Seasonal,annual, and spatial variation in the development of hard bottom communities

The development of hard bottom communities has been studied on test panels in Helgoland Harbour (German Bight) since 1977. Settlement and growth of epibenthic species was examined monthly. Natural variation in different seasons, years, and at three stations (the latter, only in 1981 and 1982) was investigated. At Station A (Binnenhafen), barnacles(Balanus crenatus) and polychaetes(Polydora ciliata) were always among the first settlers in spring. They were followed by other barnacles(Elminius modestus, Balanus improvisus) and by colonial ascidians(Botryllus schlosseri). The latter species often dominated from August to October, and tended to overgrow the barnacle populations.E. modestus showed strong annual variation, probably due to extremely low winter temperatures: after the cold winter of 1978/79, its populations were less dense than in previous years. In 1981 they recovered, and settlement increased again, but the cold winter 1981/82 damaged the population again. At Station B (Nordosthafen), mussels(Mytilus edulis) soon covered barnacles and empty space. By October they had monopolized the fouling community. At Station C (Südhafen), barnacle settlement in spring was followed by an overgrowth of hydrozoans (Laomedea spec.). In summer, ascidians (Ciona intestinalis andAscidiella aspersa) settled and began to dominate. Barnacles were weaker in the competition for space as opposed to later colonizers at all three stations.     

Comparative studies on the biology of nerita atramentosa Reeve, bembicium nanum (Lamarck) and cellana tramoserica (Sowerby) (gastropoda: Prosobranchia) in S.E. Australia

Populations of three coexisting intertidal gastropods, Nerita atramentosa Reeve, Bembicium nanum (Lamarck) and Cellana tramoserica (Sowerby), were sampled from a shore in Botany Bay, New South Wales, from July 1972 to September 1973. The recruitment and growth rates of each species were analysed from size frequency distributions. Mortality of each age cohort, and longevity, were estimated from analyses of the densities of Nerita atramentosa and Bembicium nanum.Nerita atramentosa showed no significant mortality during the first two years on the shore, but high mortality (at an instantaneous rate of 0.084 deaths/individual/month) after reaching the age of reproductive maturity, which was 20 months from settlement and at a mean shell-length of 13.5 mm. No growth could be detected after reproductive maturity was reached. Longevity of N. atramentosa was estimated as 3–5.5 years. Bembicium nanum juveniles showed higher mortality (at an instantaneous rate of 0.233 deaths/individual/month) than reproductively mature animals (0.060 deaths/individual/month). Reproductive maturity was reached at a mean shell-breadth of 11.0mm, i.e., about ten months after settlement on the shore. Longevity was estimated as from 4–8 years. Cellana tramoserica showed different growth patterns depending on the time of settlement.The three species showed different patterns of growth and life history relating to variable recruitment (which was demonstrated in all three species) and different rates of mortality of age cohorts.     

Recruitment,Growth and Mortality of an Antarctic Hexactinellid Sponge,Anoxycalyx joubini

Polar ecosystems are sensitive to climate forcing, and we often lack baselines to evaluate changes. Here we report a nearly 50-year study in which a sudden shift in the population dynamics of an ecologically important, structure-forming hexactinellid sponge, Anoxycalyx joubini was observed. This is the largest Antarctic sponge, with individuals growing over two meters tall. In order to investigate life history characteristics of Antarctic marine invertebrates, artificial substrata were deployed at a number of sites in the southern portion of the Ross Sea between 1967 and 1975. Over a 22-year period, no growth or settlement was recorded for A. joubini on these substrata; however, in 2004 and 2010, A. joubini was observed to have settled and grown to large sizes on some but not all artificial substrata. This single settlement and growth event correlates with a region-wide shift in phytoplankton productivity driven by the calving of a massive iceberg. We also report almost complete mortality of large sponges followed over 40 years. Given our warming global climate, similar system-wide changes are expected in the future.     

