Factors regulating the reproductive cycles of two northeast pacific chitons,Tonicella lineata and T. insignis

The reproductive cycles of Tonicella lineata Wood and T. insignis Reeve are examined in relation to changes in physical and biotic conditions in different locations in British Columbia, Canada, over a 3 to 4 year period. A clearly annual reproductive cycle is displayed by both species, and in any one location and year reproductive events in the two species are similar. There is a progressive increase in the size of the gonads starting in the summer and continuing to mid-winter. Although no one environmental factor is likely to be correlated with this prolonged period of gonadal development, various phases of the annual temperature and photoperiod cycles do appear to coincide with particular phases of gametogenesis (such as gonial proliferation, vitellogenesis and gonadal maturation), and thus could act as external cues. There isa synchronous drop in gonadal size in T. lineata and T. insignis in the spring, as a result of spawning. The exact timing and abruptness of this event in different years can not be accounted for on the basis of temperature. However, new correlative data are presented to add further support to the hypothesis previously presented that the spring phytoplankton is the cue for natural spawning (Himmelman, 1975). Temperature may inhibit spawning in T. lineata if conditions are unseasonably cold.     

Annual reproductive cycle of the chiton Acanthopleura japonica

Annual reproductive cycle of the chiton Acanthopleura japonica (Lischke) was studied by observing both gonad index and histological features from October 1980 to October 1981. No significant changes in each factor were observed from October to April. Increased values of gonad index and spermatogenesis or vitellogenesis in gondas were observed from May to September. In the same time, gonad index decreased and sperms or oocytes in gonalds disappeared in the days around spring tide, and they were recovered around neap tide. Suggested spawning around spring tide from May to September is also supported by the occurrence of released eggs in the days around the new and full moons.     

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.     

Seasonal breeding aggregations in low-density populations of the bathyal echinoid Stylocidaris lineata

Reproduction of bathyal cidaroid sea urchins was studied between 1985 and 1991 with the aid of a manned submersible in the northern Bahamas. During May 1988 and February 1990, discrete, single-species aggregations of Stylocidaris lineata containing 2 to 6 individuals were observed between 510 and 640 m depths. Reproductive conditions of isolated and aggregated individuals were determined histologically. In February, all individuals, regardless of their spatial distribution, contained developing gametes. During the May spawning season, most isolated individuals contained spent gonads with only relict gametes, whereas individuals in discrete aggregations mostly contained ripe gonads with mature or nearly-mature gametes. Aggregations were not observed during the autumn months, when gonads were spent or immature. These data suggest that bathyal cidaroids aggregate for reproduction. An in situ survey during May indicated that pairs of S. lineata are more common than larger clumps, but no more so than predicted by chance. Homosexual and heterosexual pairs occur at the relative frequencies predicted on the basis of sex ratio. We evaluated the need for reproductive aggregation by applying Denny's (1988) model that predicts downstream sperm concentration in a turbulent boundary layer. Flow parameters were measured with dye injected from the submersible, sperm-release rate was estimated in the laboratory, and the relationship between fertilization success and sperm concentration was determined in a laboratory dilution-experiment. The model predicted that, on smooth sandy bottoms at bathyal depths, sperm concentration should remain high enough to fertilize at least some eggs several meters directly downstream from a spawning male. Aggregation may facilitate spawning synchrony, increase gamete encounter probabilities, or cause gametes to be retained at high concentrations near the adults long enough for fertilization to occur.     

Reproductive cycle of the chiton Chiton iatricus and environmental control of its gonad growth

The annual reproductive cycle of the chiton Chiton iatricus was studied for the period from September, 1977 to December, 1978. Gametogenesis is initiated in the November – December period followed by rapid gonadal growth up to March. Breeding season of C. iatricus extends up to the beginning of August. Though the emission of the gametes halts in August, the gonad never enters into the quiescent phase owing to the time required for the gametogensis. The examination of correlation between the reproductive events during the annual reproductive cycle and existing environmental factors suggests that multiple environmental factors are involved in the control of different phases of gonad growth. The collected data are discussed in the light of the reproductive physiology of C. iatricus.     

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.     

