Identifying biotic drivers of population dynamics in a benthic–pelagic community

Benthic species and communities are linked to pelagic zooplankton through life‐stages encompassing both benthic and pelagic habitats and through a mutual dependency on primary producers as a food source. Many zooplankton taxa contribute to the sedimentary system as benthic eggs. Our main aim was to investigate the nature of the population level biotic interactions between and within these two seemingly independent communities, both dependent on the pelagic primary production, while simultaneously accounting for environmental drivers (salinity, temperature, and oxygen conditions). To this end, we applied multivariate autoregressive state‐space models to long (1966–2007) time series of annual abundance data, comparing models with and without interspecific interactions, and models with and without environmental variables included. We were not able to detect any direct coupling between sediment‐dwelling benthic taxa and pelagic copepods and cladocerans on the annual scale, but the most parsimonious model indicated that interactions within the benthic community are important. There were also positive residual correlations between the copepods and cladocerans potentially reflecting the availability of a shared resource or similar seasonal dependence, whereas both groups tended to correlate negatively with the zoobenthic taxa. The most notable single interaction within the benthic community was a tendency for a negative effect of Limecola balthica on the amphipods Monoporeia affinis and Pontoporeia femorata which can help explain the observed decrease in amphipods due to increased competitive interference.     

Effects of mixing on the pelagic food web in shallow lakes

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Relative Sensitivity Distribution of Freshwater Planktonic Crustaceans to Trace Metals

Cladocerans and copepods are common inhabitants of ponds and lakes, but assessments of the toxic effects of chemicals on aquatic organisms are often based mainly on toxicity data for cladocerans of the genus Daphnia. The paucity of toxicity data for copepods and other species of cladocerans raised the question whether toxicity data derived primarily from daphniids can be extrapolated to protect other planktonic organisms. In this study, we address this question by systematically comparing between the sensitivities of cladocerans and copepods to the metals Cd, Cu, Pb, Ni, and Zn using the species sensitivity distributions. Our results indicate that cladocerans are consistently more sensitive than copepods to Cd and Cu. Most importantly, sensitivities of D. magna, D. pulex, and Ceriodaphnia dubia, the three most commonly tested cladocerans, to these metals are higher than those of most copepod species and fall within the range of those exhibited by other cladocerans. C. dubia, a cladoceran with relatively small body size, is more sensitive to all metals than most other cladoceran species. Our results imply that water quality criteria of trace metals derived from datasets dominated by Daphnia are likely also protective to copepods and most other cladocerans.     

Water Transparency Drives Intra-Population Divergence in Eurasian Perch (Perca fluviatilis)

Trait combinations that lead to a higher efficiency in resource utilization are important drivers of divergent natural selection and adaptive radiation. However, variation in environmental features might constrain foraging in complex ways and therefore impede the exploitation of critical resources. We tested the effect of water transparency on intra-population divergence in morphology of Eurasian perch (Perca fluviatilis) across seven lakes in central Sweden. Morphological divergence between near-shore littoral and open-water pelagic perch substantially increased with increasing water transparency. Reliance on littoral resources increased strongly with increasing water transparency in littoral populations, whereas littoral reliance was not affected by water transparency in pelagic populations. Despite the similar reliance on pelagic resources in pelagic populations along the water transparency gradient, the utilization of particular pelagic prey items differed with variation in water transparency in pelagic populations. Pelagic perch utilized cladocerans in lakes with high water transparency and copepods in lakes with low water transparency. We suggest that under impaired visual conditions low utilization of littoral resources by littoral perch and utilization of evasive copepods by pelagic perch may lead to changes in morphology. Our findings indicate that visual conditions can affect population divergence in predator populations through their effects on resource utilization.     