Temporal patterns of breeding and larval settlement in a temperate population of the tropical anemonefish,Amphiprion clarkii

The breeding of the anemonefish,Amphiprion clarkii, and the settlement of larvae were observed in temperate Japan. Spawning had two peaks near the first and the third quarter of the moon in 1983, but did not in 1984. The clutch size varied greatly during the breeding season, with a peak from the second half of June to the second half of July. Adult fishes inhibited larval settlement. The proportion of disappeared 0-year olds to the total was higher in 1984 than in 1983. This difference might be mainly due to the difference in the degree of crowding in each sea anemone. The period of larval settlement closely paralleled the breeding season. The larvae settled abundantly around the first quarter and the third quarter of the moon in 1983, and did from the third quarter to the first quarter in 1984. In both years, the number of settled larvae was more from the full moon to the new moon than the new moon to the full moon. The daily pattern of larval settlement did not reflect that of breeding.     

Population structure of Formica aquilonia (Hymenoptera,Formicidae) at the boundary of boreal and subboreal landscapes in the Volga River region

The data on the population structure of Formica aquilonia in the Middle Volga Area based on the variation of pilosity and coloration of workers are presented. Two population groups were distinguished by the pilosity characters: the northern group included the southern taiga population and the one located at the southern boundary of this subzone; the southern group comprised the populations of the mixed and broad-leaved forests. The pilosity characters reveal the F. aquilonia population structure at the regional level. They can be related to the ways of postglacial colonization of the territory and to the formation of the forest belt in the Quaternary. Phenotypic changes with respect to all the characters studied were revealed in the worker population of the anthill complex of F. aquilonia during 4 years of observation. They appear to have been caused by stochastic intrapopulation processes. The phenotypic homogeneity of the worker population was greater in the sites with higher settlement capacity (the sum of the areas of anthill dome bases related to the area of the settlement). Changes in the frequencies of the morphological characteristics were observed in the zones with low settlement capacity; the location of such zones has remained unchanged for some years. The stability of the spatial structure of the settlements appears to be associated with the distribution of forest biotopes suitable for dwelling of ants. The phenotypic heterogeneity of the complex has increased during the four-year period indicating the beginning of its fragmentation.     

Reef-scale failure of coral settlement following typhoon disturbance and macroalgal bloom in Palau,Western Pacific

Factors affecting coral recruitment are critical in influencing the scope and rate of reef recovery after disturbance. In December 2012, super-typhoon Bopha caused immense damage to the eastern reefs of Palau, resulting in near complete loss of coral cover. Within weeks following the typhoon, an ephemeral monospecific bloom of the foliose red macroalga Liagora (up to 40 % cover in February 2013) was recorded at impacted reefs with moderate wave exposure. Conversely, impacted and un-impacted reefs in areas of low wave exposure remained Liagora free. To quantify the effect of this ephemeral macroalgal bloom on coral recruitment, we installed settlement tiles during the major spawning period (March–April 2013) at forereefs with and without Liagora. Reefs (n = 3) with Liagora (13–24 % cover in April) experienced an almost complete failure of settlement, with only two individual corals recorded on settlement tiles (n = 90). This settlement failure was unexpected, as tiles were situated adjacent to, and not within Liagora canopies. In contrast, settlement was significantly higher on reefs that lacked macroalgae (n = 3), ranging from an average of 0.5–2.5 and 2.7–18.9 individuals 25 cm^?2 per top- and under-sided tile, respectively. Reefs with and without Liagora were in close proximity (≤8 km), and hydrodynamic models predicted that larval supply did not limit coral settlement among sites. While some differences in the community composition on the tiles were observed among sites, settlement substrate availability also did not limit coral settlement. Generalised linear mixed effects models indicated that while no settlement substrate explained more than 10 % of the variability in coral settlement, coral cover positively accounted for 26 %, and the cover of Liagora on reefs negatively accounted for more than 50 % of the observed variation. Combined, our results indicate that the typhoon induced ephemeral macroalgal bloom resulted in a reef-scale failure of coral settlement.     

Dendrochronological analyses of wood samples from a Late Bronze to early Iron Age site at Lake Luokesa,Lithuania

