Dispersal ability and relative abundance of Boeckella and Calamoecia (Copepoda: Calanoida) in Australian and New Zealand waters

Two calanoid copepod genera, Boeckella and Calamoecia have broadly similar geographic distributions in Australia but Boeckella is more common than Calamoecia. A field survey and literature search indicate that Boeckella is usually the first calanoid to colonize newly formed bodies of water, and Boeckella is more common in temporary pools and in high altitude environments. Boeckella usually produces larger clutches than Calamoecia, increasing its probability of establishment in lakes and ponds; calculations indicate that Boeckella's probability of establishment is far greater than that of Calamoecia. That Calamoecia is common on the North Island of New Zealand may be due to the presence of planktivorous fish; small species are more likely to survive in such environments. Calamoecia clutches may be able to move more often and farther than those of Boeckella but Calamoecia remains rare because its probability of establishment is low. Boeckella may be able to survive temporary pool environments not only because it can produce larger clutches, but also because many species in this genus are highly pigmented, and as such can survive brightly lighted environments. Patterns of abundance of North American diaptomids appear to be consistent with factors affecting those of Australian and New Zealand centropagids.     

Relative impacts of Daphnia grazing and direct stimulation by fish on phytoplankton abundance in mesocosm communities

• 1 Planktivorous fish were hypothesised to influence the abundance of algal biomass in lakes by changing zooplankton grazing, affecting zooplankton nutrient recycling and by direct recycling of nutrients to phytoplankton. The relative roles of direct fish effects vs. zooplankton grazing were tested in mesocosm experiments by adding to natural communities large grazing zooplankton (Daphnia carinata) and small planktivorous fish (mosquitofish or juveniles of Australian golden perch).
• 2 The addition of Daphnia to natural communities reduced the numbers of all phytoplankton less than 30 µm in size, but did not affect total biomass of phytoplankton as large Volvox colonies predominated.
• 3 The addition of Daphnia also reduced the abundance of some small (Moina, Bosmina, Keratella) and large (adult Boeckella) zooplankton, suggesting competitive interactions within zooplankton.
• 4 The addition of mosquitofish to communities containing Daphnia further reduced the abundance of some small zooplankton (Moina, Keratella), but increased the numbers of Daphnia and adult Boeckella. In spite of the likely increase in grazing due to Daphnia, the abundance of total phytoplankton and dominant alga Volvox did not decline in the presence of mosquitofish but was maintained at a significantly higher level than in control.
• 5 The addition of juveniles of golden perch to communities containing Daphnia reduced the abundance of small zooplankton (Moina), increased the abundance of large zooplankton (adult Boeckella) but had no significant effect on Daphnia and total phytoplankton abundance.
• 6 The results of the present study suggest that some planktivorous fish can promote the growth of phytoplankton in a direct way, probably by recycling nutrients, and even in the presence of large grazers. However, the manifestation of the direct effect of fish can vary with fish species.

     

A comparison of the trophic transfer of fatty acids in freshwater plankton by cladocerans and calanoid copepods

1. Analyses of zooplankton fatty acid (FA) composition in laboratory experiments and samples collected from lakes in New Zealand spanning a wide gradient of productivity were used to assess the extent to which FAs might infer their diet. We used the cladocerans, Daphnia and Ceriodaphnia, and the calanoid copepod, Boeckella, as test organisms, and monocultures of cryptophytes, chlorophytes and cyanobacteria as food. Based on reproductive success, cryptophytes were the highest food quality, chlorophytes were intermediate and cyanobacteria the poorest. 2. Several FA groups were highly correlated between zooplankton and their diets. They were monounsaturated fatty acids (MUFAs), and ω3 and ω6 polyunsaturated fatty acids (PUFAs) for cladocerans, and saturated fatty acids (SAFAs) and ω3 PUFAs for copepods. Several FAs varied significantly less in the zooplankton than in their monoculture diets, e.g. MUFAs in Daphnia, and ω3 and ω6 PUFAs in Ceriodaphnia, despite clear dietary dependency for these FAs. 3. Zooplankton collected from lakes in New Zealand had more eicosapentaenoic acid (EPA) (Daphnia), more highly unsaturated ω3 and ω6 FAs (C₂₀, C₂₂; Daphnia, Ceriodaphnia, Boeckella) and less ω3 C₁₈ PUFAs (Daphnia, Ceriodaphnia, Boeckella) and ω6 C₁₈ PUFAs (Daphnia, Ceriodaphnia) than measured in the same species reared on phytoplankton in the laboratory. 4. Analyses of FA composition of seston and freshwater zooplankton globally showed that, in general, zooplankton had a significantly higher proportion of arachidonic acid and EPA than seston, and copepods also had a higher percentage of docosahexaenoic acid than seston. 5. These results suggest that zooplankton selectively incorporate the most physiologically important FAs. This could be a consequence of preferential assimilation, selective feeding on more nutritious cells or locating and feeding within higher food quality food patches.     

