Behavioral responses of larvae of the crab Rhithropanopeus harrisii (Brachyura: Xanthidae) during diel vertical migration

Larval Rhithropanopeus harrisii (Gould) show nocturnal vertical migration. Larval behavioral responses to different rates of increase and decrease in light intensity were measured in an apparatus with a natural angular light distribution. A central objective was to establish whether phototaxis actually participates in vertical migration. At sunset the level of light adaptation controlled the readiness of the larvae to migrate, while an ascent was initiated by a preductable relative decrease in intensity (e.g. 4.0x10^-3s^-1). Rates of relative decrease around sunset would evoke continuous upward swimming. Gravity was the orienting cue and there was no change in swimming speed during the ascent. At sunrise, the larval descent was initiated by exposure to an absolute light intensity of about 0.23 log unit above the lower visual threshold. Light served as an orienting cue, as larvae descended by a negative phototaxis. Thus, phototaxis is not a laboratory artifact and does participate in vertical migration. A consideration of behavioral responses of other crustacean zooplankton indicates there is considerable variation in the initiating and directing cues for vertical movements. The variety of behavioral responses of R. harrisii suggests that a synthesis of hypotheses about migration may provide the proper basis for explaining the mechanisms underlying diel vertical migration.     

Comparative study of crustacean larval photoresponses

Ovigerous females of four brachyuran (Cancer gracilis, Lophopanopeus bellus bellus, Hemigrapsus oregonensis and Scyra acutiforns) and two anomuran (Pagurus beringanus and P. granosimanus) species, which live as adults in coastal areas, were collected near Friday Harbor, Washington, USA, in 1985, and spectral sensitivity and phototactic pattern of their larvae were measured. Responses were compared with previous measurements on estuarine species to determine whether responsiveness varies with adult habitat. Estuarine and coastal species have similar photoresponses. Spectral sensitivity of the test brachyran species had two maxima, one near 400 to 420 nm and another around 500 nm. The anomuran species were similar, but had an additional peak in the region of 580 to 620 nm. This sensitivity is adapted to daytime light conditions in the adult environment, and available spectra at the time of larval movement during diel vertical migration. Upon light adaptation and stimulation with a narrow light field, all six species showed positive phototaxis to high light intensities and a pronounced negative response to low intensities. Only the negative response will occur in natural underwater light conditions, and it is part of a predator-avoidance shadow-response which operates in areas of higher light intensity. The same phototactic pattern is observed in all species when darkadapted except H. oregonensis. If nocturnal vertical migration occurs, this negative response may be responsible for the descent at sunrise and depth maintenance during the day.     

Photosensitivity of the calanoid copepod Acartia tonsa

The light intensity and spectral sensitivities of the calanoid copepod Acartia tonsa Dana were determined by measuring phototactic responses. Adult females displayed only positive phototaxis. The dark-adapted copepod, which possesses a single naupliar eye, perceived light at intensities as low as 2.8x10¹¹ photons m^-2 s^-1. The action spectrum for positive phototaxis had no clear maxima but rather showed a broad range of greatest sensitivity from 453 to 620 nm. This sensitivity encompassed those wavelengths that are maximally available at the depth where the copepod is found during the day. This spectral overlap, coupled with the finding that the copepod requires light cues for nocturnal vertical miration, suggests that broad spectral sensitivity is an adaptive mechanism to maximize light intensity sensitivity during migration.     

Effects of brackish water incursions and diel phasing of tides on vertical excursions of the keystone predator Pisaster ochraceus

The physical factors that constrain the vertical foraging excursions of the keystone predator, the sea star Pisaster ochraceus, hold considerable interest because they indirectly shape the vivid patterns of zonation of rocky shore communities by impeding or enhancing the ability of P. ochraceus to traverse the intertidal zone. In this paper, we describe a study conducted in the Pacific Northwest of North America in which we examined, in the field and laboratory, the abiotic factors that can affect vertical excursions by P. ochraceus. Our field observations revealed that the extreme upward reach and average shore level height reached by P. ochraceus were significantly lower for daylight high tides than nocturnal high tides. Based on diver observations following a severe storm, it would also appear that these diurnal movements can be impeded by freshwater incursions into the intertidal zone; a regularly occurring event in the Pacific Northwest. As part of an experimental investigation into this phenomenon, we observed that sea stars maintained in tall cylindrical aquaria, without tidal flux, remained near the bottom during daylight and moved to the top of the column at night, suggesting that photoperiod alone can influence the cycle of vertical movement. Adding a freshwater layer to the aquaria restricted these vertical excursions. Our results suggest that on rocky coastlines susceptible to fresh water incursions, the suppression of foraging may be an important factor in the spatial and temporal variation in the intensity of predation. Furthermore, given the relative increase in frequency and intensity of freshwater incursions in the Pacific Northwest and the intolerance of P. ochraceus to lowered salinity, there is the long-term potential to significantly alter patterns of species zonation in this essential marine habitat.     

