Parasites determine a predator's optimal feeding strategy

Summary The diet selected by three-spined sticklebacks (Gasterosteus aculeatus) depends on the degree of parasitization by one or both of two parasite species (Schistocephalus solidus, Glugea anomala). Uninfested fish prefer the more profitable of two different size classes of prey (Daphinia magna). Fish parasitized by Glugea or Schistocephalus attack both prey types equally often, whereas sticklebacks infested by both parasite species prefer to attack the less profitable prey. The diet selected is optimal under the condition that parasites decrease their host's competitive ability.     

Love thy neighbour? Social nesting pattern,host mass and nest size affect ectoparasite intensity in Darwin’s tree finches

Social nesting behaviour is commonly associated with high prevalence and intensity of parasites in intraspecific comparisons. Little is known about the effects of interspecific host breeding density for parasite intensity in generalist host–parasite systems. Darwin’s small tree finch (Camarhynchus parvulus) on Santa Cruz Island, Galápagos Islands, nests in both heterospecific aggregations and at solitary sites. All Darwin finch species on Santa Cruz Island are infested with larvae of the invasive blood-sucking fly Philornis downsi. In this study, we test the prediction that total P. downsi intensity (the number of parasites per nest) is higher for nests in heterospecific aggregations than at solitary nests. We also examine variation in P. downsi intensity in relation to three predictor variables: (1) nest size, (2) nest bottom thickness and (3) host adult body mass, both within and across finch species. The results show that (1) total P. downsi intensity was significantly higher for small tree finch nests with many close neighbours; (2) finches with increased adult body mass built larger nests (inter- and intraspecific comparison); (3) parasite intensity increased significantly with nest size across species and in the small tree finch alone; and (4) nest bottom thickness did not vary with nest size or parasite intensity. These results provide evidence for an interaction between social nesting behaviour, nest characteristics and host mass that influences the distribution and potential impact of mobile ectoparasites in birds.     

Female great tits do not alter their yolk androgen deposition when infested with a low-transmittable ectoparasite

In birds as in many other taxa, parasites can have deleterious effects on offspring development. Therefore, avian mothers have evolved responses to counteract parasite virulence in offspring via transgenerational defense mechanisms that is the transfer of immune-enhancing substances such as antibodies to their eggs. Another maternal pathway is suggested by the finding that infested great tit mothers produced eggs with lower androgens, since these yolk androgens are immunosuppressive and potentially affect parasite susceptibility of the nestlings. However, whether this pathway is a specific adaptation to infestation with parasites that affect the offspring or an epiphenomenon of lower androgen production in the female due to the parasite effects on the mother itself is as yet unclear. In this study we infested female great tits (Parus major) with sheep ticks (Ixodes ricinus), which are nonnidicolous ectoparasites with low vertical transmission capability, and evaluated the effects on yolk androgen deposition. Tick-infested females did not significantly reduce their deposition of androgens (androstenedione (A₄) and testosterone) compared to tick-reduced females, which is in contrast to a previous study showing a lowered deposition of A₄ and testosterone when females were exposed to the nidicolous hen flea. Thus, females alter their hormone deposition, and thus likely offspring phenotype, when exposed to parasites that also form the parasitic environment of their offspring, but not when temporarily infested with the field-dwelling sheep ticks with low transmission capability. This suggests that selection favored the evolution of an adaptive transgenerational effect by acting mainly on the parasite-induced maternal effect.     

Genetic relatedness and eusociality: parasite-mediated selection on the genetic composition of groups

Summary Contrary to the expectations of kin selection theory, intracolony relatedness in eusocial insects is often low. We examined the idea that associations of low relatedness (high genetic variability) may be advantageous because of negative frequency-dependent selection on common host phenotypes by rapidly evolving parasites and pathogens. Using the natural host-parasite system of the bumble bee Bombus terrestris and its intestinal trypanosome Crithidia bombi, we investigated the transmission properties of parasites in host groups. Within naturally infested nests and in artificially constructed groups of workers, prevalence of infestation increased with time of exposure (Table 1). The susceptibility of isolated groups of workers to the parasites to which they were exposed differed with identity and natural infestation of their nest of origin (Table 2). In addition, those workers that were related to the individual introducing an infection to their group were more likely to become infested than were unrelated workers (Table 3). Although the bumble bee workers in experimental boxes appeared to differ in behavior toward kin and non-kin, making more physical contacts with kin, we found no discernible relationship between number of physical contacts and prevalence of infestation in a group. Therefore, we conclude that differences in parasite transmission reflected interactions among different host and parasite phenotypes. This system thus demonstrates the factors necessary for negative frequency-dependent selection by parasites on common host phenotypes - variability for susceptibility and infectiousness in host and parasite populations and similarity for these traits among related individuals. If, as we show here, high genetic relatedness within groups enhances parasite transmission, kin directed altruism may increase the risk of contracting parasites and infectious diseases. Therefore, parasites and pathogens may be an important force moderating the genetic structure of social groups. Offprint requests to: J.A. Shykoff at the present address     

Why are larvae of the social parasite wasp <Emphasis Type="Italic">Polistes sulcifer</Emphasis> not removed from the host nest?

