Vertical, lateral and longitudinal movement of zooplankton in a large river

1. The spatial distribution and movement patterns of zooplankton in large rivers are little known compared with those in lake environments. We conducted a series of studies in the Ohio River (U.S.A.) during the low flow period to assess diel vertical (DVM), longitudinal and lateral movement of crustacean zooplankton. 2. The dominant large zooplankter, the copepod Eurytemora affinis, showed a consistent vertical migration pattern of daytime ascent and night‐time descent during all sampling periods – the reverse of the most common migratory pattern of zooplankton in lakes. The cladoceran Bosmina migrated in a similar way in two of the three sampling periods. Surveys taken longitudinally in the river showed similar trends for both taxa. 3. During the lateral surveys, E. affinis was significantly more abundant in the shallow littoral zone during the night than in the daytime. The combination of vertical and lateral movement patterns along with the diel distribution of zooplanktivorous fish suggest that these movements are a predator‐avoidance mechanism. 4. Sampling programmes in large rivers should consider that larger zooplankton such as E. affinis may not be randomly distributed in the river channel and behaviours such as diel vertical migration may be just as evident in river habitats as in lakes.     

千岛湖浮游甲壳动物垂直分布与昼夜垂直移动

2004年6月、9月、12月及2005年3月于千岛湖温馨岛站点(29°38′10.5″N,119°01′54.1″E)进行浮游甲壳动物垂直分布分层采样,分析千岛湖浮游甲壳动物的垂直分布特征及优势种昼夜垂直移动状况。结果显示,千岛湖浮游甲壳动物主要分布在10～21 m水层,不同季节中心水层分布深度12月9月6月3月;浮游动物昼夜垂直移动春(3月)、夏(6月)两季幅度较大,秋(9月)、冬(12月)季节幅度较小;浮游甲壳动物优势种在各个水层都有分布,密集区大都集中在中上水层(10～21 m),不同种类昼夜垂直移动幅度有所不同;昼夜垂直移动显著的种类有特异荡镖水蚤(Neutrodiaptanus incongruens)、近邻剑水蚤(Cyclops vicinus)、球状许水蚤(Schmackeria forbesi)、透明溞(Daphnia hyaline)、长额象鼻溞(Bosmina longirostris),不显著的种类为右突新镖水蚤(Neodiaptomus schmackeri)、短尾秀体溞(Diaphanosoma brachyurum)。光照、温度、饵料是影响千岛湖浮游甲壳动物垂直分布及昼夜垂直移动的主要因素。     

What goes down must come up: symmetry in light-induced migration behaviour of Daphnia

During a short period of the year, Daphnia may perform a phenotypically induced diel vertical migration. For this to happen, light-induced swimming reactions must be enhanced both at dawn and at dusk. Enhanced swimming in response to light intensity increase can be elicited by fish-associated kairomone in the laboratory, if food is sufficiently available. However, during the light change at dusk the Daphnia are still in the hypolimnion, where no fish kairomone is present and both temperature and food availability is low. Still, what goes down must come up. This raises questions about how Daphnia tunes its light-induced swimming behaviour to prevailing conditions such that a normal diel vertical migration can be performed. We investigated the symmetry in behavioural mechanism underlying these diel vertical migrations in the hybrid Daphnia galeata × hyalina (Cladocera; Crustacea), with special interest for the environmental cues that are known to affect swimming in response to light increase. That is, we tested whether fish- associated kairomone, food availability, and temperature affected both swimming in response to light intensity increase and decrease similarly. We quantified swimming behaviour during a sequentially increased rate of light change. Vertical displacement velocity was measured and proved to be linearly related to the rate of the light change. The slope (PC) of the function depends on the value of the factors kairomone concentration, food availability, and temperature. The changes of the PC with kairomone concentration and with temperature were similar both at light intensity increases and decreases. The PC also increased with food concentration, although during light increases in a different way from during light intensity decreases. Low food availability inhibited swimming in response to light intensity increase, but enhanced swimming in response to light intensity decrease. Hence, ascent from the deep water layers with low food concentration at dusk is facilitated. These causal relations are part of a proximate decision-making mechanism which may help the individual Daphnia to tune migration to predation intensity and food availability.     