Tree rings of 184 archaeological wood samples from two Late Bronze to early Iron Age lake sites at Lake Luokesa (Luokesai e?eras), Lithuania, Moletai district, were analyzed. Despite severe difficulties with synchronization, Pinus (pine), Quercus (oak) and Alnus (alder) yielded some cross-datable series. The general picture is that the settlers chose small trees as timber, which they used in their natural round shape. The trees did not grow in homogeneous even-aged stands, but show very different ages and growth levels. Despite the generally low numbers of tree rings in the individual samples, the strong archaeological framework allowed cross-dating of some series and the building of chronologies for single structures. Based on these attempts, a 90 year long first floating chronology of the settlement structures is presented. Luokesa Site 2 (L2) was mainly built within the relative year 53. Luokesa Site 1 (L1) was certainly in use from the relative year 74 onwards. All fences at L1 show their main building activity in the relative year 81, four years after the main building activities in the village itself. It can be concluded that the settlement L1 was in use for at least 16 years. Because of the lack of a standard dendrochronological curve for the Baltic region, wiggle-matching was applied to obtain an absolute date for both settlements. The data clearly show that all samples relate to the Late Bronze–early Iron Age. The period where all wiggle matching results overlap is the period between 625 and 535 bc (the 2σ ranges are given). Based on the dating, duration and timber characteristics of the occupation, comparisons with Polish early Iron Age sites are made, which indicate a close resemblance in terms of wood use and settlement concept.     

Timing of Parturition, Planktonic Duration, and Settlement Patterns of the Black Rockfish, Sebastes inermis

The microstructure of otoliths was used to examine the relationship among timing of parturition, duration of planktonic stage, and temporal patterns of settlement in young-of-the-year (YOY) Sebastes inermis collected during the settlement seasons of 1998–2001 in a seagrass bed at the northwestern coast of Japan. Parturition occurred in winter, predominantly around the new and full moons, and parturition dates were distributed mainly from early January to late February. In the 4 years YOY S. inermis settled into distinctive groups at early spring, synchronized mainly to a semi-lunar cycle at new and full moon phases; except for 1998, in which settlement occurred around the first and third quarter moons. In all years settlers extruded earlier in the parturition season encompassed the first settlement groups that appeared in the bed and vice versa. Planktonic period ranged from 53 to 113 days, and its mean values varied significantly among settlement groups and years. Mean size at settlement showed less variability among settled groups and between years (range: 18–26mm total length), which suggested an ideal size for settling and active migration. On the basis of our findings we discuss the extent to which the 'tidal control', 'settlement linkage', and 'competent size' hypotheses explain the temporal pattern of settlement.     

Prehistoric settlement activities changed soil pH, nutrient availability, and growth of contemporary crops in Central Europe

Background and aims

How prehistoric human settlement activities have changed soil chemical properties, plant nutrition and growth of contemporary crops is a question that has not been satisfactorily addressed. The aim of this paper was to study to what extent nutrient availability in the soil, together with nutrition and growth of spring barley (Hordeum vulgare), improved on sites of former sunken buildings (cropmarks) in comparison to their surroundings (controls) 1,700 years after abandonment of the buildings.

Methods

In the Czech Republic, a unique prehistoric settlement with many sunken buildings was discovered during aerial reconnaissance from cropmarks in stands of cereals. Soil and biomass samples were collected from cropmarks and controls in a barley crop in June 2012.

Results

A substantially higher content of organic matter, higher pH and concentrations of plant-available (Mehlich III) P, Ca, Mg, Cu and Zn were recorded in the sub-soil layer in cropmarks compared with controls, indicating the accumulation of wood ash and organic waste. In the arable layer, pH and concentrations of P, Ca and Mg were generally very high in both positions. Cropmarks were characterised by barley plants that were twice as tall as the controls, with significantly higher Ca, Mg and P concentrations.

Conclusions

Prehistoric settlement activity affected nutrient availability and plant growth in the previously settled area even after 1,700 years. We conclude that the chemical signature of prehistoric settlement activity can be detected from chemical analysis of the sub-soil layer as well as analysis of the contemporary arable layer and crop biomass.     

The phlebotomine fauna (Diptera: Psychodidae) of Guaraí, state of Tocantins,with an emphasis on the putative vectors of American cutaneous leishmaniasis in rural settlement and periurban areas