Chlorophyll fluorescence as a proxy for microphytobenthic biomass: alternatives to the current methodology

Pulse amplitude modulated (PAM) fluorescence has been used as a proxy of microphytobenthic biomass after a dark adaptation period of 15 min to stabilise the minimum fluorescence yield (F _o ¹⁵). This methodology was investigated for in situ migratory and ex situ engineered non-migratory biofilms, comparing dark adaptation to low (5% ambient) and far-red light treatments over different emersion periods. Far-red and low light reduced potential errors resulting from light history effects, by reversal of non-photochemical quenching after 5 min of treatment, compared to over 10 min required by conventional dark adaptation. An in situ decline of minimum fluorescence yield over 15 min was observed during the dark adaptation for migratory biofilms, but was not observed in the non-migratory biofilms indicating that the major cause of decline was downward vertical migration of cells into the sediment. This pattern occurred in far-red light after 10 min, but not for the low light treatment, indicating that low light maintained the biomass at the surface of the sediment. It is therefore concluded that low light treatment is a better option than conventional dark adaptation for the measurement of minimum fluorescence as a proxy of microphytobenthic biomass.     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in June 1989, when the water column was thermally weakly stratified. Because of fewer eggs and less variation in their abundance, nocturnal spawning was not apparent. A pronounced upward migration occurred in NIII. NIII to CIII resided in the upper 20 m layer throughout the day, and from CIV on their median depths descended. CV and adult females underwent significant diel vertical migration, whereas adult males did not migrate. By integrating the results from the present study and those from our previous investigations (in August–September 1988, November 1988 and March 1989), we review seasonal variation in the ontogenetic diel vertical migration of C. sinicus. Spawning was largely nocturnal, reaching its maximum level around dawn, but spawning depth and fecundity changed seasonally. The distribution of pre-feeding stages, NI and NII, was similar to that of eggs. A pronounced upward migration always occurred in the first feeding stage, NIII, and late nauplii and early copepodites always resided in the food-rich upper layer, indicating that upward migration by NIII is feeding migration. As the stages progressed, they extended their vertical distribution range, and CV and adult females usually underwent diel vertical migration. However, the pattern and strength of this migration differed seasonally. Their day depths increased with the increase of relative biomass of planktivorous fish, indicating that predator avoidance induces their diurnal downward migration. High chlorophyll a concentrations in the upper layer (<15 m deep) relative to the lower layer (>20 m deep) amplified their diel vertical migrations. Diel vertical migration of C. sinicus is a phenotypic behavior.     

Effect of microclimatic factors and tide on vertical migrations of the mangrove crab Sesarma leptosoma (Decapoda: Grapsidae)

The tree crab Sesarma leptosoma Hilgendorf migrates up the mangrove trunks twice a day, both in the morning and in the evening, leaving its refuge near the tree base to feed on fresh leaves. In spite of the 150 cm of tide excursion at the base of the trees, the timing of the migrations seems largely controlled by nontidal factors. The timing of the morning upward migration is controlled by the light level alone. The morning downward migration takes place as soon as the temperature rises and relative humidity falls, with no relationship to tidal cycle or light level. Only the evening upward flow seems primarily controlled by the tide, while the evening downward flow, which ends at twilight, seems to be related to the timing of the evening upward migration, to the tide and again to light levels. During the day, crabs abandon the canopy to avoid low relative humidity, usually coupled with the higher wind speeds, which would quickly dehydrate them. At night, crabs stop migrating, probably because their movements and orientation capability along the trunk and in the canopy are largely vision dependent. Nontidal factors have never previously been shown to influence the migratory behaviour of an intertidal crustacean species so extensively. Received: 14 June 1997 / Accepted: 17 July 1997     

Vertical distribution and nocturnal migration of zooplankton in relation to the development of the seasonal thermocline in Patraikos Gulf