Plasticity of vertical distribution of crustacean zooplankton in lakes with varying levels of water colour

Zooplankton that are vulnerable to predation by planktivorousfish typically remain in deeper, darker water during the day.Vertical position may be affected by profiles of temperatureand oxygen concentration, but light levels are thought to becritically important. Water transparency in temperate lakesis strongly dependent on the concentration of dissolved organiccarbon (DOC). Therefore, different patterns of zooplankton verticalposition might be expected in lakes that vary in their levelof DOC. The importance of three factors (light, temperatureand oxygen) that might affect vertical position of major zooplanktongroups was evaluated in 10 small lakes located in central Ontario.The lakes, encompassing a wide range of water colour, were sampledin May, June and August of 2000. Small cladocerans were foundto be always higher in the water column than large cladocerans,while copepods were deeper. Small cladocerans had a weak responseto the predictive factors, but copepods and large cladoceranswere strongly affected. The vertical position of copepods wasconsistently dependent solely on Secchi depth. Surprisingly,for large cladocerans, temperature gradient was decisive inMay and June, and only in August was water transparency themost important variable. Overall, water transparency, as determinedby level of water colour, was the most important variable but,depending on taxonomic group and time of year, temperature andoxygen were also major determinants of vertical position.     

Climate control on the long-term anomalous changes of zooplankton communities in the Northwestern Mediterranean

In marine ecosystems, pelagic copepods, chaetognaths and jellyfish play a key role in matter and energy flow. While copepods support most food webs and the biological pump of carbon into the deep ocean, chaetognaths and jellyfish may affect the strength of the top-down control upon plankton communities. In this study, we show that the main events in the long-term variability of these functional groups in the Northwestern Mediterranean were tightly linked to changes of climate forcing of the North Atlantic sector. Large-scale climate forcing has altered the pelagic food-web dynamics through changes in biological interactions, competition and predation, leading to substantial changes manifested as bursts or collapses in zooplankton populations, and consequently to a major change ca. 1987. These events become more frequent in the 1980s and the early 1990s in the studied zooplankton functional groups suggesting a shift in the functioning of the pelagic ecosystem. The environmental modifications and the results reported here are therefore, indicators of a regime change pointing to a more regeneration-dominated system in the study area. We suggest a chain of mechanisms, whereby climate variation has modified the long-term dynamics of pelagic copepods, chaetognaths and jellyfish in the Ligurian Sea.     

A matter of timing: heat wave impact on crustacean zooplankton

1. Climate change has affected zooplankton phenology and abundance in many freshwater ecosystems. The strong temperature anomalies that characterise summer heat waves make these events particularly suitable to study the effects of different seasonal warming patterns on zooplankton. Since heat waves are expected to occur more frequently under continuing climate change, they may also allow us to investigate how freshwater systems will be affected in the future. 2. Using a long‐term data set (1991–2007) from a shallow, eutrophic lake in Germany, we identify time periods in spring and summer during which cyclopoid copepods and bosminids are particularly sensitive to changes in water temperature. Based on this knowledge, we consider why summer populations responded differently to recent heat wave events that occurred at different times in the season. 3. Linear regressions of moving averages suggested that water temperatures shortly before and shortly after the clear‐water phase (CWP) were crucial for summer development of bosminids and cyclopoid copepods, respectively. Algal food availability (diatoms and cryptophytes) in the first weeks after the CWP also strongly influenced the summer populations of the two zooplankton groups. 4. Inter‐annual differences in water temperature during the critical time periods at least partly explained the contrasting responses of cyclopoid copepods and bosminids to heat wave events. 5. Our findings indicate that the zooplankton response to climate warming, particularly to heat wave events, is critically dependent on the temporal pattern of elevated water temperatures. Beyond that, we show that the summer zooplankton populations react to periods of warming in relation to events in the plankton annual cycle (such as the CWP in eutrophic lakes) rather than to warming at a fixed time in the season.     

