Omnivorousness in marine planktonic copepods

Adult females of Centropages furcatus and Temora stylifera hadsignificantly different feeding behaviours. At equal averagefood concentrations, C. furcatus ingested more nauplii of thecalanoid copepod Pseudodiaptomus coronatus and fewer diatomsthan T. stylifera. At 20°C and 2.0 µg of nitrogen.1.1.^–1of each food source, C. furcatus ingested 17.2% and 4.4% ofits body nitrogen daily as nauplii and diatoms, respectively.Ingestion rates of the same food sources by T. stylifera were5.1% and 14.5%, respectively. At the environmental concentrationssimulated in these experiments, the presence of diatoms didnot affect the ingestion of nauplii nor did the presence ofnauplii affect diatom ingestion. The behaviour of predator andanimal prey largely dictates the types and quantities of preyingested.     

Motion behavior of nauplii and early copepodid stages of marine planktonic copepods

The goal of this study was to quantify periods of activity andvelocities of late naupliar and early copepodid stages of planktoniccopepods occurring regularly on the southeastern continentalshelf of the USA. We obtained quantitative information on eightspecies, including adult females of Oithona plumifera. All studieswere conducted at food concentrations near or above satiationlevels. Activities ranged from 0.85% (adult females of O.plumifera)to 100% of time (nauplii and copepodids of various calanoidspecies). Motion velocities (excluding escape motion) coveredmore than one order of magnitude: from 0.39 mm s^–1 fornauplii of Temora stylifera to 5.24 mm s^–1 for naupliiof Oncaea mediterranea. Ranges of activities of species rangefrom occasional for early juveniles to adult females of O.plumiferato 100% for the same range of T.stylifera, the latter creatinga feeding current from N III onwards, the former not at all.Of notable interest is Centropages velificatus which moves intermittentlyas a late nauplius, continuously as an early copepodid and intermittentlyas an adult. All observed calanoid late nauplii and copepodidsmove in three dimensions, excluding copepodids of the shelfbreak/oceanicParacalanus aculeatus. The results indicate not only significantdifferences in motion behavior between cyclopoids and calanoids,but also between calanoid species. Yet, some calanoid speciesshow little ontogenetic changes at all.     

The effect of food limitation on gonad development and egg production of the planktonic copepod Calanus finmarchicus

Calanus finmarchicus, one of the dominant copepods species of the North Atlantic, often encounters low food concentration or quality during the reproductive period; however, our knowledge on the effect of these conditions on reproduction processes is scarce. The present study combines experiments with histological observations to describe the response of C. finmarchicus to limited food focussing on (1) oocyte maturation processes, (2) gonad morphology, (3) egg production rates as a function of spawning frequency and clutch size, and (4) the fuelling of egg production by measuring carbon and nitrogen content of the females. In the laboratory, C. finmarchicus females were exposed to 0, 10, 50, 150 or >300 μg C l⁻¹ for several days. To account for food quality and season, reproductive activity was compared in April and July 1999 between females feeding on diatoms or dinoflagellates. The effect of feeding history was studied in February with females fed and starving prior to the experiment. Feeding conditions had severe effects on oocyte maturation process. Hence, egg production varied significantly with food concentration and quality, season and feeding history due to variation in both clutch size and spawning frequency. Clutch size differed by a factor of 2-4 between food limited and well fed females, and is thus an important parameter for modelling egg production. Changes in clutch size were related to changes of the number of maturing oocytes in the females gonads indicating that the latter can be used to precise the prediction of egg production from preserved samples. The proportion of females carrying at least some mature oocytes was relatively high at low food availability. Apparently, these females used internal body reserves as the carbon and nitrogen content decreased significantly under these conditions. These results indicate that C. finmarchicus embarks on the strategy to enable reproduction in all or many females of a population at low rates when feeding conditions are unfavourable.     

Temporal variability in copepod fecundity during two different spring bloom periods in coastal waters off Plymouth (SW England)

The temporal variation in egg production of the planktonic copepodsCalanus helgolandicus, Temora longicornis and Pseudocalanuselongatus was studied during two different spring bloom periodsin 1989 and 1990 by weekly sampling at two permanent stationsin coastal waters off Plymouth (SW England). Copepod egg productionwas estimated in situ by incubating individual adult femalesin filtered seawater for 24 h (72 h until hatching for P.elongatus)at the field surface temperature. The relationship between copepodegg production rates and: (i) chlorophyll a concentration (totaland >10 µm size fraction), (ii) temperature and (iii)initial copepod gut pigment content was investigated. The springbloom periods were very different in both years, with the occurrenceof a Phaeocystis sp. bloom in 1990, which negatively affectedthe feeding and fecundity of copepods. Egg production ratesin spring 1989 were significantly correlated with chlorophylla concentration (particularly with the >10 µm fraction),field temperature and copepod gut pigment contents. In spring1990, egg production rates were also correlated with copepodgut pigment contents, but no significant correlations were obtainedwith temperature or with chlorophyll a concentration, as a consequenceof the lower egg production rates obtained during the Phaeocystissp. dominance period. These results show that food availabilityis the factor which mainly affects the fecundity of neriticcopepods in short time periods.     

