Food limitation and seasonal population declines of rotifers

Summary Keratella cochlearis exhibited consistent seasonal abundance patterns during a four-year study in Little Rock Lake, Wisconsin, U.S.A. In each year, spring population peaks were followed by strong summer reductions. Concomitant with population declines, there were reductions in rotifer egg ratios. Keratella taurocephala abundance patterns were similar to those of K. cochlearis during 1984 and 1985, but not in 1986 and 1987, when spring peaks and summer declines were not apparent. However, summer declines in the egg ratio of K. taurocephala were observed during each year. The reduction in rotifer populations simultaneously with decreased egg ratios suggested that population declines were caused by food limitation. Food-addition experiments conducted in situ in small enclosures indicated that food was limiting for K. cochlearis when its populations were declining, but not during other periods of the year. Keratella taurocephala did not show a consistent response to food addition.     

Observations on egg production rates and seasonal changes in the internal morphology of Mediterranean populations of Acartia clausi and Centropages typicus

Egg production rates in wild populations of Acartia clausi and Centropages typicus, sampled biweekly in the Gulf of Naples from October 1985 to July 1987, showed marked seasonal fluctuations with maximum values in early spring that proceeded the annual maxima for adult female densities in summer. A positive correlation between chlorophyll a concentrations and egg production was evident only during the early spring phytoplankton bloom. A strong diminution in egg deposition occurred later in spring and continued throughout the summer notwithstanding high chlorophyll concentrations. In winter, when population abundances for adult females were lowest, egg production rates were always higher than in summer. Differences in egg production rates coincided with pronounced morphological changes between summer and winter populations of both species. The most striking of these changes consisted, in winter, in the presence of a dark brown fluid-like mass of granular material that seemed to freely bathe the gonads. The presence of this substance only during periods of elevated egg production suggests that it may enhance egg production rates when the adult population reaches minimum annual levels. Such a mechanism of self-regulation may operate to dampen the effects of environmental variability thereby contributing to maintain a conservative structure in coastal copepod communities.     

Local variation in overwintering stages ofPanonychus akitanus Ehara (Acarina: Tetranychidae)

The overwintering behaviour ofPanonychus akitanus Ehara was compared in a population (Sapporo) overwintering as the egg and the adult female and in a population (Tomakomai) overwintering in the egg stage only. In both populations, winter eggs were larger in size and darker in colour than summer eggs, and females laying winter eggs showed clear differences in body colour and behaviour from summer females. Both populations produced winter eggs in response to short photoperiods, and the critical photoperiod was between 13.0 and 13.5 h in the Sapporo population and between 13.5 and 14.0 h in the Tomakomai population. The hatchability of winter eggs of the Sapporo population collected from the field began to increase in November, and that of the Tomakomai population in December. In the Sapporo population, a 40-day or more exposure to 5°C promoted hatching of winter eggs, and a 50-day exposure of females which had previously laid some winter eggs to 5°C induced the production of summer eggs. Females of the Sapporo population laid winter eggs in autumn, and then themselves overwintered as well, and produced summer eggs in the following spring. These females were called winter-habit females.The winter eggs of both populations were diapausing. However, the winter-habit females of the Sapporo population were not reproductively inactive and their condition of diapause was not certain. The ecological significance of the winter-habit females in the Sapporo population is discussed.     

The reproductive biology of Euchaeta antarctica Giesbrecht (Copepoda: Calanoida) at South Georgia

The reproductive biology of the predatory calanoid copepod Euchaeta antarctica Giesbrecht was investigated in two interconnected fjord systems at South Georgia. Counts of the number of spermatophores attached to adult females and the number of egg sacs encountered, indicated probable peaks of reproduction in summer and winter. Patterns of spermatophore placement were examined and compared with data for E. norvegica (Boeck) from boreal waters. Elemental analysis indicated a high proportion of carbon and a low proportion of nitrogen in adult females and egg sacs from both sites.High winter carbon levels in adults seem related to their predatory feeding habits allowing high food intake throughout the year whereas in egg sacs it probably reflects extended development times and/or non-feeding naupliar stages.Mean adult female and egg clutch dry weights were higher in Cumberland East Bay during winter than in Moraine Fjord during summer. These differences are discussed in the context of relationships between fjord morphology and production levels.     

