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1.
  1. Understanding changes in macroinvertebrate communities is important because they play a large role in stream ecosystem functioning, and they are an important food resource for fish. Beaver-induced changes to stream morphology could alter macroinvertebrate communities, which in turn could affect food webs and ecosystem function. However, studies investigating the effects of North American beaver activities on macroinvertebrates are rare in the inter-mountain west, an area with high potential for beaver-assisted restoration.
  2. The aim of this study was to quantify differences in the macroinvertebrate community between unaltered segments of streams and within beaver ponds in north-eastern Utah, U.S.A. We assessed macroinvertebrate species richness, biomass, density, functional feeding group composition, mobility group composition, and macroinvertebrate habitat characteristics to test the hypothesis that macroinvertebrate communities will differ among habitat types (undammed stream segments and beaver ponds) in beaver-occupied streams.
  3. Beaver pond communities significantly differed from lotic reach communities in many ways. Beaver ponds were less diverse with 25% fewer species. Although there was variability among streams, in general, beaver ponds had 75% fewer individuals and 90% lower total macroinvertebrate biomass compared to lotic reaches.
  4. Regarding functional feeding groups, beaver ponds contained more engulfers, while lotic reaches contained more scrapers, filterers, and gatherers. For mobility groups, beaver ponds had more sprawlers, while lotic reaches had more clingers. Swimmers were also more prevalent in lotic reaches, although this is probably due to the abundance of Baetis within lotic reaches. More beaver pond taxa were classified as lentic-dwelling insects, while more lotic reach taxa were categorised as preferring lotic habitats.
  5. The creation of ponds by beavers fundamentally altered the macroinvertebrate community in north-eastern Utah streams. Such changes to stream macroinvertebrate communities suggest that recolonisation of beavers across North America may be altering stream functioning and food webs. Our study highlights the need to further investigate the effects of beaver recolonisation on stream communities.
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2.
Our study found that beaver activity affects macroinvertebrate assemblages of both beaver ponds and downstream sites. The percentage composition of the invertebrate faunae of beaver ponds was strikingly different from the invertebrate faunae of upstream forested and downstream sites. The number of EPT (ephemeropteran, plecopteran, trichopteran) taxa in the upstream forested sites in all streams was higher than in beaver pond and downstream sites. Statistically significant differences were found in absolute and relative abundances of EPT and Chironomidae between different streams sites. The absolute and relative abundance of pollution-sensitive EPT was significantly higher in forested sites than in beaver pond and downstream sites in all measured streams. Beaver ponds had a significantly higher absolute and relative abundance of Chironomidae compared with upstream forested and downstream sites. We found that Plecoptera and Coleoptera were absent from beaver pond sites. The absolute abundance of Plecoptera was significantly higher in upstream forested sites than in downstream sites in all three streams. Gatherers were the dominant functional feeding group in relative abundance in all three habitat types. The percentage of gatherers was higher in beaver ponds than in forested and downstream sites.  相似文献   

3.
Differences among lake morphologies often explain variation in characteristics of lentic ecosystems. Although beaver ponds also vary in morphology, previous studies have not examined the effects of such variation on downstream ecosystems. This study evaluated downstream effects of multiple beaver ponds in the Colorado Rocky Mountains during one low and one high-flow year. Beaver pond morphology was described as the natural log transformed ratio of beaver dam height (which determines hydraulic head) to pond surface area and related to pond spillover phytoplankton and characteristics of the ecosystem downstream (nutrient concentrations, limiting nutrients, periphyton, benthic organic matter (BOM), and benthic invertebrate consumers). Nitrate concentration increased systematically downstream of beaver ponds, but only in the low flow year when groundwater influences predominated. Effects of beaver ponds on soluble reactive phosphorus concentration depended on pond morphology, increasing downstream of small ponds with high dams, but only during the low-flow year. In situ experiments showed that neither beaver activity nor pond morphology predicted periphyton-limiting nutrients downstream. Both periphyton biomass and BOM decreased downstream of small ponds with high dams but pond morphology did not predict abundance of invertebrate grazers or detritus-feeding consumers. While suspension feeding invertebrates increased downstream from small ponds with high dams, variation in chlorophyll a from water spilling over beaver dams did not follow a similar pattern. We conclude that the effects of beaver ponds on downstream nutrients, resources and consumers are rarely systematic, but instead depend on variation in pond morphology and on annual hydrologic variation.  相似文献   

