The distribution of primary production relative to a cyclonic gyre in Baie des Chaleurs

Primary production within the Baie des Chaleurs, Canada, is dominated by a cyclonic gyre system. The pattern of production fits the theoretical expectation of increased productivity associated with colder, nutrient-rich water being brought closer to the surface within the gyre. Upper reaches of the bay, west of the gyre, follow a normal esturine form of circulation with lower rates of production. East of the gyre, in the Gulf of St. Lawrence, primary production is less.Travaux sur les Pêcheries du Québec, no. 30, Direction des Pêcheries, Ministère de l'Industrie et du Commerce du Québec.     

Abundance,spatial distribution,and size structure of populations of Oreaster reticulatus (Echinodermata: Asteroidea) on sand bottoms

Three populations of Oreaster reticulatus (Linnaeus, 1758) inhabiting shallow-water (<4 m) seagrass habitats in the Grenadines (West Indies, Caribbean Sea) were associated predominantly with beds of Halodule wrightii. Occupation of fringing inshore areas of bare sand was inversely related to wave action; even where sandy patches occurred offshore, the preferred substratum was H. wrightii. The association of O. reticulatus with H. wrightii is related to the asteroid's microphagous feeding habit and the availability of food resources associated with the seagrass. O. reticulatus rarely occurred on dense beds of Thalassia testudinum, but was moderately abundant in areas of sparse cover. Differences in the occurrence of O. reticulatus among seagrass types may be related to factors afdecting foraging effort, such as the tractability of the substratum and mobility upon it. Populations of O. reticulatus exhibited an aggregated dispersion within beds of H. wrightii, possibly attributable to local substratum heterogeneity and/or reproductive behavior. Increased turbulence induced migration to deeper water and markedly increased aggregation along offshore boundaries. The populations were primarily adults, with some late juvenile stages. The paucity of juveniles and their cryptic behavior and coloration suggest that settlement and early postmetamorphic development occurs in alternate habitats, such as dense beds of T. testudinum. Interpopulation differences in size structure may be associated with differences in the quality and availability of food sources.     

Food-particle size and selection by bivalve larvae in a temperate embayment

Epifluorescence microscopy was used to analyze the stomach contents of bivalve larvae collected in the Baie des Chaleurs (western Gulf of St. Lawrence, Canada) in order to document food-particle sizes, compare feeding among taxa, and compare the diet with the in situ phytoplankton community. Stomach contents were mainly composed of small autotrophic flagellates (<5 μm) and cyanobacteria (<2 μm), reflecting the microbial food web which characterizes these waters. More than half (55%) of all veligers examined contained algal cells of 5 to 15 μm, whereas only 3% had cells of 15 to 25 μm. Differences in the size ranges of ingested algal cells among similar-sized larvae of different species suggests that veligers actively selected food particles. Among the smallest veligers (185 to 260 μm), scallops (Placopecten magellicanus) and mussels (Mytilus edulis) ingested more <5 μm and 5 to 15 μm algae than clams (Mya arenaria). Among larger veligers (261 to 405 μm), clams contained significantly more <5 μm cells than mussels, whereas mussels contained significantly more 5 to 15 μm algae than clams. Algal cells of 15 to 25 μm were preferentially ingested by mussel veligers. Feeding also differed between different-sized veligers within taxa, i.e. the smallest clam veligers ingested fewer of 5 to 15 μm algae than the larger size classes. Mussel veligers ingested significantly more 15 to 25 μm and fewer <5 μm cells as their size increased. The dominance of ultraplankton in the nearshore waters of Baie des Chaleurs and in the stomach contents suggests that veliger larvae may be an important export path for carbon produced by small phytoplankton. Received: 17 July 1996 / Accepted: 20 September 1996     

Relation of spawning of the giant scallop,Placopecten magellanicus,to temperature fluctuations during downwelling events

We examined the relation of spawning to biological and physical factors for the scallop Placopecten magellanicus (Gmelin, 1791) over 8 yr (1984 to 1991) in the Baie des Chaleurs, southwestern Gulf of St. Lawrence, Canada. Spawning was always abrupt and occurred between July and mid-September. It did not appear to be related to the abundance of phytoplankton or particulate organic carbon and nitrogen in the water. It further showed no relationship to lunar or tidal phascs or to current velocity. Spawning consistently occurred during the summer temperature maximum, but did not coincide with any critical temperature or cumulative temperature threshold. All but one of the 33 spawning events, for which temperature data were recorded, were associated with temperature changes; 25 of these were sharp temperature increases and 7 were during strong temperature fluctuations when the mean temperature was 9 to 14°C. Both types of temperature changes were caused by downwelling of warm surface water. The delay by about 1 d in time of spawning between sites coincided with the rate at which downwelling events propagated into the bay. Virtually all of the spawning events resulted in gametes being ejected into warm water masses where conditions are likely to favour larval development.     