Size distribution of Daphnia longispina in the vertical profile

D. longispina of the meromictic lake El Tobar is a round-headed form. It never has a helmet, but in summer a small proportion of immature individuals (0.9–1.2 mm females and males) have one or two neck teeth. The size structure of this Daphnia population, as well as the vertical distribution and migration of different size-classes, were studied in September and November of 1991 and April of 1992. The large variation in mean size and size at first reproduction, as well as the occurrence of different patterns of vertical migration are interpreted as responses to different predator situations. At the end of April, when Daphnia mortality by visually hunting predators is dominating, a typical nocturnal migration is adopted and size distribution is biased to smaller size classes. In November, when mortality is mainly attributed to the nocturnally migrating Chaoborus, Daphnia shows a reversed migration pattern. In September, when the population of Daphnia is responding to both visual (fish) and non-visual predators (Chaoborus), it adopts a pattern of twilight migration. The presence of neck teeth in vulnerable size classes in September might be an additional adaptative response to Chaoborus predation. In September, the size structure of the Daphnia population is shifted to larger classes and the vertical distribution of size classes shows a pronounced segregation between juveniles and adults. Juveniles are found closer to the surface, while adults dwell predominantly in the rich, deep waters near the oxicline. This suggests that an additional advantage of the ascent of the adult Daphnia exploiting those deep resources is the release of young in more oxygenated and warmer waters. The Daphnia population of lake El Tobar is known to be clonally diverse, and the changing frequency of genotypes could play an important part in the observed seasonal differences in behaviour and size.     

Hydroacoustic Estimates of Fish Population Depths and Densities at Increasingly Longer Time Scales

To assess the reliability of depth and density estimates of vertically migrating fish (Coregonus spp.), repeated hydroacoustic surveys were conducted in a stratified lake during the same day, during subsequent nights within a week, and over four months of a year. The reliability of the hydroacoustic method, determined by the coefficient of variance of population depth of fish, was high within nights, but decreased with increasingly longer time scales. Volumetric fish densities were likewise more variable between surveys at longer temporal distances than at short‐term scales. Night‐time population depths of fish were highly correlated to the vertical position of the thermocline. Over the diel cycle, the depth distribution of the coregonids was also correlated to light intensities at the surface. We conclude that single hydroacoustic surveys reliably reflect the depth distribution and volumetric density of the fish populations in a deep lake at temporal scales of up to a week. At longer scales, physical forcing may change the depth distribution of fish, and population dynamics may alter the density estimates substantially (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effects of aquatic insect predators on zooplankton in fishless ponds

We removed the surface-orienting aquatic insects from a fishless pond to determine their predation effects on zooplankton behavior and size structure. A second fishless pond served as the unmanipulated reference system in this two year study. In the reference pond and the treatment pond prior to manipulation, daphnids exhibited pronounced diel vertical migrations. Following the removal of surface-orienting aquatic insects from the treatment pond, daphnid migration changed to a reverse migration strategy that was significantly different from that observed in the reference system. Average daphnid body size increased significantly following predator removal in the treatment system. Our data indicate that predation by aquatic insect predators, such as notonectids and dytiscids, may affect daphnid migration behavior in fishless systems. Vertical migration by daphnids may allow coexistence with surface-orienting insects in ponds that are deep enough to provide a spatial refuge from these predators.     

Prey selection and diel feeding of the freshwater jellyfish, Craspedacusta sowerbyi

1. The diet of the invasive freshwater jellyfish, Craspedacusta sowerbyi (Lankester), was assessed by analysing its stomach contents. 2. The medusae ingested various zooplankton and benthic prey in the 0.1–3.0 mm size range. The selectivity indices for prey showed that larger zooplankton (0.4–1.4 mm) and active prey such as copepods were preferred; small potential prey, and the loricate rotifer Keratella cochlearis in particular, were almost never consumed. Measurements suggest that spacing between the tentacle branches could determine prey selection. Another possibility is that some prey are too small or slow to activate nematocysts after collision with the predator. Prey > 1.4 mm can probably escape or are otherwise too large to be handled. 3. The medusae of Craspedacusta medusae stayed in deep water during daytime and migrated upwards at night. The greatest number of prey in freshwater jellyfish stomachs was detected at night. 4. Calculation of consumption rates, based on mean stomach contents and digestion times of 4–5 h, showed that, on average, one medusa takes 190 zooplankton prey day^??1 at a mean prey density of about 100 L^??1. At the observed medusae density of 1 m^??3, the cropping rate was about 0.2 zooplankton prey L^??1 day^??1.     

