Acoustic observation of diel vertical migration and shoaling behaviour in Atlantic redfishes

Acoustic methods were used to study Atlantic redfishes Sebastes spp. vertical migration and shoaling behaviour in Newfoundland waters. Redfishes exhibited consistent patterns of vertical migration in winter, spring and summer, but pelagic shoals were not observed in winter. Pelagic daytime aggregations were generally in close proximity to dense patches of redfishes along the sea floor. Pelagic shoals exhibited high degrees of variability in size, shape and density. Attempts to explain variations in shoal density and area with features of shoal position and structure were unsuccessful. Nearest neighbour distance between fish in shoals had a lower limit near one body length. During the night, fishes were dispersed in the water column and distributions were more homogenous. Diel vertical migration appeared to be a foraging strategy, in which redfishes followed the migration of their euphausiid prey.     

The photobehaviour of Daphnia spp. as a model to explain diel vertical migration in zooplankton

Many pelagic animal species in the marine environment and in lakes migrate to deeper water layers before sunrise and return around sunset. The amplitude of these diel vertical migrations (DVM) varies from several hundreds of metres in the oceans to approx. 5–20 m in lakes. DVM can be studied from a proximate and an ultimate point of view. A proximate analysis is intended to reveal the underlying behavioural mechanism and the factors that cause the daily displacements. The ultimate analysis deals with the adaptive significance of DVM and the driving forces that were responsible for the selection of the traits essential to the behavioural mechanism. The freshwater cladoceran Daphnia is the best studied species and results can be used to model migration behaviour in general. Phototaxis in Daphnia spp., which is defined as a light-oriented swimming towards (positive phototaxis) or away (negative phototaxis) from a light source, is considered the most important mechanism basic to DVM. A distinction has been made between primary phototaxis which occurs when light intensity is constant, and secondary phototaxis which is caused by changes in light intensity. Both types of reaction are superimposed on normal swimming. This swimming of Daphnia spp. consists of alternating upwards and downwards displacements over small distances. An internal oscillator seems to be at the base of these alternations. Primary phototaxis is the result of a dominance of either the upwards or the downwards oscillator phase, and the direction depends on internal and external factors: for example, fish-mediated chemicals or kairomones induce a downwards drift. Adverse environmental factors may produce a persistent primary phototaxis. Rare clones of D. magna have been found that show also persistent positive or negative primary phototaxis and interbreeding of the two types produces intermediate progeny: thus a genetic component seems to be involved. Also secondary phototaxis is superimposed on normal swimming: a continuous increase in light intensity amplifies the downwards oscillator phase and decreases the upwards phase. A threshold must be succeeded which depends on the rate and the duration of the relative change in light intensity. The relation between both is given by the stimulus strength versus stimulus duration curve. An absolute threshold or rheobase exists, defined as the minimum rate of change causing a response if continued for an infinitely long time. DVM in a lake takes place during a period of 1-5-2 h when light changes are higher than the rheobase threshold. Accelerations in the rate of relative increase in light intensity strongly enhance downwards swimming in Daphnia spp. and this enhancement increases with increasing fish kairomone and food concentration. This phenomenon may represent a ‘decision-making mechanism’ to realize the adaptive goal of DVM: at high fish predator densities, thus high kairomone concentrations, and sufficiently high food concentrations, DVM is profitable but not so at low concentrations. Body axis orientation in Daphnia spp. is controlled with regard to light-dark boundaries or contrasts. Under water, contrasts are present at the boundaries of the illuminated circular window which results from the maximum angle of refraction at 48–9° with the normal (Snell's window). Contrasts are fixed by the compound eye and appropriate turning of the body axis orients the daphnid in an upwards or an obliquely downwards direction. A predisposition for a positively or negatively phototactic orientation seems to be the result of a disturbed balance of the two oscillators governing normal swimming. Some investigators have tried to study DVM at a laboratory scale during a 24 h cycle. To imitate nature, properties of a natural water column, such as a large temperature gradient, were compressed into a few cm. With appropriate light intensity changes, vertical distributions looking like DVM were obtained. The results can be explained by phototactic reactions and the artificial nature of the compressed environmental factors but do not compare with DVM in the field. A mechanistic model of DVM based on phototaxis is presented. Both, primary and secondary phototaxis is considered an extension of normal swimming. Using the light intensity changes of dawn and the differential enhancement of kairomones and food concentrations, amplitudes of DVM could be simulated comparable to those in a lake. The most important adaptive significance of DVM is avoidance of visual predators such as juvenile fish. However, in the absence of fish kairomones, small-scale DVMs are often present, which were probably evolved for UV-protection, and are realized by not enhanced phototaxis. In addition, the ‘decision-making mechanism’ was probably evolved as based on the enhanced phototactic reaction to accelerations in the rate of relative changes in light intensity and the presence of fish kairomones.     

