Effects of seasonality and population density on the reproduction of the Indo-Pacific echinoid<Emphasis Type="Italic"> Echinometra mathaei</Emphasis> in Kenyan coral reef lagoons

Reproduction in the widely distributed tropical sea urchin Echinometra mathaei (de Blainville) was examined in three Kenyan reef lagoons that differed in substrate cover and E. mathaei population density. Histological examination of the gonads and gonad index measurements for 3 years showed a seasonal reproductive cycle with gametogenesis commencing in July, when temperature and light are at their lowest, and spawning commencing in December and peaking in February to May, when temperature and light reach their annual maxima. Monthly gonad indices correlated significantly with seawater temperature and light intensity. Male urchins had mature gametes for a longer period (8 months) than females (6 months), possibly an adaptive strategy that increases the probability of fertilization. Lunar periodicity was not observed, as male and female gonads were full of gametes on all days of the lunar cycle during the period of spawning. The peak in spawning activity coincides with the peak in phytoplankton abundance, which could ensure high food availability for the planktonic larvae. Gonad weights were significantly higher relative to urchin weight at the reef having the largest mean sizes and lowest population of urchins, indicating availability of food resources for growth and reproduction. E. mathaei at the reef with the highest density had the smallest urchins and high relative gonad sizes, indicative of a tradeoff between growth and reproduction when food is limited.Communicated by O. Kinne, Oldendorf/Luhe     

Temporal and spatial variability in settlement of the sea urchin<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> in the NW Mediterranean

Settlement into the benthic habitat may be an important process in regulating sea urchin abundance, which potentially modifies the structure of benthic communities. Strong settlement events may increase sea urchin abundance beyond a certain threshold, leading to the formation of coralline barrens (overgrazed communities with a dominance of encrusting coralline algae). To understand the role of settlement in regulating sea urchin populations we first need to determine settlement variability. Temporal variation in settlement of the sea urchin Paracentrotus lividus was monitored at three sites in the Medes Islands, NW Mediterranean, during three settlement seasons (March 1998 through October 2000). Spatial variation in settlement was studied in 1999 at 50 sites along a gradient of exposures to waves and currents, inside and outside the archipelago, and separated by distances from tens to thousands of meters. Bathymetric distribution of settlement was also studied in 2000 at six sites at 5, 10, 15 and 20 m depths. Settlement of P. lividus occurred in a single annual peak within 3 weeks in May–June. Differences in settlement between years were more than two orders of magnitude. Spatial variability was found at all scales investigated, showing strong patchiness at the smallest spatial scales (tens of meters). Sea urchins settled preferentially at depths between 5 and 10 m. Substratum type, level of protection, and adult population densities were not significant in determining settlement. However, settlement was found to be related to the degree of exposure to waves and currents, indicating that physical processes are very important at the spatial scales investigated. This greatly variable settlement is a necessary, although not sufficient, condition to create gradients of adult P. lividus abundance. Further studies should be designed to investigate the interaction between settlement strength and post-settlement mortality.Communicated by O. Kinne, Oldendorf/Luhe     

Field depuration and biotransformation of paralytic shellfish toxins in scallop<Emphasis Type="Italic"> Chlamys nobilis</Emphasis> and green-lipped mussel<Emphasis Type="Italic"> Perna viridis</Emphasis>

