Effects of substrate type on growth and mortality of blue mussels (<Emphasis Type="Italic">Mytilus</Emphasis><Emphasis Type="Italic">edulis</Emphasis>) exposed to the predator <Emphasis Type="Italic">Carcinus</Emphasis><Emphasis Type="Italic">maenas</Emphasis>

Structure and complexity of the substrate are important habitat characteristics for benthic epifauna. The specific growth and mortality rates and inducible defence characters on medium-sized blue mussels (Mytilus edulis L.) exposed to shore crabs (Carcinus maenas L.) were examined on three different substrate types in combined field and laboratory experiments. The experiments showed that complexity of the substrate increased blue mussel survival significantly, through a decrease in predation pressure. However, increased intraspecific competition for food on the complex substrate resulted in significantly lower growth rates of the mussels. Inducible defence characters were also influenced by substrate type. Blue mussels were more affected by predators on the structurally simple substrate, where they developed thicker shells and a larger posterior adductor muscle.     

Mechanisms of range expansion in the intertidal brown alga <Emphasis Type="Italic">Fucus</Emphasis><Emphasis Type="Italic">serratus</Emphasis> in northern Spain

The intertidal brown alga Fucus serratus is naturally expanding its distributional range along the north shore of Spain. Shores being colonised by F. serratus harbour saturated algal communities, with almost no free space. Processes responsible for the successful expansion were investigated in a locality where the alga was experimentally introduced. After a short period, the number of F. serratus plants increased exponentially at a monthly intrinsic rate of 0.111. At early and intermediate stages of invasion, colonisation occurred by the growth of patches of the invader around the transplant points. F. serratus is a bad disperser, with most of propagules settling in near proximity to parent plants. Colonisation of manipulated and control areas was influenced by the proximity of propagule sources. Experimental removals of canopies, irrespective of whether the turf was removed or left intact, accelerated the expansion, as this component of the community had a negative influence on recruitment of both F. serratus and F. vesiculosus. The unmanipulated algal beds were, however, invasible, as some control quadrats close (<10 m) to F. serratus patches had a 100% cover of F. serratus at the end of the study. Prior or concurrent to increases of F. serratus, a decrease in cover of F. vesiculosus and other resident macroalgae was evident. No formal competition experiment was done but results suggest that F. serratus outcompetes F. vesiculosus and Bifurcaria bifurcata at mid-tidal levels. In scraped quadrats, in which both Fucus species recruited at large densities, F. serratus grew at a higher rate than F. vesiculosus and eventually monopolised the experimental surfaces.     

Behavioral adaptations of the cubozoan medusa <Emphasis Type="Italic">Tripedalia cystophora</Emphasis> for feeding on copepod (<Emphasis Type="Italic">Dioithona oculata</Emphasis>) swarms

The cubozoan medusa Tripedalia cystophora preys on dense swarms of the copepod Dioithona oculata in the mangrove prop-root habitat of Puerto Rico. The copepod swarms form in shafts of sunlight that penetrate the mangrove canopy during the day. T. cystophora are found primarily within the same illuminated areas, feeding heavily on the dense swarms of copepods. Laboratory studies were conducted to examine the behavioral adaptations of T. cystophora that allow them to remain within these dense copepod swarms. In the presence of vertical light shafts, T. cystophora showed both a significant change in swimming speed and a significant increase in turning rate. T. cystophora turn abruptly when they leave a vertical shaft of light and quickly re-enter. The presence or absence of copepod prey, either in swarms or dispersed, had no significant effect on T. cystophora swimming speed or turning behavior. Feeding rates of T. cystophora on the copepod D. oculata increase rapidly with copepod density, and maximum rates are not achieved until the high densities associated with swarms are reached. It seems doubtful that these small medusae could survive without their ability to encounter and exploit these dense swarms of copepods.     