The lack of vertical migration in Daphnia: the effect of homogenously distributed food

Neither Daphnia hyalina, Daphnia cristata, nor Daphnia cucullata vertically migrated in Lake Mikoajskie and Lake Majcz. We suggest that even under strong fish predation pressure there is no reason to migrate when seston is relatively homogenously distributed throughout the water column.     

Vertical distribution and diel migration of rotifers in a Parana River floodplain lake

The vertical distribution of zooplankton rotifers in the open waters of Laguna El Tigre was investigated. Rotifers showed a relatively uniform distribution throughout the water column. This pattern of distribution was maintained during the year and did not show variations in relation to hydrologic phases of inundation and isolation of the lake. Diel vertical migration of rotifers from the limnetic and the littoral area was investigated too. In littoral area rotifers exhibited a reverse migration, whereas in the limnetic the movements were less conspicuous. Horizontal migration was observed too, and there were interactions between horizontal and vertical distribution. Predation and competition offer a possible explanation.     

Absence of typical diel vertical migration in Daphnia: varying role of water clarity, food, and dissolved oxygen in Lake Eymir, Turkey

Diel vertical migration (DVM) is a complex and dynamic behaviour against predation because the reaction of migrating organisms to light intensity plays a primary role, but is modified by other factors. In the relatively shallow but thermally stratified Lake Eymir, Daphnia pulex de Geers utilized vertical refugia afforded by the hypolimnion during both day and night. Differences in general vulnerability to fish predation determined the differences in their mean residence depths (MRDs) of different population categories such as most conspicuous and vulnerable individuals of adult with eggs inhabited the deepest depth, whereas juveniles stayed close the thermocline. In late spring, profoundly high amplitude of displacement within the hypolimnion, probably due to the hypolimnion being well-lit and relatively well-oxygenated for the fish and rather unsafe for the large-sized daphnids, was recorded. Therefore, the large-sized daphnids daytime refuge was close to the bottom whereas at night they moved upward to benefit from warmer water temperature along with food availability in the presence of fish predation but still remained below the thermocline. In summer, the insignificant amplitude of the hypolimnetic, which later became epilimnetic, displacements were probably due to the near-anoxic condition found below the thermocline. This might have deterred the fish, thus providing a safer refuge for daphnids in the below thermocline, which afterwards became the above thermocline. Low oxygen availability was regarded as the summer proximate factor. The abundant food and warmer water conditions found in the below/above thermocline also accounted for absence of DVM in summer. Consequently, this study suggests that DVM by Daphnia is an adaptation that is plastic to changing environmental conditions.     