A challenge for parasites is how to evade the sophisticated detection and rejection abilities of potential hosts. Many studies have shown how insect social parasites overcome host recognition systems and successfully enter host colonies. However, once a social parasite has successfully usurped an alien nest, its brood still face the challenge of avoiding host recognition. How immature stages of parasites fool the hosts has been little studied in social insects, though this has been deeply investigated in birds. We look at how larvae of the paper wasp obligate social parasite Polistes sulcifer fool their hosts. We focus on cuticular hydrocarbons (CHCs), which are keys for adult recognition, and use behavioral recognition assays. Parasite larvae might camouflage themselves either by underproducing CHCs (odorless hypothesis) or by acquiring a chemical profile that matches that of their hosts. GC/MS analyses show that parasite larvae do not have lower levels of CHCs and that their CHCs profile is similar to the host larval profile but shows a reduced colony specificity. Behavioral tests show that the hosts discriminate against alien conspecific larvae from different colonies but are more tolerant towards parasite larvae. Our results demonstrate that parasite larvae have evolved a host larval profile, which overcomes the host colony recognition system probably because of the lower proportion of branched compounds compared to host larvae. In some ways, this is a similar hypothesis to the odorless hypothesis, but it assumes that the parasite larvae are covered by a chemical blend that is not meaningful to the host.     

Larval ecology of the great barracuda, <Emphasis Type="Italic">Sphyraena barracuda</Emphasis>, and other sphyraenids in the Straits of Florida

The great barracuda (Sphyraena barracuda) is a widespread, ecologically and socioeconomically important coastal fish, yet very little is known about its larvae. We examined spawning and larval ecology of Western Atlantic sphyraenids using monthly ichthyoplankton samples collected over 2 years along a transect spanning the east–west axis of the Straits of Florida (SOF). Samples were dominated by the great barracuda (92.8%) and sennets (Sphyraena borealis and Sphyraena picudilla; 6.6%). While larval sennets and S. barracuda displayed similar vertical distributions (majority in upper 25 m), horizontal and temporal patterns of abundance suggested a spatial and temporal species replacement between larval S. barracuda and sennets that tracks adult ecology. The diet of both taxa consisted largely of copepods, with inclusion of fish larvae at 8 mm SL, and in S. barracuda alone, a switch in the wet season to exclusive piscivory by 12 mm SL (18 days post-hatch). A lack of piscivory in S. barracuda larvae captured in the dry season corresponded to slower larval growth than in the wet season. Larval growth was also related to size-at-hatch and larval age such that larvae that were larger at hatch or larger (older) at capture grew faster at earlier ages, suggesting faster larval growth, and indirectly larger hatch size, conveys a survival advantage. Unlike larval growth, instantaneous mortality rate did not differ with season, and no lunar cyclic patterns in spawning output were identified. Our results provide insight into the pelagic phase of sphyraenids and highlight the importance of both diet and hatch size to the growth and survival of fish larvae in low latitude oceanic environments.     

Otolith size-at-hatch reveals embryonic oxygen consumption in the zebrafish, <Emphasis Type="Italic">Danio rerio</Emphasis>

Otoliths have frequently been used to reconstruct growth histories in larval, juvenile and adult fish. However, there is growing evidence that otolith growth is directly determined by metabolic intensity and, consequently, only indirectly related to somatic growth. By performing measurements of oxygen consumption rate and other early life-history traits on individual eggs of zebrafish (Danio rerio), we found that oxygen consumption explained residual variance in otolith size that is not accounted for by egg size. Total oxygen consumption during the embryonic stage explained 34% of the variance in sagitta size at hatch, whereas larval size at hatch (as a proxy for growth during the embryonic period) was not significantly correlated with sagitta size. This strongly suggests that otolith growth is directly related to metabolic rate, and yields a mechanism that may explain recent observations of a link between otolith size at hatch and viability in larval fish.     