Evidence of a spawning area of <Emphasis Type="Italic">Anguilla</Emphasis><Emphasis Type="Italic">marmorata</Emphasis> in the western North Pacific

Leptocephali of the tropical eel Anguilla marmorata have been consistently collected in the same area of the North Equatorial Current (NEC) in the western North Pacific during three consecutive cruises in June and July of 1991 (N=28) and 1994 (N=20), and July and September of 1995 (N=27), indicating that this is a spawning area of this species. These leptocephali were collected from 130°E to 142°E and 12°N to 20°N, to the west of the Mariana Islands, in 20 tows in 1991, in 13 tows in 1994 and in 17 tows in 1995, indicating a widespread presence, but a relatively low abundance. Six of these specimens (16.3-36.0 mm total length) from the 1995 cruise, which were of the typical size range of these leptocephali, were genetically confirmed to be A. marmorata in a previous study. The consistent presence of recently spawned A. marmorata leptocephali (9-20 mm) in all 3 years, suggests that the western region of the NEC is the spawning area of the northern population of A. marmorata that was identified in a recent population genetics study. These leptocephali would thus be transported westward by the NEC and then transported north into the Kuroshio Current and toward Taiwan and Japan, or south toward the southern Philippines and into the Celebes Sea by the Mindanao Current. Available evidence indicates that A. marmorata may have potentially year-round spawning, and the presence of a spawning area of this species in the same region as that of Anguilla japonica suggests that the northern population of A. marmorata has evolved a spawning migration from East Asia, the Philippines and the Celebes Sea region to the NEC area, but differs from A. japonica by having some individuals that recruit to more southern areas.     

Life on a hot dry beach: behavioural,physiological, and ultrastructural adaptations of the littorinid gastropod <Emphasis Type="Italic">Cenchritis</Emphasis> (<Emphasis Type="Italic">Tectarius</Emphasis>) <Emphasis Type="Italic">muricatus</Emphasis>

The distribution and abundance of the littorinid gastropod Cenchritis (Tectarius) muricatus were determined on an exposed shore at Lee Stocking Island, Bahamas in July/August 1995 and June 1999. Activity patterns were observed on two occasions shortly before a hurricane passed near the island in 1995. C. muricatus extended from the low water mark to 3.6 m vertical height above it. Highest densities (20/m²) were found just above the high tide mark but the animal remained relatively abundant (8/m²) at the highest point. Small animals were only found at high shore levels. With the exception of animals very near to the water's edge C. muricatus was inactive by day and on dry nights. Many individuals of the high shore populations became active at night during periods of high humidity and rainfall, moving distances of up to 4 m overnight. At lower levels, extensive pairing and copulation was associated with these periods of rainfall. Abundance changed dramatically after the passage of the hurricane. At the level of maximum abundance numbers were reduced from 20/m² to 2.8/m². Long-term experimental studies of water loss rate and haemolymph concentration showed that both water loss rate and increase in haemolymph concentration are greatest in the early period of desiccation. The concentration of the blood after 20 weeks is 250% of the original and the water lost is of the order of 22.5%. After an initial increase there appears to be evidence of regulation of haemolymph concentration possibly by exchange with free water in the shell itself. The considerable reserve capacity in tolerance is no doubt in part due to the structure and functioning of the kidney. Adaptive features of the excretory system identified from ultrastructural examination include the replacement of podocyte-containing filtration chambers in the auricle by extracellular tubules permeating the walls of both auricle and ventricle, which suggests a reduction in the rate of primary urine formation. This is also indicated by a reduced surface area of the nephridial gland, which resorbs organic solutes from the urine. Excretory cells contain single large vacuoles in which arise large multi-layered concretions composed of phospholipids and calcium salts typical of prosobranchs in which excretory products may be stored for long periods.     

Cardiac output during exposure to <Emphasis Type="Italic">Chattonella</Emphasis><Emphasis Type="Italic">marina</Emphasis> and environmental hypoxia in yellowtail (<Emphasis Type="Italic">Seriola</Emphasis><Emphasis Type="Italic">quinqueradiata</Emphasis>)

