Egg size and reproductive adaptations among Arctic deep-sea copepods (Calanoida,<Emphasis Type="Italic"> Paraeuchaeta</Emphasis>)

Reproductive strategies of the four congeneric and sympatric calanoid copepods Paraeuchaeta glacialis, P. norvegica, P. barbata, and P. polaris were studied in the Arctic Greenland Sea. Females of all species produce egg sacs and carry their brood attached to the genital opening until the offspring hatch. However, egg size and lipid content as well as clutch size and the fraction of females carrying egg masses show characteristic differences among the four species. P. glacialis and P. norvegica produce large numbers (37 to more than 50) of relatively small eggs, whereas P. barbata and P. polaris rely on small numbers (10 to 19 and 4 to 6, respectively) of large eggs with a high energy content. There is no correlation between female body size and egg size or clutch size, respectively. Females of the smallest species, P. polaris, produce relatively large eggs and show the highest energetic investment per egg. In contrast, energetic investment per clutch is highest in P. glacialis. Reproductive adaptations appear to be strongly related to the depth range inhabited by the respective species. In the central Greenland Sea P. glacialis and P. norvegica occur in the epipelagial and upper mesopelagial, whereas the other two species inhabit lower mesopelagic to bathypelagic depths. Thus, egg size increases with depth of occurrence, whereas clutch size is inversely correlated with depth. This observation leads to the hypothesis that the evolution of large eggs in deep-sea copepods may enable hatchlings to rely on a lecithotrophic development and thus represents a successful adaptation to cope with the limited food supply at great depths, whereas high offspring numbers in epipelagic species compensate for higher predation risks in the euphotic zone.Communicated by H.-D. Franke     

Seasonal variations in copepod body length: a comparison between different species in the Lagoon of Venice

The seasonal variation in length of the most abundant copepod species in the lagoon of Venice were compared and the relative influence of temperature, chlorophyll concentration and salinity examined. Temperature seems to be the primary factor influencing copepod body length in the lagoon of Venice, but the different species vary markedly in their response. Calanoid copepods (Acartia clausi, A. tonsa, Paracalanus parvus and Centropages ponticus) showed a more definite trend of size variation with temperature than the cyclopoids (Oithona nana and O. similis) and the harpacticoid Euterpina acutifrons. The size of the poecilostomatoids Oncaea media and O. subtilis was not affected by temperature, and was almost constant over time. Differences in the metabolic rates, longevity and specialization of calanoid, cyclopoid and harpacticoid copepods could explain their different responses to temperature.     

The seasonal dynamics of Copidodiaptomus numidicus (Gurney, 1909) and Thermocyclops dybowskii (Lande, 1890) in Castelo-do-Bode Reservoir

We investigated the population dynamics of Copidodiaptomus numidicus (Copepoda: Calanoida) and Thermocyclops dybowskii (Copepoda: Cyclopoida) in Castelo-do-Bode Reservoir (Portugal). Both species in the reservoir were regulated mainly by food availability during the summers of 1993 and 1994. C. numidicus was always more abundant than T. dybowskii in the reservoir. C. numidicus produced two generations during the sampling period of both years whereas T. dybowskii produced two and three generations in 1993 and 1994, respectively. Because of high temperatures and low rainfall in 1994, the reservoir was more eutrophic than in 1993. Higher clutch size and a higher percentage of ovigerous females suggest that both species were benefiting from better feeding conditions in 1994. Despite this, population growth was reduced in 1994 relatively to 1993. Vertebrate predation and predation by copepods seem to have been minor factors in explaining this decline. However, the presence of Mesostoma sp. in the reservoir may have contributed to the reduction of both copepod populations during specific periods in 1994 although the calanoid was more affected than the cyclopoid by this predation.     