Calamoecia lucasi (Copepoda,Calanoida) and other Zooplankters in Two Rotorua,New Zealand,Lakes

The seasonal cycles of the zooplankton in 1967–1969 in Lakes Rotorua and Rotoiti in the North Island of New Zealand are outlined. Bosmina meridionalis had several peaks in abundance during the summer and low winter numbers, whilst Ceriodaphnia dubia generally had only a single period of real abundance during the summer. The rotifer Asplanchna priodonta was present throughout the year in low numbers except for two brief but striking peaks in the spring-summer period. The only cyclopoid copepod present was Macrocyclops albidus which was never abundant. The annual cycles of the calanoid copepod Calamoecia lucasi were studied in some detail. Total numbers were high but very variable during spring-autumn and low in winter, the seasonal cycles being very similar in both lakes. Breeding was continuous although egg clutches were small (1.1–6.6), and the development of at least 5 cohorts per year could be followed from data on breeding intensity, population age-structure and adult size. Laboratory determined egg development rates were used to calculate birth rates in the populations so that production and losses could be estimated. Survival rates appeared to be higher in Rotoiti than in Rotorua. Birth and death rates were low, the annual means for both linear and exponential rates ranging from around 0.04 to 0.07, giving turnover times for individuals of 12–18 days in Rotorua and 18–24 days in Rotoiti. Population turnover rates were higher in Rotorua (19–28 days), which had a similar mean population per square metre but higher clutch sizes, than in Rotoiti (18 days).     

Intermittent fasting during winter and spring affects body composition and reproduction of a migratory duck

We compared food intake, body mass and body composition of male and female black ducks (Anas rubripes) during winter (January-March). Birds were fed the same complete diet ad libitum on consecutive days each week without fasting (control; nine male; nine female) or with either short fasts (2 day x week(-1); nine male; nine female), or long fasts (4 day x week(-1); eleven male; twelve female). We continued treatments through spring (March-May) to measure the effect of intermittent fasts on body mass and egg production. Daily food intake of fasted birds was up to four times that of unfasted birds. Weekly food intake of males was similar among treatments (364 g x kg(-1) x week(-1)) but fasted females consumed more than unfasted females in January (363 g x kg(-1) x week(-1) vs. 225 g x kg(-1) x week(-1)). Although both sexes lost 10-14% body mass, fasted females lost less mass and lipid than unfasted females during winter. Total body nitrogen was conserved over winter in both sexes even though the heart and spleen lost mass while the reproductive tract and liver gained mass. Intermittent fasting increased liver, intestinal tissue and digesta mass of females but not of males. Fasting delayed egg production in spring but did not affect size, fertility or hatching of the clutch. Females on long fasts were still heavier than controls after laying eggs. Thus black ducks combine flexibility of food intake with plasticity of digestive tract, liver and adipose tissue when food supply is interrupted during winter. Females modulate body mass for survival and defer reproduction when food supply is interrupted in spring.     

On the relationship between food, reproduction and survival of two carabid beetles: Calathus melanocephalus and Pterostichus versicolor

Abstract.

• 1 The relative influences of temperature and availability of food on reproduction, survival and growth of all developmental stages of two carabid beetle species are discussed with special reference to the suggested relationship between availability of food, size of egg production and survival of adults from one breeding season to the next.
• 2 Temperature as well as food supply influence the length of larval growth and adult body size. Beetles grown at low temperatures and low amounts of food are smaller than those grown at higher temperature and with more food.
• 3 The number of eggs laid per female was correlated with the amount of food gathered. There was no inverse relationship (trade-off) between reproductive output and survival in the field until the next breeding season.
• 4 In 1980 no significant relationship was found between winter mortality and the amounts of food gathered by beetles in the period after reproduction and before winter diapause. However, in 1981 in C. melanocephalus a lower number of starved beetles survived the winter than the fed ones and‘field’beetles.
• 5 Only in the first part of the feeding activity period in autumn can enough food be gathered by C.melunocephalus for successful hibernation. In the second part of this period there is not enough food to build up the fat reserves needed to survive the winter.
• 6 Difference in population fluctuations of both species are discussed in relation to their life histories.