Sexual reproduction of the scyphomedusa Aurelia aurita in relation to temperature and variable food supply

The effects of food availability and temperature on sexual maturation and female reproductive output of the scyphomedusa Aurelia aurita was examined in two populations from the contrasting environments of Southampton Water and Horsea Lake, England. Trends in oogenesis and subsequent reproductive output differed markedly between the two populations. In Southampton Water, the onset of sexual maturation occurred earliest in the larger medusae, but eventually all females became ripe, the smallest being 45 mm bell diameter (BD). The decrease in minimum size at maturity was correlated with increasing temperature. In A. aurita from Horsea Lake, size at maturity varied on a seasonal basis, with the smallest ripe female being only 19 to 20 mm BD. There were spring and autumn periods of sexual maturation in this population. During the autumn period, it is likely that food limitation was playing a more critical role in determining medusa size, with decreasing temperature indirectly affecting A. aurita by limiting primary and secondary production. In similar-sized ripe medusae, fecundity was greater in Southampton Water, but the planula larvae produced were significantly smaller than those in Horsea Lake. It is suggested that in Horsea Lake, the quality of the larvae are greater in terms of biochemical content to ensure survival of the few gametes produced (i.e. K-strategy). Comparison of the reproductive effort of the two A. aurita populations revealed that medusae from Southampton Water, which experience greater food availability, are able to direct more energy to reproduction than Horsea Lake medusae. In the latter, A. aurita medusae appear to partition the available food resources into either somatic growth (and therefore increased future fecundity) when food is abundant, or reproductive growth when food is scarce. Received: 24 June 1997 / Accepted: 23 March 1998     

Experimental examination of movement of the giant scallop,Placopecten magellanicus

We quantified movements of tagged giant scallops, Placopecten magellanicus (Gmelin, 1791), (40 to 115 mm in shell height) released in 1991 and 1992 at nine stations in Port Daniel Bay, Baie des Chaleurs, Gulf of St. Lawrence, Canada, for periods of 10 to 52 d. Both the mean distances moved per day and dispersion distances from release points were usually greater at the two stations with sand substratum, low scallop densities and high rock crab, Cancer irroratus, densities. Movement rate and dispersion of scallops from the release points at the other seven stations were similar, even though there were marked differences in substratum type (gravel, cobble or bedrock), predator density, and scallop density. Most mean dispersal directions were random, and scallops did not appear to migrate from unsuitable to suitable habitats. Although movement did reduce predation rate, scallop movement was weakly correlated with the abundance of only one predator, C. irroratus.     

Seasonal variations in condition,reproductive activity and biochemical composition of the flat oyster,Ostrea edulis,from San Cibran (Galicia,Spain)

The influence of environmental parameters (temperature, salinity and available food) on the condition, reproductive activity and biochemical composition of a native population of Ostrea edulis L. in San Cibran (Galicia, Spain) was studied between September 1988 and December 1989. Histological preparations of gonads showed that gametogenesis started when temperature was at its lowest in winter. The water temperature in San Cibran never fell low enough to interrupt it. Gametogenesis proceeded slowly and spawning took place in May–June, although the predicted time of ripening was early March. Salinity in San Cibran was relatively stable throughout the year; it did not seem to have any influence on gametogenesis. Available food appeared to be a very important factor in controlling gonad growth, once gametogenesis was initiated. The major concentration of suspended organic particulate matter was present in the spring at the time of rapid gonadal maturation. Only one spawning period was observed. Larvae were released when the food in the water was high. Seasonal changes in the main biochemical components of this oyster were determined for a standard individual. Lipids and carbohydrates presented a similar time-course whereas proteins were constant. When food was abundant, energy reserves were built up. Spawning produced a decrease in biochemical constituent levels, and recovery coincided with the phytoplankton bloom. The stored reserves, mainly lipids, were used to overcome a state of energy imbalance in late autumn associated with low food availability. Results show this oyster to be an opportunist organism which concentrates its reproductive effort during a short period of favourable conditions and which is directly dependent on nutritive availability in the environment.     

