Temperature-mediated changes in zooplankton body size: large scale temporal and spatial analysis

Climate warming has been linked with changes in the spatiotemporal distribution of species and the body size structure of ecological communities. Body size is a master trait underlying a host of physiological, ecological and evolutionary processes. However, the relative importance of environmental drivers and life history strategies on community body size structure across large spatial and temporal scales is poorly understood. We used detailed data of 83 copepod species, monitored over a 57-year period across the North Atlantic, to test how sea surface temperature, thermal and day length seasonality relate to observed latitudinal-size clines of the zooplankton community. The genus Calanus includes dominant taxa in the North Atlantic that overwinter at ocean depth. Thus we compared the copepod community size structure with and without Calanus species, to partition the influence of this life history strategy. The mean community body size of copepods was positively associated with latitude and negatively associated with temperature, suggesting that these communities follow Bergmann's rule. Including Calanus species strengthens these relationships due to their larger than average body sizes and high seasonal abundances, indicating that the latitudinal-size cline may be adaptive. We suggest that seasonal food availability prevents high abundance of smaller-sized copepods at higher latitudes, and that active vertical migration of dominant pelagic species can increase their survival rate over the resource-poor seasons. These findings improve our understanding of the impacts that climate warming has on ecological communities, with potential consequences for trophic interactions and biogeochemical processes that are well known to be size dependent.     

Annual changes in the population size of the salmon louse Lepeophtheirus salmonis (Copepoda: Caligidae) on high-seas Pacific Salmon (Oncorhynchus spp.), and relationship to host abundance

The population size of the salmon louse, Lepeophtheirus salmonis, was monitored annually in the summers of 1991–1997 by examining six species of Pacific salmon (Oncorhynchus spp.) caught by surface long-lines in oceanic offshore waters of the North Pacific Ocean and Bering Sea. The annual copepod population size on all salmonids caught was estimated by combining the calculated number of copepods carrying on each salmonid species. The copepod population fluctuated markedly from year to year, which resulted largely from marked annual changes in abundance of pink salmon (O. gorbuscha). Since pink salmon were most frequently and heavily infected and since their abundance changed every year, the copepod population was high in the years when this salmonid species was abundant, but low when it was rare. On the contrary, chum salmon (O. keta) did not show high prevalence and intensity of infection, but the annual abundance of this host species was consistently high, i.e. chum salmon carried many copepods every year. Copepods on other salmonid species (sockeye salmon O. nerka, coho salmon O. kisutch, chinook salmon O. tshawytscha, and steelhead trout O. mykiss) constantly formed a small percentage of the total copepod population. Both chum and pink salmon are the most important hosts in terms of their substantial contribution to support the copepod population, but the importance as hosts of each species is definitely different between the species. Chum salmon is a stable important host supporting the copepod population at a relatively high level every year, while the number of copepods on pink salmon annually exhibits marked fluctuations, and this salmonid species is regarded as an unstable important host.     

Antennule shape and body size of <Emphasis Type="Italic">Bosmina</Emphasis>: key factors determining its vulnerability to predacious Copepoda

Cyclopoid copepods are common in lakes and ponds, and they have a significant predation impact on the communities of the small zooplankton species. To reduce the predation risk, some cladoceran zooplankters develop protuberant (defensive) morphologies in the presence of the copepods. In the case of the small cladoceran Bosmina, they elongate their appendages (antennule and mucrone) and change the antennule morphotype. However, information about the effectiveness of these defensive devices against copepod predation is still insufficient. In our study, to find the compositive effects of these appendages on the vulnerability of Bosmina, we exposed two bosminid species (B. longirostris and B. fatalis) of different body sizes and with appendages of different lengths and shapes to copepod (Mesocyclops) predation. The experiment revealed that the shape of the antennule is a main factor determining the bosminid’s vulnerability to copepod predation and indicated that the protection of the opened ventral carapace must be a key strategy by which Bosmina avoids copepod predation.     

