Spatial variability in habitat associations of pre- and post-settlement stages of coral reef fishes at Ishigaki Island,Japan

Successful settlement of pelagic fish larvae into benthic juvenile habitats may be enhanced by a shortened settlement period, since it limits larval exposure to predation in the new habitat. Because the spatial distribution of marine fish larvae immediately prior to settlement versus during settlement was unknown, field experiments were conducted at Ishigaki Island (Japan) using light trap sampling and underwater visual belt transect surveys to investigate the spatial distribution patterns of selected pre- and post-settlement fishes (Acanthuridae, Pomacentridae, Chaetodonidae and Lethrinidae) among four habitats (seagrass bed, coral rubble, branching coral and tabular coral). The results highlighted two patterns: patterns 1, pre- and post-settlement individuals showing a ubiquitous distribution among the four habitats (Acanthuridae) and pattern 2, pre-settlement individuals distributed in all habitats, but post-settlement individuals restricted to coral (most species of Pomacentridae and Chaetodontidae) or seagrass habitats (Lethrinidae). The first pattern minimizes the transition time between the larval pelagic stage and acquisition of a benthic reef habitat, the latter leading immediately to a juvenile lifestyle. In contrast, the second pattern is characterized by high settlement habitat selectivity by larvae and/or differential mortality immediately after settlement.     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

In situ swimming and settlement behaviour of larvae of an Indo-Pacific coral-reef fish, the coral trout Plectropomus leopardus (Pisces: Serranidae)

Late larvae of the serranid coral trout Plectropomus leopardus (Lacepède), captured in light traps, were released during the day both in open water and adjacent to two reefs, and their behaviour was observed by divers at Lizard Island, northern Great Barrier Reef. Coral trout larvae (n = 110) were present in light-trap catches from 18 November to 3 December 1997, including new moon (30 November). The swimming speed of larvae in open water or when swimming away from reefs was significantly greater (mean 17.9 cm s⁻¹) than the speed of larvae swimming towards or over reefs (mean 7.2 cm s⁻¹). Near reefs, larvae swam at average depths of 2.7 to 4.2 m, avoiding 0 to 2 m. In open water, swimming depth varied with location: larvae >1 km east of Lizard Island swam steeply downward to >20 m in 2 to 4 min; larvae >1 km west oscillated between 2.6 and 13 m; larvae 100 to 200 m east of Lizard Island oscillated between 0.8 and 15 m. Nearly all larvae swam directionally in open water and near reefs. In open water, the average swimming direction of all larvae was towards the island, and 80% (4 of 5) swam directionally (p < 0.05, Rayleigh's test). Larvae swam directionally over the reef while looking for settlement sites. The frequency of behaviours by larvae differed between two reefs of different exposure and morphology. Depending on site, 26 to 32% of larvae released adjacent to reefs swam to open water: of these, some initially swam towards or over the reef before swimming offshore. In some cases, offshore-swimming seemed to be due to the presence of predators, but usually no obvious cause was observed. Depending on the reef, 49 to 64% of the larvae settled. Non-predatory reef residents aggressively approached 19% of settlers. Between 5 and 17% of the larvae were eaten while approaching the reef or attempting to settle, primarily by lizardfishes but also by wrasses, groupers and snappers. A higher percentage of larvae settled in the second week of our study than in the first. Average time to settlement was short (138 s ± 33 SE), but some larvae took up to 15 min to settle. Average settlement depth was 7.5 to 9.9 m, and differed between locations. No settlement took place on reef flats or at depths <4.2 m. Larvae did not appear to be selective about settlement substrate, but settled most frequently on live and dead hard coral. Late-stage larvae of coral trout are capable swimmers with considerable control over speed, depth and direction. Habitat selection, avoidance of predators and settlement seem to rely on vision. Received: 7 July 1998 / Accepted: 26 January 1999     

Spatial and temporal variation in coral recruitment on the Great Barrier Reef: Implications for dispersal hypotheses