Effects of gamma-aminobutyric acid on the settlement of larvae of the black chiton Katharina tunicata

Laboratory-cultured larvae of the black chiton Katharina tunicata (Wood), collected from central California in May and June of 1979 and 1980, settled preferentially on the encrusting coralline alga Lithothamnium sp. Metamorphosis, which involves the loss of the prototrochal ciliary band, occurred within a period of 2.5 h after larvae began crawling upon the alga. In response to gamma-aminobutyric acid (GABA) at a concentration of 10^-6 M, the lecithotrophic trochophore larvae ceased swimming and settled rapidly. Developmental metamorphosis did not follow settlement in the absence of encrusting coralline alga. However, when both GABA and Lithothamnium sp. were present, larval metamorphosis occurred within 2 h of settlement. Significantly different rates of settlement exist for sibling larvae of different ages; 12 d-old larvae responded more rapily and in greater numbers to treatment with GABA than 10 d-old larvae. Differences in settling rates are attributed to the onset of metamorphic competence.     

Observations on the reproductive cycles of Monodonta lineata,Gibbula umbilicalis and G. cineraria

Monthly samples of gonads of 3 common species of intertidal trochids, Monodonta lineata (da Costa), Gibbula umbilicalis (da Costa) and G. cineraria (L.) were fixed for examination by light microscopy. Numbers and proportions of immature and mature oocytes in sections of ovaries were determined. In M. lineata and G. umbilicalis, the annual reproductive cycles were very similar, the testis and ovary of both species being empty brown structures from September to January/February, when gametogenesis began and the gonads began to swell and turn green in both sexes. As ripening continued, the ovary remained green in G. umbilicalis, but in M. lineata turned greygreen by the end of May. The testis of both species changed, to yellowish in G. umbilicalis and creamy-pink in M. lineata. Spawning occurred throughout summer and early autumn, and the gonads reverted to their former empty state. These changes were matched by the increase and subsequent decrease in proportion of ripe oocytes in the gonad. Some ripe oocytes are not shed during the spawning season, but no evidence of cytolysis of such oocytes was found. In G. cineraria, in contrast, the ovary remained green and swollen and the testis remained creamy-white and packed with sperm throughout the year. The proportion of ripe to unripe oocytes in the ovary fluctuated for the entire sampling period, and did not indicate the time of spawning. In the testis of all three species, spermatogonia were rarely seen, and spermatid stages were uncommon.     

Benthic invertebrate larval response to substrate characteristics at settlement: Shelter preferences of the American lobsterHomarus americanus

The factors controlling shelter selection by the first post-larval stage of the American lobsterHomarus americanus were studied in the laboratory in 1988 using an experimental approach. The relative importance of shelter size, light level within shelters, number of shelter entrances, and presence of algal-film cover in and around them were investigated. Post-larvae showed a strong preference for opaque versus transparent shelters. No difference in post-larval selectivity was observed either for shelters of different widths or heights or between shelters with different numbers of entrances. Post-larvae showed no preference for microhabitats colonized by microorganisms compared to others without colonization. The intensity of the response measured suggests a possible hierarchical organization of the different cues involved in shelter selection, the light level within the shelter apparently being the most important. Simple linear regression was used to assess the effect of shelter area on the number of post-larvae occupying a shelter. Up to four post-larvae were observed simultaneously in a shelter of 40 cm² basal area, indicating that they can cohabit at settlement. The minimum shelter area necessary to the first post-larval stage was estimated to be about 3.25 cm². The results are discussed with reference to recent hypotheses concerning the mechanisms controlling larval settlement in other benthic species with relatively weak mobility during the larval and the post-larval phases.Contribution to the programmes of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec) and of the Institut Maurice-Lamontagne     

Settlement of larval blacklip abalone, Haliotis rubra, in response to green and red macroalgae

Surfaces from the habitat of adult Haliotis rubra were tested as inducers of larval settlement to determine the cues that larvae may respond to in the field. Settlement was high on the green algal species Ulva australis and Ulva compressa (Chlorophyta), the articulated coralline algae Amphiroa anceps and Corallina officinalis, and encrusting coralline algae (Rhodophyta). Biofilmed abiotic surfaces such as rocks, sand and shells did not induce settlement. Ulvella lens was also included as a control. Treatment of U. australis, A. anceps and C. officinalis with antibiotics to reduce bacterial films on the surface did not reduce the settlement response of H. rubra larvae. Similarly, treatment of these species and encrusting coralline algae with germanium dioxide to reduce diatom growth did not significantly reduce larval settlement. These results suggest that macroalgae, particularly green algal species, may play an important role in the recruitment of H. rubra larvae in the field and can be used to induce larval settlement in hatchery culture.     

