the remaining organic matter, indicating increased degradation. The cave appeared to be sharply divided into two distinct sections — a twilight outer section whose waters contained slightly lower amounts of particulate organic matter than the open sea, and a dark inner section, 8 to 10 m higher, separated from the outer section by a steep rise and with waters of low organic matter content. The water in the twilight section appeared to be in thermal equilibrium with the open sea, and that in the dark inner section displayed thermal stratification. These differences indicated the presence of two distinct water bodies with contrasting average residence times, estimated as 1 d in the outer twilight section and 8 d in the dark inner section. The joint action of sedimentation and degradation resulted in an abrupt depletion of particulate organic matter in the dark inner section accompanied by a decrease in the benthic fauna. The decline in benthic heterotroph populations is probably related to the abrupt transition to oligotrophic conditions.