Vertical distribution and nocturnal migration of zooplankton species in relation to the development of the seasonal thermocline in the shallow waters (90 m) of Patraikos Gulf (Ionian Sea, Greece) were investigated using a WP-2 closing net. Juvenile and adult copepods accounted for a mean of 91% of the total collected in three sampling periods, i.e. May, July and September 1985.Ctenocalananus vanus, Paracalanus parvus andOithona plumifera were the dominant copepods. The majority of the zooplankton tend to aggregate at the thermocline layer. Among copepods the two congeneric speciesClausocalanus pergens andC. furcatus exhibited different migratory responses to the development of the thermocline.C. pergens occurred in the lower part of the thermocline andC. furcatus in the upper region or above. The diel vertical migration of all species could be divided into four types: (1) no vertical migration; (2) upward migration at night; (3) occasional migration; and (4) reverse migration (down at night). In July when the strongest thermocline developed, most zooplankters rose close to the surface at night. For most species, temperature discontinuity did not limit their diel migration.Please address all correspondence and requests for reprints to Dr J.J. Lykakis     

Nocturnal movements of phototactic zooplankton in shallow waters

Downwardly directed light-traps were used by night, over seagrass and coral sand to investigate the existence and significance of nocturnal peaks in phototactic zooplankton. Samples each half-hour from sunset to sunrise showed that small invertebrates, mostly copepods, were trapped in large numbers once or twice a night. Each period of high catches lasted only 1 to 2 h and was often associated with periods of failing natural light (evening twilight, moonset). This pattern and its variations are explained in terms of nocturnal vertical migration, in which plankton move towards light in response to decreasing light intensity, move away from light in response to increasing light intensity, and disperse at constant light intensity. Some morning peaks are unexplained.     

Critical thermal maxima and upper lethal temperatures for the calanoid copepods Acartia tonsa and A. clausi

Determination of the critical thermal maxima and upper lethal temperatures for Acartia tonsa Dana and A. clausi Giesbrecht revealed the relative temperature tolerance of the two species. A. tonsa exhibited a wider range of acclimation temperatures compared to A. clausi. The response of A. tonsa is related to its ubiquity in Atlantic Ocean estuaries (temperate and tropical), and its capacity to live in temperate waters throughout most of the year. Differences between upper lethal temperatures and critical thermal maxima were very narrow for A. tonsa from tropical and subtropical areas. This suggests that, in warm climates, this copepod lives close to its upper lethal temperature limits.Based in part on a dissertation submitted in partial fulfillment for the Ph.D. degree at the University of Rhode Island.     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. II. Proximate role of exogenous light cues and endogenous rhythms

The marine copepod Calanopia americana Dahl undergoes twilight diel vertical migration (DVM) in the Newport River estuary, North Carolina, USA, in synchrony with the light:dark cycle. Copepods ascend to the surface at sunset, descend to the bottom around midnight, and make a second ascent and descent before sunrise. Behavioral assays with C. americana in the laboratory during fall 2002/2003 and summer 2004 investigated aspects of three hypotheses for the proximate role of light in DVM: (1) preferendum hypothesis (absolute irradiance), (2) rate of change hypothesis (relative rates of irradiance change), and (3) endogenous rhythm hypothesis. Results suggest that C. americana responds to exogenous light cues consistent with its DVM pattern; changes in absolute irradiance evoked swimming responses that would result in an ascent at sunset and descent at sunrise, while relative rates of irradiance decrease at sunset (–0.0046 s^–1) evoked an ascent response, and relative rates of irradiance increase at sunrise (0.0042 s^–1) evoked a descent response. Furthermore, C. americana expressed an endogenous rhythm in vertical migration that was positively correlated with field observations of twilight DVM. Collectively, these results indicate that both exogenous light cues and endogenous rhythms play a proximate role in twilight DVM of C. americana, providing redundancy in the causes of its vertical migration.Communicated by J.P. Grassle, New Brunswick     

Calibration of the sun compass by sunset polarized light patterns in a migratory bird

Summary Migrating birds derive compass information from the sun, stars, geomagnetic field and polarized light, but relatively little is known about how these multiple sources of directional information are integrated into a functional orientation system. We found that migratory warblers exposed to a rotated polarized light pattern at sunset oriented at a constant angle to the axis of polarization. When polarized light cues were eliminated, this shifted orientation was maintained relative to the setting sun. Polarized light patterns, thus, appear to provide a calibration reference for the sun compass in nocturnal migrants, and may also play a role in calibrating other compass systems. Correspondence to: J.B. Phillips     

Light quality and concentration of proteins,RNA, DNA and photosynthetic pigments in two species of marine plankton algae

Chlorophyll, protein, DNA and RNA concentrations in cultures ofCyclotella nana (Hustedt) andDunaliella tertiolecta (Butcher) were higher in blue light and lower in green light than in white light of the same intensity. Total carotenoid concentrations were highest in green light. Total pigment concentrations were highest in cells grown in blue light, lowest in those from green light. The relative changes in pigment concentrations associated with differences in spectral composition of the radiation are comparable to those often observed in phytoplankton from various depths in stratified natural water. Light adaptation in algae as a response to light quality is suggested. The chemical composition of plankton algae can be expected to vary considerably with depth.     