Prey handling time and ingestion probability for Mesocyclops sp. predation on small cladoceran species Bosmina longirostris, Bosminopsis deitersi, and Scapholeberis mucronata

We compared the vulnerability of small cladoceran species (Bosminopsis deitersi, Bosmina longirostris, and Scapholeberis mucronata) to predation by Mesocyclops sp. in the laboratory based on prey handling time and ingestion probability. We also estimated the effects of ontogenetic changes on handling time by testing prey of various body sizes. All tested prey species showed an increasing pattern of handling time along with growth (increase of body size). For juveniles smaller than 0.3mm, the prey handling time was similar for B. deitersi and B. longirostris, but markedly longer for S. mucronata. Mesocyclops sp. rejected large (>0.35mm) B. longirostris and S. mucronata soon after capture and consequently showed a low probability of successful ingestion, whereas most large individuals of B. deitersi were consumed. The shorter handling time and higher ingestion probability of small juveniles indicate that the copepods ingest small juveniles more efficiently than larger individuals. Thus, it seems that the vulnerability of small cladocerans to copepod predation differs ontogenetically, with the greatest vulnerability being during the juvenile period. The ontogenetic reduction in the vulnerability of B. longirostris and S. mucronata was shown in the rapidly reduced ingestion probability as well as by the increase in prey handling time for large individuals. The results show that the vulnerability to predation by copepods of B. longirostris and S. mucronata reduces more quickly with increasing size than does that of B. deitersi, and it seems that the former two species are more resistant to copepod predation.     

Temperature- and predator-induced phenotypic plasticity in Bosmina cornuta and B. pellucida (Crustacea: Cladocera)

SUMMARY 1. Clones of Bosmina cornuta and B. pellucida (B. longirostris species complex) were derived from samples collected from Scheuermühlenteich and Lake Windsborn(westernGermany). Experimental temperature change (to 10 °C and 20 °C) and exposure to Acanthocyclops vernalis copepods (12 L^?1) significantly altered external morphology in laboratory cultures of the two species. Morphological traits were derived from eight log₁₀‐transformed and standardised morphometric distances by factorial analysis: factor 1 represented body size, factor 2, size of appendages and factor 3, the head size. 2. Acclimation of clones to cold water (10 °C, >14 days) led to an increase in body, antennule and mucrone size in B. cornuta and B. pellucida. Moreover, at 10 °C, B. cornuta cultures usually collapsed within a few weeks. Compared to the trials at 10 °C, acclimation to 20 °C (the two species) and to 15 °C (B. pellucida only) left the size of body appendages unchanged. Individuals were unequivocally assigned to each species by discriminant functions. Conspecific individuals that were acclimated to different temperatures between 10 and 20 °C also differed in external morphology, but discriminant analysis yielded misclassification rates of 5.3–23.3%. 3. Morphological response to the presence of copepod predators was weaker than that caused by temperature change. Long‐term exposure of clones to copepod predators induced a significant increase in size of appendages in the two species but left body size unaffected. Again, species identification by discriminant functions could be made without any error, whereas conspecific controls and experimentals were misclassified at rates between 19.4 and 29.5%. 4. It is suggested that temperature is the main proximal cue for Bosmina cyclomorphosis. The distinct response to temperature of B. pellucida and B. cornuta may also account for seasonal differences in abundance observed in field.     

Is diatom size selection by harpacticoid copepods related to grazer body size?

Copepods are known as important consumers of primary production and are eaten by larger animals. They therefore form a main link to higher trophic levels. While feeding pathways and specificity of planktonic copepods have been well studied, the selectivity of the benthic harpacticoid copepods is far less documented. A better knowledge of the functional ecology of harpacticoids as important grazers on primary producers may have consequences for the re-evaluation of basic energy flow in benthic ecosystems.We tested whether size selectivity for diatoms exists in harpacticoid copepods. We hypothesized that size selectivity of harpacticoid copepod species is strongly related to body size. Because of morphological constraints, we expected smaller copepods to prefer smaller diatoms while larger copepods should be able to consume both small and large diatoms. We tested this hypothesis in four harpacticoid copepod species of varied body size: Tigriopus brevicornis, Harpacticus obscurus, Amphiascus minutus and Paramphiascella fulvofasciata. As food source we used two ¹³C labelled strains of the benthic diatom Seminavis robusta with a four-fold difference in cell biovolume.Three out of four harpacticoid species showed size selectivity: H. obscurus and A. minutus preferred the larger Seminavis cells, while P. fulvofasciata selected the smaller Seminavis cells. Based on monoclonal treatments, there was no clear preference found for T. brevicornis although there was a small preference for large cells in the mixed treatments. Except for P. fulvofasciata, all species showed a lower uptake when offered the mixed diet (both small and large cells). Although most species showed a size selectivity, our results suggest that this selectivity was not related to their body size. However, the only species that ate significantly more of small diatoms was characterised by comparatively small mouthparts in relation to its body size.     

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.     