浮游动物昼夜垂直迁移机理的主要假说及其研究进展

对有关浮游动物昼夜垂直迁移(DVM)机理的实验、假说以及理论模型方面的研究进.展进行了综述。昼夜垂直迁移通常指常规迁移(傍晚上升,拂晓下降),其行为过程不仅影响浮游动物的垂直分布,而且也间接地影响其水平分布及生活史特征,对浮游动物在一定水域的种群维持和补充具有重要的意义。与垂直迁移机理有关的假说主要有光驱动假说、逃避捕食者假说、能量和资源利用假说等,其中逃避捕食者假说已得到大量的实验证实。其理论模型研究一般在两个时间尺度上进行,一个是短期的行为机制研究,另一个从长期的生活史策略的角度考虑。随着实验技术的进步和理论及建模工具的发展,这两个时间尺度的研究正逐渐统一到一个框架下进行。     

The effect of different zooplankton grazing patterns resulting from diel vertical migration on phytoplankton growth and composition: a laboratory experiment

Diel vertical migration (DVM) of herbivorous zooplankton is a widespread behavioural phenomenon in freshwater ecosystems. So far only little attention has been paid to the impact of DVM on the phytoplankton community in the epilimnion. Some theoretical models predict that algal population growth in the epilimnion should depend on the herbivores migration and grazing patterns: even if migrating zooplankton consume the same total amount of algae per day in the epilimnion as non-migrating zooplankton, nocturnal grazing should result in enhanced algal growth and favour algal species with high intrinsic growth rates over species with lower intrinsic growth rates. To test these hypotheses we performed experiments in which several algal species were confronted with different feeding regimes of Daphnia. In the experiments algal growth did not only depend on the absolute time of grazing but was comparatively higher when grazing took place only during the night, even when the grazing pressure was the same. Furthermore, algal species with higher intrinsic growth rates had higher advantages when being grazed upon only discontinuously during the night than algal species with a smaller intrinsic growth rate. The grazing pattern itself was an important factor for relative algal performance.     

Diurnal vertical migration of Daphnia hyalina Leydig, 1860 (Crustacea: Cladocera) in Lake Bled (Slovenia) in relation to temperature and predation

Diurnal vertical migration (DVM) of Daphnia hyalina in Lake Bled was most intense during summer stratification. The extent of DVM varied with the size of the animal and its reproductive state. Migration distances were shortest in immature specimens and longest in ovigerous females. During daytime, ovigerous females stayed deeper in the water column than females without ova or immatures. The daytime temperature of water at the median depth of the ovigerous females did not exceed 10 °C, even in the warmest season. At night they migrated upward to an environment which was warmer by as much as 9 °C.Laboratory observations indicate that specimen's size and water temperature determine the velocity of passive sinking, such that morning descent of the different groups of Daphnia can be explained by passive sinking alone.Our hypothesis is that the distribution of different groups of D. hyalina in Lake Bled is influenced by two types of predators: fish (Perca fluviatilis L. and Rutilus rutilus (L.)) and larvae of Chaoborus flavicans (Meig.), the latter appearing in the epilimnion during the night. Fish predation has a key-role at the beginning of thermal stratification. Supposing that in spring the gene pool of Daphnia consists of a mix of different genotypes, distributed at different depths during the day, fish predation combined with a presence of fish chemicals favored genotypes with a lower day-depth during the spring/summer period.     

In situ grazing pressure and diel vertical migration of female Calanus euxinus in the Black Sea

Gut pigment and abundance of the female Calanus euxinus (Hulsemann) weremeasured from several water layers (defined by density values), with3–5 h intervals during 30 h and 21 h at a station in the southwesternBlack Sea in April and in September 1995, respectively. The female C.euxinus was observed to begin migration to the upper phytoplankton-richlayer approximately 3 or 4 hours before the sunset. Only a fraction of thefemale Calanus population (0.2% in April and 3.6% inSeptember) did not migrate but remained at the depth of the oxygen minimumzone during the nighttime. The migrating population was determined to havespent 7.5 h in the euphotic zone in April and 10.5 h in September. Thegrazing rate of female Calanus euxinus was measured from the gut contentdata collected from the layers which contain the euphotic zone. Thepercentage of primary production grazed by the female C. euxinus wascalculated as 14.5% in April and 9.5% in September. This revised version was published online in July 2006 with corrections to the Cover Date.     