Reconstructing marine plankton food web interactions using DNA metabarcoding

Knowledge of zooplankton in situ diet is critical for accurate assessment of marine ecosystem function and structure, but due to methodological constraints, there is still a limited understanding of ecological networks in marine ecosystems. Here, we used DNA‐metabarcoding to study trophic interactions, with the aim to unveil the natural diet of zooplankton species under temporal variation of food resources. Several target consumers, including copepods and cladocerans, were investigated by sequencing 16S rRNA and 18S rRNA genes to identify prokaryote and eukaryote potential prey present in their guts. During the spring phytoplankton bloom, we found a dominance of diatom and dinoflagellate trophic links to copepods. During the summer period, zooplankton including cladocerans showed a more diverse diet dominated by cyanobacteria and heterotrophic prey. Our study suggests that copepods present trophic plasticity, changing their natural diet over seasons, and adapting their feeding strategies to the available prey spectrum, with some species being more selective. We did not find a large overlap of prey consumed by copepods and cladocerans, based on prey diversity found in their guts, suggesting that they occupy different roles in the trophic web. This study represents the first molecular approach to investigate several zooplankton–prey associations under seasonal variation, and highlights how, unlike other techniques, the diversity coverage is high when using DNA, allowing the possibility to detect a wide range of trophic interactions in plankton communities.     

Zooplankton Dynamics of Lake Escondido (Rio Negro,Argentina), with Special Reference to a Population of Boeckella gracilipes (Copepoda,Calanoida)

The zooplancton dynamics and the populations of Boeckella gracilipes Daday of Lake Escondido (Rio Negro, Argentina) were studied from March 1987 to March 1988. Rotifers and Cladocerans showed high densities during spring and summer, while copepods were densest in autumn. The vertical distribution study showed that rotifers prefer upper levels of the water column, while cladocerans and copepods prefer lower levels, especially during thermal stratification. Boeckella gracilipes had a sex ratio biassed toward female, though changes in the sex ratio were observed. The clutch size was high in winter when the proportion of ovigerous females was low. The instantaneous birth rates were low during winter and high in summer. This result is related to the thermal cycle of the lake, by the influence of temperature on egg development time. The mortality rates showed a similar pattern, and maximum values were registered in spring and summer.     

高山林线变化的更新受限机制研究进展

全球林线位置对气候变暖的响应表现为上升、无变化或下降等截然不同趋势,表明影响林线位置及动态的因子十分复杂,除了较普遍认为的低温调控机制外,还存在其它控制林线位置变化的机制。林线向上迁移开始于种子向林线以上的传播及幼苗在林线以上的定居,这些过程中的限制因子均会影响林线的位移,因此研究更新过程及其限制因子对理解高山林线对气候变化的响应具有重要的科学意义。主要从种子和幼苗两个关键阶段综述高山林线森林更新的研究进展。在种子阶段,夏季积温不足导致种子产量和活力下降,风速过低和浓密灌丛限制种子向林线以上传播,近地表的霜冻/水分胁迫和灌木释放的化感物质会阻碍种子在林线以上萌发。在幼苗阶段,除冬季低温外,生长季内较大的温度日振幅和偶然出现的冻害事件也是导致幼苗死亡的重要原因,而低温环境下的强烈光照引起的低温光抑制会显著降低生长季的光合作用;土壤低温、由土壤温度昼夜变化引起的冻举事件、夏季土壤干旱可能会导致幼苗光合作用下降和死亡率上升;积雪太浅会导致生长季早期幼苗水分供应的严重缺乏,但积雪太深会导致幼苗感染真菌的可能性增加;浓密的灌木和草本植物以及植食动物的啃食也会降低林线以上的幼苗存活率。气候变暖对林线幼苗定居的影响复杂且具有很大不确定性,需要进一步研究气候变暖导致的环境因子变化对林线更新各关键阶段的影响。未来气候变暖无疑会导致生长季起始日提前,结束日推迟,这很可能会增加生长季期间尤其是早期的低温冻害事件,对高山林线树种幼苗的存活具有重要影响。在未来研究中,需要找出定义生长季冻害事件的温度阈值,利用长期气象观测数据分析增温背景下生长季早期冻害事件特征的变化趋势,并进一步开展野外模拟增温实验以深刻理解林线树种的种子萌发和幼苗定居与生长季冻害事件的关系,加强对不同地区林线树种的繁殖策略研究,这将有助于人们进一步理解不同区域林线的形成机制并预测未来气候变化条件下林线的动态变化趋势。     