Where to stay by night and day: Size-specific and seasonal differences in horizontal and vertical distribution of Chaoborus flavicans larvae

1. Data on the distributions of pelagic and benthic Chaoborus flavicans larvae were gathered in 1994 and tested for their agreement with the predator avoidance hypotheses. The development of all Chaoborus life stages, as well as the horizontal and vertical distribution in the four larval instars, was followed from May until October. We expected the largest larvae to dwell deeper by day, thus avoiding predation by visually foraging fish.
2. In agreement with this prediction body size increased with daytime depth, and this was true both between and within instars. The migration amplitude consequently increased with larval instar.
3. There was also evidence for horizontal migration, mainly in the third but also in the fourth instar.
4. Along a horizontal transect with increasing depth, locations with many benthic larvae had fewer pelagic larvae. Oxygen concentration was a good predictor of maximum benthic larval depth for most of the season but failed to predict their distribution in autumn.     

On the combined analysis of proximate and ultimate aspects in diel vertical migration (DVM) research

Although evolutionary ecologists agree that proximate and ultimate aspects are two sides of one coin, they are seldom interested in studies on physiological and behavioural mechanisms at the base of ecological phenomena. Nevertheless, these mechanisms are objects of selection and evolved to realise adaptive significances. This paper is a plea to bring both fields closer together, and, by means of an example of Diel Vertical Migration of Daphnia, some proximate and ultimate aspects are discussed. It is argued that light changes, not fish kairomone, is the primary cause for an individual to swim downwards at dawn and upwards at dusk. However, what is called a causal factor might differ when ecosystems or individuals are studied. In addition, causality in ecology is not simple, and has the character of a `set of necessary conditions'. To illustrate the importance of proximate analyses in DVM, two basic response mechanisms are discussed: Photobehaviour system 1 and 2. The physiological character of these systems leads to a fixed type of migration or to a phenotypically induced DVM, respectively. The adaptive significance of the first might be a reduction of the hazardous effects of UV radiation and of the second a lowering of mortality due to visually hunting predators.     

Life-history responses of a mayfly to seasonal constraints and predation risk

Abstract. 1. An experiment was conducted in the laboratory to examine the effects of photoperiod and predation risk on life-history variation in the mayfly Ephemerella subvaria .
2. Both photoperiod and predation risk affected age at maturity significantly but neither factor affected size at maturity. Mayflies perceiving themselves to be late in the growing season matured in fewer days than those perceiving themselves to be early in the growing season. The presence of predators delayed mayfly maturity significantly.
3. These results suggest that the large variation in life-history traits observed in aquatic insects may be attributed partially to seasonality but that other biotic and abiotic factors may also underlie variation in these traits.     

Vertical distribution and diel migration of flagellated phytoplankton in a small humic lake

The vertical distributions and migrations are described of the most abundant flagellated phytoplankton species from the summer community of a small forest lake in southern Finland. The lake showed a steep and stable thermal stratification with a shallow oxygenated epilimnion. Horizontal variation of phytoplankton distribution within the lake was tested on two scales and found to be statistically significant only in the case of Mallomonas reginae. The vertical distribution of flagellated phytoplankton was assessed by reference to the distribution of a non-motile, neutrally buoyant species Ankyra judayi. Statistically significant, active vertical positioning was demonstrated for all the flagellates examined with the exception of Spiniferomonas bourrellyi. Diel vertical migrations were apparent for all species showing active positioning and the pattern of an evening descent and a morning ascent was ubiquitous. The extent and timing of diel migrations varied between species. The most extensive migrations were by Cryptomonas marssonii which crossed a temperature gradient of 14 °C and penetrated far into the anoxic hypolimnion. Several categories of competitive advantage can be gained by species undertaking such diel vertical migrations.     

Growth and survival of Daphnia in epilimnetic and metalimnetic water from oligotrophic lakes: the effects of food and temperature

SUMMARY 1. In oligotrophic lakes, phytoplankton and bacteria growing in the deep chlorophyll maximum in the cool metalimnion of lakes often dominate biomass and production, but the importance of this source of food for zooplankton is unknown.
2. During much of the day, Daphnia rosea in two mountain lakes inhabited deep chlorophyll layers where food availability was at least equal to that in the epilimnion.
3. To determine the importance of the two strata (epilimnion and metalimnion) for Daphnia , we used a cross-classified factorial experiment to measure how epilimnetic and metalimnetic food and temperature (10 and 16 °C) influenced survival, growth and reproduction.
4. Daphnia survived and grew better when fed seston from the epilimnion of one lake, although chlorophyll, particulate nitrogen and particulate carbon were 2–2.5 times greater in the metalimnion.
5. Temperature had no significant influence on Daphnia survival or growth. Similar results were obtained with food from the second lake, with Daphnia surviving and reproducing better when provided with epilimnetic, rather than metalimnetic food, although the quantities of chlorophyll and carbon in the two strata were similar.
6. Food quality, rather than quantity or temperature, appeared to be the most important determinant influencing survival, growth and reproduction, and the greater food quantity in the metalimnia was not used effectively by the Daphnia .     

