生态系统营养结构多样性的测度

生态系统层次上的生物多样性是指生物圈内生境、生物群落和生态过程以及生态系统的多样性,营养结构和营养过程的多样性是其中最重要的一个方面。本文通过宏观营养动力学的理论和方法,对生态系统营养结构进行分析,定义了反映生态系统营养结构多样性的两类指标,即营养物与流通量沿宏观营养链分布特征的多样性(D_Ⅰ),以及所有宏观营养级上营养物与流通量在不同分室上分布特征的多样性(D_Ⅱ)。这两类多样性是度量生态系统营养结构多样性的两个基本测度。     

Parasites and stable isotopes: a comparative analysis of isotopic discrimination in parasitic trophic interactions

Stable isotopes are widely used to identify trophic interactions and to determine trophic positions of organisms in food webs. Comparative studies have provided general insights into the variation in isotopic composition between consumers and their diet (discrimination factors) in predator–prey and herbivore–plant relationships while other major components of food webs such as host–parasite interactions have been largely overlooked. In this study, we conducted a literature‐based comparative analysis using phylogenetically‐controlled mixed effects models, accounting for both parasite and host phylogenies, to investigate patterns and potential drivers in Δ¹³C and Δ¹⁵N discrimination factors in metazoan parasitic trophic interactions. Our analysis of 101 parasite–host pairs revealed a large range in Δ¹³C (–8.2 to 6.5) and Δ¹⁵N (–6.7 to 9.0) among parasite species, with no significant overall depletion or enrichment of ¹³C and ¹⁵N in parasites. As previously found in other trophic interactions, we identified a scaling relationship between the host isotopic value and both discrimination factors with Δ¹³C and Δ¹⁵N decreasing with increasing host δ¹³C and δ¹⁵N, respectively. Furthermore, parasite phylogenetic history explained a large fraction (>60%) of the observed variation in the Δ¹⁵N discrimination factor. Our findings suggest that the traditional isotope ecology framework (using an average Δ¹⁵N of 3.4‰) applies poorly to parasitic trophic interactions. They further indicate the need for a scaled rather than a fixed trophic discrimination factor framework along gradients of host δ¹⁵N. We also identified several conceptual and methodological issues which should to be considered in future research to help integrate parasitic interactions into a holistic isotope ecology framework across diverse trophic interactions.     

Analysis of trophic relationships in two shallow equatorial lakes Lake Naivasha (Kenya) and Lake Ihema (Rwanda) using a multispecifies trophic model

A multispecifies trophic model called ECOPATH II, which can be used to describe the trophic relationships in aquatic ecosystems on a quantitative basis, is briefly presented. When properly used, it can help to explain the trophic relationships in ecosystems and possible evolution of fishstocks after modifications of the environment (e.g. eutrophication, introduction of a new population and/or a significant increase of the fishing effort), and to compare the trophic structure of several ecosystems. Examples are provided on two shallow lakes: Lake Ihema and Lake Naivasha. They are compared with Lake George which was previously documented.     

Rescaling the trophic structure of marine food webs

Measures of trophic position (TP) are critical for understanding food web interactions and human‐mediated ecosystem disturbance. Nitrogen stable isotopes (δ¹⁵N) provide a powerful tool to estimate TP but are limited by a pragmatic assumption that isotope discrimination is constant (change in δ¹⁵N between predator and prey, Δ¹⁵N = 3.4‰), resulting in an additive framework that omits known Δ¹⁵N variation. Through meta‐analysis, we determine narrowing discrimination from an empirical linear relationship between experimental Δ¹⁵N and δ¹⁵N values of prey consumed. The resulting scaled Δ¹⁵N framework estimated reliable TPs of zooplanktivores to tertiary piscivores congruent with known feeding relationships that radically alters the conventional structure of marine food webs. Apex predator TP estimates were markedly higher than currently assumed by whole‐ecosystem models, indicating perceived food webs have been truncated and species‐interactions over simplified. The scaled Δ¹⁵N framework will greatly improve the accuracy of trophic estimates widely used in ecosystem‐based management.     

