Relationship Patterns between Ichthyoplankton and Zooplankton: A Conceptual Model

Selected cases of plankton studies were analyzed to illustrate the main types of relationships between the zooplankton and the ichthyoplankton abundance observed in the pelagic realm. Such relationships may exhibit a positive, a negative, or a random pattern. In the conceptual model here proposed, short-term oscillations among these patterns were attributed to small-scale biological processes, such as competition, predation, and intraguild predation, acting in conjunction with water turbulence. A negative relationship between zooplankton and ichthyoplankton abundance may be caused both by predation on fish eggs and larvae, and by detrimental competition and intraguild predation interactions for fish larvae. In contrast, positive relationships emerge from the absence or low abundance of major predators on the ichthyoplankton, and from food availability for fish larvae and competing species. The random pattern may appear as a gradual transitional stage between the negative and positive patterns, or be promoted by strong water turbulence – which generates random movements of individuals. The size of zooplankters greatly influences these small-scale phenomena. Hence, their role in the trophic web, the success in competition interactions and vulnerability to water turbulence depends on their size. Intra- and interspecific competition may be reduced by variability in body size within or among fish larvae populations. Owing to a strong interaction among phenomena at different scales, these small-scale processes are also influenced by larger scale features, such as seasonal changes in zooplankton biomass, water currents, or spawning periods of fish. At the space level, some theoretical studies have emphasized the role of water currents as a vector for fish larvae to reach the nursery grounds (migration triangle hypothesis), or to allow them to remain within their own population’s distributional area (member/vagrant hypothesis). At the temporal level, the match/mismatch theory insists in a synchrony between reproductive strategies of fish and cyclical changes in abundance and size spectrum of potential prey items for their larvae. In any case, a coincidence between favorable abiotic and biotic features during the whole life-cycle of fish would assure a success in survival of larvae and their subsequent recruitment to adult population.     

A review of cnidarians and ctenophores feeding on competitors in the plankton

Predation among pelagic cnidarians and ctenophores is reviewed. The diets of semaeostome scyphomedusae and hydromedusae commonly include other gelatinous zooplanktivores. However, few species of siphonophores and ctenophores are known to consume other gelatinous species. Most of these species can be said to exhibit intraguild predation, since they consume species that potentially compete with them for food. In addition, some hydromedusan and ctenophore species may consume other gelatinous zooplanktivores exclusively. Characteristics of cnidarians and ctenophores as predators and as prey of other gelatinous species are discussed.     

GENE FLOW AND INEFFECTIVE ANTIPREDATOR BEHAVIOR IN A STREAM-BREEDING SALAMANDER

Predators often feed on prey that show ineffective antipredator behavior. Gene flow among populations may constrain evolution of effective antipredator ability in larvae of the streamside salamander, Ambystoma barbouri, a species that occupies distinctly different habitats with conflicting selection pressures. Some streams are ephemeral, where larvae should be active to feed and reach metamorphosis before stream drying. In contrast, other streams are more permanent and contain pools with predatory fish, where larvae should remain inactive to avoid fish predation. Feeding rates and predator escape behavior were assayed for laboratory-reared larvae from 15 populations. Larval survival was also compared among populations in artificial streams with natural predators. Five populations represented streams subjected to fish predation along a gradient of genetic and geographic isolation from populations without fish; the remaining 10 populations were ephemeral and without fish. Individuals from populations with fish had significantly stronger behavioral responses to fish (i.e., decreased feeding rate associated with the presence of fish and increased escape response) than individuals from fishless populations. Larvae from populations containing fish that were more isolated from fishless populations showed stronger antipredator responses than less isolated populations. Further, larvae from more isolated populations survived longer in the predation experiment, indicating that the behaviors measured were related with survival. These results suggest that gene flow between populations with conflicting selection pressures limits local adaptation in some salamander populations with fish. While previous studies have typically focused on the role of gene flow in pairs of populations, the results of this study suggest that gene flow is acting to swamp local adaptation across several populations.     

An assessment of the biomass production theory in the Baltic Sea using fish landings data

Fish landings in the Baltic Sea from 1970 to 2000 were used as a proxy for fish biomass to explore variability of total fish biomass. Total demersal (total D) and total pelagic (total P) landings proved relatively invariant over time compared with most of their component species. This was explained in terms of the energy limitation imposed on the ecosystem by its carrying capacity, forcing species interactions (predation, competition, etc.) with compensations that allowed the total biomasses to remain relatively stable. Extensive interactions were demonstrated among the Baltic fish species by linear correlation with appropriate negative signs, indicating compensatory interactions consistent with the energy limitation theory. The variances of the landings of cod, herring, sprat and total landings reflected the magnitudes of variation of their biomasses as estimated from the Virtual Population Analysis (VPA), thus justifying the use of landings data in this analysis as a proxy for biomass. Significant demersal–pelagic coupling was indicated from the landings data, which could be explained by trophic interactions. Species interactions generally explained between 17 and 66% of the variations in landings. Thus, substantial portions of the variations in the landings must be attributed to other factors: biological, fishery and environmental.     