Predation on and by pelagic Turbellaria in some lakes in Brazil

Planktonic Turbellaria are of common occurrence in both natural and man-made lakes in Brazil. Experiments were performed in 1987 and 1989 to determine which zooplankton species are consumed by predatory Mesostoma sp. from three natural lakes in the Rio Doce Valley. Experiments were also performed in 1989 with a yet unidentified flatworm from Barra Bonita reservoir. Both predators consumed Daphnia and Ceriodaphnia at a high rate: 4 individuals per day in the case of Mesostoma sp., a large species and 1.5 ind day^–1 in the case of the smaller species from Barra Bonita reservoir. Consumption of copepod nauplii, copepodids and adults was much lower, and Ostracoda were not consumed at all. Experiments on food selectivity showed a clear preference by the flatworms for cladocerans.In the lakes studied, flatworms are heavily preyed upon by larvae of Chaoborus and by Mesocyclops species.Turbellaria densities in the natural lakes were around 300 individuals per cubic meter, whereas in Barra Bonita reservoir, 1000 individuals per cubic meter was a mean value in a fifteen days study.Federal University of São Carlos - Laboratory of LimnologyUniversity of São Paulo - Canter of Water Resource and Applied EcologyPost-graduate student - University of São Paulo     

Responses to fish predation and nutrients by plankton at different levels of taxonomic resolution

1. To improve mechanistic understanding of plankton responses to eutrophication, a mesocosm experiment was performed in the shallow littoral zone of a south Swedish lake, in which nutrient and fish gradients were crossed in a fully factorial design. 2. Food chain theory accurately predicted total biomass development of both phyto‐ and zooplankton. However, separating zooplankton and algae into finer taxonomic groups revealed a variety of responses to both nutrient and fish gradients. 3. That both nutrients and fish are important for phytoplankton dynamics was seen more clearly when viewing each algal group separately, than drawing conclusions only from broad system variables such as chlorophyll a concentration or total phytoplankton biovolume. 4. In some taxa, physiological constraints (e.g. sensitivity to high pH and low concentrations of free CO₂) and differences in competitive ability may be more important for the biomass development than fish predation, grazing by herbivorous zooplankton, and nutrient availability. 5. We conclude that food chain theory accurately predicted responses in system variables, such as total zooplankton or algal biomass, which are shaped by the dynamics of certain strong interactors (‘keystone species’), such as large cladocerans, cyanobacteria and edible algae (<50 μm), whereas responses at finer taxonomic levels cannot be predicted from current theory.     

Diel migration and vertical distribution of Cladocera in Lake D. Helvecio (Minas Gerais,Brazil)

Six genera of Clad ocera (Diaphanosoma, Daphnia, Ceriodaphnia, Moina, Bosmina, Bosminopsis), each of them usually with only one species were found in Lake D. Helvecio, a natural valley lake located in the eastern part of Brazil. Diurnal migratory movements of the organisms observed in this lake showed a different pattern in different species. Closely related species, which explore the same food source, live in different layers, thus avoiding interspecific competition. The migratory behaviour of the species was studied mainly in relation to temperature and oxygen distribution in the lake. Thus, analyses were made in the summer (January, 1978) when a strong stratification occurs with the establishment of a thermocline and an oxycline. Comparisons were made also with the data obtained in winter (July, 1978), when a complete mixing of water occurs.     

Spatial overlap of a predatory copepod,Acanthocyclops robustus,and its prey in a shallow eutrophic lake

Spatial overlap between Acanthocyclops robustus, with special emphasis on the adult females, and other zooplankton in one basin of a shallow (approximate depth of 2 m) eutrophic lake was studied.Horizontal distribution patterns were analysed on two dates. On both dates, most taxa examined showed large-scale patchiness between the three sections of the lake basin (approximate length of 1.2 km). Similarly, most taxa, with the important exception of the adult female Acanthocyclops robustus, were significantly patchily distributed on the small-scale (i.e. within sections). However, the intensity of such patchiness was, in general, relatively low. There was no consistent evidence of aggregation by the adult females or copepodites and adult males (the latter two were considered together) of the predator in such small-scale prey patches.Diurnal vertical distribution patterns were studied on two 24–25 hour periods. The first period was characterized by calm weather. Adult female, and perhaps male, Acanthocyclops robustus, Chydorus sphaericus, Bosmina Coregoni, Keratella cochlearis, Asplanchna species, Polyarthra vulgaris and Pompholyx sulcata seemed to show diurnal migration patterns, while seven other taxa showed consistent preferences for particular depths. Only copepod nauplii and Daphnia species were approximately evenly distributed. Negative correlations were found between the vertical distributions of the adult female predator and seven of the seventeen potential prey recognized.The first half of the second period was characterised by strong winds which abated during the second half. Most zooplankton taxa showed inconsistent heterogeneous vertical distributions or were homogeneously distributed with vertical heterogeneity developing towards the end of the period. Only Bosmina longirostris and Daphnia species seemed to show vertical migration patterns. Thus, no consistent vertical segregation between predator and prey was detected.     