Under laboratory conditions, the scallop Chlamys nobilis and the mussel Perna viridis were exposed to N-sulfocarbamoyl toxins (C2 toxin), a paralytic shellfish toxin (PST), by feeding a local toxic strain of the dinoflagellate Alexandrium tamarense (ATDP) that produced C2 toxin exclusively. The bivalves were subsequently depurated in the field, and their depuration kinetics, biotransformation and toxin distribution were quantified. Depuration was characterized by a rapid loss within the first day, followed by a secondary slower loss of toxins. In the fast depuration phase, scallops detoxified PSTs more quickly than the mussels (depuration rate constants for scallops and mussels were 1.16 day^–1 and 0.87 day^–1, respectively). In contrast, the mussels detoxified PSTs more quickly than the scallops in the slow depuration phase, and the calculated depuration rate constants (mean+SE) from day 2 to day 13 were 0.063+0.009 day^–1 and 0.040+0.019 day^–1 for mussels and scallops, respectively. The differences in the appearances of gonyautoxins, GTX2 and GTX3, and their decarbamoyl derivatives, dcGTX2, dcGTX3 and GTX5, which are all derivatives of C2 toxin, indicated active and species-specific biotransformation of the algal toxins in the two bivalves. In both species of bivalves, the non-viscera tissue contained fewer toxins and lower concentrations than the viscera-containing tissue compartment. In scallops, very little toxin was distributed in the adductor muscle. In mussels, most of the PSTs were found in the digestive gland with significant transport of toxins into the digestive gland from other tissues during the course of depuration. The toxin profiles of scallops and mussels differed from each other and from that of the toxic algae fed. A significant fraction of GTX5 was detected in the mussels but not in the scallops. Our study demonstrates a species specificity in the depuration kinetics, biotransformation and tissue distribution of PSTs among different bivalves.Communicated by T. Ikeda, Hakodate     

Feeding selectivities of the marine cladocerans<Emphasis Type="Italic"> Penilia avirostris</Emphasis>,<Emphasis Type="Italic"> Podon intermedius</Emphasis> and<Emphasis Type="Italic"> Evadne nordmanni</Emphasis>

We conducted grazing experiments with the three marine cladoceran genera Penilia, Podon and Evadne, with Penilia avirostris feeding on plankton communities from Blanes Bay (NW Mediterranean, Spain), covering a wide range of food concentrations (0.02–8.8 mm³ l^–1, plankton assemblages grown in mesocosms at different nutrient levels), and with Podon intermedius and Evadne nordmanni feeding on the plankton community found in summer in Hopavågen Fjord (NE Atlantic, Norway, 0.4 mm³ l^–1). P. avirostris and P. intermedius showed bell-shaped grazing spectra. Both species reached highest grazing coefficients at similar food sizes, i.e. when the food organisms ranged between 15 and 70 µm and between 7.5 and 70 µm at their longest linear extensions, respectively. E. nordmanni preferred organisms of around 125 µm, but also showed high grazing coefficients for particles of around 10 µm, while grazing coefficients for intermediate food sizes were low. Lower size limits were >2.5 µm, for all cladocerans. P. avirostris showed upper food size limits of 100 µm length (longest linear extension) and of 37.5 µm particle width. Upper size limits for P. intermedius were 135 µm long and 60 µm wide; those for E. nordmanni were 210 µm long and 60 µm wide. Effective food concentration (EFC) followed a domed curve with increasing nutrient enrichment for P. avirostris; maximum values were at intermediate enrichment levels. The EFC was significantly higher for P. intermedius than for E. nordmanni. With increasing food concentrations, the clearance rates of P. avirostris showed a curvilinear response, with a narrow modal range; ingestion rates indicated a rectilinear functional response. Mean clearance rates of P. avirostris, P. intermedius and E. nordmanni were 25.5, 18.0 and 19.3 ml ind.^–1 day^–1, respectively. Ingestion rates at similar food concentrations (0.4 mm³ l^–1) were 0.6, 0.8 and 0.9 g C ind.^–1 day^–1.Communicated by O. Kinne, Oldendorf/Luhe     

Patterns of gamete incompatibility between the blue mussels<Emphasis Type="Italic"> Mytilus edulis</Emphasis> and<Emphasis Type="Italic"> M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis>