The experiments were undertaken to measure, for the first time, cardiac output in yellowtail (Seriola quinqueradiata) during exposure to a harmful red tide flagellate (Chattonella marina). The responses were compared with those during exposure to environmental hypoxia to evaluate the significance of the drop of arterial oxygen partial pressure (PaO₂) in the fish-kill mechanisms by C. marina. PaO₂ immediately decreased, whereas heart rate (HR) was maintained until shortly before death during exposure to C. marina. Suffocation developing during the exposure resulted from a decrease in blood oxygen content, but not from lowered blood flow to the tissue. Although exposure to both C. marina and hypoxia immediately decreased PaO₂, arterial oxygen content (CaO₂) and pH (pHa) were significantly lower, whereas HR and cardiac output (&Qdot;) remained significantly higher, for the C. marina-exposed fish than for hypoxia-exposed fish. Although the drop in PaO₂ appears to play a pivotal role in the mechanisms of fish death by C. marina, other physiological response(s) should also be considered.     

Dynamics of pigment degradation by the copepodite stage of <Emphasis Type="Italic">Pseudodiaptomus</Emphasis><Emphasis Type="Italic">euryhalinus</Emphasis> feeding on <Emphasis Type="Italic">Tetraselmis</Emphasis><Emphasis Type="Italic">suecica</Emphasis>

Short-term (3 h) changes in concentration of chlorophylls and their derivatives in stage V Pseudodiaptomus euryhalinus and their fecal material were followed by HPLC during a 24 h experiment. Copepodites were fed with the prasinophyte Tetraselmis suecica. Intact chlorophyll a and b were found in animals and fecal material and had similar dynamics of accumulation over time. The extent of transformation of chlorophyll a and b to colorless compounds was different with chlorophyll a being more extensively degraded. Additionally, several chlorophyll derivatives (pheophytin and pyropheophytin-like pigments) were found. Pyropheophytin a was the most abundant followed by pheophytin b, pheophytin a, and pheophorbide a. Relative amounts of pheopigments were different in copepodites and fecal material, and pheophytin a, pheophorbide a, and pheophytin b concentrations were low and variable. The amount of ingested chlorophyll recovered as chlorophyll a and its derivatives in fecal and copepodite pools was generally low (<5%), with one exception occurring after 9 h, when it accounted for >70%. These data suggest individual pheopigments are produced at different rates and that chemical or enzymatic mechanisms in the gut of copepodites act on the two chlorophylls in different ways.     

Settlement of the wreckfish (<Emphasis Type="Italic">Polyprion</Emphasis><Emphasis Type="Italic">americanus</Emphasis>)

The wreckfish (Polyprion americanus) is a panoceanic, long-lived, deep-water demersal species, characterized by an extended pelagic juvenile phase associated with floating objects. In the present study morphometric data from 435 specimens collected from Greece (Crete), Italy (Ionian Sea) and the Atlantic coast of France from September 1999 to March 2001 were analyzed to estimate the settlement size of the species and associated changes in morphology. The fishery sample included specimens from both the pelagic and the demersal stage. Length-at-settlement (TL₅₀) and 95% confidence intervals were estimated by fitting a logistic function. Eleven morphometric characters were analyzed, and the existing inflection points, reflecting alterations in body shape, were defined: (1) iteratively, as the transition point, splitting the data set into two groups, for which the reduced major axis functions, between a character and total length, of the successive groups best fit (minimum sum of squares of residuals) the combined data set and (2) where the second derivative of the fitted third-order polynomial functions to morphometric ratios equaled zero. The main size range within which wreckfish settled was 56-65 cm, and the means of the inflection points defined by the two methods were 61 and 64 cm. These results indicate changes in functional morphology associated with settlement. The monitoring of wreckfish juveniles caught in the wild and kept individually in captivity showed changes in growth and food intake at approximately the estimated settlement length and suggested that temperature was the most likely factor triggering settlement.     