4.
1. Movements between a stream reach and two adjacent lentic macrohabitats, a beaver pond and a lake, were followed for the Appalachian crayfish and two fish species, brook charr and brown bullhead, over an 85‐d period from early June to late August, and were analysed in relation to water level, maximum water temperature, photoperiod length, lunar luminosity, and age, by use of time‐series regressions. 2. Brook charr showed strong net immigration to the stream reach for underyearling (age class 0+) fish but net emigration for 1+ fish. Both immigration and emigration were positively related to water level and temperature; migratory responses to temperature were age‐specific. 3. Brown bullhead used the stream primarily as a corridor for downstream migration from the beaver pond to the lake. As with brook charr, water level and temperature had a positive effect on movement and responses were stronger in younger individuals. 4. Crayfish emigrated from the stream during the summer. Movements were positively related to increases in water level and temperature, with responses of 1+ crayfish much stronger than those of older individuals. 5. The results indicate that the stream tributary played different roles for brook charr (nursery), brown bullhead (dispersal corridor) and Appalachian crayfish (early summer refuge), and suggest that variation in water level or temperature resulting from climatic change or local anthropogenic activities might modify connectivity between macrohabitats, particularly for younger individuals.  相似文献   

5.
Increasing air temperatures are changing the arctic tundra biome. Permafrost is thawing, snow duration is decreasing, shrub vegetation is proliferating, and boreal wildlife is encroaching. Here we present evidence of the recent range expansion of North American beaver (Castor canadensis) into the Arctic, and consider how this ecosystem engineer might reshape the landscape, biodiversity, and ecosystem processes. We developed a remote sensing approach that maps formation and disappearance of ponds associated with beaver activity. Since 1999, 56 new beaver pond complexes were identified, indicating that beavers are colonizing a predominantly tundra region (18,293 km2) of northwest Alaska. It is unclear how improved tundra stream habitat, population rebound following overtrapping for furs, or other factors are contributing to beaver range expansion. We discuss rates and likely routes of tundra beaver colonization, as well as effects on permafrost, stream ice regimes, and freshwater and riparian habitat. Beaver ponds and associated hydrologic changes are thawing permafrost. Pond formation increases winter water temperatures in the pond and downstream, likely creating new and more varied aquatic habitat, but specific biological implications are unknown. Beavers create dynamic wetlands and are agents of disturbance that may enhance ecosystem responses to warming in the Arctic.  相似文献   

6.
1. The North American beaver has been studied as a model ecosystem engineer for many decades. Previous studies have documented physical, chemical and biological impacts attributed to beaver engineering in both aquatic and terrestrial environments. This study focused on the effects of ecosystem engineering by beavers on life histories of a common mayfly and on the potential consequences for mayfly populations. 2. We studied 18 montane beaver ponds of varying size and shape in western Colorado near the Rocky Mountain Biological Laboratory. Our goal was to test whether variation in beaver pond morphology (pond size and shape) explains downstream changes in stream temperature, mayfly size and timing of emergence. 3. Downstream water temperatures varied predictably with pond morphology, being colder downstream of high‐head dams and warmer downstream of low‐head dams. Pond morphology was also a significant predictor of variation in the size of mature female Baetis bicaudatus (the most abundant mayfly), with larger females emerging downstream of high‐head dams and smaller females downstream of low‐head dams. The size of male B. bicaudatus was not significantly related to pond morphology or stream temperature. There was no relationship between pond morphology and variation in the timing of emergence of Baetis (males or females) between upstream and downstream reaches. 4. Our results have implications for the effects of beaver ponds on Baetis individual fitness because large Baetis females are more fecund. Therefore, predictable female size variation associated with beaver pond morphology makes it possible to model the effects of beaver activity on local contributions of Baetis to the regional pool of reproductive adults at the catchment scale. Additionally, predictable changes in the size of emerging mayflies may have important consequences for the magnitude of aquatic to terrestrial resource subsidies in beaver‐modified systems.  相似文献   