Structure and function of the head in flabelligerid polychaetes

The functional anatomy of the head of Flabelliderma commensalis is described and compared to other flabelligerid polychaetes. Prostomial parts include the dorsal lip, the palps, two pairs of nuchal organs, four eyes and the prostomial lobe and ridge. The eyes are inverse pigment cup types with the medial portions of the sensory cells expanded to form a clear lens-like body. Peristomial parts include the median and ventral lips, the branchial membrane and the branchiae. The derivation of the nephridiopore is unknown. The spiraled branchiae of Coppingeria and the gill books of Diplocirrus are newly described variations in branchial structure. The head is retractable in some species and the anterior setigers are modified to form a protective setal cage. Two methods are employed for feeding: one for host fecal pellets and the other for detrital materials. Chemoreception, respiration, feeding and cleaning rely on a complex pattern of ciliary currents.     

Relative Sensitivity Distribution of Freshwater Planktonic Crustaceans to Trace Metals

Cladocerans and copepods are common inhabitants of ponds and lakes, but assessments of the toxic effects of chemicals on aquatic organisms are often based mainly on toxicity data for cladocerans of the genus Daphnia. The paucity of toxicity data for copepods and other species of cladocerans raised the question whether toxicity data derived primarily from daphniids can be extrapolated to protect other planktonic organisms. In this study, we address this question by systematically comparing between the sensitivities of cladocerans and copepods to the metals Cd, Cu, Pb, Ni, and Zn using the species sensitivity distributions. Our results indicate that cladocerans are consistently more sensitive than copepods to Cd and Cu. Most importantly, sensitivities of D. magna, D. pulex, and Ceriodaphnia dubia, the three most commonly tested cladocerans, to these metals are higher than those of most copepod species and fall within the range of those exhibited by other cladocerans. C. dubia, a cladoceran with relatively small body size, is more sensitive to all metals than most other cladoceran species. Our results imply that water quality criteria of trace metals derived from datasets dominated by Daphnia are likely also protective to copepods and most other cladocerans.     

Seasonal variations in copepod size: effects of temperature, food abundance, and vertebrate predation

Seasonal dry weights of female and male Cyclops bicuspidatusthomasi, Diaptomus ashlandi and Diaptomus minutus were studiedin southeastern Lake Michigan during 1975–1981. Smallestanimals occurred during summer and early fall, and largest animalsin winter and spring, a pattern reported for other copepods.The range of weights for the species and sexes decreased fromwinter to summer, and converged in summer to a value of approximately2 µg/animal. Surface water temperature and abundancesof young-of-the-year (YOY) fish were inversely correlated withweights of males and females of the three copepod species. Standingstocks of important phytoplankton groups, especially pennatediatoms and flagellates, were positively correlated with copepodweight. Seasonal change of copepod body size appears to be morethan simply a function of temperature; seasonal predation byYOY fish may be a factor influencing sizes of adult copepods.     

The relationship between larval size and fitness in the tapeworm Schistocephalus solidus: bigger is better?

In organisms with complex life cycles, fitness often increases with body size at the transition from larva to adult. The translation of larval size into fitness, however, can depend on the source of size variation, with size, per se, not always increasing adult success. In parasitic worms, many factors influence larval growth, but little is known about the consequences of this growth variation. We examined how the size of the tapeworm Schistocephalus solidus in its copepod first intermediate host affects infection success and growth in the stickleback second host. Moreover, we assessed whether the conspicuous growth variation caused by copepod size is fitness‐relevant. Using larvae of the same age, we found that larger worms had a substantially higher infection probability and they tended to still be slightly larger after several months of growth in fish. However, big larvae from bigger copepods did not have higher fitness, suggesting that being large relative to the host, but not necessarily large in general, is important. These findings clarify some aspects of the life history strategy of S. solidus (e.g. why there is a flat ontogenetic reaction norm across copepod stages), but also raise questions (e.g. why growth costs have been hard to document). More generally, our results indicate that larval size can correlate with fitness in helminths, but that not all size variation is predictive of success in the next host.     