Seasonal variation in the degree of heterospecific association of two syntopic hyraxes (<Emphasis Type="Italic">Heterohyrax</Emphasis><Emphasis Type="Italic">brucei </Emphasis>and <Emphasis Type="Italic">Procavia capensis</Emphasis>) exhibiting synchronous parturition

Adult rock hyrax (Procavia capensis) and bush hyrax (Heterohyrax brucei) allocate much of their surface activity to being vigilant for vertebrate predators. Individuals of the two species associate for thermoregulatory basking on rock outcrops (koppies) and, in the Matobo National Park, Zimbabwe, frequently constitute heterospecific groups that produce offspring synchronously. Exposure to predators during basking is thought to put a high premium on vigilance, particularly during the presence of dependent offspring in nurseries. We predicted, therefore, an increase in the association of the two species of hyraxes when offspring were present because this would lead to larger groups, thus enhancing the ability to detect predators. Koppie aggregations of hyraxes were classified as homospecific or heterospecific according to basking group composition. Heterospecific association increased when offspring were present, and heterospecific groups were larger than homospecific ones. Fifty-one percent of nurseries were heterospecific for adults/subadults. During the pre-weaning phase (<2 months after birth), 85% of P. capensis and 65% of H. brucei juveniles were in nurseries of heterospecific basking groups. Heterospecific groups had more offspring than homospecific groups. Juvenile mortality is most evident after young are weaned and heterospecific associations break down. Heterospecific aggregation, through increased group size, may result in a higher probability of avoiding predators, thereby reducing the vulnerability of young.     

Distribution of the alien ctenophore <Emphasis Type="Italic">Mnemiopsis</Emphasis><Emphasis Type="Italic">leidyi</Emphasis> in the Caspian Sea in August 2001

In this study, spatial and vertical distributions of the invasive ctenophore Mnemiopsis leidyi in the Caspian Sea were evaluated by using data collected at 41 stations during the August 2001 cruise. A comparison of data from different depths revealed that M. leidyi were generally confined to surface waters. The maximum size of the ctenophore was only 41-45 mm, and the bulk of individuals (85.5%) were <10 mm in length. The average and maximum biomasses of M. leidyi were calculated as 120 and 351 g wet weight m^-2, respectively. Whilst highest biomasses were observed in the western and central Middle Caspian Sea, hot spot areas of reproduction were present along the coasts of the western Caspian Sea, with abundance values of up to 2285 ind. m^-2. The impact of such high densities of M. leidyi is expected to be significant for the pelagic ecosystem of the Caspian Sea.     

Effects of symbiotic status,flow speed,and prey type on prey capture by the facultatively symbiotic temperate coral <Emphasis Type="Italic">Oculina</Emphasis><Emphasis Type="Italic">arbuscula</Emphasis>

Symbiotic temperate corals can supplement prey capture by the coelenterate host with autotrophic carbon production by endosymbiotic zooxanthellae. To test the relationship between heterotrophic consumption and photosynthetic energy, prey capture by symbiotic and aposymbiotic specimens of the temperate scleractinian coral Oculina arbuscula (Verrill) was measured in January-April 2001. Corals were tested in a laboratory flume at five flow speeds, using Artemia franciscana cysts and nauplii as prey. Per-polyp capture rate and feeding efficiency were independent of symbiotic condition. Capture rate increased with flow speed, while capture efficiency declined. The location of capture shifted from the upstream to downstream side of the coral as flow speed increased. Differences in capture rate, location, and feeding efficiency for cysts and live brine shrimp nauplii were likely due to prey size rather than swimming ability.     

Competition,fluctuating asymmetry and sperm transfer in male gryllid crickets (<Emphasis Type="Italic">Gryllus bimaculatus</Emphasis> and <Emphasis Type="Italic">Gryllodes sigillatus</Emphasis>)

Sperm competition theory predicts that ejaculate expenditure by males should increase with the risk of competition from rivals. The tendency to modulate expenditure, however, may reflect various constraints acting on different individuals. We test this idea here by looking at the effects of competition and fluctuating asymmetry (FA) in two species of gryllid cricket. Male Gryllus bimaculatus and Gryllodes sigillatus modulated the sperm content of their spermatophores in relation to both apparent competition and FA in limb size. Males of both species increased the amount of sperm transferred when an apparent male competitor was present, sperm number increasing regardless of the species of apparent competitor in G. bimaculatus, but significantly more with a conspecific competitor in G. sigillatus. In addition, the relative size of the spermatophylax in G. sigillatus increased in the presence of apparent competition, while spermatophore transfer time decreased in G. bimaculatus. Sperm number showed opposite relationships with limb asymmetry in the two species, decreasing with increasing asymmetry in males in G. sigillatus, but increasing with asymmetry in both males and females in G. bimaculatus. G. bimaculatus, but not G. sigillatus, also transferred more sperm when paired with a larger female. Relationships between reproductive measures and limb asymmetry were significant only when competitors were present in G. bimaculatus, but were significant regardless of competition in G. sigillatus. The differences may reflect the additional burden of producing a spermatophylax in G. sigillatus.     