Effect of temperature and food availability on reproductive responses of Streblospio benedicti (Polychaeta: Spionidae) with planktotrophic or lecithotrophic development

Streblospio benedicti (Webster) from Tar Landing North Carolina (NC), USA with either planktotrophic or lecithotrophic development were reared under two food levels and three temperature regimes (two mimicking seasonal cycles in NC and one at constant 20°C). During the eight-month experiment no females switched reproductive mode and no significant differences in survivorship or reproductive activity were observed between reproductive types. However, reproductive activity and fecundity-related parameters were subject to influence by food and temperature. Survivorship, body size, and larval production was greater in winter-spring than summer-fall regimes. Higher food levels produced increased survivorship, reproductive activity and egg production in adults with lecithotrophic development but no change in those with planktotrophic development. Body size, egg size, egg number, numbers of larvae per brood pouch, and brood size were strongly correlated in female S. benedicti and most correlation coefficients were similar (or identical) in individuals having planktotrophic and lecithotrophic development. A comparison of egg size and brood size in females from Tar Landing suggests that individuals with the two forms of development package offspring differently but expend approximately equivalent reproductive effort. Larval trophic mode is best viewed as a genetic polymorphism in S. benedicti. Individuals with planktotrophic and lecithotrophic development exhibit similar reproductive responses to environmental variation and there is no evidence for speciation.     

Energetics of growth and reproduction in cockles (Cerastoderma edule): Seasonal and age-dependent variations

Seasonal patterns of growth were compared interannually between 1983 and 1985 for different ageclasses of an intertidal population of cockles,Cerastoderma edule L., in the Mundaka Estuary (Basque Country, North Spain). Growth data and simultaneous determinations of respiratory metabolism were integrated over defined periods of time in order to compute assimilation and net growth efficiency. Consistent differences between the exponents that scale both assimilation and metabolism to body size led to growth efficiency values that were a decreasing function of age. Total growth was partitioned into three components: somatic growth, reproductive output and reserve storage, and the corresponding effort indexes were calculated. Despite important differences in total growth, reproductive output was constant between years, indicating a higher reproductive effort under poorer growth conditions. Increased costs of reproduction that appeared to be associated with this higher reproductive effort did not affect somatic maintenance, since previously stored carbohydrate reserves were primarily used to fuel gametogenesis.     

Habitat related growth and reproductive investment in estuarine waters,illustrated for the tellinid bivalve <Emphasis Type="Italic">Macoma balthica</Emphasis> (L.) in the western Dutch Wadden Sea

In estuarine areas, bivalve species can be found in a variety of environments, where they experience large differences in environmental conditions. In the present paper, the importance of different habitats (intertidal, subtidal and adjacent coastal waters) for the persistence of the population was evaluated for the bivalve Macoma balthica (L.) in the western Dutch Wadden Sea estuary. Intra-specific variation in growth and reproductive output were followed during the year and related to local abiotic conditions. Significant differences in growth and reproductive investment were found between locations. Young individuals were mostly found in the intertidal area, where growth in terms of somatic mass was good. These areas were not favourable for adult individuals, since growth in shell length was low and many individuals did not reproduce. In the subtidal, where the highest densities were found, somatic and gonadal mass indices were low. Coastal areas had the lowest densities and showed high growth in terms of shell length and body mass. The habitat with the highest reproductive effort per individual was not the most important habitat in terms of reproductive output due to differences in density and in size of the habitat type. For M. balthica, the subtidal habitat contributed the most to the reproductive output of the western Dutch Wadden Sea population although the highest reproductive output per individual was in the coastal area.     