Dispersal potential of invasive algae: the determinants of buoyancy in <Emphasis Type="Italic">Codium fragile</Emphasis> ssp. <Emphasis Type="Italic">fragile</Emphasis>

The capacity for long-distance dispersal is an important factor in determining the spread of invasive species. For algae, positive buoyancy generally is correlated with increased dispersal potential, and the light environment has been previously identified as a possible determinant of buoyancy in several species. We examined the effect of light intensity on the buoyancy of fragments of the invasive green alga Codium fragile ssp. fragile. Under natural and controlled conditions, the buoyancy of samples taken from the thallus tip was higher than those from near the holdfast. Both laboratory and field experiments also showed that buoyancy was dynamic and switched from positive to negative under reduced light intensity, but this change required several days. We also observed seasonal changes in buoyancy, presumably due to natural variations in light intensity, with the buoyancy of fragments washed up on the shore highest in mid-summer. These results show that buoyancy is a dynamic property of the C. fragile ssp. fragile thallus and suggest that buoyant fragments contribute to long-range dispersal and accelerated regional spread of this invader. This finding suggests that dispersal is more likely during conditions of high light intensity and illustrates the need to understand how variations in the natural environment can affect the dispersal potential of invasive species.     

Increased sedentariness in European Blackbirds following urbanization: a consequence of local adaptation?

Urbanization changes local environmental conditions and may lead to altered selection regimes for life history traits of organisms thriving in cities. Previous studies have reported changes in breeding phenology and even trends toward increased sedentariness in migratory bird species colonizing urban areas. However, does the change in migratory propensity simply represent a phenotypic adjustment to local urban environment, or is it genetically based and hence the result of local adaptation? To test this, we hand-raised European Blackbirds (Turdus merula) from urban and forest populations, quantified their nocturnal activity and fat deposition covering two complete migratory cycles and examined the consequences of a reduced migratory propensity for the timing of gonadal development (a physiological measure of the seasonal timing of reproduction). Although nocturnal activities differed strikingly between fall and spring seasons, with low activities during the fall and high activities during the spring seasons, our data confirm, even in birds kept from early life under common-garden conditions, a change toward reduced migratoriness in urban blackbirds. The first score of a principal component analysis including amount of nocturnal activity and fat deposition, defined as migratory disposition, was lower in urban than in forest males particularly during their first year, whereas females did not differ. The results suggest that the intrinsic but male-biased difference is genetically determined, although early developmental effects cannot be excluded. Moreover, individuals with low migratory disposition developed their gonads earlier, resulting in longer reproductive seasons. Since urban conditions facilitate earlier breeding, intrinsic shifts to sedentariness thus seem to be adaptive in urban habitats. These results corroborate the idea that urbanization has evolutionary consequences for life history traits such as migratory behavior.     

Effects of nitrate on the diurnal vertical migration,carbon to nitrogen ratio,and the photosynthetic capacity of the dinoflagellate Gymnodinium splendens

A non-thecate dinoflagellate, Gymnodinium splendens, was studied in a 12 d laboratory experiment in 2.0x0.25 m containers in which light, temperature, and nutrients could be manipulated. Under a 12 h light: 12 h dark cycle, the dinoflagellates exhibited diurnal vertical migrations, swimming downward before the dark period began and upward before the end of the dark period. This vertical migration probably involved geotaxis and a diel rhythm, as well as light-mediated behavior. The vertical distribution of nitrate affected the behavior and physiology of the dinoflagellate. When nitrate was present throughout the container, the organisms resembled those in exponential batch culture both in C:N ratios and photosynthetic capacity (P_max); moreover, they migrated to the surface during the day. In contrast, when nitrate was depleted, C:N ratios increased, P_max decreased, and the organisms formed a subsurface layer at a depth corresponding to the light level at which photosynthesis saturated. When nitrate was present only at the bottom of the tank, C:N ratios of the population decreased until similar to those of nutrient-saturated cells and P_max increased; however, the dinoflagellates behaved the same as nutrient-depleted cells, forming a subsurface layer during the light period. Field measurements revealed a migratory subsurface chlorophyll maximum layer dominated by G. splendens. It was just above the nitracline during the day, and in the nitracline during the night, which concurs with our laboratory observations.     