The biogeography of three Boeckella species (Copepoda: Calanoida) in New Zealand

The New Zealand distributions of three species of Boeckella (Copepoda, Calanoida), B. triarticulata, B. dilatata and B. hamata are mapped. B. triarticulata is primarily a pond dweller but is also found in reservoirs and shallow lakes. B. dilatata is mainly found in the deeper glacial lakes and ponds in the central region of the South Island and B. hamata has a more widespread distribution in lakes and ponds in the South Island and lower half of the North Island. Differences in temperature optima, food requirements and dispersal ability among the three Boeckella species are related to vicariant events to explain their distribution in New Zealand.     

Calanoid copepods feeding on algae and filamentous cyanobacteria: rates of ingestion, defaecation and effects on trichome length

The relative abilities of freshwater calanoid copepods to grazealgae and filamentous cyanobacteria were examined in three speciesof Boeckella (B.triarticulata, B.hamata, B.dilatata) fed Cydotella,Cryptomonas and two species of Nostoc and Anabaena. Althoughrates of ingestion were highest on Cryptomonas, all the Boeckellaspecies consumed cyanobacteria, some species at similar ratesto Cyclolella. Faecal production on the foods was highest onCydotella and, in B.triarticulata, higher on A.flos-aquae andNostoc sp. 2 than on Cryptomonas. At 1 mg l^–1 of thesecyanobacteria, female B.triarticulata voided 70% by volume ofthe material they ingested as compact faecal pellets. By theirgrazing they also decreased dramatically the trichome lengthsof A.flos-aquae, A.oscillarioides and Nostoc sp. 2, therebypossibly altering the potential susceptibility of these filamentouscyanobacteria to grazing by other zooplankton.     

Long-term zooplankton composition data reveal impacts of invasions on community composition in the Waikato lakes,New Zealand

Data collected on zooplankton community composition over longer time periods (>?10 years) are rare. We examined among-lake spatial and temporal trends of zooplankton communities from a monitoring programme undertaken in the Waikato region, New Zealand. A total of 39 lakes were sampled over a period of 12 years, between 2007 and 2019, with varying degrees of temporal effort. We focussed particularly on eight lakes, considered here as ‘long-term lakes’, where samples were collected with greater regularity (including 5 with 12 years of data). Among lakes, suspended sediment concentrations and indicators of lake trophic state were inferred to be important in determining the zooplankton distributions; as this region is dominated by shallow lakes, the relative importance of suspended sediments was high. Among the long-term lakes, the greatest dissimilarities in zooplankton community composition among years were in Lake Waahi, where the Australian Boeckella symmetrica was first detected in 2012. That is, the greatest temporal changes to zooplankton composition during the study period were due to the invasion by non-indigenous species, rather than changes in trophic state or other environmental variables; non-native species commonly dominated the individual counts of species through much of 2014 and 2015, with most samples since 2016 being again dominated by native species. Following this lake, the largest and shallowest lakes in the dataset—Whangape and Waikare—exhibited the greatest variability in community composition among years.