Resonance of plankton communities with temperature fluctuations

The interplay between intrinsic population dynamics and environmental variation is still poorly understood. It is known, however, that even mild environmental noise may induce large fluctuations in population abundances. This is due to a resonance effect that occurs in communities on the edge of stability. Here, we use a simple predator-prey model to explore the sensitivity of plankton communities to stochastic environmental fluctuations. Our results show that the magnitude of resonance depends on the timescale of intrinsic population dynamics relative to the characteristic timescale of the environmental fluctuations. Predator-prey communities with an intrinsic tendency to oscillate at a period T are particularly responsive to red noise characterized by a timescale of τ = T/2π. We compare these theoretical predictions with the timescales of temperature fluctuations measured in lakes and oceans. This reveals that plankton communities will be highly sensitive to natural temperature fluctuations. More specifically, we demonstrate that the relatively fast temperature fluctuations in shallow lakes fall largely within the range to which rotifers and cladocerans are most sensitive, while marine copepods and krill will tend to resonate more strongly with the slower temperature variability of the open ocean.     

Projected marine climate change: effects on copepod oxidative status and reproduction

Zooplankton are an important link between primary producers and fish. Therefore, it is crucial to address their responses when predicting effects of climate change on pelagic ecosystems. For realistic community‐level predictions, several biotic and abiotic climate‐related variables should be examined in combination. We studied the combined effects of ocean acidification and global warming predicted for year 2100 with toxic cyanobacteria on the calanoid copepod, Acartia bifilosa. Acidification together with higher temperature reduced copepod antioxidant capacity. Higher temperature also decreased egg viability, nauplii development, and oxidative status. Exposure to cyanobacteria and its toxin had a negative effect on egg production but, a positive effect on oxidative status and egg viability, giving no net effects on viable egg production. Additionally, nauplii development was enhanced by the presence of cyanobacteria, which partially alleviated the otherwise negative effects of increased temperature and decreased pH on the copepod recruitment. The interactive effects of temperature, acidification, and cyanobacteria on copepods highlight the importance of testing combined effects of climate‐related factors when predicting biological responses.     

The effects of turbidity on prey consumption and selection of zooplanktivorous Gasterosteus aculeatus L.

It is well documented that reduced visibility caused by elevated turbidity can affect feeding of fish, yet the extent to which selective zooplanktivory is altered in turbid conditions remains ambiguous. In this study, we examined the influence of natural sediment-induced turbidity on the overall prey consumption and selective predation of a common brackish water littoral zooplanktivore, the particulate feeding three-spined stickleback (Gasterosteus aculeatus L.). We hypothesized that the effects of turbidity on prey consumption and prey type selection would be pronounced due to the vision-oriented feeding of this species and that these effects would differ between genders. Using aquarium experiments with three different groups of cladocerans and copepods varying in size and behavior, we studied prey consumption and selectivity of this key planktivore in varying turbidity treatments. Our results indicated significantly decreased total prey consumption in the high turbidity treatments, as well as altered selective feeding on copepods and an enhanced preference for larger cladocerans. We found gender-dependent differences in prey consumption, which are consistent with observations of other visually feeding fish with sexual size dimorphism. We conclude that high turbidity, such as that occurring in shallow coastal areas, may affect selective feeding in vision-oriented zooplanktivores and that these effects may be gender-related.     

Examination of a great abundance of filefish, Pervagor spilosoma, in Hawaii

Synopsis The endemic filefish Pervagor spilosoma appeared in exceptionally large numbers throughout the major Hawaiian Islands during the spring of 1985 and remained abundant through 1988. We examined this occurrence with a perspective gained from studies of reef communities at the Island of Hawaii by one or both of us during most years from 1969 to 1994. P. spilosoma is known to vary in numbers, but such great abundances are rare. Although the species is considered a reef fish, individuals constituting the abundance were mostly in the water column, and spread from coastal reefs to the pelagic zone offshore. One might assume they represented an extraordinary recruitment of pelagic juveniles in the process of shifting to benthic habits, but our data indicate a more complex situation. Individuals in the water column tended to be larger than individuals near the reef, and although the former were feeding on plankton when collected, many had shifted to planktivory from earlier feeding on the benthos. Also, many in the water column were ailing, with moribund and dead individuals common, and these tended to carry parasites of types known to induce their hosts to rise toward the surface. We suggest that the great number of filefish we observed above near-shore reefs represented the shoreward fringe of a vast pool of individuals, varied in age, that had accumulated during the exceptionally long pelagic-juvenile period that is characteristic of tetraodontiforms.     