The roles of predation,competition, and exploitation in the trophic dynamics of a warmwater stream: a model synthesis,analysis, and application

We developed a trophic dynamic model of key populations and processes in the New River, West Virginia, to identify the mechanisms most responsible for maintaining food web structure. Key populations were represented by thirteen model components and were aquatic insects; age-1 and age-2 crayfish (three species); age-1 and age-2 hellgrammites (Corydalus cornutus larvae); non-game fishes; age-0, age-1, and adult smallmouth bass (Micropterus dolomieu); age-0, age-1, and adult rock bass (Ambloplites rupestris); and age-0, age-1 to age-3, and adult flathead catfish (Pylodictis olivaris). In this system, crayfish and hellgrammites are harvested to provide bait for the recreational fishery that extensively exploits the three predatory fish species. Predation and intraspecific regulation were represented with nonlinear algorithms, and linear terms represented fishery harvests. Interspecific competition among components occurred through predation on shared prey. Error analysis of the model suggested that predation was the most important mechanism in maintaining system structure (the disposition of biomass among system components). Further, the trophic relation between each component and its prey accounted for 34–64% of the variability in food web structure, whereas predation on each component explained 1–24% of food web structure variability. Therefore, so-called ‘bottom-up’ effects were more influential than ‘top-down’ effects. Interspecific competition and intraspecific regulation had secondary roles in maintaining New River food web structure, although intraspecific regulation was most important to aquatic insects, which were not predatory in our model. Both forms of competition are probably tempered by extensive predation and exploitation in the New River system. Exploitation was a secondary structuring agent to adult smallmouth bass, which experience a high rate of harvest in the New River.     

A call for applying trophic structure in ecological restoration

Ecological restoration projects have traditionally focused on vegetation as both a means (seeding, planting, and substrate amendments) and ends (success based upon primary productivity and vegetation diversity). This vegetation‐centric approach to ecological restoration stems from an historic emphasis on esthetics and cost but provides a limited measure of total ecosystem functioning and overlooks alternative ways to achieve current and future restoration targets. We advocate a shift to planning beyond the plant community and toward the physical and biological components necessary to initiate autogenic recovery, then guiding this process through the timely introduction of top predators and environmental modifications such as soil amendments and physical structures for animal nesting and refugia.     

Fish trophic structure in estuaries,with particular emphasis on estuarine typology and zoogeography

A comparative analysis of the fish trophic structure was undertaken on some 190 South African estuaries spanning three zoogeographic regions and incorporating three broad estuarine types. Fish biomass trophic guild compositions and biomass trophic spectrum profiles were analysed using multivariate statistical techniques and included both inter‐regional (zoogeographic) and intra‐regional (estuarine typology) comparisons. Differences in the fish trophic structure of the various estuary types within each zoogeographic region were observed; these were linked to the relative biomass contribution of the various trophic guilds and also to differences in biomass trophic spectrum profiles of the fishes in each estuary type within each region. In spite of these differences in trophic structure, all estuaries were dominated by detritivores, which suggests that the main food source (detritus) is similar in all biogeographic regions. Preliminary indications are that a similar dependence by estuary‐associated fishes on detritus food sources exists on a global basis but that detailed studies are required in order to confirm this assertion.     

The effect of trophic/contaminant interactions on chironomid community structure and succession (Diptera: Chironomidae)

The effects of contaminants and organic pollution on the chironomid communities of two basins in Lac St. Louis, a river-lake on the St. Lawrence River above Montreal, Quebec, were assessed and compared using lake classification theory techniques and morphological deformity indices based on the ligulae and antennae ofProcladius, the dominant surviving component of the south-shore communities. Contaminants from the industrial complex around Beauharnois, Quebec, have seriously degraded communities along the south-shore gradient while the introduction of untreated domestic wastes from Laprairie, Quebec, has seriously affected the communities of the Laprairie basin. The interaction between trophic and contaminant effects constitutes a classic example of the setback or deflection in the trajectory of ecological succession in biological communities hypothesized by ODUM (1981, 1985).     