Small-scale distribution and diel vertical migration of zooplankton in a shallow lake (Lake Naardermeer,the Netherlands)

Groundwater is a major influence on the hydrological, chemical and thermal regime of chalk streams in the southern U.K. However, little is currently known about the nature of the sediment delivery system within these chalk stream systems, even though sediment-related problems have been increasingly cited as a cause of habitat degradation and of declining salmonid stocks. To address this knowledge gap, suspended sediment fluxes were monitored at 4 sites within the Hampshire Avon catchment between February 1999 and August 2000. Maximum suspended sediment concentrations ranged from nearly 45 mg l^–1 to 260 mg l^–1. Over the study period, annual suspended sediment loads ranged from 644 to 6215 t yr^–1 and annual specific sediment yields ranged from 1.4 to 12.5 t km^–2 yr^–1. The results show that, relative to other U.K. rivers, the study chalk streams are characterised by low suspended sediment concentrations and loads and less episodic behaviour.     

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.     

Seasonal variations in the diel vertical distribution of phytoplankton and zooplankton in a shallow pond

The seasonal succession of phytoplankton diversity, and the variations in the diel vertical distribution of phyto‐ and zooplankton were investigated in a small shallow pond (1.7 m water depth) in 2003. It was inferred that the water tended to stratify weakly in the daytime from February to June. In February and April, the green alga Golenkinia radiata Chodat dominated the phytoplankton assemblage. The cell density of G. radiata greatly decreased in April, when rotifers increased near the bottom. The vertical mixing was attenuated in June, large populations of the euglenoids (Lepocinclis salina Fritsch, Phacus acuminatus Stokes, Trachelomonas hispida (Perty) Stein et Deflandre) developed, and the cyanobacterium Aphanizomenon flos‐aquae var. klebahnii Elenk. appeared at low density. Euglenoids and A. flos‐aquae were mostly distributed in the bottom layer. In late September, when the water was mixed throughout the day, euglenoids and A. flos‐aquae were distributed evenly throughout the water column. The zooplankton (cyclopoid copepods and rotifers) densities in September were the lowest throughout the year. The vertical mixing increased in November, and the phytoplankton community was composed of A. flos‐aquae, P. acuminatus, T. hispida and the green alga Ankistrodesmus falcatus (Corda) Ralfs. In November, at the final stage of water bloom of A. flos‐aquae, its population density decreased with depth. The two euglenoids exhibited similar cell distributions at 0.8 m and 1.6 m during 1–3 November. A. falcatus was distributed evenly throughout the water column; however, when the vertical mixing lessened, the cells at the surface started to sink. Copepod nauplii and rotifers appeared at high densities in November. Seasonal variation in the phytoplankton community structure in the pond seemed to be related to the vertical mixing of the water. In addition, zooplankton, especially rotifers, might play an important role in initiating a spring clear‐water phase and in the bloom collapse of A. flos‐aquae.     

Visibility conditions and diel period affect small-scale spatio-temporal behaviour of pike Esox lucius in the absence of prey and conspecifics

Pike Esox lucius in the absence of prey and conspecifics were shown to have the highest habitat-change activity during dusk and to decrease preference for complex habitats in turbid water. As the behaviours indicate routine responses in the absence of behavioural interactions, E. lucius spatio-temporal distributions should be directly affected and thereby more easily assessed and avoided by prey, with potential consequences for encounter rates.     

Differential effects of undernutrition during pregnancy on the behaviour of does and their kids at parturition and on the establishment of mutual recognition