Over 15 000 coral recruits were counted on settlement plates from three mid-shelf reefs and six fringing reefs in the northern section of the Great Barrier Reef during two summers (1986 and 1987) and one winter (1987). The density of coral recruits on some settlement plates from a fringing reef was up to 4.88 cm^–2, the highest value ever reported. Mean density of recruits was greater on fringing reefs (81.1 recruits/settlement plate) than on mid-shelf reefs (15.6 recruits/settlement plate), but there was greater spatial variation in abundance of recruits between the fringing reef sites. Other differences between the mid-shelf reefs and the fringing reefs were that different taxa were dominant, and that settlement orientation differed, with mid-shelf recruits settling preferentially on horizontally oriented surfaces and fringingreef recruits preferring vertical surfaces. Of the three midshelf reefs, Green Island reef recorded the highest recruitment rate for each of the two summers, despite having a depauperate adult coral population following predation by the asteroidAcanthaster planci. This suggests that coral larvae frequently travel between reefs. In contrast with an earlier study, there was no consistent difference in abundance of recruits between forereef and backreef locations. Overall, the results indicated great spatial variation in the availability of coral larvae, both on the scale of whole reefs and within-reef habitats.     

Palatability of invertebrate larvae to corals and sea anemones

Risk of larval mortality is an underlying theme in debates and models concerning the ecology and evolution of the differing reproductive characteristics among marine benthic invertebrates. In these discussions, predation is often assumed to be a major source of larval mortality. Previous studies, focused primarily on planktotrophic larvae, suggested that marine larvae generally were susceptible to, and poorly defended against, planktivorous fishes and invertebrates. Larval-planktivore interactions involving larger and more conspicuous lecithotrophic larvae that are typical of many brooding sessile invertebrates have not been well studied. This lack of data for diverse larval types has hindered testing broad generalities about marine larvae and planktivore prey-preferences. This study demonstrates that lecithotrophic larvae of many Caribbean and temperate western Atlantic invertebrates are distasteful to co-occurring corals and anemones. These larval predators frequently rejected larvae of sponges (6 of 9 species), gorgonians (7 of 9 species), corals (3 of 3 species), hydroids (2 of 2 species) and a bryozoan. Larvae of three temperate colonial ascidians were readily consumed. Frequencies of survivorship for larvae captured but rejected by corals and anemones were generally high and, in 20 of 24 assays, were not statistically different from those of unattacked control larvae. Levels of metamorphosis (when it occurred) of rejected larvae also rarely differed significantly from those of unattacked controls. These results provide further evidence that larval palatability to predators may not be as high as once thought, particularly for brooded larvae of sessile colonial invertebrates. The means by which larvae may avoid or deter predators, and the demographic consequences for marine invertebrates and for the evolution of invertebrate life-history patterns, need to be assessed.     

On predation of pelagic larvae and early juveniles of marine bottom invertebrates by adult benthic invertebrates and their passing alive through their predators

The dynamic quantitative balance between prey and predator invertebrate species inhabiting the same shallow-shelf (sublittoral level bottom) benthic communities was first discussed by Thorson (1953). Thorson considered the exact timing of larval settlement of prey and predator species possessing pelagic development and temporal supression of the adult predators' feeding activities during reproduction at the time of the preys' settlement to constitute the major factors which facilitate survival of the prey species in such communities. However, information obtained demonstrates that Thorson's “mechanism of balance between predator and prey species of benthic communities” is not always effective in securing survival from predation not only of the prey's spat but even sometimes of the predator's spat also. Because of this, the “mechanism” can not be rated as universally effective in all situations. Analysis of the data so far published demonstrates that, in marine benthic communities, especially in shallow-shelf waters, it is not uncommon for gametes, larvae, or early juveniles of different prey species to pass alive through suspension (filter)-feeding and deposit-feeding adult invertebrates preying on them. Sometimes development can even continue after excretion by predators. The hypothesis of Voskresensky (1948) and Goycher (1949) of the importance of this phenomenon for the maintenance and recruitment of the mussel Mytilus edulis and other filter-feeding lamellibranchs of nearshore waters preying on their own and other lamellibranch pelagic larvae must be rejected on the basis of accumulated data on their feeding and general biology and on the adverse influence of the mucous of their faecal pellets and pseudofaeces on the larvae excreted by them alive. The data considered here demonstrate that, although the passing alive of larvae and spat of benthic invertebrates through benthic predators is not uncommon in shallow-shelf bottom-communities, it plays no important role in the processes of maintenance and recruitment of the species and communities involved nor of the marine benthos as a whole. The actual ecological significance of predation on pelagic larvae and bottom spat of benthic invertebrate prey species by all three main trophic groups of marine benthos (suspension or filter-feeders, deposit-feeders, carnivores) and its importance to predator-prey dynamics in marine benthic communities remains open to debate until more reliable quantitative data become available.     