Effects of parental acclimation temperature and egg-incubation temperature on egg-hatching time in Acartia tonsa (Copepoda: Calanoida)

Experimental results indicate that both parental acclimation temperature and egg-incubation temperature have significant effects on the egg-hatching time of Acartia tonsa Dana between 15° and 25°C. Effects of parental acclimation temperature and egg-incubation temperature are additive if the long-term parental acclimation temperature is constant. A. tonsa are sensitive to temperature and changes of temperature during development (in vivo) and after they are laid (in vitro), up to the time of hatching. A temperature change of the parent culture for time periods of 86 h to 8 d is sufficient to change egg-hatching times. The time required for a temperature change to affect egg-hatching times depends on the magnitude and direction of the temperature change. Field collected A. tonsa demonstrate that changes in egg-hatching times occur in the field populations and can be predicted by mean weekly water temperature.     

Moon orientation in sandhoppers: effects of lighting treatments on the persistence of orientation ability

The moon orientation rhythm persists in sandhoppers removed from environmental entrainments relative to moon periodicity. In order to investigate the underlying mechanism, the crustacean amphipods Talorchestia capensis and Talitrus saltator were collected at new moon and exposed in the laboratory to an artificial light during the day, but to a variety of lighting treatments during the night: (1) continuous dark, (2) dim light in phase with the natural moon, (3) continuous dim light. The animals were tested under the moon 9–21 days later by using a new type of arena. The ability to orientate in a direction perpendicular to the shore persisted in sandhoppers under treatments 1 and 2, but not under treatment 3. A disturbance due to a phototactic tendency in Talorchestia was also observed in animals captured on the day of the test when exposed to unnatural lighting, but not when they were kept under natural light. The present findings show that the timing mechanism allowing compensation for changes in the moon’s position also persists in animals that have been long removed from entraining factors. The dependence of this orientation ability on nocturnal lighting and the disturbing effect of sudden changes in light intensity support the idea that the phasal lighting of the moon, and perhaps sunrise and sunset act as resetting factors for the moon’s orientation rhythm.     

Effect of light intensity on activity and food-searching of larval herring,Clupea harengus: a laboratory study

Larvae of Clupea harengus were reared from spawning herring caught in March 1982 and 1983 in the Firth of Clyde, Scotland. An infra0red observation technique was used to record the behaviour of larval herring both in shallow dishes using a top view and in a tank 2 m deep using a side view. The amount of time larvae spent swimming, which was minimum in complete darkness, increased with increasing light intensity and as the larvae grew. Maximum swimming speeds of feeding larvae were recorded at light intensities between 10 and 100 lux. The presence of food organisms (Artemia sp., Brazilian strain) at light intensities below the feeding threshold (0.1 lux) caused an increase in the proportion of time spent active, but light intensities above the threshold had different effects, depending on developmental stage: larvae of 12 mm increased swimming speed, but 21 mm larvae decreased speed. In the 2 m deep tank in darkness, larvae displayed inactive periods wherein they sank head first, interspersed with periods of upward swimming. As light intensity increased, vertical swimming was replaced by horizontal swimming. These results are discussed with reference to food searching and vertical migration of larval herring in the sea.     

Nocturnal feeding migrations of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) on the rocky shores at Mkomani,Mombasa, Kenya

The vertical zonation of the three common rocky shore neritids at Mkomani, Mombasa, Kenya, Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, as a function of feeding migrations and of size, was studied from 28 February to 24 March 1983. These snails perform feeding migrations at night starting at around mid-ebb tide and return to their resting positions with the flood tide. They remain in their resting positions throughout the day until the next nocturnal ebb tide. The direction of migration is sizerelated, with the larger snails of each species moving in the opposite vertical direction to the smaller ones, so that the populations as a whole exhibit no statistically significant net vertical displacement. The larger individuals of two of the species, N. plicata and N. undata, invariably move downwards to their feeding levels, while the smaller individuals move upwards; the larger individuals of N. textilis display a different pattern of migration, moving downwards on and around spring-tide days and upwards on and around neap tide days, while the smaller individuals move in the opposite directions. N. textilis rest above their feeding level around spring tides, and below that level around neap tides. It is demonstrated how these nocturnal migratory feeding rhythms are integrated into the spring-neap and seasonal cycles of the snails' daytime resting positions. The adaptive significance of these migrations is also discussed.