Physioecology of zooplankton. IV. Effects of phytoplankton concentration,temperature, and body size on the net production efficiency ofCalanus pacificus

Weight-specific rates of individual production, total metabolic expenditure and assimilation, and net production efficiencies were estimated forCalanus pacificus Brodsky of selected body weights cultured at various phytoplankton concentrations and temperatures. The weight-specific rate of individual production increased hyperbolically with food concentration, and the maximum rate of individual production decreased logarithmically with a linear increase in body weight propotionally more at high than at low temperature. The weight-specific rate of total metabolic expenditure decreased logarithmically with increasing body weight and was unaffected by changes in food concentration. The effects of food concentration and temperature on the weight-specific rate of assimilation were similar to those on the rate of individual production, but the effect of body size differed considerably. The diversity in the temperature and body-size dependence of the maximum weight-specific rates of various physiological processes suggest (1) that, except for the metabolic rate, the allometric model (log-log relation) is inadequate for describing relationships between maximum rates of physiological processes and body size within species, and (2) that the common assumption that temperature affects the rates of various physiological processes in similar ways is not justified. Net production efficiency increased hyperbolically with food concentration, and the maximum production efficiency first increased slightly and then decreased gradually with increasing body weight. Small copepods attained higher efficiency at high temperature but larger ones did so at low temperature. The critical food concentrations for production efficiency and for the rate of individual production increased with increasing temperature and body size. Because of the effects of interactions among critical food concentration, temperature, and body size on the rates of growth and individual production and on net production efficiency, early development stages ofC. pacificus optimized growth and food conversion efficiency at high temperature, but late stages, particularly at low food concentrations, grew best and transformed food more efficiently at low temperature.Contribution No. 1130 From the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Acclimatization to intertidal conditions modifies the physiological response to prolonged air exposure inMytilus edulis

Both the tolerance of air exposure and the metabolic means by which mussles,Mytilus edulis, face air exposure can be modified by intertidal acclimatization. Short-term intertidal acclimatization enhanced the tolerance of air exposure most in early summer. Prevention of air breathing increased the mortality of wild and non-acclimatized cultured (i.e., control) mussels, but only slightly enhanced that of intertidally acclimatized mussels, suggesting their increased dependence upon anaerobic metabolism. In June, anaerobic end products accumulated in the adductor muscle of cultured but not of wild mussels. Intertidal acclimatization shifted the pattern and rates of metabolite accumulation. Control mussels accumulated succinate and alanine in the adductor muscle while intertidally acclimatized mussels accumulated strombine + alanopine, albeit at far lower levels. Pyruvate kinase (PK) from adductor muscle of intertidally acclimatized mussels was gradually activated during prolonged air exposure, while that of control and wild mussels was more inhibited. In July and August, during prolonged air exposure, the PK from the most viable mussles generally was the most inhibited. Prevention of air breathing did not markedly change the time course of PK inhibition during air exposure. Cultured mussels used in the present study were obtained from growers in Québec, and wild mussels were collected from the Baie des Chaleurs (Québec, PQ) in 1988.     

Quantitative studies on gonad cycle,fecundity, reproductive output and recruitment in a baltic Mytilus edulis population

Since Mytilus edulis L. is a biomass dominant in the Baltic much interest is focused on the ecology of the species. In this paper an attempt is made to quantitatively cover the reproductive cycle of a Baltic M. edulis population in order to provide data for energy flow models and to discuss aspects of recruitment in this species. Histological preparations of gonads showed that gametogenesis started with declining temperatures in autumn and proceeded very slowly through winter. At the beginning of March when food was supplied during the spring phytoplankton bloom, rapid maturation took place. This was also revealed by an increase in meat weight of the mussels. Only one spawning period was recorded, from the middle of May until the beginning of June, due to food being strongly limited to the population during the rest of the year. The length of the larval period was estimated as being 5 to 6 wk and settlement was registered from the end of June through July. In general the large annual variations found in the Baltic with regard to temperature and food abundance give rise to a more marked annual pattern in the reproductive cycle than is encountered in other seas. Fecundity was assessed for two populations from 4-and 15-m depths from studies covering two annual cycles of the changes in the relation of shell length — meat weight. The size-related fecundity was found to be equal in both populations and related to food abundance and not to growth or age. Fecundity, expressed as weight loss at spawning, ranged from 0% in 2-mm mussels and increased from 38 to 52% in 10-to 30-mm mussels. The fecundity as percentage of biomass in full-grown Baltic M. edulis is of similar magnitude as in full-grown mussels from other areas despite the smaller size of Baltic mussels. The reproductive output for the total 160 km² research area was calculated as being 1 200 tons dry weight or 80% of the standing stock, which, due to the particular features of the Baltic M. edulis population probably represents the larger part of the total mussel production. This reproductive output, calculated as 8·10⁷ eggs·m^-2 and corresponding to 50% of the total annual zooplankton production, may thus consitute an important food source for herring larvae and carnivorous zooplankton. Recruitment was divided into two phases: (1) Recruitment of juveniles (=settlement of larvae), and (2) recruitment to breeding stock. Monitoring studies of settlement on ropes and the year round presence of high abundances of mussels <2 mm indicate that settlement is in excess of the demands for maintaining population size and that most settled mussels form a pool of competitively suppressed non-growing individuals. Not until death of an already established mussel will these become recruited to the breeding population. Thus recruitment is possible throughout the year which stabilizes the population and maintains it near the carrying capacity of the area with regard to food and space availability.     