     

Temporal variations in growth and reproduction of Tedania anhelans and Chondrosia reniformis in the North Adriatic Sea

Most works concerning growth and reproduction of Mediterranean sponges have been performed in the oligotrophic western Mediterranean while little is known about sponge dynamics in the North-western Adriatic Sea, a basin characterized by low winter temperature and eutrophy. In order to deepen our understanding of sponges in the North Adriatic Sea and verify how its peculiar trophic and physical conditions affect sponge life cycles, temporal trend of sponge cover (%) and reproductive timing of Chondrosia reniformis and Tedania (Tedania) anhelans were studied over a 1-year period looking for a possible relation with variations of temperature or food availability. In C. reniformis, although little variations of sponge cover were evidenced around the year, the number of individuals and their size increase during spring. Asexual reproduction, via drop-like propagules, mainly occurs in spring and summer, while sexual reproduction is characterized by a maximum number of oocytes in August. T. anhelans progressively grows from spring to summer and develops propagules on its surface that reach their maximum size in July. In autumn, the sponge undergoes a process of progressive shrinkage and almost disappears in winter when temperature reaches 7–8°C. Larvae occur during summer. In the North Adriatic Sea sponges have larger sizes, higher density and a wider period of oocytes production compared with the same species from the Mediterranean Sea, suggesting these differences could be due to high food availability characterizing the eutrophic Adriatic basin. On the contrary, the sharp water temperature variations and the very low winter temperature, 5–6°C lower than what has been reported for the Mediterranean Sea, regulate temporal variations in abundance and cause the disappearance of thermophile species during winter.     

Population dynamics of daphnia pulex and utilization by the rainbow trout (salmo gairdneri)

The effect of rainbow trout (Salmo gairdneri) predation on the population dynamics of the water flea,Daphnia pulex, was examined during 1976 and 1977 in Becker Lake, a small, shallow, productive reservoir in northeastern Arizona.Rainbow trout were size-biased feeders, utilizing daphnids which were 1.3 mm in size or larger. Trout predation uponDaphnia pulex occurred mainly during winter and early spring when their numbers were relatively low but their clutch size high, suggesting that trout selectedDaphnia on the basis of brood pigmentation. By far the greatest proportion ofDaphnia mortality was due to nonpredatory sources, since generally less than 6% could be attributed to trout predation.TheD. pulex population exhibited a bimodal seasonal abundance curve which was attributed to ephippial egg production and trout predation during the winter and poor food quality/quantity during the summer.This work was supported in part by the Arizona Department of Game and Fish. The authors wish to thank Jim Novy and Joe Stone of that department for their invaluable assistance in the field collecting segment of this study.     

Annual cycle in female Three-spined sticklebacks (Gasterosteus aculeatus L.) from an upland and lowland population

The annual cycles in female Three-spined sticklebacks from an upland (275 m) and a lowland (25 m) population were compared. The average length and weight of the lowland females were greater, but in both populations growth was checked in winter and again in the breeding season. Both populations also showed a winter decline in condition followed by a sharp increase in spring associated with the maturation of the ovaries. The hepato-somatic index of the females also increased during the spring, but the dry matter content of the liver declined during the spring and early summer. Lowland females had larger livers than the upland fish. In both populations average gonadosomatic index increased steadily throughout the winter, then abruptly in spring. The lowland population bred from May to early August, but the upland population bred only in May.
The energy content of the carcases of females was lowest in winter and again during the breeding season whereas the ash content showed maxima at these two periods. Lipid and glycogen analyses of the carcase, liver and ovaries also suggested that winter and the breeding season were periods of depletion from the carcase and liver, but not from the ovaries.
The observations indicate that the ovaries are to some extent insulated from fluctuations in resource availability. This supports experimental studies which indicate that egg production in the female stickleback can be subsidised by depletion from the carcase and liver when necessary.     