The effect of diapause emergence on the seasonal dynamics of a zooplankton assemblage

1. The role of seasonal phenology in the emergence of zooplankton from diapause in patterns of seasonal abundance in the water column was investigated in Oneida Lake, New York. Replicate emergence traps, placed in contact with the lake sediments at two locations (one at a shallow site and one at a deep site), were monitored between May and August.
2. Although six rotifer taxa showed a clear seasonal succession in the water column throughout the study period, all but one taxon emerged exclusively in spring. Three cladoceran and three calanoid copepod species, also present in the water column throughout the study period, again showed predominantly spring emergence. In contrast, three cyclopoid copepod species had distinct seasonal periods of emergence that corresponded, at least in part, to the timing of abundance peaks in the plankton.
3. These results for a single lake are largely consistent with patterns observed or inferred by other investigators for other lakes: variable dependence of abundance in the plankton on diapause emergence for species with long-lived diapausing eggs (i.e. rotifers, cladocerans and calanoid copepods), and much closer dependence for species with short-lived diapausing immature stages (i.e. cyclopoid copepods).     

Climate‐induced warming imposes a threat to north European spring ecosystems

Interest in climate change effects on groundwater has increased dramatically during the last decade. The mechanisms of climate‐related groundwater depletion have been thoroughly reviewed, but the influence of global warming on groundwater‐dependent ecosystems (GDEs) remains poorly known. Here we report long‐term water temperature trends in 66 northern European cold‐water springs. A vast majority of the springs (82%) exhibited a significant increase in water temperature during 1968–2012. Mean spring water temperatures were closely related to regional air temperature and global radiative forcing of the corresponding year. Based on three alternative climate scenarios representing low (RCP2.6), intermediate (RCP6) and high‐emission scenarios (RCP8.5), we estimate that increase in mean spring water temperature in the region is likely to range from 0.67 °C (RCP2.6) to 5.94 °C (RCP8.5) by 2086. According to the worst‐case scenario, water temperature of these originally cold‐water ecosystems (regional mean in the late 1970s: 4.7 °C) may exceed 12 °C by the end of this century. We used bryophyte and macroinvertebrate species data from Finnish springs and spring‐fed streams to assess ecological impacts of the predicted warming. An increase in spring water temperature by several degrees will likely have substantial biodiversity impacts, causing regional extinction of native, cold‐stenothermal spring specialists, whereas species diversity of headwater generalists is likely to increase. Even a slight (by 1 °C) increase in water temperature may eliminate endemic spring species, thus altering bryophyte and macroinvertebrate assemblages of spring‐fed streams. Climate change‐induced warming of northern regions may thus alter species composition of the spring biota and cause regional homogenization of biodiversity in headwater ecosystems.     

Predicted warming and browning affect timing and magnitude of plankton phenological events in lakes: a mesocosm study

1. Aquatic ecosystems in Northern Europe are expected to face increases in temperature and water colour (TB) in future. While effects of these factors have been studied separately, it is unknown whether and how a combination of them might affect phenological events and trophic interactions. 2. In a mesocosm study, we combined both factors to create conditions expected to arise during the coming century. We focused on quantifying effects on timing and magnitude of plankton spring phenological events and identifying possible mismatches between resources (phytoplankton) and consumers (zooplankton). 3. We found that the increases in TB had important effects on timing and abundance of different plankton groups. While increased temperature led to an earlier peak in phytoplankton and zooplankton and a change in the relative timing of different zooplankton groups, increased water colour reduced chlorophyll‐a concentrations. 4. Increased TB together benefitted cladocerans and calanoid copepods and led to stronger top‐down control of algae by zooplankton. There was no sign of a mismatch between primary producers and grazers as reported from other studies. 5. Our results point towards an earlier onset of plankton spring growth in shallow lakes in future with a stronger top‐down control of phytoplankton by zooplankton grazers.     