Phlebotomine sandflies were captured in rural settlement and periurban areas of the municipality of Guaraí in the state of Tocantins (TO), an endemic area of American cutaneous leishmaniasis (ACL). Forty-three phlebotomine species were identified, nine of which have already been recognised as ACL vectors. Eleven species were recorded for the first time in TO. Nyssomyia whitmani was the most abundant species, followed by Evandromyia bourrouli, Nyssomyia antunesi and Psychodopygus complexus. The Shannon-Wiener diversity index and the evenness index were higher in the rural settlement area than in the periurban area. The evaluation of different ecotopes within the rural area showed the highest frequencies of Ev. bourrouli and Ny. antunesi in chicken coops, whereas Ny. whitmani predominated in this ecotope in the periurban area. In the rural settlement area, Ev. bourrouli was the most frequently captured species in automatic light traps and Ps. complexus was the most prevalent in Shannon trap captures. The rural settlement environment exhibited greater phlebotomine biodiversity than the periurban area. Ps. complexus and Psychodopygus ayrozai naturally infected with Leishmania (Viannia) braziliensis were identified. The data identified Ny. whitmani as a potential ACL vector in the periurban area, whereas Ps. complexus was more prevalent in the rural environment associated with settlements.     

The interplay of substrate nature and biofilm formation in regulating Balanus amphitrite Darwin, 1854 larval settlement

The settlement of marine larvae is influenced by a wide range of physical and biological factors. It is still poorly known how the nature of substrate and the biofilm can interact in regulating settlement patterns of invertebrate larvae. Here we use laboratory experiments focused on settlement behaviour of the barnacle Balanus amphitrite. The aim of this work is to understand whether: (i) the nature of substratum can affect biofilm formation and its structure, (ii) the nature of substratum can affect B. amphitrite larval settlement, (iii) the age of the biofilms and the nature of substrate can interact in influencing larval settlement.Four kinds of substrata (marble, quartz, glass, and cembonit) were biofilmed under laboratory conditions for 5, 10 and 20 days at the temperature of 28 °C. Settlement response was investigated with 5-day-old cyprids. Biofilms were quantitatively and qualitatively analysed by scanning electron microscopy. The settlement of B. amphitrite larvae significantly differed among substrata; also, the patterns of development of biofilm assemblages changed with substrate. In addition, the larval attractiveness of different substrates tends to disappear with biofilm age.     

The role of hydrodynamics in recruitment,growth, and survival of Argopecten irradians (L.) and Anomia simplex (D'Orbigny) within eelgrass meadows

Larvae of the bay scallop Argopecten irradians (L.) and the common jingle Anomia simplex (D'Orbigny) use byssal threads to settle onto blades of eelgrass Zostera marina (L.) and other elevated substrata. A 2-yr study of the role of hydrodynamics in recruitment of these bivalves into eelgrass meadows, and in their subsequent growth and survival as juveniles, was conducted in Northwest Harbor, New York. Sea-water flux through the meadow varied significantly among sites and depended on the numerical density of eelgrass shoots. Recruitment of both bivalves varied significantly among sites in a similar manner, indicating that hydrodynamics in eelgrass meadows strongly affects bivalve recruitment. Results from experimental deployments of two types of substrata indicate that hydrodynamics in eelgrass meadows exerts a stronger influence on bivalve recruitment than does either interblade abrasion or possible differences among sites in the nature of the blade substratum. In one of the two study years, hydrodynamics in eelgrass meadows influenced growth and survival of Argopecten following settlement. Abundances and growth of Anomia following settlement appeared unaffected by hydrodynamics. Several lines of evidence suggest that predation is less important than hydrodynamics in determining abundances of recruits and early juvenile stages of these bivalves.     

A fire history of a subalpine forest in south-eastern Wyoming, USA

Fire history was determined for part of the Routt‐Medicine Bow National Forest in south‐eastern Wyoming using fire‐scar and age‐class analysis. A composite chronology of fire events was used to determine mean fire intervals (MFI) for pre‐EuroAmerican settlement, EuroAmerican settlement (before 1868 ad ), EuroAmerican settlement and modern (after 1912) periods, for all fires and stand‐replacing fires. Point‐scale MFI was also determined using grand means from individual trees. Stand‐replacing fires were reconstructed to determine fire rotation. MFI for the entire time period is 5.5–8.4 years. MFI decreased from 9.3 to 15.7–1.9–2.9 years from the preto post‐EuroAmerican settlement periods, and increased during the modern period. Point‐scale MFIs are longer than MFI of the study area. Fire rotation is 182 years for the total period of record, but increased from 127 years during the pre‐EuroAmerican settlement period to 170 years during the EuroAmerican settlement period. Fire rotation during the modern period dramatically increased to 27,035 years. Results suggest fire suppression may have influenced the fire regime. Comparison of regional fire events with fire events from this study indicate regional weather has an important influence on Rocky Mountain fire regimes.     