Phenotypic divergence in reproductive traits of a moth population experiencing a phenological shift

Allochrony that is reproductive isolation by time may further lead to divergence of reproductive adaptive traits in response to different environmental pressures over time. A unique “summer” population of the pine processionary moth Thaumetopoea pityocampa, reproductively isolated from the typical winter populations by allochronic differentiation, is here analyzed. This allochronically shifted population reproduces in the spring and develops in the summer, whereas “winter” populations reproduce in the late summer and have winter larval development. Both summer and winter populations coexist in the same pine stands, yet they face different climatic pressures as their active stages are present in different seasons. The occurrence of significant differences between the reproductive traits of the summer population and the typical winter populations (either sympatric or allopatric) is thus hypothesized. Female fecundity, egg size, egg covering, and egg parasitism were analyzed showing that the egg load was lower and that egg size was higher in the summer population than in all the studied winter populations. The scales that cover the egg batches of T. pityocampa differed significantly between populations in shape and color, resulting in a looser and darker covering in the summer population. The single specialist egg parasitoid species of this moth was almost missing in the summer population, and the overall parasitism rates were lower than in the winter population. Results suggest the occurrence of phenotypic differentiation between the summer population and the typical T. pityocampa winter populations for the life‐history traits studied. This work provides an insight into how ecological divergence may follow the process of allochronic reproductive isolation.     

Life history and production of Agapetus quadratus (Trichoptera: Glossosomatidae) in a temporary, spring-fed stream

1. The life history and trophic basis of production of the caddisfly grazer Agapetus quadratus were studied in the torrent Gorg Blau, a spring‐fed stream on the island of Majorca that dries annually during summer. 2. Quantitative random samples were taken every 2–3 weeks during an annual surficial flow period, from November 2000 to mid‐July 2001. Instars of field‐collected larvae were determined by measurements of head width and pronotum length, and the sex of all pupae was determined to study sexual dimorphism and sex ratio. 3. Stage‐frequency histograms suggested a trivoltine population, with an average cohort time of 4 months. Larval development was asynchronous, with continuous growth and overlapping generations. Recruitment peaks were identified in mid‐November, early March and late June, indicative of winter, spring and summer generations. On average, females were larger than males and the mean sex ratio was 2 : 3 (females : males). Population densities and biomasses derived from the field data were used to calculate production and turnover rate. 4. Annual production of A. quadratus in the torrent Gorg Blau (4.80 g dry mass m^?2 year^?1) was the highest ever reported for the genus, being comparable with that estimated for some insects with rapid development and multiple cohorts. 5. Estimates of production of A. quadratus were combined with foregut content analysis to estimate the fraction of total production derived from the principal food sources: algae and organic detritus. Algae supported a major proportion of the production of this grazer. 6. The low density of predators characteristic of many temporary streams, and the small amplitudes in discharge and temperature during most of the wet period that characterise the spring habitats might allow high levels of grazer production in this particular Mediterranean stream.     

Patterns of energy storage in Pseudoboeckella poppei (Crustacea,copepoda) from two contrasting lakes on Signy Island,Antarctica

The copepod Pseudoboeckella poppei (Daday) (Calanoida, Centropagidae) was sampled from Sombre and Heywood Lakes on Signy Island, Antarctica (60° S, 45° W) between January 1984 and March 1985. Sombre Lake is clear and oligotrophic with little phytoplankton and a bottom sediment low in organic content. By contrast Heywood Lake is turbid and mesotrophic; a substantial phytoplankton develops in summer and the bottom sediments are comparatively rich in organics. Both lakes freeze over for much of the year, forcing the copepods to adopt a benthic feeding strategy over winter. Adult Pseudoboeckella feed on phytoplankton when this is available, but also on detritus, diatoms and short algal filaments stirred up from the sediment. In Heywood Lake, male copepods show a smooth seasonal trend in lipid content with lipid being synthesised in early summer and utilised in late summer and winter. The summer increase in lipid content is associated with an increase in dry weight. Female lipid contents show evidence of two peaks of egg production. In Sombre Lake both male and female copepods increase in size during summer and show a wider range of lipid contents than in Heywood Lake; it is likely that this is due to the poorer winter feeding conditions which necessitate the synthesis of a much larger store of reserves during the summer. In contrast to marine calanoid copepods, lipid stores are exclusively triacylglycerol with no trace of wax ester.     