Defensive mechanisms and ecology of some tropical holothurians

The ecology, defensive behavior and toxicity of three species of reef flat holothurians (Actinopyga mauritiana, Holothuria atra and Holothuria difficilis) were studied at Eniwetok Atoll, Marshall Islands. The average diurnal population density of H. difficilis ranged from 1.4 to 32 holothurians/900 cm²; resting respiratory rates (0.05 ml O₂/g wet wt/h) were comparable during day and night; nourishment in H. difficilis may be primarily from bacteria and foraminifera in which about 2% of the dry weight of sediment consumed is utilized, and the species probably passes at least 3 g dry wt of sediment/m²/day (>1 kg/m²/year). A fundamental difference in energy flow is suggested: considerably more energy is passed from benthic algae to grazing and browsing fishes to predatory fishes on coral reefs whereas, in extra-tropical latitudes, more energy is shunted from benthic algae to invertebrates to predators. The effects of holothurin leading to death in fishes are irreversible. Holothuria difficilis is best protected from predation. Its body wall is toxic and it can accurately eject Cuvierian tubules, which are also toxic. The discharge of tubules was regulated by a circadian rhythm in May. Studies on holothurians and sponges suggest that many exposed coral reef invertebrates have evolved effective defensive mechanisms in association with high intensity predation.Supported by A.E.C. Contract AT (29-2)-226 with the University of Hawaii.     

Settlement patterns and presettlement behavior of the naked goby,Gobiosoma bosci,a temperate oyster reef fish

Dispersion, distribution, development and feeding incidence of larvae of the naked goby,Gobiosoma bosci (Lacepéde), were examined for linkages between larval behavior while near the reef surface and later patterns of settlement and recruitment. Field sampling and experiments were conducted during the summers of 1988 and 1989 in the Flag Pond oyster reef along the western shore of the Chesapeake Bay near Camp Conoy, Maryland, USA. Results indicated that prior to settlement most demersal larvae aggregate in shoals and exhibit distinct microhabitat preferences on the reef. In a field experiment, larvae settled both during the day and at night. Dispersion at settlement was aggregated, suggesting that demersal shoaling influences settlement patterns in this species. The distribution of demersal larvae also indicated that larval swimming behavior is sufficiently strong to permit active control of position on reefs. Large demersal larvae settled rapidly when brought to the laboratory, but small larvae in demersal shoals appeared to require additional growth and morphological development prior to settlement. Development of the pelvic fins, used by juveniles and adults for perching on the substrate, may be a good indicator of competence to settle in this species. The adaptive significance of demersal shoaling by small larvae of the naked goby, and the fate of these larvae, remains perplexing because the low feeding rates found for larvae shoaling near the reef surface should slow or prevent the growth and development required prior to settlement. Observations made by other authors indicate that demersal shoaling and the use of water directly overlying reefs may be common behaviors of temperate and tropical reef fishes.     

Larval period and its influence on post-larval life history: comparison of lecithotrophy and facultative planktotrophy in the aeolid nudibranch Phestilla sibogae

The variable duration of the pelagic phase of metamorphically competent larvae of benthic marine invertebrates is set by an interaction between environmental factors and larval traits that together influence the chance that a larva will encounter and respond to a suitable settlement site. In the Hawaiian aeolid nudibranch Phestilla sibogae Bergh, an extended competent larval phase resulted in a cascade of negative effects on larval and post-larval life-history traits. When raised as fed (i.e., facultatively planktotrophic) larvae, an extended larval period resulted in lower larval survival, slightly lower metamorphic success, and delayed reproduction. When raised as unfed (i.e., lecithotrophic) larvae, an extended larval period resulted in lower larval and post-larval weights, survival, metamorphic success, and reproductive output, and also resulted in a longer juvenile period and delayed reproduction. The chance nature of locating a settlement site generally spreads these negative effects over all larvae of a cohort, and so balances the relative fitness of the genetic lineages within a population.     