Zooplankton growth rates: diel egg production in the copepods Oithona, Euterpina and Corycaeus from tropical waters

Clear diel periodicity in the timing of egg laying was observedin the tropical cyclopoid cope-pods Oithona plumifera, Oithonanana, Oithona simplex and Corycaeus amazomcus, and the harpacti-coidEuterpina acutifrons in waters surrounding Jamaica, West Indies.Individual females incubated under ambient food conditions formultiple clutches displayed remarkable constancy in their clutchdurations (= egg or embryonic development time). All speciesexhibited clutch durations of 20–23 h and cycle durations(= sum of clutch and inter-clutch durations) of 24 h, exceptfor the offshore species O.plumifera where the clutch durationwas 36 h and the cycle duration 48 or 72 h. Egg laying appearedto be synchronized by photoperiod, occurring around dawn formost species. In order to avoid biases, diel periodicity mustbe accounted for in estimates of egg production rates, and futureefforts should concentrate on the responses of clutch size andinterclutch durations to natural food climates.     

Elemental and isotopic fractionation of carbon and nitrogen by marine, planktonic copepods and implications to the marine nitrogen cycle

Particle-grazing copepods, primarily Temora longicornis andT. stylifera, and seawater with natural particles were collectedfrom the northwest Gulf of Mexico. Control and ammonium-enrichedaliquots of seawater were incubated in triplicate for 2 days,copepods added and the incubation continued for 2 days. Analyseswere made of dissolved nutrients (nitrate, ammonium and phosphate),suspended particles (chlorophyll a and phaeopigments, C, N,     

Mesozooplankton community indicates climate changes in a shelf area of the inner Bay of Biscay throughout 1988 to 1990

The mesozooplankton community was monitored at both coastaland offshore sites of the Basque shelf (inner Bay of Biscay)during 1988–1990 This was considered of interest becausethe local climate change toward dry conditions was found tobe responsible for changes in shelf-water properties. In additionto seasonal changes, interannual variations in mesozooplanktonabundance and composition were evident, and primarily relatedto the year-to-year increase in water temperature and salinity.The increasing trends in copepod dominance, coupled with theincreasing abundance of species with oceanic affinity (e.g.Euchaeta hebes), were presumably a response to the increasein oceanic features in the shelf area. In the same way, thesubstantial year-to-year increase in the abun dance of specieswith summer-autumn development was attributed to the progressivewarming. Among these species, the case of Temora stylifera isdiscussed specifically because this species was very scarcein 1988, but showed unusual high abundances for the study area,and dominated the summer-autumn assemblage in 1990. We thinkthat T.stylifera can be a key species in monitoring climate-oceanologicalchanges affecting the Bay of Biscay. Results are also discussedin the context of the long climate-oceanological series forthe Bay of Biscay, and its biogeographical status.     

Experimentally induced clutch size enlargements affect reproductive success in the Pied Flycatcher

We performed a food provisioning experiment in a population of Pied Flycatchers Ficedula hypoleuca breeding at high altitude in central Spain to test if food availability before and during laying determines clutch size. Food was provided to one of two pairs with the same date of initiation of nest-building (15 dyads of subsequently reproducing pairs were thus created). Food provisioning began on the day of initiation of nest-building and ended on the day after the last egg was laid. Although laying date was unaffected by the experiment, clutch size in the experimental treatment was significantly larger. This result could indicate that food availability at laying (1) proximately constrained clutch size or (2) that females evaluated future conditions for incubating eggs and feeding nestlings based on food availability at laying. Reproductive success (proportion of eggs that resulted in fledged young) was significantly reduced in the experimental treatment. This effect suggest that supplemented females were tricked by the experiment into laying more eggs than the number of eggs they were able to incubate with success and the number of nestlings they were able to feed, a source of error in clutch size adjustment which could be common in non-experimental situations.     

