Fine-scale spatial and temporal variation in settlement of the intertidal mussel Perna perna indicates differential hydrodynamic delivery of larvae to the shore

Scales of spatial and temporal variation in the settlement of early and late plantigrades of the mussel Perna perna to artificial substrata were measured on the south coast of South Africa. One study examined daily settlement over one month at three sites of similar tidal height and aspect. The sites were separated by 100′s of m with 5 replicated artificial settlement substrata approximately 1 m apart within each site. Early (< 0.5 mm) and large late plantigrades (1 - 3.5 mm) dominated samples with few individuals of 0.5-1 mm. Settler abundance showed clear peaks, synchronised among sites and apparently following spring tides, though the relationship between tide and settlement was not significant. Two-way ANOVA indicated significant spatial variation, which varied with time, at the 100 m-scale among sites. A second study investigated the local effects of substratum aspect. Seaward facing surfaces had consistently lower settlement (but higher adult cover) than horizontal and landward facing surfaces, this effect was significant for large late plantigrades. This suggests that either juveniles migrate after settlement, or that post-settlement mortality differs markedly among aspects. The high degree of temporal variation at the scale of days and weeks was expected, but the results also suggest a degree of pulsing in settlement, with maximum intensities around spring tide. A previous study of nearshore waters indicated a common pool of larvae at scales of 100 s m. Thus, consistent ranking of sites for all size classes of settlers indicates predictably differential hydrodynamic delivery of larvae to sites separated by 100 s m. The effect of aspect on settlement highlights the importance of smaller scale hydrodynamics, suggesting that the stronger turbulence experienced by seaward facing surfaces may affect larval settlement negatively.     

Two levels of spacing and limits to local population density for settled larvae of the ascidian Clavelina moluccensis: a nearest-neighbour analysis

Among sessile marine invertebrates, the dispersion patterns of settling larvae are likely to strongly affect the nature and strength of subsequent intraspecific interactions. We use a recently developed graphical analysis technique to examine in situ spacing among settled larvae of the colonial ascidian Clavelina moluccensis. Larvae were mapped at settlement each day for 1 month at the time of their release from adults. Experimental plots measuring 90×90 mm were located on naturally occurring bare patches of wood on pier pilings which were extensively encrusted with sponges and ascidians. An inclusive one-tailed nearest-neighbour analysis applied at the time of the maximum number of individuals per plot revealed consistent fine-scale regular spacing between settled larvae. Furthermore, a two-tailed goodness-of-fit analysis indicated two modes of spacing: one at about 4.5 mm and the other at around 9 mm. No contact between larvae was observed at settlement and the spatial regularity is interpreted as a response of settling larvae to water-borne cues released from settled individuals. The two modes of spacing indicate the possibility of kin recognition as has been demonstrated for bryozoans. Water-borne cues may also limit local population density, as those plots with a high density of settlers showed a much greater decrease in settlement rate over time than low-density plots. Our findings are consistent with the hypothesis that intraspecific competitive interactions shape the patterns of dispersion of these invertebrates on hard substrata in the field.     

Coral larvae settle at a higher frequency on red surfaces

Although chemical cues serve as the primary determinants of larval settlement and metamorphosis, light is also known to influence the behavior and the settlement of coral planulae. For example, Porites astreoides planulae settle preferentially on unconditioned red substrata. In order to test whether this behavior was a response to color and whether other species also demonstrate color preference, settlement choice experiments were conducted with P. astreoides and Acropora palmata. In these experiments, larvae were offered various types of plastic substrata representing three to seven different color choices. Both species consistently settled on red (or red and orange) substrata at a higher frequency than other colors. In one experiment, P. astreoides settled on 100% of red, plastic cable ties but failed to settle on green or white substrata. In a second experiment, 24% of larvae settled on red buttons, more than settled on six other colors combined. A. palmata settled on 80% of red and of orange cables ties but failed to settle on blue in one experiment and settled on a greater proportion of red acrylic squares than on four other colors or limestone controls in a second experiment. The consistency of the response across a variety of plastic materials suggests the response is related to long-wavelength photosensitivity. Fluorescence and reflectance spectra of experimental substrata demonstrated that the preferred substrata had spectra dominated by wavelengths greater than 550 nm with little or no reflection or emission of shorter wavelengths. These results suggest that some species of coral larvae may use spectral cues for fine-scale habitat selection during settlement. This behavior may be an adaptation to promote settlement in crustose coralline algae (CCA)-dominated habitats facilitating juvenile survival.     