     

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)     

Mating behaviour and parapatry in two species of Australian reptile tick

Summary Few quantitative studies have examined the ecological consequences of similarities and/or differences in mating behaviour of parapatric species. Reproductive interference occurs between several parapatric species of Australian reptile tick, due to similarities in their mating behaviour (Andrews et al. 1982a). Attempts to determine whether reproductive interference serves to maintain parapatry between Amblyomma limbatum and Aponomma hydrosauri have been hindered because of difficulties in providing conditions conducive to conspecific mating in Amb. limbatum. The present study examined whether off-host and/or onhost temperature influenced the subsequent mating behaviour (i.e. the proportion of females that mate and the time when mating occurs) of these two species. Irrespective of the temperature experienced by ticks prior to host attachment, specific on-host temperatures were needed to induce mating in Amb. limbatum (i.e. host cloacal temperatures >32° C prior to the time of peak mating activity). Significantly more Amb. limbatum females were mated and the time taken by females to mate decreased with increasing on-host temperatures. mating in Ap. hydrosauri occurred over a wider range of on-host temperatures and the time when mating occurred did not alter at different on-host temperatures. In addition, significantly more Ap. hydrosauri males moved and each male made more moves on hosts than did Amb. limbatum males. It is suggested that Ap. hydrosauri may in consequence have a competitive mating advantage over Amb. limbatum at a boundary. Similarities in mating behaviour, on the other hand, increase the probability of reproductive interference, hence reduce the reproductive fitness of colonizing females of both species. We propose that similarities and differences in mating behaviour could play a critical role in the maintenance of parapatric boundaries.     

Effects of female deprivation on mating propensity and mate selectivity by maleRequena verticalis (Orthoptera: Tettigoniidae)

This study examined the effects of female deprivation on the mating propensity and size selectivity of male Requena verticalis,an Australian katydid in which males provide a large spermatophylax to females during mating. Specifically, I tested the hypotheses that mating readiness would increase, while mate discrimination would decrease (i. e., females would be accepted independently of their size, an indicator of fecundity) as the period of female deprivation increased. Fieldcaught males were held in isolation for 1, 5, or 14 days in the laboratory and then presented with virgin females. As expected, males held for 5 and 14 days were more likely to mate than males held only a single day. However, in none of the treatments did males discriminate among females on the basis of size. These results are compared with those obtained in a similar study on a zaprochiline katydid.     

The importance of zooplankton in the diets of three native fish species in floodplain waterholes of a dryland river,the Macintyre River,Australia

The zooplankton of freshwater systems has been recognized as an important energy resource for fish of small body size that, in turn, provide energy to piscivorous fish consumers higher up the food web. This study evaluates the importance of zooplankton to the diets of three species of fish living in floodplain waterholes of an Australian dryland river. The species selected for study represent different trophic categories in waterhole food webs: Ambassis agassizii is a microcarnivore, Leiopotherapon unicolor is an omnivore, and Nematalosa erebi is a detritivore. Dietary differences among size classes of each species were also evaluated to understand possible ontogenetic shifts in zooplankton consumption. Ambassis agassizii fed primarily on zooplankton (99.9%, made up mostly of 81.6% Calanoida and 17.4% Moinidae), regardless of the size of individual fish. Leiopotherapon unicolor fed on zooplankton (47%, mostly Daphniidae and Moinidae) and aquatic insects (46.7%). Smaller individuals of Leiopotherapon unicolor (30–49 mm TL—total length) were responsible for 36.1% of the plankton consumed by the species. Nematalosa erebi fed on detritus (84.6%) with zooplankton (Calanoida, Moinidae, and Cyclopoida) contributing only 13.7% of the mean diet. Smaller individuals (40–69 mm TL) were responsible for 98% of the plankton consumed by Nematalosa erebi, and individuals of 40–49 mm (TL) fed exclusively on zooplankton (53.8% Moinidae and 46.2% Calanoida). Although the three fish species had different diets, reflecting differences in species-specific and ontogenetic morphological and behavioral characteristics, zooplankton formed the basis of the diet of all species when young. These results confirm the importance of zooplankton as a major food resource for three fish species and smaller size classes of these species in floodplain waterholes of the Macintyre River, Australia. Guest editors: U. M. Azeiteiro, I. Jenkinson & M. J. Pereira Plankton Studies     