We investigated whether undernutrition during the second half of pregnancy impaired the behaviour of does and their kids at parturition and early mutual recognition. Twenty-two control and 22 underfed mixed-breed, multiparous dairy goats were used, together with their respective kids (control, n = 31: nine singles, 16 twins and six triplets; underfed, n = 32: 11 singles, 18 twins and three triplets). Undernutrition involved limiting protein and energy intake at 70% of the nutritional requirements for maintenance and foetal growth from day 70 of pregnancy until birth. The behaviour of mothers and their two first-born kids was observed for 90 min from the birth of each kid. Maternal olfactory recognition of the kid was assessed at 4 h post partum by testing selective nursing behaviour. Non-olfactory recognition was assessed at 8 h in a two-choice test excluding olfactory cues. In kids, preference for the mother was assessed in a two-choice test at either 12 or 24 h post partum. Bodyweight of does and kids were lower in the underfed group up to 2 weeks post partum. At parturition, licking, maternal bleating frequency and latency to nursing did not differ between nutritional groups. Control kids were faster than underfed kids to stand, search for and reach the udder, but underfed kids bleated more and tended to spend more time at the udder. Both control and underfed does accepted their own kid and rejected the alien in the selectivity test at 4 h. In contrast, at 8 h post partum, only control goats showed a significant preference for their own kid in the non-olfactory recognition test. Both control and underfed kids showed a preference for their own mother at 12 and 24 h and undernutrition during pregnancy had little influence on the performance of kids. However, 12 h-old underfed kids tended to be less active than control kids and visited their own mothers less than control kids. There were no significant correlations between the behaviour of the mother or of the kid at parturition and their performance in the discrimination tests. Overall, undernutrition in the second half of pregnancy appears to be more detrimental for the behaviour of the mother than for the kid. Furthermore, it has more impact on the establishment of maternal non-olfactory recognition than on maternal care at parturition or the establishment of maternal selectivity.     

Plasma corticosterone concentrations in males of the skink Egernia whitii during acute and chronic confinement, and over a diel period

This study examined adrenal and gonadal responses to capture and to longer term captivity in males of the scincid lizard, Egernia whitii. Animals subjected to acute capture stress in the field exhibited a rapid rise in plasma corticosterone concentrations that had not attenuated by 4 h after capture; plasma testosterone concentrations decreased significantly over this period. Animals returned to captive conditions in the laboratory showed little change in plasma corticosterone concentrations by 1 week, but concentrations were significantly lower at 4 weeks. Plasma testosterone did not vary significantly during this period. Sampling over a diel period indicated that plasma corticosterone concentrations vary little over the daylight hours; however, there was a significant decrease between the samples taken at 17:00 h and 21:00 h. These results suggest that, as in other species, the acute adrenal response to capture stress may confound assessment of other physiological parameters, although if experiments are carried out during daylight hours, time of sampling should have little influence on plasma corticosterone concentrations. The results also suggest that males of E. whitii require at least a week to adapt to captivity, although further studies investigating different captive conditions are warranted for this social species.     

Synchronous division and the pattern of diel vertical migration of Heterosigma akashiwo (Hada) Hada (Raphidophyceae) in a laboratory culture tank

Heterosigma akashiwo (Hada) Hada (Raphidophyceae) causes red tides in Osaka Bay (Japan). A clonal culture of the alga was grown in a 2 m tall culture tank on a 12: 12 LD cycle to determine patterns of vertical migration and cell division. A specific growth rate of 0.43 ln unit · day⁻¹ was obtained during complete mixing conditions. Under weakly stratified conditions (≈ΔT = 3–4°C/1.5 m), H. akashiwo in the tank grew and showed a similar pattern of vertical migration to that observed in the field for at least 6 days. Cell concentration, mean cell volume, and photosynthetic capacity, estimated by DCMU-induced fluorescence increase of H. akashiwo, were monitored in the stratified tank at 2-h intervals over 24 h at three levels in the water column. Cell ascent began shortly before the light period and vertically swimming cells were smaller in size than those sampled near the bottom of the tank. The cell division cycle and the pattern of vertical migration were phased individually by the light regime and were well synchronized with each other. This synchrony must be due to the interrelation between these two processes or the existence of a clock which controlled endogenous rhythms of both processes and was entrained by a light: dark cycle. The relative increase of fluorescence with DCMU was higher for migrating cells than for non-migrating cells.     