Effects of sediments on larval settlement of abalone Haliotis diversicolor

The effects of deposited sediments on the settlement of the abalone Haliotis diversicolor Reeve were examined through both field observations and a laboratory experiment. Occurrences of newly settled post-larvae (shell length < 500 μm) and the amount of suspended and deposited sediments were monitored at two stations (Stns 1 and 2), which experienced different sediment deposition conditions, located at Nagai on the coast of Sagami Bay, Japan. Quantities of suspended sediments at the two stations varied considerably during the survey period, but no significant difference was detected between the stations. Mean volume densities of sediments deposited on cobbles were significantly higher at Stn 2 than at Stn 1. Densities of newly settled post-larvae from the 2001 to 2004 cohorts were significantly higher at Stn 1. A laboratory experiment was conducted to assess the effects of sediment quantity and quality on larval settlement. Two substances with different physical properties, kaolin and clamshell powder, were used as sediments. Larvae were subjected to four different sediment treatments with crustose coralline algae (CCA) substrates; thin and thick treatments for both kaolin and clamshell powder. Negative (without CCA) and positive (with CCA) controls without sediments were also established. The rate of metamorphosis decreased as sediment thickness increased in both the kaolin and clamshell powder treatments. Larvae in the kaolin treatments appeared to be trapped by the kaolin, and most could not metamorphose successfully. There were no trapped larvae in the clamshell powder treatments. The results indicate that the quantity and physical properties of sediments deposited on substrata affect the settlement and behaviour of larval abalone. Experimental results suggest that the lower densities of newly settled post-larvae observed at Stn 2 may have been a result of larger quantities of deposited sediments, which reduced the availability of suitable substrate for larval settlement.     

A standard unit for monitoring recruitment of fishes to coral reef rubble

We developed a benthic standard monitoring unit for recruitment of fishes (SMURF) to sample fishes that settle in coral rubble and tested it on three fringing reefs on the west coast of Barbados, West Indies. These rubble SMURFs are inexpensive to construct and permit newly settled fishes, including cryptic, secretive and small species, to be quickly and fairly completely (> 83%) removed by divers without returning the unit to the surface. Over a 3-month period involving 9 collections, 32 units on 3 reefs yielded 948 newly settled fishes belonging to 28 taxa (3.3 ± 0.2 S.E. recruits per sample, n = 287) in addition to numerous crustaceans. Most specimens were Sparisoma spp. (Scaridae) (41.8%) and Stegastes partitus (Pomacentridae) (13.6%), with moderate numbers of Scarus spp. (Scaridae) (8.6%), Lythrypnus spp. (Gobiidae) (7.4%), and Pseudogramma gregoryi (Serranidae) (7.2%). Sampling SMURFs at 1-day rather than 11-day intervals yielded approximately twice as many fish, although some taxa were not affected by sampling frequency, indicating taxon-specific differences in post-settlement loss rates. Netting, intended to reduce predation on settlers, did not affect settlement estimates unless algae grew on it, suggesting that the SMURF design minimizes fish predation even in the absence of the netting. Over time, however, the netting resulted in substantial algal growth that was associated with an increase in abundance of Sparisoma spp. and a decrease in the abundance of S. partitus. This suggests that settler estimates are more affected by microhabitat changes from algal growth than by predation. During low settlement periods, SMURFs performed better than light-traps in assessing the daily input of new settlers of two abundant taxa. Eight SMURFs per reef were sufficient to demonstrate differences in settlement rates and temporal correlations in settlement within and among reefs. This study underscores the potential of benthic SMURFs to measure settlement of coral reef fishes at a range of spatio-temporal scales with moderate research cost and effort, and to facilitate the study of the settlement stage of several taxa.     