7.
温彬  高勤峰  董双林  宁鲁光 《生态学报》2016,36(14):4327-4336
于2012年7月至2013年4月调查了荣成靖海湾3个不同水深的刺参(Apostichopus japonicus)养殖池塘内大型底栖动物的构成,以了解不同水深对刺参养殖池塘环境条件的影响以及由此引起的大型底栖生物群落结构的改变。结果表明:3个不同水深梯度池塘(1#浅水位、2#正常水位和3#高水位)底部光照强度、叶绿素a(Chla)和总有机物(TOM)含量存在显著差异,各池塘水温差异不显著。光强、Chla和TOM含量在夏季、冬季和春季均表现为1#池塘显著高于3#池塘;秋季各池塘间光强和TOM含量差异不显著,Chla含量则表现为3#池塘显著高于1#池塘。各季节3个池塘间大型底栖动物在种类组成、丰度、生物量和多样性指数上均存在显著性差异。大型底栖动物丰度和生物量夏季均表现为1#池塘显著高于3#池塘,秋季和冬季则相反;春季1#池塘丰度显著高于3#池塘,生物量则差异不显著。这些差异主要与其各自优势种及其优势度指数大小有关。大型底栖动物多样性指数夏季和秋季均表现为1#池塘高于3#池塘,春季则相反,冬季各池塘间多样性指数差异不显著。单因子相似性分析(ANOSIM)表明,各季节3个池塘间大型底栖动物群落结构均存在显著差异,表明水深梯度对刺参养殖池塘大型底栖动物群落结构造成显著性影响。相似性百分比分析(SIMPER)显示,各季节对3个池塘间大型底栖动物群落差异起主要作用的物种为各个池塘的优势种。典范对应分析(CCA)表明,水深、Chla和TOM含量为影响大型底栖动物群落的主要环境因子。  相似文献   

8.
Aquatic predators and habitat permanence can jointly affect benthic invertebrate biomass and community composition. In 2006 I sampled fish and invertebrates in ten ponds embedded in a seasonal wetland before and after a natural drought. Drought reduced fish biomass and density leaving some ponds in a fishless condition when rains returned in July. In July, large aquatic insects and crayfish colonized and reproduced in the ponds, but did not colonize all of the ponds equally. Using measurements of fish abundance and other environmental parameters of the ponds, I conducted linear regression analyses to explore potential drivers of variable invertebrate biomass in July. Fish biomass had a negative effect on invertebrate biomass and it explained more of the variation in total invertebrate biomass and total non-shrimp biomass than fish abundance (number of fish caught). Dissolved oxygen and pond depth were both correlated with fish biomass, but were poorer predictors of invertebrate biomass. Ponds with few or no fish had 20× greater total biomass and 200× more non-shrimp biomass than ponds with high fish biomass. Shrimp dominated the invertebrate composition, and were only found in the two deepest ponds with the highest fish biomass; predatory insects and crayfish dominated the other eight ponds. When taxa were analyzed separately, fish biomass explained a large portion of the variation for predatory insects (Coleoptera, Hemiptera, and Odonata) and crayfish (Procambarus alleni), but dissolved oxygen was the best predictor of larval stratiomyid (order Diptera) biomass. These results are generally consistent with studies demonstrating negative effects of fish on large predatory invertebrates, but also suggest that more severe local droughts can seasonally enhance insect and crayfish populations by generating fishless or nearly fishless conditions. Handling editor: J. Trexler  相似文献   

9.

In recent decades, the Eurasian beaver (Castor fiber) has once again become the keystone species in small river ecosystems in Russia. In many places, beaver activity has resulted in a significant change in lotic habitats, affecting the diversity, density, and biomass of aquatic organisms, including fish. While many studies have considered the ecosystem impacts of beavers, relatively few have focused on understanding the influence of beaver activity on steppe rivers. We conducted the first quantitative study of beaver impacts on fish assemblages in beaver-influenced and beaver-free sites on two small steppe rivers in the Don River basin in Russia. The presence of beavers altered the habitats in small steppe rivers and affected the diversity, density, and biomass of fish. A comparison of the number of species, density, and biomass of fish in six types of river habitats showed that these parameters were lower in beaver ponds than at riverine sites without beaver activity. Three fish species primarily preferred a single habitat type. Barbatula barbatula was found in riffles, Misgurnus fossilis in old beaver ponds, and Eudontomyzon mariae in abandoned beaver ponds. Beavers impacted fish distribution and density by changing dissolved oxygen, pH, and water current velocity. Overall, our results showed that the presence of beavers led to a temporary homogenization of fish habitats at a local scale in the valleys of small steppe rivers because beavers occupied these rivers only for a short period. However, habitat heterogeneity may increase if the beaver population stays stable or expands in the future.