Boeckella diamantina n.sp. (Calanoida,Centropagidae), from a high Andean lake in Mendoza,Argentina

A new species of Boeckella from limnetic samples of Laguna del Diamante, a high lake in the Andes (34°10 S) is described and illustrated. The species is defined by the characters of the male fifth leg: the right two segmented endopod bears four peculiar, short, claw-like spines, the left endopod is a simple finger-like projection. This species is related to B. gibbosa, also a species from the Andes and B. vallentini from Malvinas (Falkland Islands) and other subantarctic islands. It is distinguished from them by diagnostic features of the fifth legs of the male and abdominal structure and fifth legs of the female. Some current views on the features used in the taxonomy of the genus Boeckella are discussed.     

Habitat patch size and mating system as determinants of social group size in coral-dwelling fishes

It is thought that the size and dispersion of habitat patches can determine the size and composition of animal social groups, however, this has rarely been tested. The relationship between group size, the mating system, and habitat patch size in six species of coral-dwelling gobies was examined. For all species, there was a positive correlation between coral colony size and social group size, however the strength of this relationship varied among species. Paragobiodon xanthosomus exhibited the strongest relationship and a manipulative field experiment confirmed that coral colony size limited group size in this species. For other species including Paragobiodon melanosomus and Eviota bifasciata, either a highly conservative mating system (P. melanosomus), or increased mobility (E. bifasciata) appeared to disrupt the relationship between habitat patch size and group size. There was no consistent relationship between the mating system exhibited and group size among the species investigated. These results demonstrate that habitat patch size, mobility, and mating systems can interact in complex ways to structure group size even among closely related species.     

Trophic relationships and seasonal utilization of salt-marsh creeks by zooplanktivorous fishes

Synopsis In a high salinity estuary at North Inlet, South Carolina, co-occurrence and possible competition among adults of four dominant zooplanktivorous fishes were minimized by seasonal adjustments in lateral and vertical distributions as well as in dietary preferences. In winter, Atlantic silversides, Menidia menidia, occupied the entire water column while other planktivores were rare or absent from the estuary, and they consumed large prey such as mysid shrimps and fish larvae. An immigration of bay anchovies, Anchoa mitchilli, in the spring resulted in a redistribution of species with Atlantic silversides shifting to the surface waters and bay anchovies dominating the lower half of the water column. Both fishes consumed mostly copepods in the spring, but each favored a different species. There was little similarity in the large prey items consumed by the two fishes. Striped anchovies, Anchoa hepsetus, arrived in mid-summer and were most abundant at the surface while bay anchovies continued to dominate the bottom waters. Atlantic silversides were rare in all summer collections. The diets of the two anchovies were similar, but vertical separation during the period of maximum zooplankton abundance probably minimized competition. Rough silversides, Membras martinica, which were obligate surface dwellers, shared the upper water column with striped anchovies, but the two species had very different diets during their period of co-occurrence. Although seasonal changes in fish diets reflected shifts in zooplankton composition and all fishes consumed a variety of prey types, preferences for some prey taxa and total avoidance of others were indicated. Electivity indices indicated an especially strong selection for fiddler crab megalopae by all fishes in the summer and fall. All fishes, except rough silversides, which fed almost exclusively on copepods and crab zoeae, consumed large prey items when they were available. Fine scale partitioning of the food resources was apparent in the selection of different copepod and insect species by the fishes. Spatial and temporal separation in the distribution and/or dietary preferences of the zooplanktivores fishes probably reduces the potential for resource competition. Given the high abundances and selectivity of the planktivores, significant impacts on some zooplankton populations probably result.     