Ultraviolet photosensitivity and feeding in larval and juvenile coral reef fishes

The ability of young coral reef fishes to feed using solely ultraviolet-A (UV-A) radiation during ontogeny was examined using natural prey in experimental tanks. Larvae and juveniles of three coral reef fish species (Pomacentrus amboinensis, Premnas biaculeatus and Apogon compressus) are able to feed successfully using UV-A radiation alone during the later half of the pelagic larval phase. The minimum UV radiation intensities required for larval feeding occur in the field down to depths of 90–130 m in oceanic waters and 15–20 m in turbid inshore waters. There was no abrupt change in UV sensitivity after settlement, indicating that UV photosensitivity may continue to play a significant role in benthic juveniles on coral reefs. Tests of UV sensitivity in the field using light traps indicate that larval and juvenile stages of 16 coral reef fish families are able to detect and respond photopositively to UV wavelengths. These include representatives from families that are unlikely to possess UV sensitivity as adults due to the UV transmission characteristics of the ocular media. Functional UV sensitivity may be more widespread in young coral reef fishes than in the adults, and may play a significant role in detecting zooplanktonic prey.     

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.     

Effects of selective settlement and of aggression by residents on distribution of young recruits of two tropical damselfishes

The reef fishes that settled on an array of experimental corals at Lizard Island, Queensland, Australia, were counted during a pulse of recruitment in December 1986. NeitherPomacentrus sp. norP. amboinensis showed any evidence that harassment by residentDascyllus aruanus caused a decrease in persistence during the first day after settlement. LarvalPomacentrus sp. settled selectively on corals without residentD. aruanus. The results forP. amboinensis were ambiguous. Settlers of both species positioned themselves closer to the sand on corals with residentD. aruanus than on unoccupied corals. This could reduce access to planktonic food and increase the risk of predation. Adult aggression may be less important and active selection of settlement sites by larvae may be more important to the distribution of recruits than is suggested by the literature. The presence or absence of particular species should be included among the cues that larval reef fishes use to choose settlement sites.     

Inhibition of larval settlement to a soft bottom benthic community by drifting algal mats: An experimental test

The hypothesis that drifting red algal mats inhibit settlement of planktonic larvae was tested in a field experiment in 1986. Substratum free of algae (caged boxes) was compared with substratum covered with algae (natural substratum and open boxes). Whereas settling densities of 1500 to 5500 ind. m^-2 of the bivalves Macoma balthica, Cardium glaucum and Mya arenaria were observed in sediment without algal cover, no recruits of the same species were found beneath the algae during the period of peak settlement (June to July). The same difference was observed for the polychaete Nereis diversicolor, although in this case occasional individuals were found beneath the algae. The results demonstrate that algae mats may be efficient inhibitors of larval settlement to benthic soft-substratum communities. It is concluded that algae act as a larval filter.     

Temporal patterns of larval settlement and survivorship of two broadcast-spawning acroporid corals

Acroporid corals are the main reef-building corals that provide three-dimensional habitats for other reef organisms, but are decreasing on many reefs worldwide due to natural and anthropogenic disturbances. In this study, temporal patterns of larval settlement and survivorship of two broadcast-spawning acroporid coral species, Acropora muricata and A. valida, were examined through laboratory rearing experiments to better understand the potential for larval dispersal of this important coral group. Many larvae were attached (but not metamorphosed) to settlement tiles on the first examination 3–4 days after spawning (AS). The first permanent larval settlement (i.e. metamorphosed and permanently settled juvenile polyps) occurred at 5–6 days AS, and most larval settlement (85–97% of total) occurred within 9–10 days AS. Larval survivorship decreased substantially to around 50% by the first week of the experiment and to approximately 10% by the second to third week. The rates of larval attachment, settlement, and the initial drop in survivorship of larvae suggest that effective dispersal of some acroporid species may largely be completed within the first few weeks AS.     

Influences of habitat and natural disturbances on contributions of massive Porites corals to reef communities