The adaptive value of larval behavior of a colonial ascidian

Didemnum candidum Savigny, 1816 is an abundant member of the fouling community encrusting docks in Pearl Harbor, Hawaii. In the laboratory, larvae of D. candidum are positively phototactic upon release, but become indifferent to the direction of light prior to settlement. In both the laboratory and the field, settlement is most abundant on shaded, downward-facing surfaces, the location where juvenile survival is greatest. The colonial ascidians Diplosoma listerianum Milne-Edwards, 1842 and Diplosoma sp. also settle abundantly on shaded undersurfaces in the field. Juvenile survival of the Diplosoma species is not affected by exposure to direct light, while survival is greatest on downward-facing surfaces. For Didemnum candidum, light is an important environmental cue used by larvae to locate settlement sites on shaded, downward-facing surfaces where juvenile survival is enhanced.     

Analytical expression for predicting the reduced settling velocity of small particles in turbulence

The influence of turbulence on the settling velocity of small particles remains an inconclusive research subject. Both enhanced and retarded particle settling compared to quiescent settling have been reported in previous literature, and several theories have been proposed. Among the mechanisms that account for reduced settling velocities in turbulence, the loitering effect is an important one that generally exists in various conditions. This study focuses on an analytical prediction of reduced particle settling velocities due to the loitering effect. By considering the velocity autocorrelation function as a step function analogous to the free path theory, and using a modified integral time scale to capture the essence of the loitering effect, an analytical expression is derived for predicting the reduction in particle settling velocities in turbulent flows. Calculation results of the expression are then examined by comparing with the results of a random walk model, direct numerical simulations that conditionally captured the reduction of particle settling velocities due to the loitering effect, and several representative experiments. Major possible influencing factors on the prediction of the analytical expression and applicable conditions of the expression are then further discussed. The proposed analytical expression is shown to be suitable for predicting the reduced settling velocities of small particles with relatively weak inertia in turbulent flows and could provide a reasonable explanation for reported cases in which small to moderate reductions in particle settling velocities were observed.

     

Fertilisation, embryogenesis and larval development in the tropical intertidal sand dollar Arachnoides placenta in response to reduced seawater pH

We examined the response of the tropical sand dollar Arachnoides placenta to reduced seawater pH in experiments spanning ca. 50 % of the planktonic larval duration. A. placenta inhabits intertidal sandy beaches where we observed a minimum in situ pH range 0.06 pH units (pH 8.10–8.16). The responses of gametes and larvae to seawater pH were tested in vitro in ambient (pH 8.14, pCO₂ = 525.7 μatm, total alkalinity = 2,651 μmol kg soln^?1) and three reduced pH seawater treatments (7.8–7.0). Percentage fertilisation decreased significantly with decreasing pH across a range of sperm/egg ratios (4:1 up to 4,000:1). A. placenta reached the advanced pluteus stage in 4 days, and during this time, we saw no difference in survival rate of larvae between the ambient (67 %) and pH 7.79 (72 %) treatments. Four-day survival was, however, reduced to 44 and 11 % in the pH 7.65 and 7.12 treatments, respectively. Larval development and morphometrics varied among pH treatments. Embryos reared in pH 7.12 exhibited arrested development. Larvae reared at pH 7.65 showed delayed development and greater mortality compared with those reared at pH 7.79 and 8.14. When larval morphometrics are compared among larvae of the same size, differences in larval width and total arm length between pH treatments disappear. These results suggest that variation in larval size among the three highest pH treatments at a given time are likely the result of slower development and apparent shrinkage of surviving larvae and not direct changes in larval shape. There were no differences in the percentage inorganic content (a proxy for calcification) in larvae reared in either an ambient or a pH 7.7 treatment. The responses of fertilisation and development to decreased pH/increased pCO₂ in A. placenta are within the range of those reported for other intertidal and subtidal echinoid species from colder latitudes.     