Shoaling as an anti-ectoparasite mechanism in juvenile sticklebacks (Gasterosteus spp.)

Summary In laboratory experiments, we tested the hypothesis that by living in larger shoals, juvenile threespine (Gasterosteus aculeatus) and blackspotted (G. wheatlandi) sticklebacks lower their risk of being parasitized by the crustacean ectoparasite Argulus canadensis. An increase in shoal size resulted in a lower average number of attacks received by individual fish, but had no negative effect on the attack performance (attack rate and attack success) of the parasites. In addition, more fish formed shoals and shoal sizes were larger in the presence of parasites. We conclude that ectoparasitism may have been a strong selective factor in the evolution of social behaviour in juvenile sticklebacks.     

Patterns of host switching in the fish ectoparasite <Emphasis Type="Italic">Argulus coregoni</Emphasis>

Mating is essential for sexually reproducing species to pass on their genes. For sexual parasites, this implies a need to switch the host in the search for a mate when potential partners are not available on the same host, although host switching includes risks like starvation and predation. Studies of mate-searching behavior predict that commonly only one sex searches for mating partners while the other remains stationary. In the present work, we experimentally infected rainbow trout (Oncorhynchus mykiss) with the fish ectoparasite Argulus coregoni and determined whether the sex, age and the presence of mating partners influenced the host switching activity. Our results show that increased host switching by A. coregoni occurred only after the parasite reached the adult stage and a difference between the sexes was also recorded. Host switching by mature males was enhanced when no mating partners were present on the same host, whereas females remained mainly stationary on their host regardless of male presence or absence. Our data therefore support the hypothesis that there is a strategic difference in reproductive behavior between males and females; males invest energy in mate searching while females are rather stationary and invest into body size and hence increased fecundity. Our data also showed that leaving the host to find a mate is potentially costly in terms of predation since a substantial number of free-swimming parasites were eaten by fish.     

Electrophoretic survey of genetic variation in Pecten maximus,Chlamys opercularis,C. varia and C. distorta from the Irish Sea

Food selection by young larvae of the gulf menhaden (Brevoortia patronus) was studied in the laboratory at Beaufort, North Carolina (USA) in 1982 and 1983; this species is especially interesting, since the larvae began feeding on phytoplankton as well as microzooplankton. When dinoflagellates (Prorocentrum micans), tintinnids (Favella sp.), and N1 nauplii of a copepod (Acartia tonsa) were presented to laboratory-reared, larval menhaden (3.9 to 4.2 mm notochord length), the fish larvae ate dinoflagellates and tintinnids, but not copepod nauplii. Larvae showed significant (P<0.001) selection for the tintinnids. Given the same mixture of food items, larger larvae (6.4 mm notochord length) ate copepod nauplii as well as the other food organisms. These feeding responses are consistent with larval feeding in the northern Gulf of Mexico, where gulf menhaden larvae between 3 and 5 mm in notochord length frequently ate large numbers of dinoflagellates (mostly P. micans and P. compressum) and tintinnids (mostly Favella sp.), but did not eat copepod nauplii. As larvae grew, copepod nauplii and other food organisms became important, while dinoflagellates and tintinnids became relatively less important in the diet. Since the tintinnids and nauplii used in the laboratory feeding experiments were similar in size as well as carbon and nitrogen contents, the feeding selectivity and dietary ontogeny that we observed were likely due to a combination of prey capturability and larval fish maturation and learning.Contribution No. 5575 of the Woods Hole Oceanographic Institution     

Movements of albacore tuna (Thunnus alalunga) in the South Pacific: Evidence from parasites

Parasites were collected from over 400 albacore (Thunnus alalunga) caught by surface trolling and longlining in the south-west Pacific between 1985 and 1988. Parasites found included 1 apicomplexan, 3 nematode species, 4 cestode species, 1 acanthocephalan, 12 digenean species and 3 copepod species. Twelve of these parasite species which could be accurately recognised and counted were used in the subsequent analyses. Parasite data from albacore caught around New Zealand show a decrease in prevalence of three didymozoid parasites with increasing fish length up to a fork length of 70 to 79 cm. The subsequent increase in prevalence of these didymozoids in large longline-caught fish is consistent with fish returning from spawning in tropical waters where re-infection is presumed to occur. Albacore collected at widely separate locations in the south-west Pacific have differences in parasite prevalence, supporting an hypothesis that juvenile albacore move south to New Zealand from the tropics and do not return until the onset of sexual maturity. Albacore appear to move along the subtropical convergence zone, as indicated by a decline in prevalence and abundance ofAnisakis simplex andHepatoxylon trichiuri from New Zealand to the central South Pacific. This is supported by tagging and seasonal movements of the fishery.     