Population dynamics and secondary production of the wedge clam <Emphasis Type="Italic">Donax</Emphasis><Emphasis Type="Italic">hanleyanus</Emphasis> (Bivalvia: Donacidae) on a high-energy,subtropical beach of Brazil

The population biology of Donax hanleyanus (Philippi, 1845) (Bivalvia: Donacidae) was studied by monthly sampling from June 1998 through June 2000, at Restinga da Marambaia Beach, Brazil. Two transects were established and divided into ten strata parallel to the waterline, and five replicates were taken with a 0.04 m² sampler in each stratum. The highest densities of D. hanleyanus were recorded in winter (September 1998, July 1999) and autumn (April 2000). A stratified distribution was observed: recruits were found mainly in the middle swash zone, while juveniles and adults occurred across the tidal gradient up to the retention zone. Mortality rates did not differ significantly between years. Annual production (in ash-free dry mass) ranged from 0.76 g AFDM m^-2 year^-1 (1998) to 3.67 g AFDM m^-2 year^-1 (1999), while the production-to-biomass (P/B) ratio varied from 1.45 to 1.59. Life span was ca. 17 months. Of all variables tested, only two, one biological and one physical, seem to have influenced the population dynamics of this species. A significant negative correlation between the densities of the suspension-feeders D. hanleyanus and Emerita brasiliensis (Crustacea: Decapoda) indicated possible intraguild competition, and there was a significant exponential correlation between beach slope and the mean across-shore position of D. hanleyanus. Data compiled from literature suggests a latitudinal gradient in population parameters, with mortality and renewal rate (P/B) of different species of Donax increasing and life span decreasing from temperate to tropical regions. Other factors influencing population dynamics, such as food availability, and the contributions of individual and community biomass to energy and nutrient cycling are also discussed.     

Doubly uniparental transmission of mitochondrial DNA length variants in the mussel <Emphasis Type="Italic">Mytilus</Emphasis><Emphasis Type="Italic">trossulus</Emphasis>

Doubly uniparental inheritance of mitochondrial DNA (mtDNA) is an unusual feature found in marine mussels and a few other bivalve species. In addition to a mitochondrial genome (F) inherited through the female line, heteroplasmic individuals (males) contain a second highly diverged genome (M) that is inherited through the paternal line. The Baltic mussel Mytilus trossulus is characterized by the presence of two phylogenetically close female and male genomes. We examined M. trossulus sampled from the Gulf of Gdansk, southern Baltic. Somatic tissues and gametes were surveyed for the presence of different types of mtDNA. Length variants were identified using PCR (polymerase chain reaction) amplification of the mtDNA fragment containing the major noncoding region. Seventeen length variants present in homo- and heteroplasmic individuals were found in a sample of 343 individuals analyzed. Two length variants of the major noncoding region (PCR product lengths: 1520 and 1370 bp) were the most frequent. Length heteroplasmy was found in 46.8% of males, and was mostly caused by the presence of the short length variant (PCR product length: 1370 bp) of the major noncoding region. MtDNA variants were also detected by restriction analysis of a 1.3 kb segment of coding region (ND2-COIII) amplified by PCR. This study provides evidence that the short length variant is always transmitted to sperm and has taken over the role of the M genome. The remaining length variants were present in both males and females. Longer length variants were transmitted mainly through the female line. A possibility of much higher incidence of genome role reversals in Baltic M. trossulus, in comparison with other populations of mussels is discussed.     