FINE STRUCTURE OF MISTLETOE POLLEN. IV. EURASIAN AND AUSTRALIAN VISCUM L. (VISCACEAE)

Fifteen Eurasian and Australian species of Viscum L. were examined by light, scanning, and transmission electron microscopy. Pollen characters divide the species into two groups, each correlated with differences in habit and inflorescence structure: Group I (12 spp.) characterized by psilate or slightly sculptured exines and a non-uniform ektexine pattern and Group II (3 spp.) possessing highly sculptured (echinate, rodlet) surfaces and uniform ektexine patterns. Within each of the groups, pollen characters divide the species into several subgroups. Among Group I, species V. nepalense, V. heyneanum and V. ovalifolium are particularly close. The Group I species, V. trilobatum, is placed in its own subgroup primarily because of its uniform ektexine pattern—a unique feature among Asian and Australian Viscum. Of the three Group II species, V. album and V. alniformosanae are palynologically almost indistinguishable. Pollen of the Group II V. cruciatum, though exomorphologically similar to V. album, is closest ultrastructurally to the Indian V. trilobatum. Overall, the most common and probably basic pollen characters among the Eurasian and Australian species include: subprolate, rounded convex almost spherical shapes; tricolporate apertures, and non-uniform sculpturing and ektexine patterns. Oblate-spheroidal or prolate-spheroidal shapes, prominent sculpturing, and a uniform ektexine pattern are derived characters largely restricted to the Eurasian and Asian Group II species (V. album, V. alniformosanae, V. cruciatum).     

ESTIMATES OF INTRAGAMETOPHYTIC SELFING AND INTERPOPULATIONAL GENE FLOW IN HOMOSPOROUS FERNS

Relatively little information is available on mating systems and interpopulational gene flow in species of homosporous pteridophytes. Because of the proximity of antheridia and archegonia on the same thallus, it has long been maintained that intragametophytic selling is the predominant mode of reproduction in natural populations of homosporous ferns and other homosporous plants. Furthermore, quantitative estimates of interpopulational gene flow via spore dispersal are lacking. In this paper, we examine five species of homosporous ferns (Botrychium virginianum, Polystichum munitum, P. imbricans, Blechnum spicant, and Dryopteris expansa) and present estimates of 1) rates of intragametophytic selling, 2) levels of interpopulational gene flow, and 3) interpopulational genetic differentiation (F_ST). Our data demonstrate that mating systems vary among species of ferns, just as they do among species of seed plants. The data also suggest that levels of interpopulational gene flow are generally high. The F_ST values indicate little genetic divergence among populations for all species except Dryopteris expansa, which exhibits significant levels of interpopulational genetic differentiation. Patterns of genetic diversity in the five species examined are related to the mating system and rate of interpopulational gene flow in each species. The F_ST values for all species except Botrychium virginianum are in close agreement with those predicted for an island model of population structure.     

Population structure of a vector‐borne plant parasite

Parasites are among the most diverse groups of life on Earth, yet complex natural histories often preclude studies of their speciation processes. The biology of parasitic plants facilitates in situ collection of data on both genetic structure and the mechanisms responsible for that structure. Here, we studied the role of mating, dispersal and establishment in host race formation of a parasitic plant. We investigated the population genetics of a vector‐borne desert mistletoe (Phoradendron californicum) across two legume host tree species (Senegalia greggii and Prosopis velutina) in the Sonoran desert using microsatellites. Consistent with host race formation, we found strong host‐associated genetic structure in sympatry, little genetic variation due to geographic site and weak isolation by distance. We hypothesize that genetic differentiation results from differences in the timing of mistletoe flowering by host species, as we found initial flowering date of individual mistletoes correlated with genetic ancestry. Hybrids with intermediate ancestry were detected genetically. Individuals likely resulting from recent, successful establishment events following dispersal between the host species were detected at frequencies similar to hybrids between host races. Therefore, barriers to gene flow between the host races may have been stronger at mating than at dispersal. We also found higher inbreeding and within‐host individual relatedness values for mistletoes on the more rare and isolated host species (S. greggii). Our study spanned spatial scales to address how interactions with both vectors and hosts influence parasitic plant structure with implications for parasite virulence evolution and speciation.     