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10.
1. We experimentally reduced densities of predatory fish in replicated 2 m2 areas of the littoral zone in two ponds to test whether density and biomass of invertebrates would respond to release from fish predation. The ponds are of similar size and in close proximity, but support different fish assemblages: bluegills ( Lepomis macrochirus Rafinesque) and largemouth bass ( Micropterus salmoides (Lacepede)) in one pond, and bluespotted sunfish ( Enneacanthus gloriosus (Holbrook)) and chain pickerel ( Esox niger Lesueur) in the other. Fish densities were reduced to less than 15% of ambient levels in both experiments.
2. In the bluegill–bass pond, density and biomass of most invertebrate taxa and size classes were unaffected by the fish manipulation. Total invertebrate densities did not differ significantly between fish treatments, but total invertebrate biomass was significantly greater where fish density was reduced, averaging 30% higher over the course of the study. Likewise, manipulation of fish in the bluespotted sunfish–pickerel pond had few significant effects on individual taxa and size classes. There were no significant effects on total invertebrate abundance in the bluespotted sunfish–pickerel pond.
3. Our results provide direct experimental evidence consistent with the collective evidence from previous work, suggesting that the impact of fish predation on density and biomass of invertebrate prey in littoral habitats is variable, but generally weak. Invertebrates that coexist successfully with fish in littoral systems probably are adept at taking advantage of refugia offered by the structurally complex physical environment.  相似文献   

11.
Despite a recent surge of interest in temporary lentic systems, a strong theory linking the biota to its environment has not emerged. Using data from 10 temporary ponds at Mammoth Cave National Park, Kentucky, USA, we investigated how invertebrate communities were structured along environmental gradients, both between and within ponds. Samples were collected with a benthic corer in winter and spring, and a sweep net in spring. Six between-pond and two within-pond datasets were created. Between-pond analyses yielded significant CCA’s with only one of the six data sets. The ranges of environmental variables (EV’s) within ponds were often similar to the ranges of EV’s when averaged and compared between ponds. Some taxa were aggregated in a single pond, and richness increased with pond area. The theory that richness increases with hydroperiod did not apply to these systems. Within-pond analyses yielded more consistent relationships, with both CCA’s being significant. Sample depth was the best predictor of invertebrate richness and abundance, with most taxa preferring shallow habitats. Richness and abundance were higher in both shallow ponds and shallow areas of deep ponds than in deep areas of deep ponds. Standardizing sample depth may be an effective way to remove this gradient as a confounding variable in future research. The presence of within-pond gradients, possibly coupled with the limited dispersal and random colonization of tolerant taxa, makes between-pond comparisons difficult. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorised users. Handling editor: S. Declerck  相似文献   

12.
1. We experimentally reduced densities of predatory fish in replicated 2 m2 areas of the littoral zone in two ponds to test whether density and biomass of invertebrates would respond to release from fish predation. The ponds are of similar size and in close proximity, but support different fish assemblages: bluegills ( Lepomis macrochirus Rafinesque) and largemouth bass ( Micropterus salmoides (Lacepede)) in one pond, and bluespotted sunfish ( Enneacanthus gloriosus (Holbrook)) and chain pickerel ( Esox niger Lesueur) in the other. Fish densities were reduced to less than 15% of ambient levels in both experiments.
2. In the bluegill–bass pond, density and biomass of most invertebrate taxa and size classes were unaffected by the fish manipulation. Total invertebrate densities did not differ significantly between fish treatments, but total invertebrate biomass was significantly greater where fish density was reduced, averaging 30% higher over the course of the study. Likewise, manipulation of fish in the bluespotted sunfish–pickerel pond had few significant effects on individual taxa and size classes. There were no significant effects on total invertebrate abundance in the bluespotted sunfish–pickerel pond.
3. Our results provide direct experimental evidence consistent with the collective evidence from previous work, suggesting that the impact of fish predation on density and biomass of invertebrate prey in littoral habitats is variable, but generally weak. Invertebrates that coexist successfully with fish in littoral systems probably are adept at taking advantage of refugia offered by the structurally complex physical environment.  相似文献   