中国北方桃李间作对步甲群落组成和营养级结构的影响

【目的】由于桃树和李树在生态环境要求上很相似, 在生产实际中, 在中国很多地方存在桃李间作或邻作的现象。本研究将通过试验明确桃李间作对步甲营养级结构的影响以便更全面地评价桃李间作是否有利于害虫防治。【方法】2006和2007年的4 -10月, 在山西省临汾市郊, 通过陷阱法对3块桃李间作果园和3块单一桃园内的步甲进行抽样, 每块果园的面积约0.3 hm²。比较两类桃园内步甲物种丰富度。将6种优势步甲物种, 两两单一成虫个体组合放在玻璃瓶内, 放在桃树下, 统计步甲相互攻击后的伤亡情况, 比较各物种的相互攻击能力, 确定主要步甲所属的营养级。统计分析两类桃园步甲营养级结构的差异。【结果】桃李间作园和单作桃园调查到的步甲物种均为24种, 估测步甲物种数在两类果园的差异也不显著（P≥0.38）。6种优势步甲可以分为4个营养级： 大盗步甲Lesticus magnus （Motschulsky）为高级捕食者, 大锹步甲Scarites acutides Chaudoir为次级捕食者, 双斑青步甲Chlaenius bioculatus Chaudoir和躅步甲Calathus halensis （Schaller）为初级捕食者, 毛婪步甲Harpalus griseus （Panzer）和直隶婪步甲Harpalus tschiliensis Schauberger为植食者。各营养级步甲相对多度均是桃李间作果园大于单一桃园, 独立T检验结果表明, 植食者步甲相对多度差异达极显著水平（P≤0.002）; 而捕食者步甲相对多度, 除2006年初级捕食者步甲相对多度差异显著（P=0.013）外, 在调查的两年中其他捕食者营养级步甲相对多度差异均不显著（P≥0.085）。【结论】结果表明, 桃李间作对步甲群落的物种组成和丰富度没有影响, 但是对果园内各营养级步甲相对多度有影响, 其中, 对植食者营养级步甲相对多度影响显著, 而对捕食者营养级步甲相对多度影响多不显著。     

An experimental demonstration of trophic interactions affecting water quality of Rice Lake, Ontario (Canada)

Eight cylindrical enclosures (3 m diameter, 2.7 m long, V = 20m³) were installed in eutrophic Rice Lake (Ontario, Canada) in late spring of 1987. Fish (yearling yellow perch (Perca flavescens) and macrophytes (Potamogeton crispus) presence and absence were set at the beginning of the experiment to yield four combinations of duplicate treatments. The purpose of the experiment was to determine if the phytoplankton, zooplankton, macrophytes and fish species resident in the lake interact to influence water quality (major ions, phosphorus, algal densities and water clarity).The presence of fish was associated with: (1) decreased biomass of total zooplankton, (2) decreased number of species in the zooplankton, (3) decreased average size of several zooplankton taxa, (4) higher total phosphorus concentrations, (5) higher phytoplankton and chlorophyll a concentrations, (6) lower water clarity, (7) lower potassium levels during macrophyte die-back, (8) lower pH and higher conductivity in the presence of macrophytes. Biomass of large Daphnia species (but not total zooplankton) was highly correlated with the algal response (r ² = 0.995) and was associated with reduced biomass of several algal taxa including some large forms (Mougeotia, Oedogonium) and several colonial blue-green algae. However, no significant control of late summer growth of the bloom-forming blue-green alga Anabaena planctonica Brun. was achieved by the Daphnia presence-fish absence treatment. Release of phosphorus to the water column during the die-back of P. crispus was not an important phenomenon.     

Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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Diet and trophic niche of Antarctic silverfish Pleuragramma antarcticum in the Ross Sea,Antarctica

