Differentiated phenotypic plasticity in larvae of the cannibalistic salamander Hynobius retardatus

Alternative phenotypes in natural populations can arise from either genetic polymorphism or an environmentally induced phenotype, that is, polyphenism. Evolutionary models of polyphenism developed by theoretical studies predict that polyphenism is favored when there are environment-dependent fitness trade-offs between alternatives and that the threshold frequency for a facultative switch between alternative phenotypes is adjusted in accordance with different selection regimes. The broad-headed (alternative) larval morph of Hynobius retardatus, which is induced by crowding with conspecifics or heterospecific anuran (Rana pirica) larvae, is a representative example of cannibalistic polyphenism. Morph induction by such proximate factors must reflect evolutionary (conditional frequency-dependent) processes. To clarify the role of frequency-dependent processes in polyphenism, I investigated the occurrence rate of the broad-headed morph under experimental crowding conditions (low conspecific, high conspecific, and high heterospecific densities) using larvae from eight natural populations with different larval densities of conspecifics and heterospecifics, and found interpopulational differences in the expression of the morph. Thus, there is a larval density-dependent equilibrium frequency of the morph in each pond, suggesting that the local switch point for morph induction was modified by selection to produce evolved differences between ponds. The evolution of such interpond differences has three necessary conditions: (1) There are pond-dependent fitness trade-offs between alternatives, (2) The maintenance of the morph is costly, and (3) The presence of conspecific or, especially, heterospecific larvae provides a reliable cue to the receiver.     

A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment.     

Kin recognition by juvenile predatory mites: prior association or phenotype matching?

Kin recognition, the biased treatment of conspecific individuals based on genetic relatedness, is a widespread phenomenon in animals. The most common mechanisms mediating kin recognition are prior association (familiarity) and phenotype matching. Recognition based on prior association allows identifying familiar individuals. Recognition based on phenotype matching is an extension of prior-association-based recognition and allows identifying familiar and unfamiliar individuals due to a shared phenotypic trait. I investigated which of the two mechanisms is used by cannibalistic juvenile predatory mites, Phytoseiulus persimilis. Protonymphs that were associated with either siblings or non-kin discriminated familiar and unfamiliar larvae and preferentially cannibalized the latter irrespective of genetic relatedness. In contrast, despite previous association with either siblings or non-kin, protonymphs did not discriminate unfamiliar sibling and unfamiliar non-kin larvae. Association in the larval stage therefore mediated kin recognition based on familiarity, but not phenotype matching in cannibalistic P. persimilis protonymphs. Furthermore, in the presence of a familiar prey individual, sibling cannibalism occurred significantly sooner than non-kin cannibalism. This quick sibling cannibalism may have been the consequence of preferential association of siblings and/or may indicate the occurrence of an alternative cannibal phenotype. I discuss the adaptive significance of prior-association-based recognition for P. persimilis juveniles and emphasize the ability of P. persimilis to use multiple recognition mechanisms in dependence of the ontogeny and the ecological context.     

The selective advantages of cannibalism in a Neotropical mosquito

We investigated the benefits of larval cannibalism in the Neotropical mosquito Trichoprosopon digitatum. The clutch size of the mosquito in the field was strongly correlated with adult female size, indicating a fitness advantage to being large. In controlled laboratory experiments, we compared the survivorship and eventual adult sizes of larvae that were given the opportunity to cannibalise conspecifics throughout their lifetimes with the survivorship and adult sizes of larvae that were prevented from cannibalising. Since the benefits of cannibalism are likely to depend on the context in which it occurs, the experiment was conducted at two levels of alternative food availability. When food availability was high most larvae survived to adulthood, females cannibalised more than males and there was no measurable advantage to cannibalism in terms of survival rate, emergence time or adult size. Larvae were significantly more cannibalistic when food availability was lower, although under these conditions no larvae survived to adulthood. Nevertheless, under low food an important fitness benefit to cannibalism was revealed: individuals which had the opportunity to cannibalise survived significantly longer as larvae than those which did not. This increased longevity is likely to provide an important advantage to mosquito larvae when they wait for the input of unpredictable food sources. Received: 7 October 1995/Accepted after revision: 13 April 1996     

Induction of larval settlement and metamorphosis of the sea urchin Strongylocentrotus intermedius by glycoglycerolipids from the green alga Ulvella lens