Breeding of the bush fly,Musca vetustissima Walker,in an over-wintering area during winter and spring

Field experiments in an area of south-western Australia where bush fly (Musca vetustissima) adults occur permanently, showed that it overwintered by continuous breeding; but only two to three generations occurred between June and September. Low survival of eggs and larvae in the food (cattle dung) from May to August was associated with rainfall rather than low temperatures. High egg-adult survival occurred in late August; lower egg and larval survival in September and early October was attributed partly to dung fauna. Egg and larval survival was high in late October until December but predation or parasitization caused low survival of puparia outside the dung. Experimental and wild adult flies were largest and therefore most fecund in early spring; smaller wild flies in late spring appeared to result from larval competition for food. Simultaneous high egg-adult survival, high fecundity and rapidly decreasing generation times in late winter and early spring provide a basis for explaining the major increase in adult bush fly abundance observed in some areas in mid-spring.     

Effects of food limitation,notonectid predation,and temperature on the population dynamics of Daphnia carinata

The relative contributions of invertebrate predation (Notonectidae: Anisops spp.), food limitation, and certain abiotic factors in driving the population dynamics of Daphnia carinata were studied for a two year period in a large farm dam in southern Victoria, Australia. Detailed measurements were made on the population densities of Daphnia and Anisops spp., the amount of food available, the nutritional status, and the size-specific fecundity of Daphnia. The density of the Daphnia population at the field site oscillated closely with water temperature. The amplitude of the population fluctuations varied seasonally, being much greater during the warmer months of the year and switching to fluctuations with low peaks when water temperature dropped below approximately 15°C. Anisops spp. densities were greatest in winter and declined during the spring of each year. Nymphs appeared in late spring and early summer and numerically dominated the population. During the warmer periods of the year, the daphnid population went through a series of rapid growth phases leading to over-exploitation of food resources and subsequent population collapses. Daphnia population densities were not correlated with Anisops numbers suggesting that predation was not a major regulatory factor during the warmer periods of the year. When water temperatures fell below ca. 15°C daphnid population densities remained low despite high food levels. During this period the impact of Anisops may have been greater. Two distinct phases were identified: a warm water period when food limitation was the main regulatory factor, and a cool water period when Anisops predation may have been the paramount factor. Low oxygen concentrations were associated with heavy rainfall in the spring and may have had a limiting effect on Daphnia for short periods. Daphnia may have had an important role in sustaining the Anisops population in the pond over each winter.     

Population dynamics of Daphnia spp. and implications for trophic interactions in small, monomictic lake