Variation in recruitment of the subtidal colonial ascidian Podoclavella cylindrica (Quoy & Gaimard): the rôle of substratum choice and early survival

The rôle of substratum choice and subsequent survival in determining the patterns of recruitment of the colonial ascidian Podoclavella cylindrica (Quoy & Gaimard) (Clavelinidae : Aplousobranchia) were examined. Larval settlement preferences on five substrata were determined by following individual larvae underwater and observing settlement behaviour on their first encounter with a substratum. The survival of settling larvae was monitored daily for 30 consecutive days in quadrats on the five substrata. Larvae settled preferentially on substrata that showed reduced mortality of settlers, hence recruitment was correlated with settlement. In almost all cases larvae rejected the unfouled surfaces of sponges, preferring those with epifauna, and settled in highest densities on bare space. Mortality of new recruits varied markedly between substrata but was very low on bare space. Juvenile mortality resulted from overgrowth by neighbouring invertebrates and from the removal of epifauna, associated with sponges, on which juveniles had settled.     

The importance of flow and settlement cues to larvae of the abalone, Haliotis rufescens Swainson

In flow tank experiments, I tested the relative importance of active and passive processes to larvae settling on manufactured casts that were hydrodynamically rough at a small scale (mm to <1 cm). I predefined two distinct regions of small-scale flow that I used to manipulate larval settlement behaviour of the red abalone Haliotis rufescens Swainson. The larvae show a stringent settlement response associated with coralline red algae. Haliotis rufescens larvae settled preferentially to an inducer regardless of the flow conditions, as expected. However, the ability of H. rufescens larvae to show this stringent behaviour was altered by changing the small-scale flow. When the free-stream velocity was low, the larvae responded to a settlement cue regardless of the small-scale hydrodynamics. When free-stream velocity was higher, the larvae acted increasingly as passive particles in their deposition, but settled only in response to an inducer. The results were consistent in two flow tanks, across 2 years and between different batches of larvae.     

Predator-prey interactions from in situ time-lapse observations of a sublittoral mussel bed in the Gulf of Trieste (Northern Adriatic)

Hexaplex trunculus (Linnaeus, 1758) is one of the most abundant and widespread muricid gastropods in the Northern Adriatic Sea, but relatively little is known about the feeding ecology of this predator. We examined the activity of H. trunculus on a sublittoral mussel bed at 24 m depth through in situ time-lapse observations and bulk samples. The camera photographed a 0.25 m² section of the mussel bed at 6-min intervals for ~ 23 h. Photos were examined frame-by-frame for gastropod movement and activities, especially interactions between H. trunculus and Mytilus galloprovincialis (Lamarck, 1819). Our survey indicates high activity-levels of H. trunculus on the sea floor: all gastropods made minor movements, most made major movements, and most left the field of view during the study-interval. On average, individuals remained stationary for only 7.3 h. Two predation attempts on Mytilus involving conspecific competition were documented, and one Hexaplex was consuming a mussel at the onset of the deployment. Additionally, 487 M. galloprovincialis from four diver-taken 0.25 m² quadrates were measured and examined for traces of marginal chipping and drilling predation. Mytilus from surface samples ranged from 11.1 mm to 95.5 mm in length, and one of the four samples had a significantly different average shell length from the others. 114 H. trunculus were collected and measured. Hexaplex ranged from 22.1 mm to 86.1 mm and the mean shell length did not differ among samples, though they were overwhelmingly medium and large. Predation frequency (the ratio of successfully preyed upon bivalves to the total number of bivalves sampled) is high at the studied site (> 55%), and large gastropods preferred a chipping mode of predation to drilling, supporting earlier laboratory studies showing a preference for M. galloprovincialis and this predation strategy. Prey effectiveness (the ratio of failed predatory attacks to total predatory attacks) is also high (63.8%), and no evidence of a size refuge was found. Feeding in H. trunculus is highly facultative, calling for caution when using drill holes to estimate predation intensities; whenever possible, traces of multiple predation modes should be considered.     

Larval settlement of the green sea urchin, Strongylocentrotus droebachiensis, in the southern Gulf of Maine

Abstract. Settlement timing and differential settlement for the larval stage of the green sea urchin, Strongylocentrotus droebachiensis , in the southern Gulf of Maine was studied during the summer of 1996. Settlement densities on astroturf panels were highest in June and early July (13 to 37 m ⁻² d⁻¹), and peaked in mid-June (199 m⁻² d⁻¹). Settlement was low to nonexistent from mid-July through August (0 to 2 m⁻² d⁻¹). During the peak in settlement, no selection for substrate type was observed. In the remainder of the settlement period, differential settlement occurred, with a preference for substrate covered with live coralline algae. Test diameter of newly settled urchins varied among the substrates, with urchins settling on live coralline algae having the largest test diameter (0.43 ± 0.01 mm). There were no differences in test diameter among the different weeks in which sampling was done. Sustained onshore winds occurred only during peak settlement, suggesting that wind drift currents may concentrate larvae and influence patterns of larval settlement.     