Previous research on gametic incompatibility in marine invertebrates suggests that for highly dispersive marine invertebrate species, barriers to fertilization among closely related taxa are often incomplete and sometimes asymmetric. The nature of these barriers can dramatically affect the patterns of gene flow and genetic differentiation between species, and thus speciation. Blue mussels, in the genus Mytilus, are genetically distinct in allopatry yet hybrids are present wherever any two species within the group co-occur. The present study sampled M. edulis (L.) and M. trossulus (Gould) in May and June 2001 from the East Bay section of Cobscook Bay, Maine, USA (latitude 44°56′30″N; longitude 67°07′50″W), where the two species are sympatric. Gamete incompatibility was investigated in a series of laboratory fertilizations carried out in July 2001. The proportion of fertilized eggs typically exceeded 80% at sperm concentrations of 10³–10⁴ ml^?1 among intraspecific matings (n=18), but was <30% even at sperm concentrations in excess of 10⁵–10⁶ ml^?1 for interspecific matings (n=13). Further analysis indicated that approximately 100- to 700-fold higher sperm concentrations were required to achieve 20% fertilization in interspecific matings relative to intraspecific matings, indicating strong barriers to interspecific fertilization. The proportion of fertilized eggs did not follow this general pattern in all matings, however. The eggs from two (out of five) M. edulis females were almost as receptive to M. trossulus sperm as they were to M. edulis sperm. In contrast, the eggs from all M. trossulus females (n=3) were unreceptive to M. edulis sperm, suggesting that fertilization barriers between these species may be asymmetric. Given the experimental design employed in this study, the results are also consistent with a strong maternal or egg effect on the level of interspecific gamete compatibility in M. edulis.     

Gonadal development in the coral reef damselfish<Emphasis Type="Italic"> Dascyllus flavicaudus</Emphasis> from Moorea,French Polynesia

The present study investigated the sexual pattern of the yellow-tailed dascyllus, Dascyllus flavicaudus (Randall and Allen), through histological examination of the process of gonadal differentiation and maturation and through male-removal experiments in the laboratory. The study was conducted in Moorea, French Polynesia (17°32S; 149°50W), from 31 December 1994 to 11 March 1995. For gonadal histology, two populations were sampled, one consisting of large aggregations (n=54) and the other consisting of small- to medium-sized groups around isolated corals (n=55). An additional small sample (n=21) was also collected from a population that consisted of small groups around isolated corals. After an initially undifferentiated state, gonads of D. flavicaudus developed an ovarian lumen, followed by oocytes in the primary growth stage. From this ovarian state or from more developed ovaries with cortical-alveolus stage oocytes, some gonads developed into testes through degeneration of oocytes and development of spermatogenic tissue. In all three populations, spermatogenic tissue developed only in gonads that contained pre-vitellogenic oocytes (termed as mixed stage 1–2 gonads). The two main populations did not differ in their expression of sex change despite differences in their social organization. In both populations, size of individuals with mixed stage 1–2 gonads overlapped mainly with the size range of immature females, which suggests that functional female-to-male sex change was rare. The hypothesis that D. flavicaudus is primarily gonochoristic is further supported by removal experiments in laboratory groups, in which removal of a dominant male(s) failed to induce sex change in any of the high-ranking females. Yet, in all three field populations, some fish with mixed stage 1–2 gonads were found within the size ranges of mature females, which indicates possible occurrences of functional sex change. These conflicting results indicate that it may be premature to draw any definitive conclusions about the sexual pattern of D. flavicaudus.Communicated by T. Ikeda, Hakodate     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Effect of sunlight intensity and albinism on the covering response of the Caribbean sea urchin<Emphasis Type="Italic"> Tripneustes ventricosus</Emphasis>

At Discovery Bay, Jamaica, Tripneustes ventricosus lives in beds of the turtle grass Thalassia testudinum. Especially during daylight hours, it covers its aboral surface with fragments of this plant and other objects. Normally pigmented, wild-type sea urchins covered themselves significantly less with Thalassia when sunlight was experimentally decreased to 66% or 32% ambient intensity. Consistent with this result, naturally occurring sea urchins exhibited significantly less covering at a deep (3.5 m) site than at a shallow (1 m) site, where light intensities at the bottom were 619 and 946 mol s^–1 m^–2, respectively. The graded covering response to light intensity suggests that covering is a defense against damaging solar radiation. Albino sea urchins covered themselves significantly more with Thalassia than wild-type sea urchins in both full and 66% sunlight. In addition, at the shallow site where they accounted for about 4% of the population, they showed significantly greater covering than wild-type urchins. The greater covering response of albino sea urchins suggests a greater susceptibility to solar radiation.Communicated by P. W. Sammarco, Chauvin     