Reproductive and life cycle strategies in egg-carrying cyclopoid and free-spawning calanoid copepods

Egg-carrying cyclopoid copepods have lower fecundity and feedingrates, and longer egg hatching times, than free-spawning calanoidcopepods. Simple demographic considerations suggest . that thelower feeding and fecundity of egg-carrying cyclopoids are adaptationsto the potentially elevated mortality of ovigerous females,while the shorter egg hatching time and higher feeding and fecundityrates found in free-spawning calanoid copepods represent adaptationsto the very high mortality rates experienced by suspended eggs. ¹Present address: Instituto Nacional de Investigatin y DesarrolloPesquero, INIDEP, CC 175, Play a Grande, Mar del Plata, Argentina     

Body size regulation in copepod crustaceans

Body size affects survival probabilities, reproductive output and individual fitness in many organisms. In freshwater zooplankton, traits ranging from demographic rates to community composition depend on body size, and predation is most often identified as the selection pressure determining body size. We examined the extent to which stage-specific growth trajectories and body sizes of copepod crustaceans are constrained, independent of selection. We used exuviae shed at each molt to quantify the relationship between size at molting and growth during the subsequent instar for two common, herbivorous calanoid copepods, Boeckella triarticulata and Diaptomus leptopus. Individuals of both species were raised under diets of different food quality or quantity, and at different temperatures. Size at molting varied little among individuals of both species, as a consequence of a persistent negative relationship between size at molting and subsequent (absolute) growth increment. Individuals that were small when they molted grew more during the subsequent instar than individuals that were large. This relationship was statistically significant for nearly all instars of both species raised in different food or temperature conditions, and not affected by food quality, food quantity or temperature. Our results indicate that body size is constrained or regulated over much of the copepod life cycle, independent of the effects of environmental conditions (food, temperature, predation). Received: 26 December 1998 / Accepted: 13 September 1999     

Reproductive allocation and reproductive ecology of seven species of Diptera

1. Variation in resource allocation to egg size and number was investigated in seven sympatric species of Piophilidae that oviposit on carcasses or discarded cervid antlers: Liopiophila varipes (Meigen), Prochyliza xanthostoma Walker, Protopiophila latipes (Meigen), Protopiophila litigata Bonduriansky, Stearibia nigriceps (Meigen), and two unidentified species of Parapiophila McAlpine. 2. Following optimal reproductive allocation theory, relatively larger, fewer eggs were expected in (1) species that oviposit on antlers, where larvae probably experience lower risk of predation and greater competition than larvae in carcasses, and (2) species with aggressive males and male-biased sex ratios on the oviposition substrate, where risk of injury during oviposition may have favoured females laying fewer eggs. 3. Variation in reproductive allocation strategies could not be explained by known differences in larval or adult environment, but congeneric species clustered by reproductive allocation patterns. The Parapiophila species produced larger, fewer eggs than the other species, and egg number increased slowly with body size. The Protopiophila species did not deviate from expected egg sizes and numbers, and egg number increased steeply with body size. 4. An interspecific egg size–egg number trade-off resulted in a tight linear scaling of ovary volume to body size, suggesting common physiological constraints on relative ovary mass. 5. Within each species, egg size was nearly constant whereas egg number increased with female body size, suggesting species-specific stabilising selection on egg size.     

Some evidence for different ecological pressures that constrain male and female body size

The dwarf morph of the Lake Tanganyika cichlid Telmatochromis temporalis uses empty snail shells as shelters and breeding sites in shell beds, in which many empty shells exist. Here, we assessed selection forces regulating body size in this fish. Field observations showed that large males tended to have a greater number of females in their territories, suggesting that sexual selection favours large males. Nonetheless, a transplant experiment suggested that male body size was limited by the ability to hide in empty shells from large piscivorous fish. In females, the number of ovarian eggs increased with body size, suggesting that fecundity selection favours large females. However, females are smaller than males. Females spawned eggs close to the apices inside the shells. The small space there would decrease the risk of egg predation by egg predators, and small body size of females may be a result of adaptation to spawn eggs in the small, safe spaces. This study provides support for the idea that male and female body sizes have been limited by different ecological pressures (predation on adult fish in males, predation on eggs in females), which has not been reported previously in any animal.     