The effect of bacterial and diatom biofilms on the settlement of the bryozoan Bugula neritina

Biofilms of marine bacteria and diatoms and their combinations were examined in laboratory choice assays to determine their effects on the attachment and successful metamorphosis of the larvae of the bryozoan Bugula neritina (Linnéus). The larval settlement in response to unfilmed surfaces, a natural biofilm (NBF) and adsorbed cells of three strains of bacteria, five strains of pennate diatoms and combinations of the two at different densities. Bacterial and diatom strains showed different effects on the larval settlement of B. neritina. Bacterial monospecific strains of an unidentified α-Proteobacterium and Vibrio sp. mediated the same percentage of settlement as a filtered seawater control. Biofilms of Pseudoalteromonas sp. caused significantly lower larval settlement. Larval settlement of B. neritina was negatively correlated with increasing densities of Pseudoalteromonas sp. The highest percentages of settlement were mediated by the biofilms of the diatom species Achnanthes sp., Amphora cofeaeformis, Amphora tenerrima, Nitzschia constricta and a 5-day-old natural biofilm, while the lowest settlement was found on a N. frustulum film. A three-way analysis of variance demonstrated that the density of bacteria and the presence of particular species of diatoms and bacteria in combined biofilms, significantly affected the settlement of B. neritina larvae. High settlement of larvae (50-90%) at all treatments indicated that B. neritina larvae are much more indiscriminate settlers than previously expected. Hence, using this species as a monitoring organism to trace ecologically relevant subtle changes of settlement cues in the natural environment should be carefully re-examined.     

The larval and juvenile ecology of the temperate octocoral Alcyonium siderium Verrill. I. Substratum selection by benthic Larvae

The Alcyonacean octocoral Alcyonium siderium Verrill forms dense stands on vertical rock walls (6–18 m subtidal) along the coast of Massachusetts and further north into the Gulf of Maine. Larvae are brooded until their release as crawling demersal planulae in August. Each colony produces several hundred larvae; most crawl onto the nearby substratum although others may be carried away by wave surge.Experimental enclosures were used to examine substratum selection in the field. Planulae metamorphosed in greater numbers than expected by percent cover on the fleshy red crustose alga Waernia mirabilis Wilce (proposed). The crustose coralline algae Phymatolithon rugulosum (Foslie) and Lithothamnium glaciale Kjellman were settled on commonly but less often than expected by their percent cover. Bare rock surfaces were also used, but least frequently. The encrusting invertebrate colonies that cover much of the available substratum were never used as sites of settlement and metamorphosis. These results agree with laboratory evidence that settlement and metamorphosis of Alcyonium planulae can be induced by Waernia and by coralline algae.     

The chemical cue tetrabromopyrrole from a biofilm bacterium induces settlement of multiple Caribbean corals

Microbial biofilms induce larval settlement for some invertebrates, including corals; however, the chemical cues involved have rarely been identified. Here, we demonstrate the role of microbial biofilms in inducing larval settlement with the Caribbean coral Porites astreoides and report the first instance of a chemical cue isolated from a marine biofilm bacterium that induces complete settlement (attachment and metamorphosis) of Caribbean coral larvae. Larvae settled in response to natural biofilms, and the response was eliminated when biofilms were treated with antibiotics. A similar settlement response was elicited by monospecific biofilms of a single bacterial strain, Pseudoalteromonas sp. PS5, isolated from the surface biofilm of a crustose coralline alga. The activity of Pseudoalteromonas sp. PS5 was attributed to the production of a single compound, tetrabromopyrrole (TBP), which has been shown previously to induce metamorphosis without attachment in Pacific acroporid corals. In addition to inducing settlement of brooded larvae (P. astreoides), TBP also induced larval settlement for two broadcast-spawning species, Orbicella (formerly Montastraea) franksi and Acropora palmata, indicating that this compound may have widespread importance among Caribbean coral species.     