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.     

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.     

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.     

贵州典型峰丛洼地区域近50年村落演变规律及驱动机制

揭示当前多种因素驱动下聚落演变模式与功能转型的地域差异具有重要意义。通过长时间序列的高清影像数据,研究典型峰丛洼地区不同自然和社会经济条件下,近50年来,尤其最近10年乡村聚落在多重背景下的演变模式、内在机制,及其地域差异。结果发现:(1)研究区聚落分布分部主要受耕地、地形和道路的影响,聚落面积增长热点和聚落斑块增长热点的耦合关系存在区域差异;(2)聚落格局演变存在集聚与均匀分布两种趋势;(3)研究区聚落特征和演变的阶段性地域差异十分明显;(4)喀斯特山区聚落的规模变化、演变阶段和演变模式既存在一些共同性的特征,又有各自的差异性;(5)外部因素和聚落农户的自我调适是演化的主要驱动力。     

Reproductive seasonality,settlement, and post-settlement mortality of Pocillopora damicornis (Linnaeus), at Lizard Island,Great Barrier Reef

Pocillopora damicornis (Linnaeus) has seasonal gametogenesis and planula release at Lizard Island, Great Barrier Reef, in contrast with several previous reports on the species at other locations. The number of planulae released and gonad development varied considerably among colonies sampled at the same time, but reproductive activity occurred predominantly in winter. P. damicornis planulae settled preferentially on algal-covered substrata, rather than bare coral substrata, but showed subsequent mortality inversely related to this settlement preference. Competition with algae and biological disturbance contribute to spat mortality at different stages of settlement and growth.     

Positive and negative effects of a dominant competitor on the settlement, growth, and survival of competing species in an epibenthic community

Because dominant competitors can monopolize resources, any positive effects they have on other species can have large community impacts. The solitary tunicate Ascidia ceratodes is a dominant competitor in the fouling community in Bodega Harbor, CA. This tunicate preempts primary substratum from competitors, but its thick tunic also allows other species to grow on its surface. The net effect of Ascidia on the community as a whole therefore depends on the balance between competitive and facilitative effects. In this study we evaluate the facilitative effects of Ascidia on different life stages of common competing species. We quantified larval settlement onto Ascidia compared to unoccupied space; we quantified the growth rate on Ascidia of small colonies of two common species; and we measured whether established colonies could escape overgrowth by Ascidia by growing onto its tunic. We found evidence for high rates of settlement on Ascidia, with some species showing higher and others lower settlement relative to that observed on free space. The growth rate of settlers was generally lower on Ascidia compared to primary substratum. We also found that colonial species established on primary space commonly escape overgrowth from Ascidia by growing onto Ascidia, as this occurs in about half of all encounters. This study indicates that the total effect of Ascidia on the community will depend on species-specific and life stage-specific effects, with likely long-term consequences for the diversity and composition of the community.     

Effects of GABA and epinephrine on the settlement and metamorphosis of the larvae of four species of bivalve molluscs

Competent larvae of different marine bivalve species were treated with GABA and epinephrine at different concentrations and times of exposure to test the ability of the drugs to induce settlement and metamorphosis. GABA induced both settlement and metamorphosis in the mussel Mytilus galloprovincialis, the clams Venerupis pullastra and Ruditapes philippinarum and the oyster Ostrea edulis. Maximum induction of settlement (>39%) was achieved after exposure of V. pullastra larvae to 10⁻⁴ M GABA; this concentration of GABA also induced the highest percentages of metamorphosis in the four species studied. Epinephrine was identified as an active inducer of settlement and metamorphosis in bivalve molluscs. Exposure to 10⁻⁵ M epinephrine induced significant levels of settlement in Mytilus, Venerupis and Ostrea. In contrast, epinephrine failed to induce settlement behaviour in Ruditapes. Maximum induction of metamorphosis was produced by 10⁻⁵ M epinephrine in mussels, clams and oysters; Ruditapes showed the highest percentage of metamorphosis (>78%). This is the first report in which the involvement of GABA in the settlement and metamorphosis of bivalve molluscan larvae is demonstrated. It was also recognised that epinephrine plays a role not only in inducing metamorphosis but also in initiating settlement.