Age and sexual differences in the exploitation of two anthropogenic food resources for an opportunistic seabird

The availability of food resources has been suggested as a major factor in the substantial increase in reproductive output, survival, recruitment and, ultimately, population growth rates in most organisms. In fact, the artificial increase in food availability resulting from human activities has been suggested as a factor in the substantial increase in population size of several seabirds in recent decades. In the present study, our primary aim was to estimate the importance of the main natural prey and two alternative feeding resources, fishery discards and the invasive American crayfish Procambarus clarkii, for an opportunistic seabird, the Audouin’s gull Larus audouinii. We also assessed the influence of age and sex in the use of those three types of food. For this purpose, we compared the analyses of δ¹⁵N and δ¹³C in blood of male and female adults of known age and chicks with those in their potential prey. Our results reveal sex-related and age-related differences in the consumption of fish discards, small pelagic fish and American crayfish. Differences in the diet of males and females and also between adults and chicks could be related to different nutritional requirements. Age differences were probably related to their different foraging proficiency and the tendency of young breeders to opportunistically exploit anthropogenically derived food. This study illustrates the importance of considering the age and sex of individuals to obtain feasible dietary information and to understand how the exploitation of food of human origin could affect population growth.     

Multi-year lactation and its consequences in Bornean orangutans (Pongo pygmaeus wurmbii)

In most mammals, females pay for reproduction by dramatically increasing net energy intake from conception to mid- or late lactation. To do this, they time their reproductive events in relation to environmental cycles so that periods of peak food availability coincide with peak demand or are used to build energy stores. This timing is not possible in species with slow development in which lactation is prolonged over a multi-year period with fluctuating food availability. Here, mothers are expected to sustain a stable but generally lower level of nutrient transfer. In a sample of over 1,050 complete follow days of eight mother–infant pairs collected over 7 years, we document maternal effort for wild Bornean orangutans (Pongo pygmaeus wurmbii) over their average 6.5-year lactation period. As predicted, maternal feeding time was independent of the age of her growing offspring, indicating a stable sustained “plateau” effort of ≤ 25 % above baseline level, instead of a short peak lactation as seen in seasonal breeders. Infant orangutans started to regularly supplement milk with self-harvested food when they were 1–1.5 years old, indicating milk intake was insufficient from this age onwards, even though maternal effort did not decrease. We expect the same regulation of sustained maternal effort in other large and large-brained mammals with slow infant development. We also predict that mother–infant conflict over suckling may show another peak at the onset of the milk?+?solid food phase, in addition to the well-known conflict around the endpoint of lactation (weaning), which is reached after a long and gradual increase in solid food intake by the infant.     