Self-fertilisation in Celleporella hyalina

The accumulation of parasites in a fish host is modelled as a function of the total amount of prey consumed. The accumulated parasite load is then expressed as a function of fish length so that the asymptotic growth, L , of any population of commonly infected fish can be estimated. Estimates of L are obtained for orange roughy (Hoplostethus atlanticus), albacore tuna (Thunnus alalunga) and the New Zealand southern arrow squid (Nototodarus sloanii).     

Evidence for differential selection and potential adaptive evolution in the worker caste of an inquiline social parasite

Social parasites exploit the socially managed resources of social insect colonies in order to maximise their own fitness. The inquilines are among the most specialised social parasites, because they are dependent on being fully integrated into their host's colony throughout their lives. They are usually relatives of their host and so share ancestral characteristics (Emery's rule). Closely related inquiline-host combinations offer a rare opportunity to study trade-offs in natural selection. This is because ancestral adaptations to a free-living state (e.g. the production of a worker caste) become redundant and may be replaced by novel, parasitic traits as the inquiline becomes more specialised. The dynamics of such processes are, however, unknown as virtually all extant inquiline social parasites have completely lost their worker caste. An exception is Acromyrmex insinuator, an incipient permanent social parasite of the leaf-cutting ant Acromyrmex echinatior. In the present study, we document the size distribution of parasite and host workers and infer how selection has acted on A. insinuator to reduce, but not eliminate, its investment in a worker caste. We show that the antibiotic producing metapleural glands of these parasite workers are significantly smaller than in their host counterparts and we deduce that the metapleural gland size in the host represents the ancestral state. We further show experimentally that social parasite workers are more vulnerable to the general insect pathogenic fungus Metarhizium than are host workers. Our findings suggest that costly disease resistance mechanisms are likely to have been lost early in inquiline evolution, possibly because active selection for maintaining these traits became less when parasite workers had evolved the ability to exploit the collective immune system of their host societies.     

Infestation dynamics of gnathiid isopod juveniles parasitic on the coral-reef fish Hemigymnus melapterus (Labridae)

The dynamics of infestation by parasitic juveniles of gnathiid isopods were investigated at Lizard Island in the summer of 1997/1998 to determine when, and at what rate, they infest fishes. Variation in gnathiid abundance on wild-caught fish (Hemigymnus melapterus) between dawn and sunset was examined, and unparasitized H. melapterus in cages were exposed to gnathiids in the field for 4 h (8 h for fish sampled at 06:00 h) at five different times of the day and night (10:00, 14:00, 18:00, 22:00, 06:00 hrs) on three reefs. To control for any potential effect of the cleaner-fish Labroides dimidiatus, which consumes large numbers of gnathiids each day, sampling was carried out on reefs from which all cleaner-fish had been removed. Gnathiid abundance decreased during the day. Standardized abundance per wild-caught fish was 1.9 times higher at dawn than at sunset. Gnathiids successfully infested fish in cages. Sixty-one percent of the fish in cages were infested with ≥1 gnathiids, with 51% of the fish having between 1 and 4 gnathiids after 4 h exposure. Gnathiids infested fish both day and night. The log₁₀ (abundance +1) of gnathiids per caged fish varied significantly between time periods, with higher gnathiid numbers on fish sampled at 18:00, 22:00, and 06:00 hrs than on fish sampled at 10:00 and 14:00 hrs. In contrast to gnathiid abundance on wild-caught fish, the number of gnathiids on caged fish sampled at sunset was as high as that at dawn. The estimated mean (SE) cumulative number of gnathiids per caged fish [mean size = 13.7 cm (±0.25)] per day was 7.8 (1.1); this is similar to the estimated mean number of 7.3 gnathiids on similar-sized wild-caught fish at 6:00 hrs. The high infestation rate of gnathiids on caged fish in the late afternoon contrasted with the low numbers on wild-caught fish at this time, suggesting that factors other than infestation behaviour may be responsible for the low numbers of gnathiids on wild-caught fish in the afternoon. Received: 19 August 1998 / Accepted: 1 June 1999     

An experimental evaluation of transgenerational isotope labelling in a coral reef grouper