Recent invasion by a non-native cyprinid (common bream Abramis brama) is followed by major changes in the ecological quality of a shallow lake in southern Europe

We present an example of how an invasion by a non-native cyprinid (common bream, Abramis brama (Pisces: Cyprinidae), hereafter bream) in a natural shallow lake in southern Europe (Lake Montorfano, northern Italy) may have adversely affected the state of the lake’s ecosystem. In less than two decades, bream became the most abundant species and characterized by a stunted population with asymptotic length 33.5 cm, an estimated mean length at first maturity of 19.6 cm, a total mortality rate of 0.64 year^?1 and a diet overwhelmingly dominated by microcrustaceans. Following bream establishment, nutrients and phytoplankton biomass rose, the proportion of Cyanobacteria by numbers increased markedly and water transparency decreased. Total zooplankton abundance increased with a marked increase in small cladocerans and copepods, whereas the abundance of large herbivorous cladocerans did not change. The coverage of submerged macrophytes declined, as did the abundance of native pelagic zooplanktivorous fish. The composition of the fish community shifted towards a higher proportion of zoobenthivorous species, such as bream and pumpkinseed (Lepomis gibbosus). Our results indicate that bream affected water quality through bottom-up mechanisms, while top-down effects were comparatively weak. Selective removal of bream and perhaps stocking of native piscivores might improve the ecological status of the lake.     

Zooplankton dynamics in the upstream part of Stratos reservoir (Greece)

The investigation of the zooplankton community in the upstream part of Stratos reservoir during a 24 months survey (September 2004–August 2006) revealed 26 invertebrate species (14 rotifers, 6 cladocerans, 5 copepods and one mollusk larva). The mean abundance of the total zooplankton was higher in the first sampling period (2004–2005) and ranged between 8.81 and 47.74 ind. L⁻¹, than the second period (2005–2006) when fluctuated between 1.91 and 43.09 ind. L⁻¹. The seasonal variation was strongly influenced by the presence of rotifers, which accounting on average for 68.4% in total. Among them Keratella cochlearis and the order Bdelloidea were numerically the most important, while Macrocyclops albidus prevailed among the copepods and Bosmina longirostris among the cladocerans. Dreissena polymorpha was the only mollusk found in the zooplankton community. Rotifers, copepods and cladocerans showed a seasonal succession with the former preceding in the abundance having their first maximum in spring, while copepods and cladocerans followed, having peaks of abundance in early summer and in autumn, respectively. No seasonal succession among the cladoceran species was observed. The intense water flow in the upstream part of the reservoir, as well as temperature, conductivity, DO, pH, phosphates and silicates, were significant parameters controlling abiotic and biotic elements of the ecosystem and consequently influencing the seasonal variation and the dynamics of the zooplankton community.     

Rehabilitation of shallow lakes: time to adjust expectations?

We investigated the relative importance of flood defenses and other environmental variables for the cladoceran and copepod communities in floodplain water bodies in Southeastern Norway. The water bodies covered gradients of size, distance to the river and water chemistry, and half of them were located behind flood defenses. The effects of environmental variables on the communities were analyzed using redundancy analysis. The set of environmental variables accounted for more of the explained variation in the cladoceran community than in the copepod community. Water quality was much more important than flooding-related variables for both communities. Although cladoceran species richness was slightly higher in water bodies outside flood defenses, total nitrogen, total organic carbon (TOC), and water body area were the most important factors for the cladocerans. Macrophyte coverage was the most important variable for both species richness and community structure of copepods. Although our results show that water quality and spatial/structural variables are more important than flood defenses in structuring the communities, the effect of TOC on the cladoceran community could likely be mediated through effects of flood defense on TOC. Prospects for a continued anthropogenic pressure on river floodplains raise concern for the future of these unique ecosystems.