In aquaculture centers of the northern region of Japan, "Nami-ita" (waved polycarbonate plates), on which the green alga Ulvella lens Crouan frat. (Chaetophoraceae: Chaetophorales) was cultured, are used to promote larval settlement and metamorphosis of the sea urchin species Strongylocentrotus intermedius (A. Agassiz) and S. nudus (A. Agassiz). We investigated chemical inducer(s) for larval settlement and metamorphosis of these sea urchins with extracts of U. lens. Bioassay-guided separation of the methanol extract using a combination of column and thin-layer chromatography led to the isolation of several active compounds, the chemical structures of which were determined by spectral and chemical methods. These active compounds were identified as glycoglycerolipids, all comprising several molecular species: sulfoquinovosyl monoacylglycerols (SQMGs), sulfoquinovosyl diacylglycerols (SQDGs), monogalactosyl monoacylglycerols (MGMGs), monogalactosyl diacylglycerols (MGDGs), digalactosyl monoacylglycerols (DGMGs) and digalactosyl diacylglycerols (DGDGs). Among these glycolipids, SQMGs, MGMGs, MGDGs and DGMGs induced larval metamorphosis of the sea urchin S. intermedius. SQMGs and MGDGs induced larval metamorphosis at a concentration of 5 µg ml^-1, whereas SQDGs and DGDGs only induced larval settlement. These glycoglycerolipids are new congeners of chemical inducers to settlement and metamorphosis of planktonic larvae of sea urchins. The findings would provide a better understanding of larval settlement and metamorphosis in sea urchins.     

Recruitment in the intertidal limpet Lottia digitalis (Patellogastropoda: Lottiidae) may be driven by settlement cues associated with adult habitat

The ribbed limpet, Lottia digitalis, is found high in rocky intertidal habitat throughout its geographic range. In order to identify likely natural settlement locations for larvae of this species, laboratory-reared larvae were settled onto substrata collected from within and near an adult L. digitalis habitat. Of larvae exposed to rock chiseled from within high-intertidal adult habitat, 31.0% and 23.3% underwent metamorphosis during two separate experiments. Similarly, an unidentified filamentous green alga that was isolated from this rock induced metamorphosis in 26.6% and 8.7% of larvae during additional experiments. In contrast, larvae did not metamorphose upon bare rocks or upon rocks encrusted with a crustose corraline alga (CCA) that were collected from lower intertidal zones, nor did they metamorphose upon the macroalgae Ulva sp., Enteromorpha contorta, Alaria marginata, or Polysiphonia sp. The presence of mucus from adult conspecifics during these experiments did not enhance metamorphosis onto rock taken from adult habitat, but it did induce metamorphosis in an average of 13.5% and 7.0% of larvae introduced to the mid-intertidal bare rock and CCA substrata, respectively. Finally, 38.0% and 34.4% of larvae from two experiments underwent metamorphosis when exposed to the high-intertidal barnacle Pollicipes polymerus. These results suggest that larval settlement, rather than differential post-settlement mortality and/or migration, drives recruitment of L. digitalis in high-intertidal habitats.     

Settlement patterns of newly settled plaice (Pleuronectes platessa) in a non-tidal Swedish fjord in relation to larval supply and benthic predators

In demersal fish species with a pelagic larval stage, settlement patterns may be a consequence of variations in larval supply, habitat selection at settlement, and processes acting between the time of settlement and the time of benthic sampling. This study describes temporal (1994-1998) and spatial variation in plaice (Pleuronectes platessa L.) settlement densities in four semi-isolated nursery areas with similar habitat characteristics, in the non-tidal Gullmarsfjord on the west coast of Sweden. Juvenile abundance varied by a factor of ten, both among years and among nursery grounds. For the 3 years when larval sampling was undertaken (1994-1996) and all nursery areas, there was a significant positive relationship between larval supply and juvenile abundance (linear regression: r²=0.45, n=24, P<0.001). On the southern side of the fjord, a significant positive relationship between larval and juvenile abundance was found in one area (r²=0.62, n=6, P<0.05). The absolute mortality rate of plaice after settlement was related to the initial settlement density (r²=0.95, n=20, P<0.001), and to the abundance of predatory shrimps Crangon crangon (r²=0.44, n=20, P<0.01). Plaice otoliths were found in 6% of the shrimp stomachs analysed from an area with high density (13.3 m^-2) of newly settled plaice. The present study suggests that the density of juvenile plaice was limited by larval supply to the nursery grounds. Consistency in the relative abundance of juveniles among nursery grounds between years also suggested that some nursery areas may be in the settlement shadow of others. The irregular nature of the coastline in combination with larval depletion could thereby cause small-scale (10³-10⁴ m) variation in settlement densities of the same order of magnitude as the inter-annual variability in recruitment to individual nursery grounds.     