Transgenerational isotope labelling (TRAIL) using enriched stable isotopes provides a novel means of mass-marking marine fish larvae and estimating larval dispersal. The technique, therefore, provides a new way of addressing questions about demographic population connectivity and larval export from no-take marine protected areas. However, successful field applications must be preceded by larval rearing studies that validate the geochemical marking technique, determine appropriate concentrations and demonstrate that larvae are not adversely affected. Here, we test whether injection of enriched stable barium isotopes (¹³⁵Ba and ¹³⁷Ba) at two dose rates produces unequivocal marks on the otoliths of the coral reef grouper Epinephelus fuscoguttatus. We also assess potential negative effects on reproductive performance, egg size, condition and larval growth due to injection of adult female fish. The injection of barium isotopes at both 0.5 and 2.0 mg Ba/kg body weight into the body cavities of gravid female fish was 100% successful in the geochemical tagging of the otoliths of larvae from the first spawning after injection. The low-dose rate produced no negative effects on eggs or larvae. However, the higher dose rate of 2 mg Ba/kg produced small reductions in yolk sac area, oil globule area, standard length and head depth of pre-feeding larvae. Given the success of the 0.5 mg Ba/kg dose rate, it is clearly possible to produce a reliable mark and keep the concentration below any level that could affect larval growth or survival. Hence, enriched Ba isotope injections will provide an effective means of mass-marking grouper larvae.     

The response of an insect parasitoid,Ormia ochracea (Tachinidae), to the uncertainty of larval success during infestation

Ormia ochracea is a parasitoid fly which lays its larvae on its hosts, the field crickets Gryllus integer and Gryllus rubens, in two distinct modes: (1) directly on the host and (2) around the host. In the field, 12.7% of male crickets were parasitized and 3.2% were super-parasitized. Despite the disadvantages of parasitizing infested hosts, there was no evidence that O. ochracea avoided superparasitism. This and other experiments suggest that the host assessment ability of O. ochracea is less than that reported for many hymenopteran parasitoids. by manipulating the number of larvae in each cricket, we determined that four to five larvae per host resulted in the largest number of adult flies. However, as larval number per host increased from one to six, pupal size, and hence adult size, declined. In the field, hosts were found with a mean of 1.7 ± 1.0 (SD) larvae per cricket, suggesting that there may be some selection pressure against larger clutch sizes. Nevertheless clutch sizes larger than the host can support were sometimes found in the field. During the first mode of larviposition, gravid flies deposited no more than three larvae directly onto the host. Larvae deposited directly on the host had a high probability of infesting it. During the second mode of larviposition, gravid flies laid a larger number of larvae around the host (6.1 ± 5.2). Larvae that were laid around the host were less likely to infest a cricket than were larvae that were deposited directly onto it. O. ochracea is unique in that its two different modes of larviposition have different probabilities of larval success. Even though the success rate for larvae laid during the second mode of larviposition was low, the possibility of parasitizing more hosts appears to have selected for flies laying more larvae (e.g. increasing clutch size) than is optimal if all the larvae successfully entered a single host.     

Network structure and parasite transmission in a group living lizard,the gidgee skink, <Emphasis Type="Italic">Egernia stokesii</Emphasis>

Gidgee skinks (Egernia stokesii) form large social aggregations in rocky outcrops across the Flinders Ranges in South Australia. Group members share refuges (rock crevices), which may promote parasite transmission. We measured connectivity of individuals in networks constructed from patterns of common crevice use and observed patterns of parasitism by three blood parasites (Hemolivia, Schellackia and Plasmodium) and an ectoparasitic tick (Amblyomma vikirri). Data came from a 1-year mark-recapture study of four populations. Transmission networks were constructed to represent possible transmission pathways among lizards. Two lizards that used the same refuge within an estimated transmission period were considered connected in the transmission network. An edge was placed between them, directed towards the individual that occupied the crevice last. Social networks, a sub-set of same-day only associations, were small and highly fragmented compared with transmission networks, suggesting that non-synchronous crevice use leads to more transmission opportunities than direct social association. In transmission networks, lizards infested by ticks were connected to more other tick-infested lizards than uninfected lizards. Lizards infected by ticks and carrying multiple blood parasite infections were in more connected positions in the network than lizards without ticks or with one or no blood parasites. Our findings suggest higher levels of network connectivity may increase the risk of becoming infected or that parasites influence lizard behaviour and consequently their position in the network. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Comparison of larval fish assemblages in three large estuarine systems, KwaZulu-Natal, South Africa