Oogenesis and gonad maturation in the copepodCalanus finmarchicus and the prediction of egg production from preserved samples

 Oogenesis and gonad development of Calanus finmarchicus during the winter spring transition in a fjord in Northern Norway were studied. A combination of observations on whole animals (macroscopic), taking advantage of their transparency, and histological sections was used to establish a system of gonad maturation stages, to describe the spawning cycle and to estimate egg production rates from preserved samples. During their development, oocytes change their shape and size, the morphology of the nucleus, and the appearance of the ooplasm. Four oocyte development stages were identified and related to a macroscopic system of four gonad developmental stages (GS) that can be applied to whole stained animals. During gonad development, the macroscopic morphological modifications of the gonads are mainly manifested in an increase in number and size of the oocytes and their distribution in diverticula and oviducts. The maturation processes during the spawning cycle (the period between egg depositions) were described from histological sections of females preserved at regular intervals after egg deposition. The macroscopic GS did not vary during the spawning cycle. This GS was a highly significant indicator for females, which spawned within 24 h after collection at 5°C. Another histological indicator for the last third of the spawning cycle is the appearance of distinct chromosomes, which, however, are only seen in histological sections. For the prediction of egg production rate of a Calanus population, in addition to the number of females spawning, knowledge of clutch size and the interval of clutch deposition is required. Clutch size was determined by counting the number of the oocyte stages to be released during the next spawning event in serial sections of the gonads. There was no significant difference in clutches laid by the females during 5 days before preservation. From the observations it seems that GS 4 predicts females that are ready to spawn within the minimum spawning interval characterisitic for a given temperature. If this holds true, it should be possible to predict in situ egg production rates solely from preserved samples. Received: 18 August 1995/Accepted: 6 January 1996     

Pigment ingestion and egg production rates of the calanoid copepod Temora longicornisr. implications for gut pigment loss and omnivorous feeding

We examined the relationship between egg production rate (E)and pigment ingestion rate (I, from gut content corrected for33% loss) for adult female Temora longicornis in Long IslandSound on 47 occasions. Linear regression of E on I [both variablesexpressed in mass of nitrogen (N) female^–1 day^–1]was: E_N = 0.0016 + 0.770 x I_N. The slope, 0.77, is the apparentgross efficiency of egg production, equivalent to the grossgrowth efficiency (GGE) assuming that females partition allnitrogen for growth into egg production. Published work suggeststhat a GGE of 0.37 would be expected for herbivorous copepods.The discrepancy between the expected value of 0.37 and observedvalue of 0.77 could result from unquantified losses of gut pigmentor because T.longicomis ingested a significant amount of nitrogenby feeding as a carnivore. We suggest that if T.longicomis femalesderive all of their nitrogen for growth by feeding on phytoplankton,and if no correction for pigment loss is employed, then thegut pigment method underestimates pigment ingestion by no morethan a factor of two.     

ASPECTS OF THE BIOLOGY OF THE GIANT FORM OF STHENOTEUTHIS OUALANIENSIS CEPHALOPODA: OMMASTREPHIDAE) FROM THE ARABIAN SEA

An investigation was carried out on the recently discovered‘giant’ extra large (XL) form of the squid Sthenoteuthisoualaniensis from the Arabian Sea. The sample consisted of 2males, which have not been previously described, and 13 females.Diet composition, parasite loading, sucker ring dentition, biolumi-nescenceand sexual dimorphism were examined and compared to known parametersof the medium (M) form. Reproductive strategy, potential fecundity,egg size distribution in the ovary and oviducts were examinedin mature XL females. Evidence of multiple spawning in the giantform was also investigated. Overall body shape, bioluminescentstructures and coloration of the giant form were similar tothe M form, though the XL form had a smaller fin angle thanthe M form. The mature female XL form has a dorsal mantle lengthabout twice that of a mature female M form. Adult females ofthe XL form have a dorsal mantle length about twice that ofadult males of the same form. Differences between males andfemales were found in arm sucker ring dentition and parasiteload, suggesting a difference in diet. This could be linkedto size differences between the sexes. A strong correlationbetween ovary mass and mantle length was found (r² = 0.64).Poor correlation was found between mantle length and oviductmass (r² = 0.128) and potential fecundity (r² = 0.07). Potentialfecundity ranged between 2–5 million eggs and the holdingcapacity of the oviducts was approximately 300, 000 eggs. Thiscombined with the presence of spermatangia and the presenceof food in the stomach suggest that the XL form is a multiplespawner. S. oualaniensis appears to have a plastic phenotypeand has adapted to the Arabian Sea conditions by evolving thecapacity to grow to a giant size. (Received 6 November 1996; accepted 15 February 1997)     

Individual quality and food availability determine yolk and egg mass and egg composition in tree swallows Tachycineta bicolor