Calanoid copepods from South Georgia,with special reference to size dimorphism within the genus Pseudoboeckella

Summary Three species of calanoid copepods, Boeckella michaelseni, Parabroteas sarsii and Pseudoboeckella poppei were recorded from 6 freshwater localities in the Husvik area, South Georgia. Of the latter species two distinct size morphs occurred, with no overlap in size even in closely situated populations. The large morph was recorded in lakes, the small was found in ponds. The small morph did not coexist with the large predatory P. sarsii, and we suggest predation pressure from this species as the major cause for the observed distribution of these morphs. The pronounced size segregation as well as small morphological dissimilarities suggest that these morphs are reproductively isolated. While the large morphotype corresponds to that of P. poppei, the taxonomic affinities of the small morph are uncertain.     

Freshwater Copepods and Rotifers: Predators and their Prey

Three main groups of planktonic animals inhabit the limnetic zone of inland waters and compete for common food resources: rotifers, cladocerans and copepods. In addition to competition, their mutual relationships are strongly influenced by the variable, herbivorous and carnivorous feeding modes of the copepods. Most copepod species, at least in their later developmental stages, are efficient predators. They exhibit various hunting and feeding techniques, which enable them to prey on a wide range of planktonic animals from protozoans to small cladocerans. The rotifers are often the most preferred prey. The scope of this paper is limited to predation of freshwater copepods on rotifer prey. Both cyclopoid and calanoid copepods (genera Cyclops, Acanthocyclops, Mesocyclops, Diacyclops, Tropocyclops, Diaptomus, Eudiaptomus, Boeckella, Epischura and others) as predators and several rotifer species (genera Synchaeta, Polyarthra, Filinia, Conochilus, Conochiloides, Brachionus, Keratella, Asplanchna and others) as prey are reported in various studies on the feeding relationships in limnetic communities. Generally, soft-bodied species are more vulnerable to predation than species possessing spines or external structures or loricate species. However, not only morphological but also behavioural characteristics, e.g., movements and escape reactions, and temporal and spatial distribution of rotifer species are important in regulating the impact of copepod predation. The reported predation rates are high enough to produce top-down control and often achieve or even exceed the reproductive rates of the rotifer populations. These findings are discussed and related to the differences between the life history strategies of limnetic rotifer species, with their ability to quickly utilize seasonally changing food resources, and adjust to the more complicated life strategies of copepods.     

Short communication. On the free-spawning reproductive behaviour of the copepod Clausocalanus lividus (Frost & Fleminger 1968)

Unlike other species of Clausocalanus, which are egg carriers, Clausocalanus lividus lays free eggs into the water. As far as we know, this is the first observation of such a difference in reproductive strategy within a genus of calanoid copepods.      

Predation by juvenile roach on the calanoid copepod Eudiaptomus gracilis and the cyclopoid copepod Cyclops vicinus: a laboratory investigation with mixed and single prey

We tested the outcome of predation by juvenile roach on the calanoid copepod Eudiaptomus gracilis and the similar sized cyclopoid copepod Cyclops vicinus in laboratory experiments with mixed and single-prey. When ovigerous females and adult non-ovigerous females of the calanoid copepod and the cyclopoid copepod were offered in equal numbers to the fish in the mixed-prey experiments, the cyclopoid copepods were significantly more heavily preyed upon than the calanoids. Between 14 and 16 females of the cyclopoid copepod and only between 2 and 4 calanoids had been consumed after the observation period of two hours. The single-prey experiments revealed that the conspicuousness of the calanoid vs the cyclopoid depended on sex and gravidity. Ovigerous females of the cyclopoid copepod were slightly earlier detected by the fish than ovigerous females of the calanoid, probably a result of the highly visible egg-sacs carried by the cyclopoid females. Females without eggs and males of the cyclopoid copepod were recognized later by the predator than females without eggs or males of the calanoid, probably a result of the different behaviours of the different copepod taxa. Cyclopoids frequently congregated near the aquarium bottom while calanoids were closer to the water surface where they were better visible. The calanoid copepod could better escape the fish's attacks than the cyclopoid copepod. Egg-bearing females of both the calanoid and the cyclopoid copepod could significantly better escape than non-ovigerous females or males. Probably ovigerous females react very early to water disturbances caused by the predator. Activity measurements showed that the cyclopoid copepod displayed 2 to 4 times more hops per time unit than the calanoid. Probably the high number of jerky movements displayed by the cyclopoid attracted attention of the predator and contributed to its greater vulnerability.     