We compared densities, distributions and size frequencies of massive corals in the genus Porites on five relatively exposed, mid-shelf reefs (50 km offshore) in the central Great Barrier Reef with those on a sheltered inshore reef (10 km offshore). Data included various transect and mapping studies between 1984 and 1990, estimates of size-dependent damage from the crown-of-thorns starfish Acanthaster planci, estimated densities of herbivorous sea urchins (potential predators of juveniles), and observations of size-specific effects of tropical cyclones. Assemblages of Porites spp. on mid-shelf reefs were dominated by small colonies (2 to 10 cm diam) established either from planula larvae or from small tissue remnants that had survived A. planci predation in the early to mid-1980s. Large colonies (up to 10 m diam) were scarce, except for localized aggregations on terraces at the base of reef slopes (6 to 12 m deep). Extensive space suitable for settlement by coral larvae can be attributed to recurrent cyclones and A. planci outbreaks. Despite low sea urchin predation, the slowly growing Porites juveniles are likely to die from overgrowth by numerous, much faster growing corals. On the sheltered inshore reef, the coral community was dominated by very large (>5 m diam) Porites colonies, several centuries old; recruitment was mainly by fragmentation of large colonies; there was little space available for settlement, and probabilities of juvenile mortality from grazing urchins were high. Differences in settlement and early survival of Porites spp. are exacerbated by different regimes of storm damage. A model is proposed that links wave climate with the size and age reached by corals before dislodgement by storm waves, and which is consistent with observed densities and size-frequency distributions of Porites in sheltered and exposed areas.     

Transmission distance of chemical cues from coral habitats: implications for marine larval settlement in context of reef degradation

The present study (Ishigaki Island, Japan) explored the distance of transmission of chemical cues emitted by live versus dead coral reefs (Exp. 1: High performance liquid chromatography (HPLC) analyses with water sampling station at 0, 1, and 2 km away from the reef) and the potential attraction of these chemical cues by larval fish, crustaceans, and cephalopods (Exp. 2: choice flume experiment conducted on 54 Chromis viridis larvae, 52 Palaemonidae sp larvae, and 16 Sepia latimanus larvae). In the experiment 1, HPLC analyses highlighted that the live coral reef (and not the dead coral reef) produced different and distinct molecules, and some of these molecules could be transported to a distance of at least 2 km from the reef with a reduction of concentration by 14–17-fold. In the experiment 2, C. viridis, Palaemonidae sp, and S. latimanus larvae were significantly attracted by chemical cues from a live coral reef (sampling station: 0 km), but not from a dead coral reef. However, only C. viridis larvae detected the chemical cues until 1 km away from the live coral reef. Overall, our study showed that chemical cues emitted by a live coral reef were transported farthest away in the ocean (at least 2 km) compared to those from a dead coral reef and that fish larvae could detect these cues until 1 km. These results support the assumption of a larval settlement ineffective in degraded coral reefs, which will assist conservationists and reef managers concerned with maintaining biodiversity on reefs that are becoming increasingly degraded.     

Comparison of laboratory rates of predation of five species of marine fish larvae by three planktonic invertebrates: effects of larval size on vulnerability

Rates of predation by the invertebrates Aurelia aurita, Thysanoessa raschi and Euchaeta norvegica on larval stages of cod (Gadus morhua L.), flunder (Platichthys flesus L.), plaice (Pleuronectes platessa L.), herring (Clupea harengus L.), and turbot (Scophthalmus maximus L.) were determined. Experiments were conducted in late winter and early spring 1982 with predators collected in Loch Etive, Scotland and prey obtained from several locations in Great Britain. Early stages of the smallest species, cod, flounder and turbot, tended to be most vulnerable to all three predators, while the early stages of the larger species, plaice and herring, and older stages of all species, were less vulnerable. For all stages and species of larvae, predation rates by the three predators were most closely related to larval length and escape swimming speed. Larval length itself was closely correlated to indices of larval escape ability. Low predation rates on large larvae by E. norvegica could be due to handling difficulties, whereas for A. aurita and T. raschi these low rates were due to escape abilities of the larger larvae. Prey movement is an important stimulus eliciting predation in E. norvegica but not in A. aurita or T. raschi.     

Safety in numbers and the spatial scaling of density-dependent mortality in a coral reef fish

In coral reef fishes, density-dependent population regulation is commonly mediated via predation on juveniles that have recently settled from the plankton. All else being equal, strong density-dependent mortality should select against the formation of high-density aggregations, yet the juveniles of many reef fishes aggregate. In light of this apparent contradiction, we hypothesized that the form and intensity of density dependence vary with the spatial scale of measurement. Individual groups might enjoy safety in numbers, but predators could still produce density-dependent mortality at larger spatial scales. We investigated this possibility using recently settled juvenile bluehead wrasse, Thalassoma bifasciatum, a small, aggregating reef fish. An initial caging experiment demonstrated that juvenile bluehead wrasse settlers suffer high predation, and spatial settlement patterns indicated that bluehead wrasse juveniles preferentially settle in groups, although they are also found singly. We then monitored the mortality of recently settled juveniles at two spatial scales: microsites, occupied by individual fish or groups of fish and separated by centimeters, and sites, consisting of approximately 2400-m2 areas of reef and separated by kilometers. At the microsite scale, we measured group size and effective population density independently and found that per capita mortality decreased with group size but was not related to density. At the larger spatial scale, however, per capita mortality increased with settler density. This shift in the form of density dependence with spatial scale could reconcile the existence of small-scale aggregative behavior typical of many reef fishes with the population-scale density dependence that is essential to population stability and persistence.     