The diet of Antarctic silverfish Pleuragramma antarcticum was evaluated by examining stomach contents of specimens collected in the Ross Sea (71°–77° S; 165°–180° E) in January to March 2008. Pleuragramma antarcticum (50–236 mm standard length, L_S) and prey items were analysed for stable‐isotopic composition of carbon and nitrogen. According to index of relative importance (I_RI), which incorporates frequency of occurrence, mass and number of prey items, the most important prey items were copepods (81%I_RI over all specimens), predominantly Metridia gerlachei and Paraeuchaeta sp., with krill and fishes having low I_RI (2·2 and 5·6%I_RI overall). According to mass of prey (M) in stomachs, however, fishes (P. antarcticum and myctophids) and krill dominated overall diet (48 and 22%M, respectively), with copepods being a relatively minor constituent of overall diet by mass (9·9%M). Piscivory by P. antarcticum occurred mainly in the extreme south‐west of the region and near the continental slope. Krill identified to species level in P. antarcticum stomachs were predominantly Euphausia superba (14·1%M) with some Euphausia crystallophorias (4·8%M). Both DistLM modelling (PRIMER‐permanova+) on stomach contents (by I_RI) and stepwise generalized linear modelling on stable isotopes showed that L_S and location were significant predictors of P. antarcticum diet. Postlarval P. antarcticum (50–89 mm L_S) consumed exclusively copepods. Juvenile P. antarcticum (90–151 mm L_S) consumed predominantly krill and copepods by mass (46 and 30%M, respectively). Small adult P. antarcticum (152–178 mm L_S) consumed krill, fishes and copepods (37, 36 and 15%M, respectively). Large adult P. antarcticum (179–236 mm L_S) consumed predominantly fishes and krill (55 and 17%M, respectively), especially in the north (near the Ross Sea slope) and in the SW Ross Sea. Amphipods were occasionally important prey items for P. antarcticum (western Ross Sea, 39%M). General concordance between stomach contents and trophic level of P. antarcticum and prey based on δ¹⁵N was demonstrated. Pleuragramma antarcticum trophic level was estimated as 3·7 (postlarval fish) and 4·1 (fish aged 3+ years).     

Fish species and trophic diversity of haplochromine cichlids in the Kyoga satellite lakes (Uganda)

Prior to the 1980s, lakes Kyoga and Victoria previously supported an exceptionally diverse haplochromine fish fauna comprising at least 11 trophic groups. The species and trophic diversity in these lakes decreased when the introduced Nile perch depleted haplochromine stocks. From December 1996 to October 1998, we studied species and trophic diversity of haplochromine fishes in six satellite lakes without Nile perch in the Kyoga basin and compared them with the Kyoga main lake against historical data from Lake Victoria where Nile perch were introduced. Forty‐one species were found in the study area, of which, the Kyoga satellite lakes contributed 37 species in comparison to only 14 from the Kyoga main lake. Analysis of trophic diversity based on 24 species that contained food material revealed seven haplochromine trophic groups (insectivores, peadophages, piscivores, algal eaters, higher plant eaters, molluscivores and detritivores) in the Kyoga satellite lakes in comparison to two trophic groups (insectivores and molluscivores) in the Kyoga main lake. Many of the species and trophic groups of haplochromines depleted by the introduced Nile perch in lakes Kyoga and Victoria still survive in the Kyoga satellite lakes. This is attributed to the absence of Nile perch in those lakes. Nile perch has been prevented from spreading into the satellite lakes by swamp vegetation that separate them from the main lakes. If these swamps prevent Nile perch from spreading into the lakes, it is possible to conserve fish species, especially haplochromines, which are threatened by introduction of Nile perch in the main lakes.     

Diet and trophic interactions of a circumglobally significant gelatinous marine zooplankter,Dolioletta gegenbauri (Uljanin, 1884)

Gelatinous zooplankton play a crucial role in marine planktonic food webs. However, primarily due to methodological challenges, the in situ diet of zooplankton remains poorly investigated and little is known about their trophic interactions including feeding behaviour, prey selection and in situ feeding rates. This is particularly true for gelatinous zooplankton including the marine pelagic tunicate, Dolioletta gegenbauri. In this study, we applied an 18S rRNA amplicon metabarcoding approach to identify the diet of captive‐fed and wild‐caught D. gegenbauri on the midcontinental shelf of the South Atlantic Bight, USA. Sequencing‐based approaches were complimented with targeted quantitative real‐time polymerase chain reaction (PCR) analyses. Captive‐fed D. gegenbauri gut content was dominated by pico‐, nano‐ and micro‐plankton including pico‐dinoflagellates (picozoa) and diatoms. These results suggested that diatoms were concentrated by D. gegenbauri relative to their concentration in the water column. Analysis of wild‐caught doliolids by quantitative real‐time PCR utilizing a group‐specific diatom primer set confirmed that diatoms were concentrated by D. gegenbauri, particularly by the gonozooid life stage associated with actively developing blooms. Sequences derived from larger metazoans were frequently observed in wild‐caught animals but not in captive‐fed animals suggesting experimental bias associated with captive feeding. These studies revealed that the diet of D. gegenbauri is considerably more diverse than previously described, that parasites are common in wild populations, and that prey quality, quantity and parasites are likely all important factors in regulating doliolid population dynamics in continental shelf environments.     