Variation in egg size and composition can have important consequences for the quality of offspring. We investigated the factors influencing the yolk mass and egg mass of tree swallows breeding in Ithaca, NY. Using a nondestructive technique to estimate yolk mass via standardized digital-candler photographs, we compared yolk size and egg size of tree swallows Tachycineta bicolor in response to variation in food availability and individual quality. Insect availability one to three days prior to laying, but not four to six days, predicted yolk mass, while insect availability two to three days prior to laying predicted total egg mass. This suggests that, while eggs are formed over longer periods, food availability closer to time of laying has the greatest influence on egg size. These results were supported by collected eggs, as yolk rings revealed that tree swallow eggs are formed over 5–6 days. There was an influence of female quality as well, with early laying birds, independent of food availability, laying larger eggs. Eggs laid later in the laying sequence had larger yolks and greater egg mass. Overall, variation in egg quality appears to be due to a combination of environmental conditions, reflected in food resources, individual quality, and allocation tradeoffs during the laying period.     

Egg production of the marine,planktonic copepod,Calanus pacificus Brodsky: Laboratory observations

When acclimated to a continuous, superabundant food supply and constant temperature, Calanus pacificus Brodsky females produce eggs at a weight-specific rate ranging from 0.13 · day^?1 at 8°C to 0.21 · day^?1 at 15°C. Maximum weight-specific egg production rates do not change with seasonal changes in female body size. The relationship between egg production rate and food concentration is hyperbolic, with threshold and critical concentrations that are high relative to other species for which data are available. Food concentration and temperature influence spawning frequency (i.e., the time required for oocytes to mature) much more than the number of eggs in a single spawning event (i.e., clutch size). Clutch size is significantly related to female body size.     

Supplementary feeding increases Common Buzzard Buteo buteo productivity but only in poor‐quality habitat

Temporal heterogeneity in the effects of food supply during the breeding season on the productivity of the Common Buzzard Buteo buteo was investigated in a supplementary feeding experiment. Pairs were fed artificially (1) before egg‐laying, (2) after chicks hatched and (3) continuously throughout the season, and compared with (4) unfed controls. Pairs fed before egg‐laying had marginally larger clutches than those not fed, but lay date, egg volume and weight, brood size and hatching success were unaffected. Territorial quality had far greater effects, with pairs nesting in low‐quality habitats (bog, scrub and semi‐natural grassland) laying later and having lower hatching success, smaller broods and fewer fledglings than those in more productive agricultural landscapes. Supplementary feeding after egg hatching neutralized the negative effect of poor habitat, resulting in fed birds having significantly more fledglings. This study emphasizes the importance of food availability when provisioning chicks in suboptimal habitats and has implications for the success of diversionary feeding in reducing game losses to Buzzards.     

Great tits provided with ad libitum food lay larger eggs when exposed to colder temperatures

The amount of nutrients deposited into a bird egg varies both between and within clutches of the same female. Larger eggs enhance offspring traits, but as a tradeoff, laying large eggs also infers energetic costs to the female. Income breeders usually lay larger eggs later in the season, when temperatures and food availability are higher. Egg size is thus affected by the daily amount of energy available to produce an egg under cold conditions, but it is less well known in how far temperature exerts direct effects on egg size. We show that great tit females Parus major with access to ad libitum food and breeding in climate‐controlled aviaries varied their egg investments. The size of an individual egg was best predicted by mean temperatures one week pre‐laying, with females laying larger, rather than smaller, eggs under colder conditions. Eggs increased in size over the season, but not significantly over the laying sequence. The degree of daily temperature fluctuation did not influence egg size. In addition to a substantial between‐female variation, sisters were more similar to each other than unrelated females, showing that egg size does also reflect heritable intrinsic female properties. Natural variation in egg size is thus not only determined by energy‐limitation, but also due to females allocating more resources to eggs laid in colder environments, thus increasing early survival of the chicks. That the positive correlation between temperature and egg investments that is found in a natural population is reversed under ad libitum food conditions demonstrates that wild great tits tradeoff own condition with survival prospects of their chicks as a function of available food, not ambient temperature.     

Temperature impact on reproduction and development of congener copepod populations

The goal of this study was to relate the temperature response of all developmental stages and reproductive biology of two congener copepod pairs inhabiting different biogeographic regions to their geographic distribution patterns. Survival of adult females and egg production, embryonic development and hatching success of the genera Centropages and Temora from two stations, in the North Sea and the Mediterranean, were studied in laboratory experiments in a temperature range from 2 to 35 °C. Postembryonic development was determined from cohorts raised at temperatures between 10 and 20 °C with surplus food. Tolerance limits and optima of female survival, reproduction and development distinguished the northern species Centropages hamatus and Temora longicornis from the southern T. stylifera, while C. typicus, which is found in both regions, was intermediate. Thus, thermal preferences could in part explain distribution patterns of these species. While C. hamatus and the two Temora species showed distinct temperature ranges, C. typicus was able to tolerate different temperature conditions, resulting in its wide distribution range from the subarctic to the tropics. However, the thermal range of a species did not necessarily correlate with the optimal temperatures in the experiments. Optima of egg production and stage development were surprisingly low in T. stylifera, which has a mere southern distribution.     