Larval settlement in the serpulid polychaete Hydroides elegans in response to bacterial films: an investigation of the nature of putative larval settlement cue

Larval settlement in the marine polychaete Hydroides elegans (Haswell) is induced by certain bacteria in marine biofilms. The exact nature of the settlement cue that larvae of H. elegans receive from these bacteria remains unknown. In this study, we revealed some properties of the bacterially derived larval settlement cue by investigating the larval settlement inductive activity of two bacterial strains after various treatments. These two bacterial strains, Roseobacter sp. and an α-subclass Proteobacteria, are highly inductive to larval settlement of H. elegans. The larvae responded similarly to Roseobacter and Proteobacteria in all the larval settlement bioassays, suggesting that the larval settlement-inducing substances produced by these bacteria may share common characteristics. First of all, the larvae did not settle in the seawater conditioned by the bacteria attached as a film or by the bacteria that were freely suspended in seawater. The results suggest that the putative larval settlement cue is not released into seawater and, therefore, should be associated with the surface of the bacteria. Secondly, formaldehyde treatment entirely eliminated the larval settlement induction activity of the bacterial films, and streptomycin treatment reduced the percentage of larval settlement on the bacterial films in a concentration-dependent manner. Since both treatments can kill bacteria with little damage to the surface chemistry of bacterial cells, the decline in larval settlement is suggested be due to a reduction of the viable bacterial population in the bacterial films. In fact, the reduction of larval settlement in the streptomycin treatments coincided with the decrease in viable bacterial populations in broth cultures containing respective concentrations of streptomycin. These results suggest that the viability of Roseobacter and Proteobacteria is important to their settlement induction effect. Since the larval settlement induction activity of the bacterial strains appears to correlate with their viability, we suggest that the putative larval settlement cue is derived from a metabolic pathway in the bacteria and that the cue is exported to and concentrated at the extracellular polymer matrix of the bacterial cell, at which the larvae establish contact with the bacteria. The larval settlement cue may be highly susceptible to degradation so that a metabolically active bacterial film is needed to maintain the putative cue at a concentration that surpasses the threshold for induction of larval settlement. Received: 14 October 1998 / Accepted: 5 September 2000     

The role of the generally unrecognised microprey source as food for larval fish in the Irish Sea

Few studies have studied the food supply to larval fish in the Irish Sea; thus, we have assessed the full prey-field available to larval fish, ranging from protozoa to copepods. Specifically we assessed if fish larvae feed on protozoa, as suggested by others, and if densities of the protozoa and the appropriate size of metazoan prey were previously underestimated. By examining the gutcontents of fish larvae, the prey available to them, and the potential accessibility of prey to fish, we develop a simple food web, presented as a box-model. By doing so, we indicate that the lack of focus on small metazoa and protozoa has underestimated the food available to fish larvae; without these, we might have concluded that prey levels were too low to support the growth of the larval fish assemblage. Our methods were as follows. Sampling was at two sites, off the Isle of Man, with distinct physical and biological structures, soon after fish spawning: the southwest coast, where many species occur in spring-summer (23 April; 6, 19 May; 1, 22 June; 12 July) and the east coast, where only herring larvae occur in September–November (12, 28 October). Microplankton (15–200 μm), mesozooplankton, and larval fish were collected at 1, 15, and 25 m: microplankton with 1.5 L bottles and a 64 μm-mesh net; mesozooplankton and larval fish with a Gulf VII high-speed sampler (280 μm mesh). The 64 μm mesh net, mounted on the Gulf VII, provided simultaneous hauls. Fixed samples were evaluated to determine species composition, abundance, and biomass. Larval fish diet was determined from fish collected by short net hauls: fixed guts were examined and prey, including protozoa, analysed. Using physical data as a guide, plankton data were integrated through the water column to determine standing stocks. Size-based food availability to larval fish was estimated from the gut contents. The role of protozoa was examined, assuming that they are digested at the same rate as metazoan and if they are digested 2.5–10 times faster; increased digestion rates indicated that they contributed substantially to the larval fish diet.     