Experimental assessment of trophic impacts from a network model of a seagrass ecosystem: Direct and indirect effects of gulf flounder, spot and pinfish on benthic polychaetes

Trophic cascades are predicted to occur when the abundance of predators is increased, directly reducing the abundance of the intermediate prey and indirectly increasing the abundance of the prey at the base of a food web. Mixed trophic impact analysis of a network model developed for Apalachee Bay, near St. Marks, FL, USA predicted such a trophic cascade, in that increased abundance of juvenile gulf flounder Paralichthys albigutta (x¯ = 149 mm SL, effective trophic level 3.9) should have a negative impact on juvenile spot Leiostomus xanthurus (x¯ = 30 mm SL, effective trophic level 2.9) and a positive impact on benthic polychaetes (effective trophic levels 2.3 for deposit feeders and 3.0 for predatory polychaetes) in Halodule wrightii seagrass beds. We tested the predictions of the mixed trophic impact analysis by manipulating the abundance of the high trophic-level species (juvenile gulf flounder) in a cage-exclusion study in the North River, near Harkers Island, NC, USA. We compared the polychaete communities in St. Marks, FL and Harkers Island, NC, and showed that they are 51% similar (Jaccard's Index) at the family level, with the same eight dominant families (Nereidae, Capitellidae, Syllidae, Spionidae, Cirratulidae, Terebellidae, Sabellidae, and Maldanidae) present in both locations. We used 24 open-bottom cages to enclose the benthos and its seagrass-associated animal communities. We manipulated each cage by assigning it to one of the following treatments: (1) inclusion of fishes in upper and intermediate trophic levels (1 juvenile gulf flounder and 10 juvenile spot, the flounder + spot treatment); (2) inclusion of the intermediate predator (10 juvenile spot with no gulf flounder, the spot-only treatment); and (3) no fish added (unmanipulated controls). Core samples taken within the cages provided pre- and post-experimental measures of polychaete density and biomass, and the difference in density and biomass were used as response variables. At the end of the experiment, we collected, weighed, and analyzed the gut contents of all juvenile spot present in the cages. Juvenile pinfish (Lagodon rhomboides, x¯ = 30 mm SL) were present at the end of the study, having arrived as larvae or being trapped during cage set-up, and these fish were also examined, because they also eat polychaetes and their natural densities exceeded our introduced spot densities. Significant differences among treatments were detected for the polychaete family Terebellidae for both the change in density and biomass (pre-experiment − post-experiment). Densities of the Terebellidae changed in the direction predicted by the network model's impact analysis, declining in the cages with spot added compared with the control cages. Analyses of the other response variables (post-experiment spot and pinfish densities and biomass, difference between pre- and post-experiment polychaete densities and biomass for other families, and post-experiment spot and pinfish stomach content biomass) showed no significant differences among treatments. Several variables (Nereidae densities, pinfish densities and biomass, and pinfish stomach content biomass) varied between cages with low and high seagrass cover (significant blocking effect, P < 0.001). Nereidae densities declined significantly in cages with high (73%) rather than with low coverage (31% cover) of seagrass. Pinfish density and biomass were significantly greater in the high seagrass cages at the end of the experiments (P < 0.001), suggesting that dense seagrass attracted them. We conclude that the high density of pinfish in dense seagrass was responsible for the decline in density of the Nereidae. The direct effect of intermediate predators (pinfish feeding on polychaete prey) can be influenced by preferential recruitment of fishes to structurally complex habitats. The direction of change of indirect effects, but not the magnitude, in multi-trophic-level food webs can be predicted by the mixed trophic impact analysis of network models. However, these indirect effects are likely to be small in magnitude relative to direct effects and may be difficult to detect experimentally, especially in low-power experimental caging studies with natural fluctuations in recruitment rates of competitor species.     