Changes in landscape composition influence the decline of a threatened woodland caribou population

1. Large-scale habitat loss is frequently identified with loss of biodiversity, but examples of the direct effect of habitat alterations on changes in vital rates remain rare. Quantifying and understanding the relationship between habitat composition and changes in vital rates, however, is essential for the development of effective conservation strategies. 2. It has been suggested that the decline of woodland caribou Rangifer tarandus caribou populations in North America is precipitated by timber harvesting that creates landscapes of early seral forests. Such habitat changes have altered the predator-prey system resulting in asymmetric predation, where predators are maintained by alternative prey (i.e. apparent competition). However, a direct link between habitat condition and caribou population declines has not been documented. 3. We estimated survival probabilities for the threatened arboreal lichen-feeding ecotype of woodland caribou in British Columbia, Canada, at two different spatial scales. At the broader scale, observed variation in adult female survival rates among 10 distinct populations (range = 0.67-0.93) was best explained by variation in the amount of early seral stands within population ranges and population density. At the finer scale, home ranges of caribou killed by predators had lower proportions of old forest and more mid-aged forest as compared with multi-annual home ranges where caribou were alive. 4. These results are consistent with predictions from the apparent competition hypothesis and quantify direct fitness consequences for caribou following habitat alterations. We conclude that apparent competition can cause rapid population declines and even extinction where changes in species composition occur following large scale habitat change.     

Reproductive potential of Daphnia galeata in relation to food conditions: implications of a changing size-structure of the population

A field study was conducted in the highly eutrophic, biomanipulated Bautzen Reservoir to investigate the influences of food conditions on fecundity of Daphnia galeata during spring and early summer in four consecutive years. By defining a relation between body size and maximum brood size, the realised reproductive potential (RRP) of Daphnia was expressed as percentage of actual egg numbers of the population in relation to calculated maximum egg numbers. This parameter is independent of the size structure of the population. It was found to correlate generally with mean clutch size. However, deviations occurred if size at maturity (SAM) declined. Extremely low SAM values (0.8 mm) were recorded in 2 years, corresponding with increasing food concentrations. Low clutch sizes in these situations can be explained by small body sizes of ovigerous females, whereas RRP increased, thus reflecting good food conditions. From the comparison of different reproductive parameters, it is concluded that RRP is the most reliable one with regard to bottom-up influences in case of extremely variable size structure and possible predation impacts. Fecundity of D. galeata was clearly food limited during the clear-water stage. Relatively high POC-threshold concentrations of 0.26 mg C l^–1 suggest that food quality was low and may be equally important as food quantity in limiting egg production.     

Changes in the timing of spring and autumn migration in North American migrant passerines during a period of global warming

Butler (2003) used first arrival dates (FADs) of 103 migrant birds in northeastern USA and found that both long-distance migrants (LDMs; wintering south of the USA) and short-distance migrants (SDMs; wintering in the southern USA) arrived earlier in the second half of the 20th century than they had in the first, consistent with scenarios of global warming; the trend was stronger in SDMs. Using FADs to characterize migration systems can be problematic because they are data from one tail of a distribution, they comprise a mostly male population and they may not correlate well with the balance of the migration period. FADs also provide no information about autumn migration. This paper uses a banding dataset from Long Point Bird Observatory, Ontario, for 14 passerines for a period of global warming (1975–2000), taking these issues into account. The data were filtered to minimize effects of unequal netting effort (147 491 resulting records), and the passage dates then calculated in each season of each year for the 1st, 2nd and 3rd quartiles for regression analysis. Only two of 13 species analysed in the spring showed significantly earlier passage times, although the overall trend was towards earlier spring migration, especially among SDMs. Autumn responses were more prevalent, however, and in some cases more dramatic with six of 13 species showing delayed migration (four SDMs, two LDMs). Two LDMs exhibited earlier autumn migration. Where earlier spring migration occurred, both sexes appeared to contribute to the change. Where delayed migration occurred in autumn, both sexes and both adults and hatch-year birds appeared to contribute in at least some cases. The spring FAD results are consistent with those of Butler, but when the whole migration is considered, change is far from universal in spring and is in fact more substantial and complex in autumn.