Responses of cyanobacteria to herbivorous zooplankton across predator regimes: who mows the bloom?

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Zooplankton of Fishless Ponds of Northern Patagonia: Insights into Predation Effects of Mesostoma ehrenbergii

The turbellarian predator Mesostoma ehrenbergii, a common inhabitant of fishless ponds of northern Patagonia, can consume prey larger than 1 mm. Because the feeding strategy of M. ehrenbergii includes mucus trapping and external digestion, this predator may exploit a broad range of prey sizes. We hypothesize that M. ehrenbergii could exert a strong effect on zooplankton composition and body size spectra in Patagonian fishless ponds. We investigated this hypothesis by analyzing the crustacean zooplankton composition and size spectra in five fishless ponds of northern Patagonia, and we carried out experiments to assess predation rates of M. ehrenbergii on potential prey species from 0.8 mm to 6 mm. These ponds were colonized by macrophytes, which favored habitat heterogeneity, especially in the smaller ponds that had higher species richness. The surveyed ponds showed distinctive crustacean zooplankton assemblages and sizes, but all were dominated by calanoid copepods of the genus Boeckella. Our results indicated that M. ehrenbergii consumed the whole size range of offered prey, from ∼0.8 mm (Ceriodaphnia dubia) to ∼6 mm (Parabroteas sarsi). Predation rates were higher for intermediate‐bodied copepods (∼1.5 mm) and C. dubia (0.8 mm), but we did not find conclusive evidence that variations in size spectra of crustacean zooplankton are a result of M. ehrenbergii predation. We suggest that an interaction between prey body size and its evasion tactic might be important to determine the true effect Mesostoma on zooplankters. The structural complexity created by macrophytes in Patagonian fishless ponds may also help account for the lack of a strong predation effect of Mesostoma in the field survey. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Coexistence of two similar copepod species, <Emphasis Type="Italic">Eudiaptomus gracilis</Emphasis> and <Emphasis Type="Italic">E. graciloides</Emphasis>: the role of differential predator avoidance

Habitat choice in relation to environmental factors of two coexisting calanoid copepod species, Eudiaptomus gracilis and E. graciloides, was studied in a mesotrophic lake and in large indoor mesocosms. Both species and sexes showed pronounced diel vertical migration (DVM) in the field. In 12 m deep mesocosms with free ranging fish DVM was observed and species increased day depth over time. No changes were observed in copepod day depth over time in experiments with fish kairomone. It is hypothesized that fish kairomone acts as an early warning system to copepods which respond by moving deeper, but only as far as the thermocline. For full DVM, a nearby mechanical stimulus is necessary. Thus, as fish go deeper to feed, copepods retreat. The response of copepods to fish predation, in the presence of low and high numbers of Daphnia,shows that copepods effectively use Daphniaas living shields to avoid predation. The two species adopt different vertical migration strategies depending on whether there are high or low numbers of Daphniapresent. A dominant feature of mesocosm experiments was the night time aggregating (lekking) of E. gracilis males at the surface. When the spring and autumn percentages of risk takers in the epilimnion were compared, E. gracilis, particularly males, suffered the greatest cost.     