Responses of zooplankton body size and community trophic structure to temperature change in a subtropical reservoir

Understanding the effects of global warming on trait variation and trophic structure is a crucial challenge in the 21st century. However, there is a lack of general patterns that can be used to predict trait variation and community trophic structure under the ongoing environmental change. We investigated the responses of body size and community trophic structure of zooplankton to climate related factors (e.g., temperature). Isotopic niche breadth was applied to investigate the community trophic structure across a 1‐year study from a subtropical reservoir (Tingxi Reservoir) in southeastern China. Body size and community isotopic niche breadth of zooplankton were larger during water mixing than stratification periods and correlated significantly with water temperature change along the time series. The contributions of intra‐ and intertaxonomic components to body size and community trophic structure variation showed significant relationships with the temperature change going from the mixing to stratification periods. Water temperature imposed direct effect on body size, while direct and indirect effect on the community trophic structure of zooplankton occurred through trophic redundancy along time series. Water temperature and community properties (e.g., body size, trophic redundancy, or trophic interaction) showed complex interactions and integrated to influence community trophic structure of zooplankton. Our results can expand the knowledge of how elevated temperature will alter individual trait and community trophic structure under future climate change.     

Determining trophic state in Lake Whatcom,Washington (USA), a soft water lake exhibiting seasonal nitrogen limitation

We evaluated an eleven year data set to assess trophic state and nutrient limitation in Lake Whatcom, an oligotrophic, soft water, chain lake located in the Puget Sound lowlands of Washington (U.S.A.). Although total phosphorus (TP) and soluble reactive phosphate (SRP) concentrations were relatively low throughout the lake, there were significant differences between the northern basin (Site 1) and the other sampling sites (Sites 2–4). Nonparametric correlation coefficients (Kendall's ) were highest between chlorophyll (CHL), Secchi depth (SD), total nitrogen (TN), and dissolved inorganic nitrogen (DIN). Late summer algal biomass correlated best with DIN and TP. Trophic State Indices based on TP, TN, CHL and SD revealed that although algal growth was most likely phosphorus limited throughout the year, the northern basin of the lake may have developed nitrogen co-limitation during late summer and fall. During this period, N/P ratios were often less than 20, and in 1998 the epilimnetic DIN concentrations dropped below 20 g l^–1 while DIN/TP ratios fell below 4. Reviews of the literature suggest that while co-limitation by phosphorus and nitrogen is fairly common in unproductive lakes, the patterns seen in Lake Whatcom were more similar to those reported for eutrophic lakes experiencing secondary nitrogen limitation resulting from excess phosphorus loading.     

Measurement of trophic niche breadth using occurrence frequencies

A modification of Gladfelter-Johnson's index to measure trophic niche breadth using occurrence frequencies is proposed, in order to make it more sensitive to different resource use patterns. The new index ranges from 0 to 1 and measures niche breadth in a guild context.     

Day and night trophic variations of dominant fish species in a lagoon influenced by freshwater seeps

The aim of this study was to determine the trophic structure and nycthemeral variations in the diet of dominant fish species (Ariopsis felis, Bairdiella chrysoura, Micropogonias undulatus, Eucinostomus gula, Eucinostomus argenteus, Lagodon rhomboides and Sphoeroides testudineus) in Celestun Lagoon, a biosphere reserve located in the southern Gulf of Mexico, and influenced by freshwater seeps. A total of 1473 stomachs were analysed and nine trophic groups were recorded. Bray–Curtis analyses with analyses of similarity (ANOSIM) statistical tests were used to determine two groups of feeding guilds: zoobenthivores and omnivores, with significant differences between time and habitat. The relationships between fish feeding habits, size class and environmental variables were investigated using canonical correspondence analysis (CCA). Most of the species showed a low niche breadth with high specialization towards amphipod consumption, with the exception of L. rhomboides (0·60), which indicated generalist feeding. This study in a protected area is an important source of information for drawing up conservation policies in relation to the management of aquatic resources, and will aid in the establishment of priority areas for conservation.