Feeding ecology of the mangrove crab <Emphasis Type="Italic">Ucides cordatus</Emphasis> (Ocypodidae): food choice,food quality and assimilation efficiency

Food preferences, consumption rates and dietary assimilation related to food quality were investigated for the large semi-terrestrial and litter-consuming mangrove crab Ucides cordatus cordatus (Ocypodidae, L. 1763) in northern Brazil. Stomach contents were composed of mangrove leaves (61.2%), unidentified plant material and detritus (28.0%), roots (4.9%), sediment (3.3%), bark (2.5%), and animal material (0.1%). U. cordatus prefers Rhizophora mangle over Avicennia germinans leaves despite a higher nitrogen content, lower carbon to nitrogen (C/N) ratio and lower tannin content of the latter. Also, assimilation rates for senescent R. mangle leaves (C: 79.3%, N: 45.4%) were higher than for A. germinans leaves (C: 40.6%, N: 9.1%). Faeces composition indicates that A. germinans leaves were more difficult to masticate and digest mechanically. The leaf-ageing hypothesis, according to which crabs let leaves age in burrows to gain a more palatable and nutritive food, was rejected for U. cordatus since N content, C/N ratio and the abundance of microorganisms did not differ significantly between senescent leaves and leaves taken from burrows. The low microbial biomass on leaf surfaces and in the sediment indicates its minor importance for the nutrition of U. cordatus. It is concluded that high ingestion and assimilation rates of a R. mangle diet together with the consumption of algae allow for a high intake of C, N, and energy. The data suggest that the digestibility of mangrove leaves by U. cordatus is not hampered by tannins. This may have provided a competitive advantage over other leaf-consuming invertebrates unable to digest mangrove litter with high tannin concentrations. Due to the large stock biomass of U. cordatus in the study area, a great amount of finely fragmented faeces is produced (about 7.1 ton dry matter ha⁻¹ year⁻¹ in a R. mangle forest) which is enriched in C, N and bacterial biomass compared to the sediment. The decomposition of mangrove litter, and thus nutrient and energy transfer into the sediment, is greatly enhanced due to litter processing by U. cordatus.     

Foraging responses in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae) to floral scents

Summary. For butterflies to be efficient foragers, they need to be able to recognize rewarding flowers. Flower signals such as colours and scents assist this recognition process. For plant species to attract and keep butterflies as pollinators, species-specific floral signals are crucial. The aim of this study is to investigate foraging responses to floral scents in three temperate butterfly species, Inachis io L. (Nymphalidae), Aglais urticae L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae), in behavioural choice bioassays. The butterflies were allowed to choose bet-ween flower models varying in scent and colour (mauve or green). Flowers or vegetative parts from the plants Centaurea scabiosa L. (Asteraceae), Cirsium arvense (L.) (Asteraceae), Knautia arvensis (L.) (Dipsacaceae), Buddleja davidii Franchet (Loganicaeae), Origanum vulgareL. (Lamiaceae), Achillea millefolium L. (Asteraceae), and Philadelphus coronarius L. (Hydrangiaceae) were used as scent sources. All visits to the models — those that included probing and those that did not — were counted, as was the duration of these behaviours. Both flower-naive and flower-experienced (conditioned to sugar-water rewards, the colour mauve, and specific floral scents) butterflies were tested for their preference for floral versus vegetative scents, and to floral scent versus colour. The butterflies were also tested for their ability to switch floral scent preferences in response to rewards. Flower-naive butterflies demonstrated a preference for the floral scent of the butterfly-favourable plants C. arvense and K. arvensis over the floral scent of the non-favourable plants Achillea millefolium (Asteraceae), and Philadelphus coronarius cv. (Hydrangiaceae). Most of the butterflies that were conditioned to floral scents of either C. arvense, K. arvensis, or B. davidii readily switched theirfloral scent preferences to the one most recently associated with reward, thus demonstrating that floral scent constancy is a result from learning. These findings suggest that these butterflies use floral scent as an important cue signal to initially identify and subsequently recognize and distinguish among rewarding plants. Received 2 September 2001; accepted 9 September 2002.     