Effects of Predation Risk on Habitat Selection by Water Column Fish, Benthic Fish and Crayfish in Stream Pools

Predation risk can affect habitat selection by water column stream fish and crayfish, but little is known regarding effects of predation risk on habitat selection by benthic fish or assemblages of fish and crayfish. I used comparative studies and manipulative field experiments to determine whether, (1) habitat selection by stream fish and crayfish is affected by predation risk, and (2) benthic fish, water column fish, and crayfish differ in their habitat selection and response to predation risk. Snorkeling was used to observe fish and crayfish in, (1) unmanipulated stream pools with and without large smallmouth bass predators (Micropterus dolomieui >200 mm total length, TL) and (2) manipulated stream pools before and after addition of a single large smallmouth bass, to determine if prey size and presence of large fish predators affected habitat selection. Observations of microhabitat use were compared with microhabitat availability to determine microhabitat selection. Small fish (60–100 mm TL, except darters that were 30–100 mm TL) and crayfish (40–100 mm rostrum to telson length; TL) had significantly reduced densities in pools with large bass, whereas densities of large fish and crayfish (> 100 mm TL) did not differ significantly between pools with and without large bass. Small orangethroat darters (Etheostoma spectabile), northern crayfish (Orconectes virilis), and creek chubs (Semotilus atromaculatus) showed significantly greater densities in pools without large bass. The presence of large smallmouth bass did not significantly affect depths selected by fish and crayfish, except minnows, which were found significantly more often at medium depths when bass were present. Small minnows and large and small crayfish showed the greatest response to additions of bass to stream pools by moving away from bass locations and into shallow water. Small darters and sunfish showed an intermediate response, whereas large minnows showed no significant response to bass additions. Response to predation risk was dependent on prey size and species, with preferred prey, crayfish and small minnows, showing the greatest response. Small benthic fish, such as darters, are intermediate between small water column fish and crayfish and large water column fish in their risk of predation from large smallmouth bass.     

Neutral markers, ecologically relevant traits, and the structure of genetic variation in Daphnia

Zooplankton, Daphnia in particular, are increasingly used as model organisms to investigate general evolutionary biological questions. I here discuss some recent insights into the patterns and processes determining genetic diversity within and genetic differentiation among natural populations of cyclically parthenogenetic Daphnia. I focus on three aspects: (1) the interplay of phenotypic plasticity and genetic polymorphism in explaining variability in ecologically relevant traits, (2) the patterns of genetic variation revealed by neutral markers and ecologically relevant traits, and (3) the evolutionary ecological importance of hybridization events in Daphnia. The need for studies on the evolutionary ecology of sexual reproduction and dispersal via ephippial eggs in Daphnia is stressed.     

夏季南黄海浮游动物的垂直分布与昼夜垂直移动

对 2 0 0 1年 8月南黄海青岛外海至济州岛断面 ,浮游动物的日垂直变化、断面垂直分布及其与海区水文特征的关系进行了研究。共鉴定浮游动物 40种。断面浮游动物的平均丰度为 13 62 ind/ m 3 ;种类丰富度与暖水种数呈正相关 ,分布趋势为测区东部高、西部低 ;优势种为中华哲水蚤 (Calanus sinicus)、小拟哲水蚤 (Paracalanus parvus)、拟长腹剑水蚤 (Oithona similis)、强壮箭虫 (Sagitta crassa)、近缘大眼剑水蚤 (Corycaeus affinis)。在不同水层 ,浮游动物丰度分布不同 ,随水深增加而减少 ,表层丰度最高 (3 2 2 1ind/ m3) ,温跃层和底层分别为 743 ind/ m3、43 8ind/ m3;种类组成也不相同 ,表层数量最多的种类为鸟喙尖头(Penilia avirostris)、温跃层及其以下水体为中华哲水蚤 ,其它主要种类亦表现不同水层取向。在连续观测站 ,根据浮游动物主要种类的昼夜垂直移动与温跃层关系 ,将之分为表层分布 ,近温跃层分布 ,底层分布及全水层分布 4种类型