Effects of temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis

To assess the effects of both temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis (O.F. Müller), individuals collected December 1996 (winter experiment) and June 1997 (summer experiment) were maintained for 3 months in one of four experimental treatments: (1) 3 °C and fed ad libitum (high ration), (2) 3 °C and fed one-seventh of the maximum ration (low ration), and (3) 12 °C and fed the high ration; (4) 12 °C and fed the low ration. All individuals were fed an artificial diet and exposed to only 1 h of light every day. At the end of both experiments, mean gonad indices of sea urchins fed the high ration had increased significantly (11–24% and 6–19% in the winter and summer experiments, respectively), while the gonad indices of individuals fed the low ration did not change. At the high ration (both experiments), the increase in gonad index of sea urchins occurred primarily as the result of a significant increase in the mass of nutritive phagocytes, as revealed by histological analyses. Primary oocytes were significantly larger in individuals held at 3 °C than at 12 °C throughout the winter experiment, regardless of food ration; during the summer experiment, primary oocytes were significantly larger in individuals receiving the high ration, regardless of the temperature at which they were held. These results suggest that: (1) food availability is the most important factor regulating energy storage and the relative size of gonads throughout the year, (2) temperature affects the rate of growth and maturation of primary oocytes during the later stages of oogenesis, and (3) once gametogenesis has been initiated, mature ova will be produced, even under conditions of low food availability. Conditions of high food availability in summer and low temperature in winter would thus favor reproductive output in sea urchin populations. Received: 1 March 2000 / Accepted: 4 October 2000     

Age- and state-dependent reproductive effort in male mountain goats, Oreamnos americanus

In mammalian polygynous mating systems, male reproductive effort consists mainly of male–male competition and courting of females, which entail substantial somatic costs. Males are thus expected to adjust their reproductive effort according to their age and condition. In this study, we examined how activity budgets of male mountain goats (Oreamnos americanus), a polygynous ungulate, varied with age in a marked population over two periods: (1) summers 1995–2006 and (2) ruts 2004–2006. We then assessed if the proportions of time spent in male–male competition and courtship behaviors were influenced by age-specific body mass and social rank during the rut. Males spent most of their time foraging and resting during summer, and rested more and foraged less with increasing age. During the rut, pronounced shifts in activity budgets occurred as juveniles (1–2 years) increased time spent foraging, whereas adults (≥3 years) increased standing and time spent in social interactions at the expense of foraging. At old age, reproductive effort either stabilized or decreased slightly, providing weak support for the ‘mating strategy–effort’ hypothesis, predicting that courtship behaviors should peak in prime-aged males. Age-specific body mass did not affect time spent in male–male competition, but was positively related with time spent in courtship behaviors, providing support for the ‘individual quality’ hypothesis, predicting that males with more resources at the start of the rut should spend more time in mating-related activities. Age-specific social rank did not affect reproductive effort. Surviving to prime age while increasing mass each year should thus allow male ungulates to gain greater ability to court estrus females.     

Seasonal variation in the effects of food availability on gametogenesis in the purple urchin (<Emphasis Type="Italic">Strongylocentrotus purpuratus</Emphasis>)

The amount of food and when it is available affect both the timing of reproduction and the number of gametes produced by purple sea urchins, Strongylocentrotus purpuratus. To investigate this further, the effects of food availability on feeding rates, gonad growth, and gamete development were examined in S. purpuratus collected from the Point Loma kelp forest near San Diego, California, USA (32.69° N, 117.26° W) in September (Fall) 2007, and February (Spring) and July (Summer) 2008, using laboratory mesocosms. Each seasonal laboratory feeding experiment lasted 3 months, and different levels of food availability were established with different feeding frequencies (from 1 to 7 days week⁻¹). Gonad tissues of male and female urchins were staged at the end of each experiment using histological analyses. Reduced food availability resulted in increased daily consumption rates, especially in the Fall when gamete development began. Food limitation at this time resulted in failure to produce viable gametes, suggesting there is a critical period early in gonad development when food limitation affects reproductive competency. Food limitation later in gonad development did not stop viable gamete production, although it did reduce gamete output.     

Effects of age and food availability on diel vertical migration of Calanus pacificus

Age-specific differences in diel vertical migration behavior of Calanus pacificus were investigated in a 58 d (30 April–26 June, 1981) experiment in the Scripps Institution of Oceanography Deep Tank, La Jolla, California, USA; the experiment spanned three successive generations of copepods. The onset of vertical migration behavior occurred in the first feeding stage, Nauplius III. The amplitude of vertical migration gradually increased with age, becoming maximal in the late copepodite stages. Night depths remained constant with age while daytime depths increased. The migratory behavior of late copepodite stages was influenced by food availability. When phytoplankton was abundant and individual ingestion rates were high, copepodites performed high-amplitude migrations. As food availability declined, however, and the competition for food increased, migration amplitudes decreased and then ceased altogether so that copepodites remained in the relatively food-rich surface waters at all times. We suggest that hunger is the primary factor controlling vertical migration behavior.