Early growth and mortality of the Caribbean king crab Mithrax spinosissimus reared in the laboratory

The berried females of the Caribbean king crab Mithrax spinosissimus (Lamarck) used in this study were collected from canals on Big Pine Key, Sugarloaf Key and Lower Matecumbe Key (south Florida, USA) on 9 August, 8 October and 15 November 1986. Viable spawns hatched as first zoeae and molted to second zoeae within ca. 10 to 12 h. Most of the larvae reached the megalopa stage 1 d later, and molted to first crab 4 to 8 d after hatching (water temperature: 27.2° to 28.8°C). Low water temperature and/or early lack of food had a negative effect not only on stage duration, but also on the size of the early crab stages. Successful molt to first crabs occurred, however, in the absence of food. The growth rate (carapace length) between molts in early crab stages varied between ca. 20 and 30%. When provided with good water exchange, stocking density could be very high (>22 500 individuals m^-2), with no increase in mortality. The highest mortality rate was recorded when the larvae molted to first crab, and the highest rates of survival were always recorded when feeding was not initiated until after 5 to 8 d after hatching. No cannibalism was observed among larvae, and cannibalism was low in early crab stages. The study indicates that to achieve viable hatches and high larval survival in rearing M. spinosissimus, a continuous and adequate supply of high-quality seawater is a prerequisite both in larviculture and in maintaining brooding females.Contribution No. 93, Department of Oceanography and Ocean Engineering, Florida Institute of Technology     

Effects of algal and larval densities on development and survival of asteroid larvae

Effects of larval and algal culture density and diet composition on development and survival of temperate asteroid larvae were studied in the laboratory at Santa Cruz, California, USA, during summer and fall of 1990. Larvae of Asterina miniata were reared at two densities, 0.5 or 1.0 ml^-1, and fed one or two species of cultured phytoflagellates — Dunaliella tertiolecta alone or mixed with Rhodomonas sp. — at three concentrations of 5x10², 5x10³, and 5x10⁴ total cells ml^-1. Algal concentration strongly influenced larval development; however, larval density also had a marked effect. Development progressed further with increasing algal concentration. Larval growth and differentiation were sometimes uncoupled; i.e., growth measures were directly related to food level, while differentiation indicators were less so. At the lowest food level, growth was negative and differentiation was arrested at early precompetent stages; these larvae never formed juvenile rudiments or brachiolar attachment structures. Development times of larvae given more food ranged from 26 to 50 d and depended directly on food availability. Development time to metamorphosis at the highest food concentration was similar for siblings fed D. tertiolecta alone or mixed with Rhodomonas sp. In contrast, when food level was an order of magnitude lower, larvae fed the algal mixture metamorphosed significantly earlier than larvae fed the unialgal diet. This suggests interactive effects of food quantity and food quality. Survival was little affected by larval or food density, except at the lowest ration. Feeding experiments in well-controlled laboratory conditions are useful to predict and compare the physiological or developmental scope of response of larvae to defined environmental factors; however, results from such studies should not be extrapolated to predict rates and processes of larval development in nature.     

Strange parental decisions: fathers of the dyeing poison frog deposit their tadpoles in pools occupied by large cannibals

Parents may increase the probability of offspring survival by choosing suitable rearing sites where risks are as low as possible. Predation and competition are major selective pressures influencing the evolution of rearing site selection. Poison frogs look after their clutches and deposit the newly hatched tadpoles in bodies of water where they remain until metamorphosis. In some species, cannibalism occurs, so parents deposit their tadpoles singly in very small pools. However, cannibalism also occurs in species that deposit tadpoles in larger pools already occupied by heterospecific or conspecific larvae that could be either potential predators or competitors. Here, I test the hypothesis that, given the choice, males of Dendrobates tinctorius would deposit their newly hatched tadpoles in low-risk sites for their offspring. I characterised the pools used by D. tinctorius for tadpole deposition, conducted experiments to determine the larval traits that predict the occurrence of and latency to cannibalism, and tested whether parents deposit their tadpoles in low-risk pools. I found that (1) neither pool capacity nor the presence of other larvae predict the presence/absence or number of tadpoles; (2) cannibalism occurs often, and how quickly it occurs depends on the difference in size between the tadpoles involved; and (3) the likelihood of males depositing their tadpoles in occupied pools increases with the size of the resident tadpole. I suggest that predation/cannibalism is not the only factor that parents assess when choosing deposition sites, and that the presence of larger conspecifics may instead provide information about pool quality and stability.     