Feeding response of a benthic copepod to ciliate prey type,prey concentration and habitat complexity

1. Protozoans are important consumers within microbial food webs and, in turn, they represent potential prey for small metazoans. However, feeding interactions within these food webs are rarely characterised and this is especially true for freshwater sediments. 2. We aimed to quantify the feeding links between a freshwater meiofaunal copepod and ciliates in two laboratory experiments. The first experiment addressed the response of Eucyclops serrulatus towards ciliate density and type (two ciliate species of the same genus differing in terms of body size). A second experiment assessed the effect of habitat structure on feeding rates by introducing different structural complexity into the feeding arena. In contrast to the first experiment, which was run only for one time period, this experiment also tested three different total feeding times (4, 7 and 9 h). 3. Eucyclops serrulatus exhibited high ingestion rates, with 3–69 ciliates copepod^?1 h^?1 consumed depending on food concentration, food type and habitat complexity. Copepods exhibited a preference for the smaller ciliate when total ciliate concentration was low, but selected both ciliates equally when food concentrations were medium or high. However, at very high food concentration, Eucyclops preferred the larger ciliate (which was 1/3 of its own body size), suggesting that the longer handling times of the larger prey are rewarding when the large prey is present in high numbers. In terms of total numbers consumed, copepods fed on more small ciliates, but in terms of carbon units both ciliates were selected equally when total prey concentration was low or medium. However, copepods derived more carbon from the larger prey at high and very high prey concentrations (up to 0.7 μgC out of a maximum of 1.1 μgC copepod^?1 h^?1). Habitat complexity influenced the feeding of copepods when it was observed over time. 4. The copepod–ciliate link is well known from the pelagic zone of both marine and freshwater habitats. We have shown its potential importance within the benthos, where it can be influenced by food identity, food quantity and possibly by habitat complexity.     

A new species of Enterognathus (Copepoda, Cyclopoida, Enterognathidae) collected from the Seto Inland Sea, western Japan

A new species of the endoparasitic copepod Enterognathus (Cyclopoida, Enterognathidae) is described from a crinoid host in the Seto Inland Sea, western Japan. This is a third species of the genus and its first occurrence in the Pacific Ocean. The new species is distinguished from two previously known congeners by the morphology of the body somites, caudal rami, antennae and legs. Crinoid parasites belonging to Enterognathus and the closely related genus Parenterognathus have a broad distribution from the northeastern Atlantic through the Red Sea to the West Pacific.     

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     

Egg-brooding, body size and predation risk in planktonic marine copepods

We investigated the interacting effects of copepod body size and the presence or absence of egg masses on the risk of predation by a visual predator. We conducted selection experiments involving three-spined sticklebacks (Gasterosteus aculeatus) and copepods ranging in body mass from 0.5 to 740 μg C: Oithona similis, Corycaeus anglicus, Pseudocalanus newmani, P. moultoni, Pseudodiaptomus marinus, and Paraeuchaeta elongata. We found that sticklebacks selected ovigerous females of the two smallest-bodied species of copepods (Oithona similis and Corycaeus anglicus). In contrast, the fish showed no significant selection for ovigerous females of the remaining, larger-bodied species. Unexpectedly, egg mass position (i.e., in a ventral, dorsal or lateral location on the urosome) appeared to influence predation risk more than did body size, resulting in higher predation risk for the cyclopoid and poecilostomatoid species than for the calanoid species we tested. Although the sticklebacks showed no statistically significant preference for ovigerous females of any of the four calanoid species, for each species the overall proportion of ovigerous females ingested was slightly greater than 0.50. Thus, whether body size influences the susceptibility of egg-brooding calanoid copepods to predation remains an open question. Received: 24 August 1998 / Accepted: 6 July 1999     

Feeding ecology of two sillaginid fishes and trophic interrelations with other co-existing species in the southern part of South China Sea

Feeding ecology of Sillago sihama and Sillago ingenuua and interrelationship with other 15 fish species, including Stolephorus commersonii, Escualosa thoracata, Lutjanus russelli, Terapon puta, Ambassis kopsii, Ambassis vachellii, Dendrophysa russelli, Gerres filamentosus, Leiognathus decorus, Leiognathus splendens, Leiognathus equulus, Thryssa hamiltonii, Carangoides praeustus, Lethrinus lentjan, and Pomadasys maculatum, utilizing coastal waters off the southern part of the South China Sea were investigated. Dietary contents from 4308 guts of those species collected during July 2003–May 2005 were examined. They were significantly grouped, based on similarity coefficient, into five feeding clusters (P < 0.01). Of these, most of the diets were dominated by calanoid copepods, shrimps, polychaetes and gammarid amphipods. Comparative study on two sillaginid species showed that both Sillago sihama and Sillago ingenuua were carnivores fed mainly on polychaetes and other benthic organisms. Dietary ontogenetic change of Sillago sihama was very distinctive where small sized classes started feeding mainly on calanoid copepods before changing almost completely to polychaetes. The two species applied both sharing and partitioning strategies for food, depending on size classes, to live with each other.     