 The structure of the larval fish assemblages in three large estuarine systems on the KwaZulu-Natal coast of South Africa was examined using a combination of univariate, distributional and multivariate techniques. The database was comprised of a full annual set of larval fish samples taken from each estuarine system: Durban Harbour, Richards Bay Harbour and St Lucia Estuary. The mean monthly densities of each species in each system were used in the species matrix, and the mean monthly values of salinity, temperature and turbidity were used in the environmental matrix. The mean species diversity and evenness index were significantly higher in Durban Harbour (H′ = 1.03, J′ = 0.65) than in the other two systems. The cumulative dominance curve showed that St Lucia Estuary has a high dominance of a few species, with Richards Bay Harbour intermediate and Durban Harbour being the most diverse. Classification and MDS (multiple-dimensional scaling) analyses of larval fish densities in all three systems grouped together into three main clusters on the basis of system. The species similarity matrix (inverse analysis) clustered into five groups at the 25% similarity level. The MDS analysis of the same matrix showed that the groups separated out according to the degree of estuarine association of a species and hence habitat type. The species most responsible for system groupings were: Glossogobius callidus, Gilchristella aestuaria, Stolephorus holodon, Croilia mossambica and Gobiid 12. The “best fitting” of the environmental variables to explain the larval fish community patterns in each system was turbidity on its own (weighted Spearman's rank correlation, ρ_w = 0.55). The relationship of larval densities to environmental conditions was shown to be species-specific with estuarine species (e.g. G. callidus and G. aestuaria), having a strong positive correlation with temperature and turbidity but negative correlations with salinity. In summary, much longer term studies with more sites within each system are needed to assess whether the larval fish assemblages are stable or at an equilibrium (both spatially and temporally) and whether these assemblages are indicative of the relative “health” and nursery function of the system. Received: 28 April 1999 / Accepted: 30 May 2000     

Distribution and abundance of fish larvae off the coast of West Pakistan

The material was collected during 4 different cruises on the shelf off the coast of W. Pakistan in November–December, 1964 and in March, 1967 and 1968. The hydrography as well as the zooplankton-biomass distribution are briefly described. A total of 5777 larval fish were taken in 33 positive vertical hauls by an Indian Ocean Standard Net. The number of larvae in positive hauls ranged from 2 to 1262 larvae haul^-1. The most productive area of the ichthyoplankton was the waters south of Karachi, covered during the Machera Cruise, where 828 to 1262 larvae haul^-1 were obtained. The larvae were identified to species, genera or family. Sardinella sindensis (specific identification uncertain) larvae dominated in abundance. The other abundant larvae belonged to Benthosema spp., Amentum commersonii, Vinciguerria spp. and Diaphus spp. Larval distributions and abundances are described. Aggregation and spawning of adult Sardinella (sindensis) in the waters off the coast of W. Pakistan in November/December, 1964, are discussed. Presumably, the waters south of Karachi represent good feeding grounds for s. (sindensis) larvae. Displacement volume, length and weight relationships of various larval size groups have been studied and a high rate of larval mortality (at a length of 5.0 to 8.5 mm) observed.     

Foraging behavior in early Atlantic cod larvae (Gadus morhua) feeding on a protozoan (Balanionsp.) and a copepod nauplius (Pseudodiaptomussp.)

Food limitation is likely to be a source of mortality for fish larvae in the first few weeks after hatching. In the laboratory, we analyzed all aspects of foraging in cod larvae (Gadus morhua Linnaeus) from 5 to 20 d post-hatching using protozoa (Balanion sp.) and copepod nauplii (Pseudodiaptomus sp.) as prey. A camera acquisition system with two orthogonal cameras and a digital image analysis program was used to observe patterns of foraging. Digitization provided three-dimensional speeds, distances, and angles for each foraging event, and determined prey and fish larval head and tail positions. Larval cod swimming speeds, perception distances, angles, and volumes increased with larval fish size. Larval cod swam in a series of short intense bursts interspersed with slower gliding sequences. In 94% of all foraging events prey items were perceived during glides. Larval cod foraging has three possible outcomes: unsuccessful attacks, aborted attacks, and successful attacks. The percentage of successful attacks increased with fish size. In all larval fish size classes, successful attacks had smaller attack distances and faster attack speeds than unsuccessful attacks. Among prey items slowly swimming protozoans were the preferred food of first-feeding cod larvae; larger larvae had higher swimming speeds and captured larger, faster copepod nauplii. Protozoans may be an important prey item for first-feeding larvae providing essential resources for growth to a size at which copepod nauplii are captured. Received: 20 April 1999 / Accepted: 12 January 2000