Does cannibalism in Spodoptera frugiperda (Lepidoptera: Noctuidae) reduce the risk of predation?

The incidence of cannibalism of larval Spodoptera frugiperda (Lepidoptera: Noctuidae) on maize under field conditions was investigated using field cages. Cannibalism was found to account for approximately 40% mortality when maize plants were infested with two or four fourth-instar larvae over a 3-day period. Field trials examined the effect of larval density on the prevalence of natural enemies of S. frugiperda. The abundance of predators (earwigs, staphylinids, other predatory beetles, and Chrysoperla spp.) was significantly greater on maize plants with higher levels of larval feeding damage, while the relationship between predator abundance and number of S. frugiperda larvae per plant was less clear. As larval damage is probably a more reliable indicator of previous larval density than numbers collected at an evaluation, this indicates that predation risk will be greater for larvae living in large groups. Parasitism accounted for 7.1% mortality of larvae in sorghum, and involved six species of Hymenoptera and Tachinidae. There was no effect of larval density or within-plant distribution on the probability of larval attack by parasitoids. The selective benefits of cannibalism, in relation to the risk of predation and parasitism, are discussed. Received: 23 March 2000 / Accepted: 24 June 2000     

Development of the escape response in larval walleye pollock (Theragra chalcogramma)

The development of the escape response of walleye pollock (Theragra chalcogramma) larvae from attacks by macrozooplanktonic and small-fish predators was quantified in laboratory experiments. Behavior was recorded using video cameras with silhouette illumination from infrared-emitting diodes and by visual observation. Laboratory-reared larvae of 1, 3, 8, 10, 12, 18, 22, 27, 42 days post-hatching, ranging in size from 4 mm to 10 mm total length, were used in the experiments. Even the youngest larvae were observed to exhibit a fast startle response. The percentage of successful larval escapes from the different predators increased as the larvae developed. Euphausiids (Thysanoessa raschii) and amphipods (Calliopiella pratti) often touched larvae but the larvae were usually able to escape and no successful captures of larvae over 22 days old were observed. Although successful escape from initial attacks by three-spine sticklebacks (Gasterosteus aculeatus) increased ontogenetically, sticklebacks were able to consume most larvae, even of the oldest age group, by repeated attacks. Day-old larvae had the lowest percent of escapes after encounters with jellyfish (Sarsia sp.), but the percentage of escapes increased dramatically for 3-day-old larvae. Escape speeds after an attack also increased with age, and tended to be higher after stickleback attacks and lower after jellyfish attacks. This study revealed that the escape response of larval pollock to attack by predators improves rapidly with development during the early larval stage.     

Ecology of kin and nonkin larval interactions in Tribolium beetles

Summary The larvae of flour beetles Tribolium castaneum and T. confusum were reared in two kinds of groups: full siblings and unrelated individuats. These kin and nonkin groups were reared in open cultures, in which emigration was permitted (both species) and in closed cultures, in which emigration was prohibited (only T. confusum). We measured larval development time and survivorship, weight of pupae, and time of larval emigration from open cultures. The effects of age structure were investigated by establishing open cultures of larvae of uniform age (larvae hatched from eggs laid within 72 h) and cultures of larvae of variable age (eggs laid within 240 h). In closed cultures of siblings, T. confusum larvae pupated on an average 2.2 days earlier than larvae reared in nonsibling groups. In T. castaneum, more small and medium size and fewer large size larvae emigrated from groups of siblings compared to groups of nonsiblings. Males that emigrated and pupated remained with their sibs for a shorter time than did similar males raised with unrelated larvae. In T. castaneum, age structure variation reduced the sibs tendency to migrate, but did not influence interactions among unrelated larvae. The genetical effects of kinship and the ecological effects of age structure were shown to affect the interactions of Tribolium larvae reared in groups. Reducing the similarity between individuals, either genetically or demographically (using mixed broods or mixed age cohorts), changed the pattern of larval interactions. Upon occasion, the effects of kin interactions may well be the mechanical consequences of the coexistence of similar individuals rather than the effects of altruistic behavior.     