Impact of whitefish on an enclosure ecosystem in a shallow eutrophic lake: selective feeding of fish and predation effects on the zooplankton communities

Bag-type enclosures (75 m³) with bottom sheets and tube-type enclosures (105 m³) open to the bottom sediment were stocked with exotic whitefish (Coregonus lavaretus maraena) to study their predation effects on the plankton community. The fish fed mainly on adult chironomids during the period of their emergence (earlier part of the experimental period). Thereafter, the food preference was shifted to larvae of chironomids and crustacean zooplankters. The predation effects on the plankton community were not evident in the bag-type enclosures where zooplankton densities were consistently low. The fish reduced the crustacean populations composed ofBosmina fatalis, B. longirostris andCyclops vicinus in the tube-type enclosures where the prey density was high (above ca. 50 individuals 1⁻¹). The results suggested that the intensity of predation depended on the prey density. Rotifers increased in the fish enclosure, probably becauseCoregonus reduced the predation pressure byCyclops vicinus on rotifers and allowed the latter to increase. In the fish enclosures, no marked changes in species composition were observed. Zooplankton predated by the fish seemed to be distributed near the walls of the enclosures. Problems of enclosure experiments for examining the effects of fish predation on pelagic zooplankton communities are discussed.     

Behavioral linkage of pelagic prey and littoral predators: microhabitat selection by Daphnia induced by damselfly larvae

Only recently ecologists started treating the previously separately considered benthic, littoral and pelagic zones of lake ecosystems as closely connected compartments. Here we study a link between organisms belonging to a different compartment – namely the pelagic and the littoral – through behavior in a series of laboratory experiments. Waterfleas of the genus Daphnia are inhabitants of the pelagic zone and suffer a high predation pressure from syntopic vertebrate predators (mainly fish). Presumably to escape this predation, they sometimes migrate in the day to the littoral to seek refuge within macrophytes and return to the pelagic at night. Zygopterans from the genus Ischnura do commonly co-occur in ponds with Daphnia and are known as opportunistic predators of Daphnia . In two initial experiments in microcosms in the lab we showed that Ischnura larvae are littoral predators strongly associated with macrophytes. Although we found that predation rates of individual Ischnura larvae on Daphnia are approximately 1.5 fold lower in macrophytes compared to open water, total predation from Ischnura on Daphnia per unit area is tenfold higher within macrophytes than in open water, making the open water a safer place for Daphnia with regard to Ischnura predation. In a third microcosm experiment we monitored horizontal distribution of Daphnia in the absence, presence and odor only of Ischnura larvae. After 2 hours, on average 10% less Daphnia remained within the vegetation when Ischnura larvae or only their odor were present compared to when Ischnura or their odor were absent. We interpret this as a behavioral anti-predation response of Daphnia to the presence of Ischnura larvae that seems primarily chemically mediated. The observed horizontal migration of the pelagic prey driven by the littoral predator may couple both lake compartments and may interact with the predator–prey relationships within the pelagic.     

How Nemo finds home: the neuroecology of dispersal and of population connectivity in larvae of marine fishes

Nearly all demersal teleost marine fishes have pelagic larval stages lasting from several days to several weeks, during which time they are subject to dispersal. Fish larvae have considerable swimming abilities, and swim in an oriented manner in the sea. Thus, they can influence their dispersal and thereby, the connectivity of their populations. However, the sensory cues marine fish larvae use for orientation in the pelagic environment remain unclear. We review current understanding of these cues and how sensory abilities of larvae develop and are used to achieve orientation with particular emphasis on coral-reef fishes. The use of sound is best understood; it travels well underwater with little attenuation, and is current-independent but location-dependent, so species that primarily utilize sound for orientation will have location-dependent orientation. Larvae of many species and families can hear over a range of ~100-1000 Hz, and can distinguish among sounds. They can localize sources of sounds, but the means by which they do so is unclear. Larvae can hear during much of their pelagic larval phase, and ontogenetically, hearing sensitivity, and frequency range improve dramatically. Species differ in sensitivity to sound and in the rate of improvement in hearing during ontogeny. Due to large differences among-species within families, no significant differences in hearing sensitivity among families have been identified. Thus, distances over which larvae can detect a given sound vary among species and greatly increase ontogenetically. Olfactory cues are current-dependent and location-dependent, so species that primarily utilize olfactory cues will have location-dependent orientation, but must be able to swim upstream to locate sources of odor. Larvae can detect odors (e.g., predators, conspecifics), during most of their pelagic phase, and at least on small scales, can localize sources of odors in shallow water, although whether they can do this in pelagic environments is unknown. Little is known of the ontogeny of olfactory ability or the range over which larvae can localize sources of odors. Imprinting on an odor has been shown in one species of reef-fish. Celestial cues are current- and location-independent, so species that primarily utilize them will have location-independent orientation that can apply over broad scales. Use of sun compass or polarized light for orientation by fish larvae is implied by some behaviors, but has not been proven. Use of neither magnetic fields nor direction of waves for orientation has been shown in marine fish larvae. We highlight research priorities in this area.     