Adult fiddler crabs Uca pugnax (Smith) enhance sediment-associated cues for molting of conspecific megalopae

Adult-associated chemical cues can stimulate settlement and metamorphosis of invertebrate larvae into habitats with an enhanced likelihood of juvenile and adult survival. For example, sediments from adult fiddler crab habitat stimulate fiddler crab megalopae to metamorphose (molt) sooner than sediments without adult cues. A similar stimulation of molting occurs after exposure to waterborne chemical cues from adult habitats and to exudates and extracts of adult crabs. We tested whether sediments from habitats without adult Uca pugnax (Smith), which do not stimulate molting of their megalopae, could become stimulatory through brief exposure to adult crabs. Sediments were collected from tidal flats at several distances (∼ 1 m, ∼ 50 m, and ∼ 5.4 km) from adult habitats, and incubated for 24 h with or without adult crabs. Molting rates of laboratory-reared megalopae exposed for 48 h to adult-conditioned sediments were compared to those for untreated controls. Sediments collected in or within 1 m of adult habitat elicited the highest molting rates, and natural sediments from 50 m and 5.4 km had little or no effect on molting. However, incubating sediments collected away from adult habitat with adult crabs produced a higher molting response, and the magnitude of the enhancement increased with distance from adult habitat. Results suggest that the chemical cues that adult crabs release are retained by sediments and consequently stimulate molting of megalopae, regardless of the nature of the sediments themselves. Lack of chemical cues may retard colonization of newly created or heavily disturbed habitats that are otherwise suitable settlement and adult habitat.     

Settlement of bivalve spat on artificial collectors in Eyjafjordur, North Iceland

The seasonal pattern of bivalve spat settlement in Eyjafjordur, North Iceland, was investigated using artificial collectors of monofilament netting over 14 months (March 1998–January 2000) at 5, 10 and 15 m depth. SCUBA divers replaced the collectors at 4-weekly intervals. Twelve bivalve species settled on the collectors but only Mytilus edulis and Hiatella arctica were present throughout the year; they were the most abundant bivalve taxa. Of the remaining species, only Chlamys islandica, Heteranomia spp., Arctica islandica, Serripes groenlandicus and Mya spp. were sufficiently abundant to enable statistical analysis. All settled in late summer and autumn. Peak settlement of M. edulis, in September, consisted mainly of primary settlers (0.25–0.5 mm) although secondary settlers (>0.5 mm) were present in all samples. Mytilus edulis settled mostly at 5 m depth, especially larger individuals, possibly reflecting stronger currents at shallower depth and the proximity of mussel beds in the intertidal zone. Primary (<1 mm) and secondary H. arctica settlers (>1 mm) were present in most months, with the former being most numerous in September, 1999; settlement was equally abundant at 5 and 10 m depth. Primary settlement of C. islandica and S. groenlandicus occurred in autumn (mainly in September), and secondary settlers were very scarce and only seen in winter. Arctica islandica, Heteranomia spp. and Mya spp. settled mainly in September 1999 at 10 m depth, except for A. islandica, which was more numerous in August.     

The effects of larval abundance, settlement and juvenile mortality on the depth distribution of a colonial ascidian

The colonial ascidian Didemnum candidum (Savigny) is more abundant at shallow depths on floating docks than at greater depths along pilings in Pearl Harbor, Oahu, Hawaii. To compare the effects of selective settlement and postsettlement mortality on adult distribution, I determined if settlement was nonrandom relative to depth, if differences in adult abundance were responsible for the depth distribution of settlement, and if juvenile mortality varied with depth. A plankton pump was used to measure variation in larval abundance with depth. Acrylic settling plates were suspended at different depths and sampled nondestructively to measure settlement intensity and juvenile mortality. Settlement and mortality of the didemnid ascidians Diplosoma listerianum Milne-Edwards and Diplosoma sp. were also measured in the field and compared to that of D. candidum. Settlement of D. candidum was indeed nonrandom over depth. Both planktonic larvae and settled juvenils were more numerous at 0.5 than 3 or m. Settlement intensity on vertical plates was greatest within 1–2 cm of the water surface in the laboratory and within 2–3 cm in the field. Directly adjacent to the floating dock, where adults were equally abundant, settlement on horizontal plates was greater at shallow depths, suggesting directed movement of larvae upward. However, the ratio of settled juveniles to planktonic larvae (number of settlers: number of larvae) did not significantly differ with depth, suggesting that larvae were not more likely to settle at a particular depth. Settlement of the Diplosoma species was also heaviest near the surface. Juvenile mortality was greater at 0.5 than at 3 or 6 m for both D. candidum and the Diplosoma species. Experimental settlement showed that mortality, per se, of D. candidum was independent of depth. Rather, mortality was density-dependent, and the higher mortality near the surface was due to the greater number of larvae settling there. At this location, nonrandom settlement appears to determine the adult distribution of D. candidum, despite greater juvenile mortality at shallow depths. The pattern of settlement over depth is largely determined by adult proximity, rather than active larval behavior.     