Global phylogeography of the fissiparous sea-star genus <Emphasis Type="Italic">Coscinasterias</Emphasis>

The fissiparous starfish genus Coscinasterias (Verrill) is represented in shallow waters around many of the world's continents. This wide distribution could be explained by dispersal, vicariance, or translocation associated with shipping, and represents an excellent system for marine biogeographic research. We conducted a global phylogeographic analysis of 42 Coscinasterias mtDNA cytochrome oxidase 1 sequences (15 haplotypes) from 18 sites, including representatives of all four recognised species. Phylogenetic analysis yielded a robust phylogeny, with strong support for the monophyly of the genus (90% bootstrap support) and of each separate species (99-100%). Haplotypes exhibited strong phylogeographic structure, with robust mtDNA clades often associated with distinct land masses. A general lack of genetic differentiation within sites may reflect fissiparity. However, shared haplotypes over larger distances (e.g. across Japan), and the presence of related haplotypes on adjacent land masses (e.g. Tasmania, New Zealand; 1.6-1.8%) suggest that long-distance dispersal is an important biogeographical process for Coscinasterias. The 4.0-4.4% divergence between Japanese and South African sister groups may relate to transequatorial dispersal around the onset of the Pleistocene. Divergent Atlantic and Mediterranean populations of C. tenuispina (maximum 1.5%) may deserve subspecific recognition, and high divergences within Australian C. muricata (maximum 8.0%) suggest a species complex.     

Multiple-front iron-mineralisation in chiton teeth (<Emphasis Type="Italic">Acanthopleura</Emphasis><Emphasis Type="Italic">echinata</Emphasis>: Mollusca: Polyplacophora)

The mineralisation of iron in the major lateral teeth of the chiton Acanthopleura echinata has been examined using light and electron microscopy, together with energy dispersive and Raman spectroscopy. The deposition of magnetite in the posterior region of the tooth cusp has been shown to occur simultaneously on two fronts, from both the anterior and posterior sides. In contrast, in the lepidocrocite region, the mineral is formed by aggregation over the whole region. In addition, evidence is presented for the existence of a thin veneer of ferrihydrite over the surface of the magnetite region. The veneer preserves this region from oxidation prior to use in the mature tooth. Data also suggest that the junction zone plays a vital role in the overall biomineralisation process, contributing large amounts of iron at critical stages of development.     

Development and biology of <Emphasis Type="Italic">Periphylla</Emphasis><Emphasis Type="Italic">periphylla</Emphasis> (Scyphozoa: Coronatae) in a Norwegian fjord

The globally distributed coronate scyphomedusa Periphylla periphylla (Peron and Lesueur, 1809) occurs in permanent and extremely high abundance in some Norwegian fjords. Studies on the species in Lurefjorden, 40 km north of Bergen, have revealed a holopelagic life cycle with direct development. We distinguished 14 successive developmental stages, characterised by size and morphology. Eggs and early stages are non-mobile, neutrally buoyant, and found mainly at intermediate depths. Rearing studies indicated a development time of 2-3 months from fertilisation to stage 9, when the medusa starts feeding and becomes motile. The fjord population usually shows a strong diel vertical migration, with aggregations at shallower depths during the night. The development includes a gradual increase in pigmentation (porphyrins), starting with the stomach, and thereafter extending to tentacles and the whole exumbrella. This pigment is photodegraded by natural light. Rearing experiments in the laboratory have shown a lethal effect of light. Here, development stops at stage 5. Pigments become visible in stage 7. Older medusae are severely damaged and killed within a few days by exposure to daylight. All stages of P. periphylla are bioluminescent, with this capacity increasing with growth, probably in connection with the development of the nervous net. The light reaction starts at the point of stimulation and spreads at two different speeds corresponding to the two nerve nets of the medusae. The coronal furrow has a central function in transmission of the reaction. It is hypothesised that the bioluminescence is used mainly as a warning to some potential predators, signalling that the porphyrin-containing medusa is unpalatable. P. periphylla has the same two categories of nematocysts as other coronates, with a total of six different types. Nematocyst abundance, distribution and morphology indicate their function. The giant euryteles (capsule up to 100 µm and tubule up to 1160 µm) are the largest ones known among Scyphozoa, and are unique in that this size increases during medusa development.