In Lake Oglethorpe, Georgia, USA, herbivorous crustacean zooplankton are abundant and dominate zooplankton biomass in winter, but are scarce throughout most of the summer. We used a 3.5 year study of Daphnia population dynamics to infer when food, predators or temperature constrained growth of this population. Transitions between winter and summer consumer assemblages are concurrent with seasonal changes in water temperature, thermal structure (stratification/destratification), resources (autotrophic/heterotrophic-domained production), and predator abundance and activity (e.g. Lepomis macrochirus and Chaoborus punctipennis. We sampled at weekly or less intervals from April 1992 to September 1995, and determined population abundances for all cladoceran species. For the Daphnia population (Daphnia ambigua + Daphnia parvula, we measured clutch size and length for all individuals. We used average water column temperature (where dissolved oxygen is >1 mg l^-1) to estimate egg development time from an empirical model. Estimates of Daphnia population birth and death rate were thus generated from abundance, egg ratio and temperature/dissolved oxygen data. We compared observed birth rate (bobs) with expected birth rate (bexp>95% CI; predicted for food-saturated conditions at ambient temperature). For variable (1-13 week) periods between later November and March, 1992-1995, water temperature was the primary factor constraining Daphnia population growth (bobs = bexp). From about April to early November, bobs < bexp suggested food-limited population growth. In spring, summer and early fall (March-October), population densities were several-fold lower than in late fall and winter (November-February). However, all else being equal, egg ratio and population birth rate data would have predicted that Daphnia abundance fluctuates over equivalent ranges in spring and fall. We interpret this discrepancy as evidence for increased rates of extrinsic mortality during the growing season and a seasonal shift in the relative importance of resource and predator regulation. The duration of predator suppression of crustacean population abundance in Lake Oglethorpe and other warm-latitude lakes (36N-27°S) is longer (3-7 months) than that observed in north temperate lakes (1-2.5 months; 41-52°N).      

Seasonal life cycle and autogeny in the mosquitoAedes togoi in northern Kyushu,Japan, with experimental analysis of the effects of temperature,photoperiod and food on life-history traits

We studied seasonal changes in the larval population structure, adult size, and autogeny (egg production without a bloodmeal) of the mosquitoAedes togoi on the seacoast of northern Kyushu, Japan. The effects of temperature, photoperiod and food conditions on larval development, adult size and autogeny were studied in the laboratory.Aedes togoi overwintered in both the egg and larval stages and was multivoltine. Adult size was greater in spring and autumn and smaller in summer. Autogeny occurred in spring and autumn but not in summer. Autogenous females were larger than anautogenous females, and larger autogenous females produced more eggs. Laboratory experiments showed that autogeny was promoted under low temperature and short-day conditions which corresponded with spring/autumn conditions in the field. Experimental food limitation in the larval stage greatly reduced adult size and autogenous egg production. In the field, large inter-pool variations in adult size and autogeny rate probably resulted from variations in nutrient and crowding conditions. Seasonal autogeny ofAedes togoi was discussed in terms of life-history strategy under fluctuating environmental conditions where relative advantages of autogenous and anautogenous reproduction alternate seasonally.     

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.     

Autotrophic Picoplankton: Community Composition,Abundance and Distribution across a Gradient of Oligotrophic British Columbia and Yukon Territory Lakes

Autotrophic picoplankton communities were examined in eleven oligotrophic lakes from a broad geographic region of western Canada, representing a variety of physico-chemical and biological conditions. During our study, several of the lakes were treated with additions of inorganic nitrogen and phosphorus fertilizers. Picoplankton communities in most lakes were dominated (>70%) by unicellular or colonial coccoid cyanobacteria, provisionally identified by morphological and autofluorescence properties as Synechococcus. Also common in some lakes were red-fluorescing cyanobacteria and Chlorella-like eucaryotes. Autotrophic picoplankters contributed from 36-63% to total chlorophyll, from >2-26% to total phytoplankton carbon, and from 29–53% to total photosynthesis. Average populations ranged from >5-10,000 cells·ml⁻¹ in winter and early spring to 65-75,000 cells · ml⁻¹ in summer and fall. Peak densities in most lakes occurred in August-September and most populations were within the epilimnion or metalimnion/hypolimnion boundary. Subsurface peaks were prevalent only in untreated, strongly stratified lakes. Eucaryotic picoplankters became dominant in acidic (pH < 6.2), humic lakes. Colonial picoplankters were more common in more productive interior lakes in August, and though present, were uncommon in coastal systems. Picoplankton populations exhibited large increases under ice in a Yukon lake, and their abundance and seasonal distribution showed little relation to temperature or to light. Fertilization of lakes resulted in picoplankton population increases (>2x) and the elimination of subsurface peaks. Nutrients were considered to be one of the major factors controlling population abundance in these oligotrophic lakes with average pH < 6.5.     

Saving for the future: Pre‐winter uptake of algal lipids supports copepod egg production in spring

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