Development of the pH in the intestinal tract of larval turbot

The pH in the gut of turbot larvae and juveniles of turbot was studied from day 11 until the completion of metamorphosis. Dietary effects on the gut pH were estimated when larvae were offered live feed, a microdiet, only microalgae or no feed. The pH in the gut was weakly alkaline until day 24 after hatching with no differences between the foregut, midgut and hindgut. The foregut contents started to turn acidic from day 28 after hatching when the larvae were already weaned successfully, which indicates that an acidic pH is not necessary for the digestion and utilisation of formulated feed. During the following 20 days the pH in the foregut/stomach decreased further to a minimum of pH 3.5, while the pH in the midgut and the hindgut increased slightly to a maximum of pH 9.0. Larvae receiving live feed, microdiet or microalgae had a similar pH in the midgut on day 11, while starved larvae exhibited a lower gut pH. This suggests bicarbonate secretion from the larval pancreas stimulated by ingested microalgae or feed particles.     

The role of images of conspecifics as visual cues in the development and behavior of larval anurans

Tadpoles can alter their behavior, morphology, and life history in response to habitat change. Although chemical signals from conspecifics or predators play an important role in tadpole habitat assessment, little is known about the role of visual cues and the extent to which tadpoles rely on their vision for intraspecific social assessment. The aim of our experiments was to determine whether larval anurans use visual images of other tadpoles as indicators of density and to analyze how, and to what extent, images of conspecifics alone affect tadpole development, growth, and behavior. To assess this, we raised both Rana sylvatica and Bufo americanus tadpoles in aquaria with either quarter- or half-mirrored walls. Both physically increased density and increased density simulated with mirrors decreased tadpole growth and developmental rates, and increased activity in Rana tadpoles. Bufo tadpoles did not significantly alter their growth and development in response to visually increased density. Only true, i.e., physically, increased density had an effect on growth and activity in Bufo tadpoles. Our data show that images of conspecifics are used as visual cues by Rana tadpoles and can induce phenotypically plastic changes in several traits. This response to visual cues is taxon-specific. An erratum to this article can be found at     

Assessment of patch quality by ladybirds: relative response to conspecific and heterospecific larval tracks a consequence of habitat similarity?

Summary. Aphid colonies can reach high levels of abundance but last for short periods of time. The larvae of aphidophagous ladybirds (Coleoptera: Coccinellidae) that feed on these colonies might therefore suffer from starvation, which favours the occurrence of cannibalism and intraguild predation. Thus, the assessment of patch quality becomes crucial and it has been shown that female ladybirds refrain from laying eggs in the presence of an oviposition deterring semiochemical deposited by their larvae. Adalia bipunctata (L.), Adalia decempunctata (L.) and Coccinella septempunctata L. are 3 sympatric species of ladybirds, which can co-occur in aphid colonies. As a consequence, their eggs and larvae are under threat, not only from cannibalism but also intraguild predation. Females should, therefore, also use the tracks deposited by heterospecific larvae to assess the quality of aphid colonies as oviposition sites. The expectation is that: 1- the strength of the reaction to each other’s larval tracks should be correlated with percentage habitat overlap and that 2- the reaction to conspecific larval tracks should be stronger than to heterospecific tracks. In order to test these hypotheses, females’ oviposition behaviour was analysed and a chemical analysis of the tracks of their larvae undertaken. The results show that oviposition behaviour is not related to habitat overlap. Both species of Adalia react to tracks of their own larvae and those of C. septempunctata, but A. decempunctata reacted more strongly than A. bipunctata. C. septempunctata reacted very slightly to its own tracks but not to those of either species of Adalia. The larval tracks are mainly composed of alkanes. Those of the two species of Adalia are qualitatively 100% similar and 60% so when the quantitative results are compared. They are, however, only 24–29% similar to those of Coccinella septempunctata.