Experimental evidence for predation risk sensitive oviposition by a mosquito, Culiseta longiareolata

1. Females should choose to oviposit in habitats where risk of predation and competition are low. The ovipositional responses of a mosquito, Culiseta longiareolata, to a predator and to species sharing the same trophic level as this mosquito (controphic species) were assessed experimentally in outdoor artificial pools. 2. The predator, larval Anax imperator, which strongly reduced larval C. longiareolata survival, resulted in a 52% reduction of C. longiareolata egg rafts. The controphic species (primarily Daphnia magna), which had a small but statistically significant negative effect on the survival of C. longiareolata larvae, did not have a statistically significant influence on the number of egg rafts. 3. Laboratory trials indicated that only a small fraction of the reduced number of egg rafts seen in predator pools may be due to consumption of the egg rafts by A. imperator. 4. The experimental evidence indicates that the reduced number of C. longiareolata egg rafts found in the presence of A. imperator is due largely to oviposition habitat selection, i.e. C. longiareolata females choose pools with low risk of predation for their offspring.     

Effect of adult females' predation risk on oviposition site selection in a water strider

In many egg‐laying species, females avoid ovipositing at sites where the predation risk is high. Previous studies have mainly focused on the risk for offspring. The effect of predation risk for the females has been considered in some taxa in which parents spend much time at an oviposition site for parental care or mating (e.g., birds, amphibians). In species in which females do not perform activities other than oviposition at sites, the effect of predation risk for females on oviposition site selection has been rarely investigated. We examined whether the predation risk for ovipositing females affects the decision on oviposition in the water strider Aquarius paludum insularis (Motschulsky) (Heteroptera: Gerridae). Adult A. paludum are preyed upon by a backswimmer, Notonecta triguttata (Motschulsky) (Heteroptera: Notonectidae), but N. triguttata does not prey upon eggs or early instars of A. paludum. We allowed female A. paludum to oviposit under one of three conditions: in presence of N. triguttata, in presence of its chemicals, and in absence of the predator or its cues (control). Female A. paludum less frequently oviposited in presence of N. triguttata than in its absence. Oviposition frequency did not differ between females in presence of chemicals of N. triguttata vs. those in the control. Female A. paludum recognized the predation risk upon themselves from the presence of N. triguttata and avoided ovipositing. This study is the first to directly show that the predation risk upon ovipositing females changes oviposition site selection in species in which the time spent at an oviposition site is short.     

Egg production rates of eight calanoid copepod species during summer 1997 off Newport, Oregon, USA

Measurements of hydrography, water transparency, chlorophyll (Chl) a and egg production rates (EPRs) by females of Calanus marshallae Frost, Calanus pacificus Brodsky, Eucalanus californicus Johnson, Epilabidocera longipedata Sato, Pseudocalanus mimus Frost, Centropages abdominalis Sato, Acartia longiremis Lilljerborg and Paracalanus parvus (Claus) were estimated at weekly intervals between 17 July and 2 September 1997. Production of eggs was determined in 24 h incubations to examine the effects of environmental variability on EPR, to detect the possibility of food limitation of EPR, and to evaluate the hypothesis that growth rates of females are size dependent. During the study, an anomalous downwelling event occurred, possibly in response to the 1997 El Niño, which allowed us to determine how El Niño events affect EPRs of coastal copepods. The larger copepods Calanus marshallae, Calanus pacificus and Centropages abdominalis showed the highest egg production and specific growth rates during the period of active upwelling (18 July-13 August, water temperatures 8-13C, Chl a concentration 4.7-16.2 l^-1 and water transparency 3-5 m). After 27 August, the 1997-98 El Niño arrived off Oregon, creating a downwelling situation. Upwelling winds ceased, the thermocline intensified, temperature and transparency increased (to >18°C and 16 m), and Chl a declined to <2 g l^-1. Densities of the common coastal species declined greatly as well. Paracalanus parvus became the dominant species, and Eucalanus californicus, Epilabidocera longipedata and Corycaeus anglicus became common in our samples. EPRs for the larger boreal copepods (Calanus and Centropages) declined greatly during El Niño;; the smaller copepods, Pseudocalanus mimus, A.longiremis and Paracalanus parvus, showed low but relatively constant egg production and specific growth rates during both upwelling and downwelling events. Over the entire study period, only three species produced eggs at its maximum rate (24-28 eggs female^-1 day^-1), Pseudocalanus mimus which averaged 4 eggs day^-1. All other species had EPRs that were two or five times below their maximum EPR. Thus, EPRs were not related to body size, contrary to our expectations. Hatching success was variable among species. Values as low as 20-40% were found for all species at least once during the study period, suggesting that occasionally a substantial portion of egg production may not be viable.      