A metacommmunity approach to co-occurrence patterns and the core-satellite hypothesis in a community of tropical arboreal ants

Inferring mechanisms of community assembly from co-occurrence patterns is difficult in systems where assembly processes occur at multiple spatial scales and among species with heterogeneous dispersal abilities. Here, we demonstrate that local scale analysis of co-occurrence patterns is inadequate to fully describe assembly mechanisms and instead utilize a metacommunity and core-satellite approach. We generated a co-occurrence and life-history data set for a community of twig-nesting ants on coffee plants across 36 sites within a tropical agroecosystem to test the following three hypotheses: (1) twig-nesting ant species compete for nest-sites, (2) they are structured as a metacommunity, and (3) core species show segregated patterns, while satellite species show random patterns of co-occurrence. Species were divided into four groups: core species that are well distributed regionally and dominant locally, regional dominants that are well distributed regionally but do not dominate locally, local dominants that are dominant locally but are not widely distributed, and satellites that are neither widely distributed nor dominant locally. Only the most abundant species in the community, Pseudomyrmex simplex, was classified as a core species. Regional dominants, local dominants, and satellite species show random patterns of co-occurrence. However, when P. simplex is included in the co-occurrence matrix, patterns become aggregated for all three species groupings. This suggests that P. simplex “assembles” the community by providing a core metapopulation that other species track. Analyzing co-occurrence patterns among candidate subsets of species, at multiple spatial scales, and linking them to species traits substantially improves the explanatory power of co-occurrence analyses in complex metacommunities.     

Composition,size, and biomass of zooplankton in large productive Florida lakes

Crustacean zooplankton data were compiled from long-term observational studies at seven large shallow Florida lakes, to determine whether there are general characteristics in regard to species composition, body size, and biomass. In particular, we examined whether patterns in body size and species richness fit empirical models developed by Stanley Dodson. The lakes included range in size from 125 to 1730 km² and encompass mesotrophic to hyper-eutrophic conditions. We found that zooplankton biomass was strongly dominated by one species of calanoid copepod—Arctodiaptomus dorsalis. Large daphnids were absent, and Cladocera assemblages were dominated by small taxa such as Ceriodaphnia, Chydorus, and Eubosmina. The total number of species of pelagic cladocerans (8–12) was consistent with Dodson’s predictions based on lake area. The average size of crustacean zooplankton in Florida lakes is small in comparison with temperate communities. A. dorsalis is the smallest calanoid copepod in North America, and the mean length of Cladocera (0.6 mm) is consistent with Dodson’s results that size decreases from temperate to tropical zones. Total biomass of crustacean zooplankton was very low, ratios of zooplankton to phytoplankton biomass (0.01–0.1) are among the lowest reported in the literature, and the zooplankton displayed short-lasting early spring peaks in biomass. Cladocera were almost entirely absent in spring and summer. Factors known to occur in Florida lakes, which appear to explain these characteristics of biomass, include intense fish predation and high summer water temperature.     

Mating frequency and interspecific matings in some freshwater cyclopoid copepods

Laboratory investigations with Cyclops vicinus and Metacyclops minutus documented that mating frequency decreased significantly after the initial mating, i.e. re-mating is relatively rare in cyclopoid copepods. Re-mating is unnecessary because females are able to fertilize multiple clutches of viable eggs from one insemination. Similar sized Cyclops vicinus and Cyclops furcifer interbreed frequently. Interbreeding does not occur when size differences and taxonomic differences are as great as between C. vicinus and M. leuckarti. In C. vicinus and M. leuckarti, the duration of the last mating phase (spermatophore transfer until release of the female) and consequently the duration of the entire mating process, is different. I conclude that the low re-mating frequency in cyclopoid copepods probably evolved to accelerate clutch production in unpredictable environments and to reduce predation risk. The occurrence of interspecific mating discourages the co-existence of similarly sized, related species. Mating behaviour may have a considerable influence on zooplankton community structure.