The behavioral ecology of cannibalism in cane toads (<Emphasis Type="Italic">Bufo marinus</Emphasis>)

Laboratory studies show that predatory cane toads (Bufo marinus) exhibit specialized toe-luring behavior that attracts smaller conspecifics, but field surveys of toad diet rarely record cannibalism. Our data resolve this paradox, showing that cannibalism is common under specific ecological conditions. In the wet–dry tropics of Australia, desiccation risk constrains recently metamorphosed toads to the edges of the natal pond. Juvenile toads large enough to consume their smaller conspecifics switch to a primarily cannibalistic diet (67% of prey biomass in stomachs of larger toads). Cannibalistic attack was triggered by prey movement, and (perhaps as an adaptive response to this threat) small (edible-sized) toads were virtually immobile at night (when cannibals were active). Smaller metamorphs were consumed more frequently than were larger conspecifics. The switch from insectivory to cannibalism reflects the high dry season densities of small conspecifics (in turn, due to desiccation-imposed constraints to dispersal) and the scarcity of alternative (insect) prey during dry weather. Our study pond (102 m in circumference) supported >400 juvenile toads, which consumed many metamorphs over the course of our study. Toads appear to be low-quality food items for other toads; in laboratory trials, juvenile toads that fed only on conspecifics grew less rapidly than those that ate an equivalent mass of insects. This effect was not due to parotoid gland toxins per se. Thus, cane toads switch to intensive cannibalism only when seasonal precipitation regimes increase encounter rates between large and small toads, while simultaneously reducing the availability of alternative prey.     

Feeding ecology of the three juvenile phases of the spiny lobster<Emphasis Type="Italic"> Panulirus argus</Emphasis> in a tropical reef lagoon

The three juvenile phases of the spiny lobster Panulirus argus (algal phase: 5-15 mm carapace length, CL; postalgal phase: 15-45 mm CL, and subadults: 45-80 mm CL) occur in the reef lagoon at Puerto Morelos, Mexico. The algal phase abounds in this lagoon, which is covered by extensive seagrass-algal meadows, but the density of postalgal and subadult juveniles is low, owing to the scarcity of crevice-type shelters suitable for these phases. The feeding ecology of the three juvenile phases was investigated to examine whether spatial or temporal differences in food intake, diet composition, or nutritional condition occurred among phases and could partially account for the low abundance of the larger juveniles. Juveniles were collected by divers at night, from January to November 1995, throughout the mid-lagoon and back-reef zones. Percent stomach fullness, relative weight of the digestive gland (RWDG, an index of nutritional condition), percent frequency of occurrence and percent volume of food categories in the diet were compared between sexes, juvenile phases, molt stages (postmolt, intermolt, premolt), seasons, and sampling zones (mid-lagoon and back-reef zones). Significant differences in stomach fullness occurred only among molt stages, mainly because postmolt individuals had emptier stomachs. The main food categories in all juvenile phases were crustaceans (mostly hermit crabs and brachyurans) and gastropods, but the food spectrum was wide, including many other animal taxa as well as plant matter. In June 1995, the epibenthic macrofauna was sampled in five sites in the lagoon that differed in their amount of vegetation. The most abundant taxa in all sites were decapods and gastropods, but density and diversity measures showed that the distribution of these potential prey taxa for juvenile P. argus was rather patchy. Diet overlap in juvenile lobsters was high between sexes, juvenile phases, sampling zones, seasons, and molting stages, indicating that all juveniles fed on the same general food categories throughout time. The only factor that affected the RWDG was the juvenile phase. RWDG was significantly lower in subadults than in algal and postalgal phases, suggesting a poorer nutritional condition in the largest juveniles. This may be related to the scarcity of suitable shelters for large juveniles throughout the lagoon, which may preclude subadults from exploiting food resources in areas of the lagoon where shelter is limited.     

Leptocephalus energetics: assembly of the energetics equation

The principles of energetics were used to examine the energetic requirements of leptocephali. Respiration and excretion rates and daily growth rates combined with proximate composition were used to examine the allocation of energy into each of the three main components of energetics: metabolism, excretion and growth. The daily energetic requirements for leptocephali, referred to as type 2 larvae based upon their unique developmental strategy, were compared to the requirements of non-leptocephalus larvae, known as type 1. Leptocephalus daily energetic requirements were also compared to the energy available from the leptocephalus' proposed food sources. The four species of eel larvae selected were all from the order Anguilliformes: Paraconger caudilimbatus (Poey), Ariosoma balearicum (Delaroche), Gymnothorax saxicola Jordan and Davis, and Ophichthus gomesii (Castelnau). The allocation of energy to each of the components of energetics as well as the total energetic requirements for the leptocephali proved to be very different from those of type 1 larvae. Metabolism received the majority, 60-92%, of the energy required per day. Growth and excretion were allocated 4-39% and <1-21%, respectively, of the total energy needed per day. Leptocephali required <50% of the energy needed by type 1 larvae of equal dry mass. The unique growth strategy used by leptocephali allows them to increase rapidly in size while allocating the majority of their energy, not to growth as in most larval fish, but to metabolism.     