Water renewal in reservoirs,and its relation to body size and egg numbers in copepods

Summary In a study of four species of centropagid copepods, particularly Boeckella minuta Sars, occurring in south-east Queensland, Australia, body length was found to correlate with the rate of water renewal in the reservoir. Ravera & Tonolli's explanation for interpopulational differences in lengths of copepods has thus been confirmed. There was also a positive correlation between body length and egg number in ovigerous females.Sex ratio (females/males) increased with the rate of water renewal, this providing for relatively more females to be present when population numbers were smaller.

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Zusammenfassung In einer Untersuchung von vier Spezies zentropagischer Kopepoden, besonders Boeckella minuta Sars, die sich bei Südostqueens-land, Australien, finden, wurde entdeckt, daß die Körperlänge mit dem Wassererneuerungsgrad in dem Reservoir in Beziehung steht. Ravera & Tonolli's Erklärung für die Verschiedenheiten in Körperlängen von Kopepoden zwischen der Bevölkerung ist, auf diese Weise, bestätigt worden. Es gab auch eine positive Wechselbeziehung zwischen der Körperlänge und der Anzahl Eier in eiertragenden Weibchen.Das Geschlechtsverhältnis (Weibchen/Männchen) wurde grösser mit dem Wassererneuerungsgrad, wodurch etwas mehr Weibchen anwesend Bind, wenn die Bevölkerungszahl kleiner war.

     

Inter‐ and intraspecific variation in body‐ and genome size in calanoid copepods from temperate and arctic waters

The tendency of ectotherms to get larger in the cold (Bergmann clines) has potentially great implications for individual performance and food web dynamics. The mechanistic drivers of this trend are not well understood, however. One fundamental question is to which extent variation in body size is attributed to variation in cell size, which again is related to genome size. In this study, we analyzed body and genome size in four species of marine calanoid copepods, Calanus finmarchicus, C. glacialis, C. hyperboreus and Paraeuchaeta norvegica, with populations from both south Norwegian fjords and the High Arctic. The Calanus species showed typical interspecific Bergmann clines, and we assessed whether they also displayed similar intraspecific variations—and if correlation between genome size and body size differed between species. There were considerable inter‐ as well as intraspecific variations in body size and genome size, with the northernmost populations having the largest values of both variables within each species. Positive intraspecific relationships suggest a functional link between body and genome size, although its adaptiveness has not been settled. Impact of additional drivers like phylogeny or specific adaptations, however, was suggested by striking divergences in body size – genome size ratios among species. Thus, C. glacialis and C. hyperboreus, had fairly similar genome size despite very different body size, while P. norvegica, of similar body size as C. hyperboreus, had the largest genome sizes ever recorded from copepods. The inter‐ and intraspecific latitudinal body size clines suggest that climate change may have major impact on body size composition of keystone species in marine planktonic food webs.     

Feeding patterns of post-larval and juvenile notothenioids in the southern Weddell Sea (Antarctica)

Summary The food of 163 juvenile specimens of 13 species of notothenioid fishes collected in the southern Weddell Sea (Antarctica) was analyzed. Investigated fish size range was 3–13 cm SL. Principal food items were calanoid copepods Metridia gerlachei, Calanoides acutus, and Calanus propinquus; all developmental stages of Euphausia crystallorophias, and post-larval nototheniid fish Pleuragramma antarcticum. Diet of juvenile channichthyids is limited to few species of euphausiids and fish in the size > 10 mm, but does not include significant numbers of copepods. Pelagic stages of nototheniids feed on copepods and/or larval euphausiids smaller than 10 mm. At similar size, nototheniids and bathydraconids take smaller prey items than channichthyids.