Larval settlement of the common Australian sea urchin Heliocidaris erythrogramma in response to bacteria from the surface of coralline algae

Bacterial biofilms are increasingly seen as important for the successful settlement of marine invertebrate larvae. Here we tested the effects of biofilms on settlement of the sea urchin Heliocidaris erythrogramma. Larvae settled on many surfaces including various algal species, rocks, sand and shells. Settlement was reduced by autoclaving rocks and algae, and by treatment of algae with antibiotics. These results, and molecular and culture-based analyses, suggested that the bacterial community on plants was important for settlement. To test this, approximately 250 strains of bacteria were isolated from coralline algae, and larvae were exposed to single-strain biofilms. Many induced rates of settlement comparable to coralline algae. The genus Pseudoalteromonas dominated these highly inductive strains, with representatives from Vibrio, Shewanella, Photobacterium and Pseudomonas also responsible for a high settlement response. The settlement response to different bacteria was species specific, as low inducers were also dominated by species in the genera Pseudoalteromonas and Vibrio. We also, for the first time, assessed settlement of larvae in response to characterised, monospecific biofilms in the field. Larvae metamorphosed in higher numbers on an inducing biofilm, Pseudoalteromonas luteoviolacea, than on either a low-inducing biofilm, Pseudoalteromonas rubra, or an unfilmed control. We conclude that the bacterial community on the surface of coralline algae is important as a settlement cue for H. erythrogramma larvae. This study is also an example of the emerging integration of molecular microbiology and more traditional marine eukaryote ecology.     

Larval settlement preferences of Acropora palmata and Montastraea faveolata in response to diverse red algae

Settlement specificity can regulate recruitment but remains poorly understood for coral larvae. We studied larvae of the corals, Acropora palmata and Montastraea faveolata, to determine their rates of settlement and metamorphosis in the presence of ten species of red algae, including eight species of crustose coralline algae, one geniculated coralline and one encrusting peyssonnelid. Twenty to forty percent of larvae of A. palmata settled on coralline surfaces of Hydrolithon boergesenii, Lithoporella atlantica, Neogoniolithon affine, and Titanoderma prototypum, whereas none settled and metamorphosed on Neogoniolithon mamillare. Larvae of M. faveolata had 13–25 % settlement onto the surface of Amphiroa tribulus, H. boergesenii, N. affine, N. munitum, and T. prototypum, but had no settlement on the surface of N. mamillare, Porolithon pachydermum, and a noncoralline crust Peyssonnelia sp. Some of these algal species were common on Belizean reefs, but the species that induced the highest rates of larval settlement and metamorphosis tended to be rare and primarily found in low-light environments. The shallow coral, A. palmata, and the deeper coral, M. faveolata, both had increased larval settlement rates in the presence of only a few species of red algae found at deeper depths suggesting that patterns of coral distribution can only sometimes be related to the distribution of red algae species.     