A new species of Leptodiaptomus (Copepoda, Diaptomidae) from Northwestern Mexico with comments on the distribution of the genus

A new species of the freshwater planktonic copepod genus Leptodiaptomus is described for a small pond in Northwestern Mexico. Leptodiaptomus dodsoni n. sp. can be easily distinguished mainly by the presence of an unusually large sinusoid spine on male antennular segment 13, and by the features of the fifth legs of both sexes. This genus is known to be distributed mainly in North America with 19 recognized species. Of these, six ocur in Mexico, and the new species seems to be closely related to most of them. It is probable that this group of species (including the new one) represents the southwards radiation of the genus from North America. Compared to the Caribbean and South American, the North American influence seems to be the most relevant for diaptomid copepods in Mexico. At least two Mexican species of Leptodiaptomus, including L.dodsoni, are restricted in distributional range to high-altitude temporal ponds, and both could be considered endemics.      

The peculiarities of parasitism in the copepoda and rhizocephala

In the parasitology it is accepted to subdivide the parasites into two categories based on their spatial relations with the organism of a host. To first category--ectoparasites--includes organisms, which live "on external covers, on a skin, on gills". To the second category--endoparasites--includes organisms living "in internal cavities, tissues and cells of the host" (Dogiel, 1941). Feizullaev (1971) offered the third category--mesoparasites--where he placed "the parasites inhabiting places, which are connected to cavities being open to external environment". According to him the places are oral, nasal, eye cavity, cloaca etc. Since some parasitic copepods living in certain kind of interrelation with a host can be referred neither to ectoparasites nor to endoparasites, the author offers these following criteria for the mesoparasitism definition. 1. Morphological and anatomic subdivision of the body of the parasite into two main parts: ectosoma located in the external environment (environment of the second order) and endosoma located inside the organism of the host (environment of the first order). 2. Functional subdivision of parasite body into two parts: endosoma which performs the trophic function and ectosoma with genital organs performs the reproduction function. 3. Existence of the parasite on the border of two environments (of the first and second order) which have an influence on parasite directly and simultaneously. The mesoparasites are known among the parasitic Copepoda and Rhizocephala only. As for the copepods all members of the family Herpyllobiidae may be placed in this category; the members of the families Phyllodicolidae and Chitonophilidae may be also included in it, albeit tentatively (because their anatomy has not yet been adequately studied), as well as five genera of the family Nicothoidae (Aspidoecia, Cephalorhiza, Diexanthema, Nicorhiza and Rhizorhina), and Bradophyla pigmaea.     

Some ecological aspects of the copepod parasites of the common hake, merluccius hubbsi, from the argentineuruguayan coasts

A total of 2164 common hake Merluccius hubbsi captured in the Argentine-Uruguayan Common Fishing Zone was examined for parasitic copepods. The fish were infested with Chondracanlhus palpifer and Neobrachietia insidiosa f. lageniformis, the former parasitizing the buccal cavity and the latter the branchial arches. C. palpifer showed an increase in both prevalence and intensity in relation to the host size; infestation with N. insidiosa f. lageniformis decreased with increasing length of hake. Attachment site preferences and variations in the parasite distribution patterns as related to the host biology were observed. Evidence of negative association between copepod species and seasonal changes in the parasite composition was also found.     

Phytoplankton Cell Size: Intra- and Interspecific Effects of Warming and Grazing

Decreasing body size has been suggested as the third universal biological response to global warming after latitudinal/altitudinal range shifts and shifts in phenology. Size shifts in a community can be the composite result of intraspecific size shifts and of shifts between differently sized species. Metabolic explanations for the size shifts dominate in the literature but top down effects, i.e. intensified size-selective consumption at higher temperatures, have been proposed as alternative explanation. Therefore, we performed phytoplankton experiments with a factorial combination of warming and consumer type (protist feeding mainly on small algae vs. copepods mainly feeding on large algae). Natural phytoplankton was exposed to 3 (1^st experiment) or 4 (2^nd experiment) temperature levels and 3 (1^st experiment: nano-, microzooplankton, copepods) or 2 (2^nd experiment: microzooplankton, copepods) types of consumers. Size shifts of individual phytoplankton species and community mean size were analyzed. Both, mean cell size of most of the individual species and mean community cell size decreased with temperature under all grazing regimes. Grazing by copepods caused an additional reduction in cell size. Our results reject the hypothesis, that intensified size selective consumption at higher temperature would be the dominant explanation of decreasing body size. In this case, the size reduction would have taken place only in the copepod treatments but not in the treatments with protist grazing (nano- and microzooplankton).