Jellyfish as Prey: Frequency of Predation and Selective Foraging of Boops boops (Vertebrata,Actinopterygii) on the Mauve Stinger Pelagia noctiluca (Cnidaria,Scyphozoa)

In recent years, jellyfish blooms have attracted considerable scientific interest for their potential impacts on human activities and ecosystem functioning, with much attention paid to jellyfish as predators and to gelatinous biomass as a carbon sink. Other than qualitative data and observations, few studies have quantified direct predation of fish on jellyfish to clarify whether they may represent a seasonally abundant food source. Here we estimate predation frequency by the commercially valuable Mediterranean bogue, Boops boops on the mauve stinger jellyfish, Pelagia noctiluca, in the Strait of Messina (NE Sicily). A total of 1054 jellyfish were sampled throughout one year to quantify predation by B. boops from bite marks on partially eaten jellyfish and energy density of the jellyfish. Predation by B. boops in summer was almost twice that in winter, and they selectively fed according to medusa gender and body part. Calorimetric analysis and biochemical composition showed that female jellyfish gonads had significantly higher energy content than male gonads due to more lipids and that gonads had six-fold higher energy content than the somatic tissues due to higher lipid and protein concentrations. Energetically, jellyfish gonads represent a highly rewarding food source, largely available to B. boops throughout spring and summer. During the remainder of the year, when gonads were not very evident, fish predation switched towards less-selective foraging on the somatic gelatinous biomass. P. noctiluca, the most abundant jellyfish species in the Mediterranean Sea and a key planktonic predator, may represent not only a nuisance for human leisure activities and a source of mortality for fish eggs and larvae, but also an important resource for fish species of commercial value, such as B. boops.     

Major changes in pelagic rotifers during natural and forced recovery from acidification

Pelagic rotifers were studied in lakes with contrasting acidification histories situated in an acid-stressed region of southern Norway. Life histories and spatial distribution varied considerably between the investigated species, and influenced the recovery processes. Most headwater lakes have experienced strongly acidified environments during the last five decades, whereas lakes close to the Skagerrak coast have been stable within the same period. Rotifer diversity and abundance were reduced in the most acidic sites and increased towards the coast. Most surveyed species are known to possess sediment egg-banks, and after chemical recovery most rotifers dispersed into the plankton from these egg-banks and produced viable populations. Some species of the genera Polyarthra and Collotheca, and the species Kellicotta longispina and Keratella serrulata showed a striking ability to tolerate acidification, and were the dominant taxa in the acidmost environments. K. serrulata characterised, but did not numerically dominate, acid rotifer communities especially in the most coloured sites, and decreased following liming. The predominantly bacteriophageous genus Conochilus exploded in numbers shortly after liming, most probably because bacteria increased strongly during this transition phase. Planktivorous fish influenced indirectly rotifer abundance by consuming invertebrate predators and important rotifer competitors such as filter feeding cladocerans. Invertebrate predators, such as larvae of Chaoborus spp. and Heterocope saliens probably influenced rotifer distributional patterns in a complex top-down manner, both during chronic acidification and liming in environments with low fish predation. Important rotifer predators such as pelagic cyclopoid copepods, Bythotrephes longimanus and Leptodora kindti, were absent from the most acidic fishless lakes. Considerable populations of large-sized Daphnia longispina probably suppressed several rotifer species in sites with low fish predation, as did large populations of Bosmina longispina and Ceriodaphina quadrangula in lakes with intense fish predation.     

Complex behaviour by coral-reef fish larvae in open-water and near-reef pelagic environments

We present the first in situ observations of the pelagic larvae of coral-reef fishes feeding, schooling and being preyed upon. In addition, we report on their behavioural interactions with adult and juvenile fishes. Observations on over 500 larvae of over 50 species (mostly from four families) near the end of their pelagic interval were made in both open water (> 1 km offshore) and near-reef environments. Nearly 10% of larvae were seen to feed in open water, but < 1% fed near the reef. Presettlement schooling was observed in five species of four families. We observed no predation upon larvae in open water except near the bottom. Near the reef, 8.5% of larvae were eaten. The main predators near and on the reef were a species of wrasse and lizardfishes. Rates of predation seem to differ among genera of pomacentrids, perhaps related to differences in behaviour when settling. When confronted with adult fishes, which happened largely near the reef, larvae reacted with a limited range of behaviours, including sheltering near the observer, swimming to the surface, slowing or stopping, or swimming offshore. The frequency of these behaviours differed among larvae of three pomacentrid genera. Interactions with reef residents, particularly pomacentrids, were common, and usually involved aggression by the resident toward settling larvae. This may act to discourage settlement during the day when such residents are active. These data show that behaviour of late larvae of coral-reef fishes is complex and can greatly influence survival and recruitment. Further, behaviour differs among taxa, showing that not only are larvae not passive, but also that a generalised behaviour of larvae does not exist.