Depth and timing of settlement of veligers from different populations of giant scallop, Placopecten magellanicus (Gmelin), in thermally stratified mesocosms

The present study was conducted to evaluate the relative roles that water column stratification intensity and possible inter-population behaviour differences play in determining depth of larval settlement of giant scallops, Placopecten magellanicus, in relation to thermoclines. Differences in timing of settlement of larvae from various populations were also examined. Two separate experiments were conducted in a 10.5-m-deep, 3.7-m-diameter, thermally stratified tank with larvae spawned from scallops collected from several adult beds located in areas with differing oceanographic regimes (Georges Bank, Mahone Bay, Passamaquoddy Bay). Previous mesocosm experiments had shown that veligers from these various populations differ in their vertical migration patterns. In the first experiment (December 1992-February 1993), larvae from all three populations were held in separate 9.5-m-deep tubes and exposed to a 1.5 °C temperature differential established over a depth interval of 1 m. The number of settled juveniles (spat) of each population collected at the end of the experiment increased with depth and showed no peak at or above the thermocline. This depth distribution of spat was most likely driven by preferential larval settlement. In the second experiment (February-May 1994), larvae spawned from Georges Bank and Passamaquoddy Bay stocks were held in 9.0-m-deep tubes and exposed to a 5 °C temperature differential established over a depth interval of 1 m. The number of settled spat of both populations was greater above this thermal boundary and increased with decreasing depth. This depth distribution was most likely driven by preferential settlement above the thermocline followed by upward post-settlement migration. The results from the two experiments indicate that larvae from various populations show similar trends in settlement patterns in response to similar thermal stratifications. Stratification intensity, however, does affect depth of larval settlement. In the second experiment, both populations of larvae settled throughout the time interval of collector deployment (larvae 32-82 days old). These results extend the range of planktonic developmental times generally reported in the literature and may be more indicative of natural planktonic development in the field. While Georges Bank larvae settled in consistent numbers through time, Passamaquoddy Bay larvae showed peaks in settlement at certain time periods, indicating that pulses of larval settlement may occur even from an individual spawning event.     

Characterization of a natural inducer of coral larval metamorphosis

External chemical signals used by scleractinian corals to recognize suitable substrata for larval settlement and metamorphosis were identified from crustose coralline red algae (CCA). A fragment of coral rubble with CCA induced larval metamorphosis of the scleractinian coral Pseudosiderastrea tayamai. A natural inducer and compounds that enhanced its effect in larval metamorphosis were isolated from the methanol extracts of coral rubble with CCA. A bromotyrosine derivative, 11-deoxyfistularin-3 (10^− 7 M) isolated from the CCA, induced the metamorphosis of P. tayamai larvae (27.5 ± 24.0%). In the presence of fucoxanthinol (10^− 9 M) and fucoxanthin (10^− 9 M), the percentage of metamorphosis induced by the bromotyrosine derivative was further enhanced to 87.8 ± 13.0 and 88.4 ± 17.8%, respectively. Both carotenoids are also found in the coral rubble with CCA. These results suggest that bromotyrosine derivative and carotenoids have a synergistic effect in the metamorphosis of P. tayamai larvae. The synergistic effect provides a higher selectivity for recruitment than a single-component natural inducer for recognizing suitable substrata for larval metamorphosis. Thus, the effect might offer a survival advantage for benthic marine invertebrates.     

Temporal scales of variation in settlement and recruitment of the mussel Perna perna (Linnaeus, 1758)

Population dynamics of many intertidal organisms are strongly affected by the abundance and distribution of larvae arriving on the shore. In particular, not only absolute numbers of settlers but also the degree of synchronisation of settlement can have a strong influence on whether density-dependent or density-independent processes shape adult shape populations. Temporal variation in rates of settlement and recruitment of the mussel Perna perna on the south coast of South Africa was investigated using a nested spatial design at different temporal scales. Variability in settlement at spring tides was examined at two temporal scales: lunar (to investigate the effect of state of the moon on settlement) and tidal (to investigate the influence of state of the tide on mussel settlement). Recruitment over neap tides was examined at one temporal scale, fortnight (to investigate the effect of date on mussel recruitment).Strong temporal variation was evident for both settlement and recruitment, but not at all time scales. Distinct peaks of settler/recruit abundance were observed during the lunar and neap tide studies. Recruitment intensity differed over the course of the year, and pulsing of recruitment was generally synchronised among locations. However, the strength of pulsing differed dramatically among locations, giving a significant interaction between fortnight and location. The finest temporal scale, investigated in the tidal study, did not reveal a significant effect of the state of the tide on settlement. The state of the moon (new or full) was not significant as a main factor (p = 0.052), although generally more settlers arrived on the shore during new moon. Phase of the moon appeared to have an effect on settler abundances, but only when and where densities were high.