13.
The relationship of fish age class to parasitism by Paulisentis missouriensis was determined by sampling at least 29 creek chubs, Semotilus atromaculatus, from Easly Creek, Richardson County, Nebraska, every month from February 1996 to March 1997. In general, mean abundance and prevalence of the acanthocephalans increased with the age or length of chubs. It is unlikely that this distribution is explained by increased consumption of intermediate hosts by older, larger fish or by predatory fish acquiring parasites from paratenic hosts. The intermediate host for P. missouriensis is the cyclopoid copepod Acanthocyclops robustus, and creek chubs do not consume more microscopic crustaceans as they age or grow. Instead, the percentage of fish in the diet of creek chubs increases. Furthermore, P. missouriensis apparently does not use paratenic hosts. In laboratory infections, P. missouriensis survived predation of its original definitive host and transferred to the predator. Postcyclically transmitted P. missouriensis survived at least 14 days in the intestine of creek chubs, where they localized around the first flexure beyond the stomach. All stages of development of both sexes were transferred successfully. Postcyclic transmission is a plausible explanation, in some cases, for the greater worm burden frequently observed in older, larger hosts and for the occurrence in top carnivores of parasites not known to have paratenic hosts. This method of transmission appears to result in distribution of acanthocephalans to groups of animals that otherwise would be inaccessible.  相似文献   

14.
Our aim was to determine how beavers affect habitats and food resources for juvenile salmon in the Kwethluk River in western Alaska.
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15.
Positive interactions are important in influencing communities. Facilitation is one such influence in which a species makes the environment more suitable for others species. Ecological engineers are powerful facilitators as they physically modify habitats in a way that alters resource availability. We studied the effect of beaver flooding on breeding success of a dabbling duck, the teal. Our long-term (1988-2001) study was conducted in a boreal area where dabbling ducks face resource limitation. We focused on two aspects of facilitation: resource enhancement and habitat amelioration. We found that pond use by teal broods systematically increased upon beaver flooding. Beaver ponds harboured more resource, i.e. aquatic invertebrates than undisturbed waters, and invertebrate abundance was reflected as enhanced teal brood density. Beaver inundated shores were shallower than those of other waters, making their habitat structure more favourable to ducklings. Consequently, teal brood mortality was lower in beaver ponds than in waters unaffected by beaver. In boreal areas beaver facilitate teal breeding by enhancing both resource abundance and habitat structure.  相似文献   

16.
Synopsis We examined the short-term effects of the natural catastrophic collapse of a beaver dam on downstream benthic insect density and fish community structure in a headwater tributary of the Mississippi River. The catastrophic collapse of the dam and ensuing flash flood resulted in a dramatic (< 90%) decrease in benthic insect density in riffle and pool habitats. Sixty days after collapse of the dam, insect densities in riffles were 62% of pre-collapse densities. Insect recolonization of pools was slower than for riffles; 60 days after collapse of the dam insect densities in pools were 8% of pre-collapse levels. Collapse of the beaver dam altered the structure of the downstream fish community by causing a short-term (2–4 days) influx of pond species, resulting in a brief increase in species richness and abundance. Fish species richness and abundance then decreased for 4–60 days to levels below those prior to the collapse.  相似文献   

17.
Beaver pond biogeochemical effects in the Maryland Coastal Plain   总被引:4,自引:0,他引:4  
The fluxes and concentrations ofmaterials from two contiguous second-order watershedsin the Coastal Plain of Maryland, U.S.A. were measuredfor six years prior to and six years subsequent to theformation of a 1.25 ha beaver pond near the bottom ofone of the watersheds. The watersheds have a clayaquiclude and were equipped with V-notch weirs andcontinuous volume-integrating water samplers. Thebeaver pond reduced annual discharge of water,total-N, total-P, dissolved silicate, TOC, and TSS by8, 18, 21, 32, 28, and 27%, respectively. Most ofthe total-N reduction was due to increased retentionof nitrate in the winter and spring and TON in thewinter and summer. Most of the total-P reduction wasthe result of retention of both TPi and TOP in thewinter and summer. Dissolved silicate retentionpeaked in the spring, while TOC and TSS retentionpeaked in the winter. Prior to the formation of thebeaver pond, concentrations of TON, TPi, TOP, TOC, andTSS had highly significant correlations with streamdischarge, especially in the winter, but subsequent tothe pond there was little or no relationship betweenthese concentrations and stream discharge. However,concentrations of nitrate in the spring and ammoniumin the summer were highly correlated with streamdischarge both before and after the formation of thebeaver pond and regressions of discharge versusconcentrations of these nutrients explained more ofthe variation in concentrations after the formation ofthe pond.  相似文献   