Egg dumping by predatory insects

Synovigenic insects resorb oocytes when food is scarce and mature oocytes when food is plentiful. These two antagonistic processes allow an optimal allocation of resources to reproduction and somatic functions. Unlike hymenopteran parasitoids, ladybirds cannot resorb mature oocytes present in the oviducts. Is the energy contained in these oocytes lost or is there a mechanism for recovering it when needed? Females of two species of ladybird beetles Adalia bipunctata (L.) and Adalia decempunctata (L.) that are starved for >24 h lay single infertile eggs, which they immediately eat, and these eggs comprise the mature oocytes in the oviducts at the onset of starvation. This behaviour has some similarities to egg dumping reported in herbivorous insects and is part, in ladybird beetles, of a process to retrieve energy invested in reproduction. Such behaviour may exist in other predatory synovigenic insects species that do not invest in maternal care.     

Variations in clutch and egg sizes in the amphidromous goby <Emphasis Type="Italic">Rhinogobius</Emphasis> sp. CB along a river course and within a spawning season

Rhinogobius sp. CB is a common amphidromous goby in Japan. I investigated egg and clutch sizes along a river course and within a spawning season in the Aizu River, Kii Peninsula, Japan. Clutch size within the same female size did not differ along the course of the river, but decreased from early to later in the spawning season. Egg size within the same female size decreased from early to later in the spawning season and from the upper to the lower reaches. These results mean energy resources allocated to the ovary decreased seasonally and locally. Several factors are presented to explain these variations, such as female somatic conditions, a tradeoff between reproduction and growth, food availability, water temperature, and presence of competitive species. Some factors explaining the seasonal and local variations in egg size, such as water temperature and food availability for offspring, are also presented. Egg size correlated well with female size. The trend in egg size variation with female size seems to be an adaptation for their migrating pattern, as larvae from larger female, which tended to inhabit the upper reaches, would have a greater endurance against starvation during their migration to the sea.     

Effects of a food supplementation experiment on reproductive investment and a post-mating sexually selected trait in magpies Pica pica

Food availability is an important factor affecting breeding success in birds. Food supplementation experiments in birds have in general focused on the effects on reproductive success in terms of female investment (laying date, clutch size, egg size), however, it is also known that the estimation of mate quality based on sexually selected signals influences female reproductive investment. In the particular case of magpies, females use nest size, a post-mating sexually selected signal, to assess male's likelihood to invest in reproduction, and accordingly adjust reproductive investment (clutch size). Then, the possible effects of food supplementation on female reproductive investment could be mediated by other variables related to parental quality, such as nest size in magpies. In the present study, we explore if higher food availability in a magpie territory affected both male sexually selected traits (i.e. nest size) and female reproductive investment (laying date, egg size, clutch size). We performed a food supplementation experiment in which we experimentally increased food availability in several magpie territories, keeping others as controls. In food-supplemented territories, males built significantly larger nests and females significantly increased egg size by 4.1% compared to control females. Results suggest that the continuous provisioning of protein rich food allowed magpie females to increase egg size. However, laying date and clutch size did not differ between control and food-supplemented magpie pairs. Food availability also affected the relationship between female reproductive investment and nest size. In control territories, females decreased their egg size in response to a larger nest, whereas a tendency for the opposite relationship was revealed in food-supplemented territories. We discuss the possibility that magpie females adopt different strategies for reproductive investment according to food availability.     

Inhibition of the onset of vitellogenesis by abnormal retention of mature eggs in the stick insect,Carausius morosus

Mature eggs stored in the lateral oviducts are unusual in Carausius morosus. When egg laying is experimentally prevented, ovarian production is inhibited by the accumulation of mature eggs within the lateral oviducts. Such storage affects only the very small terminal oocytes and not the vitellogenic follicles, which attain to term and are ovulated. The fact that unilateral retention of eggs affects only the corresponding ovary strongly suggests that the mature eggs themselves inhibit ovarian production in the stick insect without the involvement of an endocrine organ. It is likely that this mode of action, different from that suggested for the housefly or Rhodnius prolixus, is correlated with the independence of ovarian development from the corpus allatum hormone in the stick insect.