Specificity and sensitivity of microsatellite markers for the identification of larvae

The identification of larval marine invertebrates to species or even higher taxonomic levels by morphological examination is notoriously difficult. Many diagnostic features are absent or poorly formed at early stages in development. This is particularly true for the larvae of bivalve molluscs, for which a routine and accurate method of identification would prove valuable to both ecologists and fishery managers. A simple molecular genetic method to identify specifically larvae of the European oyster, Ostrea edulis L., 1758, is presented. The test is based on PCR amplification of highly species-specific microsatellite loci and is sensitive enough to register the presence of a single larval individual (~200 µm width) in a mixed sample of 20 mg wet weight plankton (approximately 250 larval animals). This work demonstrates that microsatellite loci can be used as highly sensitive and specific taxonomic indicators, for studies of planktonic larvae. Details of three novel microsatellite loci are also given for O. edulis, increasing the suite of molecular tools available for use in population genetic studies of this commercially important species.     

Planktonic larval duration and settlement marks on the otoliths of Mediterranean littoral fishes

The planktonic larval duration (PLD) was estimated for 42 species of littoral fishes from the north-western Mediterranean Sea. Daily increments and settlement marks on the otoliths (sagittae or lapilli) of new settlers and post-settlers were used to determine the larval stage duration. We also used PLD in the new settlers of some species to confirm the accuracy of the settlement marks in post-settlers. The duration of the planktonic larval stage ranges from 9 days in Symphodus ocellatus to 55 days in Xyrichtis novacula and 71 days in Lipophrys trigloides. Species in the same family did not display any clear tendency toward having similar PLDs. On the other hand, larval duration tended to be similar within a genus, with the exception of Lipophrys. Among conspecifics, the time spent in the plankton usually varied only by 2-7 days, except in Aidablennius sphynx, Lipophrys trigloides, Coris julis and Thalassoma pavo. No clear patterns were discernible in genera, with some species that settled in winter and other species that settled in summer, although we observed a certain tendency of individuals of closely related species (e.g. family Sparidae) to have a shorter larval duration in the warmer part of the year than in the colder part of the year. Settlement marks have been observed on the otoliths of all the species studied, and the PLDs in new settlers are an appropriate means of validating settlement marks. A rapid decrease in increment width over settlement (type Ia) is the most common type of mark (66.7% of the total species studied).     

Effect of prey densities on cannibalism in Cape hake (Merluccius capensis) off Namibia

The preference of large (>60 cm total length) Cape hake (Merluccius capensis Castelnau, 1861) for small conspecifics under natural conditions was analyzed off southern Namibia (S.E. Atlantic) between 1983 and 1989. The results indicate that cannibalism is not related to the density of the small conspecifics nor to the density of alternative prey, but rather that large hake have a dietary preference for small conspecifics. The main consequence of such cannibalism is a lack of density-dependent regulation of the M. capensis population.     

Feeding behavior and acquisition of zooxanthellae by planula larvae of the sea anemone Anthopleura elegantissima

Symbiotic associations between cnidarians and photosynthetic dinoflagellates (i.e., zooxanthellae) are common in the marine environment. Many symbiotic cnidarians produce offspring that are initially nonsymbiotic. These new hosts must acquire symbiotic algae from environmental sources. We examined zooxanthella acquisition by laboratory-reared planula larvae of the temperate sea anemone Anthopleura elegantissima. Larvae ingested zooxanthellae while they were feeding. However, the signal that prompted larval feeding behavior did not originate from the symbiotic algae; the addition of algal cells to larval cultures never elicited a feeding response. In contrast, the addition of macerated animal tissue from several sources invariably generated a strong feeding response, which resulted in the larvae indiscriminately ingesting any particulate matter that was present, including zooxanthellae or other unicellular algae. Ingested zooxanthellae were incorporated into endodermal cells, where they remained undigested, while all other ingested material was digested or expelled within 24 h. Our results provide evidence that one source of zooxanthellae likely to serve as a route of infection in the natural environment is zooxanthella-laden mucus egested by anemones. This egested material fulfilled both of the criteria necessary for successful infection: it prompted larvae to begin feeding and provided an abundant supply of zooxanthellae that were ingested and taken up into endodermal cells of the new host.