18.
 The objective of this study was to determine if pond permanence and vertebrate predation (by fish and waterfowl) affect invertebrate community structure in the mudflat habitat of floodplain ponds. Invertebrate communities were studied for 1 year in four Mississippi River floodplain ponds with different hydroperiods. Pond 1 experienced five dry periods, pond 2 experienced four, pond 3 dried once, and standing water remained in pond 4 for the entire year. Vertebrate predator exclusion treatments (all access, no access, small-fish access and cage controls) were placed in all ponds. As pond duration increased, predatory invertebrate richness and abundance increased while overall invertebrate richness and abundance decreased. With the exception of the cladoceran Diaphanosoma, all commonly encountered taxa were strongly affected by pond permanence in terms of abundance, biomass and, generally, individual biomass. Taxa were nearly early divided between those that were more abundant in less permanent ponds and those that were more abundant in longer-duration ponds. Invertebrate taxa richness, abundance, and total biomass were lower in the all-access treatment than in the treatments that restricted predator access, and these effects were stronger in the more permanent ponds. In general, there were no significant differences in responses to the treatments with small-fish access and no access. These results support models that predict relatively weak effects of predation in frequently disturbed habitats. Received: 30 May 1995 / Accepted: 21 June 1996  相似文献   

19.
1. Modelling the effects of climate change on freshwater fishes requires robust field‐based estimates accounting for interactions among multiple factors. 2. We used data from an 8‐year individual‐based study of a wild brook trout (Salvelinus fontinalis) population to test the influence of water temperature on season‐specific growth in the context of variation in other environmental (i.e. season, stream flow) or biotic factors (local brook trout biomass density and fish age and size) in West Brook, a third‐order stream in western Massachusetts, U.S.A. 3. Changes in ambient temperature influenced individual growth rates. In general, higher temperatures were associated with higher growth rates in winter and spring and lower growth rates in summer and autumn. However, the effect of temperature on growth was strongly context‐dependent, differing in both magnitude and direction as a function of season, stream flow and fish biomass density. 4. We found that stream flow and temperature had strong and complex interactive effects on trout growth. At the coldest temperatures (in winter), high stream flows were associated with reduced trout growth rates. During spring and autumn and in typical summers (when water temperatures were close to growth optima), higher flows were associated with increased growth rates. In addition, the effect of flow at a given temperature (the flow‐temperature interaction) differed among seasons. 5. Trout density negatively affected growth rate and had strong interactions with temperature in two of four seasons (i.e. spring and summer) with greater negative effects at high temperatures. 6. Our study provided robust, integrative field‐based estimates of the effects of temperature on growth rates for a species which serves as a model organism for cold‐water adapted ectotherms facing the consequences of environmental change. Results of the study strongly suggest that failure to derive season‐specific estimates, or to explicitly consider interactions with flow regime and fish density, will seriously compromise our ability to predict the effects of climate change on stream fish growth rates. Further, the concordance we found between empirical observations and likely energetic mechanisms suggests that our general results should be relevant at broader spatial and temporal scales.  相似文献   

20.
Winter processes such as overwinter survival and growth of individuals can have wide-ranging consequences for population dynamics and communities within and across seasons. In freshwater organisms winter processes have been mainly studied in fish despite that invertebrates also have substantial impacts on lake and pond food webs. One of the major invertebrate consumers in lake and ponds is the planktonic larvae of the dipteran insect Chaoborus spec. However, while much is known about Chaoborus feeding ecology, behaviour and structuring role in food webs, its winter ecology and how it affects its populations are poorly understood. Here size- and density-dependent winter mortality and body growth of late Chaoborus flavicans larvae were quantified over naturally occurring size and density ranges in autumn and under natural winter conditions using two field enclosure experiments. Winter mortality increased with autumn density but decreased with autumn body size while winter growth rates decreased with autumn density and body sizes. There was also a density- and size-independent background mortality component. The proportion of pupae found in spring decreased strongly and exponentially with autumn density. These results may explain the commonly observed univoltine life cycle and multi-annual density fluctuations in northern Chaoborus populations. They further demonstrate the relevance of winter processes and conditions for freshwater invertebrates and ecosystems.  相似文献   

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