Influence of substratum heterogeneity and settled barnacle density on the settlement of cypris larvae

On the Atlantic coast of Canada, Semibalanus balanoides (L.) is widely distributed in the mid-intertidal zone, whereas in the Gulf of St. Lawrence, this species is mostly limited to crevices. We tested the hypothesis of regional differences in microhabitat selection by barnacle larvae at settlement in 1984 and 1985 at St. Andrews, New Brunswick, Canada. Since larvae settle in microhabitats already colonized by adults, the relative influence of settled barnacle density and of different scales of substratum heterogeneity on settlement were evaluated experimentally at Capucins, Québec, (Gulf of St. Lawrence) and at St. Andrews, New Brunswick (Atlantic coast). On a large scale (>10 cm deep crevices) of heterogeneity, results show that, in the Gulf, cypris larvae settled nearly exclusively (93%) in natural crevices rather than on adjacent horizontal surfaces. On the Atlantic coast, settlement was more important outside than inside of crevices, when the substrata were either natural or artificial. This result is unique and contrasts sharply with all known reports on barnacle settlement in relation to surface contour. The influence of barnacle density on settlement was greater than that of large scale heterogeneity. On a small scale (<1.5 cm deep cracks), the presence of conspecifics had a stronger effect on settlement than heterogeneity in both regions. Field observations showed a relationship between larval settlement density and percentage of adult cover. Settlement increased up to 22 or 30% (Gulf and Atlantic coast) of adult cover and decreased afterwards. The results confirm the hypothesis of larval selection for cryptic habitats in the Gulf and the opposite behaviour (preferences for horizontal surfaces) on the Atlantic coast. This microhabitat selection is apparent at large scales of heterogeneity, whereas at small scales, the presence of conspecifics is the predominant factor.Contribution to the programme of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Inhibition and facilitation of bryozoan and ascidian settlement by natural multi-species biofilms: effects of film age and the roles of active and passive larval attachment

Sessile marine invertebrate larvae can recognize suitable settlement substrata by using various environmental cues, including organic/microbial biofilms. In laboratory choice assays, the effect of biofilms of varying ages on the settlement behaviour of two fouling organisms was assessed. The species included the arborescent cheilostome bryozoan Bugula flabellata (Thompson) and the solitary ascidian Ciona intestinalis (L.), both of which are characteristic of temperate sublittoral hard substratum assemblages in northwest Europe. Experiments were carried out using polystyrene petri dish substrata preconditioned with multispecies biofilms from natural laboratory-aquarium seawater for 1, 3, 6 or 12 d. Unfilmed (new, initially sterile) dishes were used as control substrata. Whereas the coronate larvae of B. flabellata generally were inhibited by biofilming, irrespective of film age, the settlement of tadpole larvae of C. intestinalis was facilitated on biofilmed substrata, and numbers of settled larvae generally increased with biofilm age: the highest mean numbers were counted on 12 d-old biofilms. In C. intestinalis, settlement and metamorphosis are processes which can be temporally separate and are possibly induced by different environmental cues. This study therefore distinguished between C. intestinalis larvae which were attached to the biofilm surface by the anterior, and those larvae entrapped by the biofilm but not settled in the conventional meaning of the term. As reported in previous studies, we did observe that such entrapped larvae could subsequently attach and develop successfully into sessile juveniles. Both the numbers of “attached” and “trapped” tadpoles increased with biofilm age. Assuming that “settlement” is essentially a process involving the active behavioural response of larvae to environmental cues, it seems that the facilitated attachment of C. intestinalis onto biofilmed substrata is due to the combined effect of active habitat selection and passive deposition/“entrapment” of larvae onto the “sticky” substratum. Received: 21 August 1996 / Accepted: 21 November 1996