Histological investigation of the reproductive cycles of the limpets <Emphasis Type="Italic">Patella </Emphasis><Emphasis Type="Italic">vulgata</Emphasis> and <Emphasis Type="Italic">Patella ulyssiponensis</Emphasis>

This study devised a staging system for, and monitored, the gonad development of the limpet species Patella vulgata and Patella ulyssiponensis on the South West coast of Ireland using histological techniques. Maturation began in the males of both species in January and in the females it began in March. There was no statistical difference in gonad development between sexes and between species. Spawning in the male P. vulgata occurred from September to December 2003 and in September and October 2004. In female P. vulgata spawning occurred from October to December 2003, no spawning of females was observed in 2004. In male P. ulyssiponensis spawning occurred in November and December 2003 and from September 2004 to December 2004. Spawning was observed from November 2003 to January 2004 and in September 2004 in female P. ulyssiponensis. Sex ratios also varied between the species and between months sampled. Nevertheless more males were observed in both species.     

Food web analysis in intertidal <Emphasis Type="Italic">Zostera marina</Emphasis> and <Emphasis Type="Italic">Zostera noltii</Emphasis> communities in winter and summer

The food web of two intertidal seagrass (Zostera marina and Zostera noltii) beds that may be influenced by the seasonal variation in food source abundance was studied in winter and in summer with δ¹³C and δ¹⁵N analysis. In spite of high relative variation of abundance of main primary producers at the two sites, the food web did not vary between winter and summer. The δ¹³C range of primary producers was wide. Zostera leaves, the most ¹³C-enriched source, were not consumed directly by grazers. Deposit and filter feeders have a similar δ¹³C and could use a mix of suspended and sedimented organic particulate matter, largely composed of detritus from macroalgae to seagrass. This trophic pathway allows the local incorporation of the high biomass produced by seagrasses. The wide δ¹⁵N range of predators was linked either to a large variety from omnivore to carnivore predators or to the also wide ranges of δ¹⁵N of primary consumers.     

Localization of infaunal prey by the sea star<Emphasis Type="Italic"> Leptasterias polaris</Emphasis>

We performed field and laboratory studies to investigate how large adult Leptasterias polaris detect and locate their major prey, large infaunal bivalves, in the sediment bottom community. A field survey using SCUBA diving showed that 95% of the locations where L. polaris dug into the sediment bottom were over bivalves and this success rate was much greater than if digging was done at random (22%). Furthermore, when sea stars were provided with a low density of randomly distributed prey in a laboratory arena, they dug exclusively in locations where a clam had been buried. These observations indicated that L. polaris locates infaunal prey prior to investing energy into digging. Studies in a laboratory flow tank showed that L. polaris readily detected and moved towards its preferred prey Ensis directus whereas its responses to less preferred prey Mya truncata and Spisula polynyma were much weaker. The degree to which it oriented towards these three common prey seemed to reflect potential energy intake relative to foraging costs (which likely increase with the depth of the different prey) and risks from interactions with other carnivores (which are greatest when feeding on large prey). This is the first study to clearly demonstrate that sea stars use prey odours to locate infaunal prey.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-004-1497-1Communicated by R.J. Thompson, St. Johns     

Larval metamorphosis of the sea urchins,<Emphasis Type="Italic"> Pseudocentrotus depressus</Emphasis> and<Emphasis Type="Italic"> Anthocidaris crassispina</Emphasis> in response to microbial films

In Japan, mass-production of sea urchin juveniles involves the culture of periphytic diatom films on plastic plates in 5- to 15-tonne tanks for the induction of larval metamorphosis. This study focused on the larval response of sea urchins, Pseudocentrotus depressus and Anthocidaris crassispina, to natural microbial films in the sea and diatom-based films formed in the tanks. The effect of diatoms and bacteria on larval metamorphosis was also examined using laboratory-cultured diatom-based films in the presence of germanium dioxide and antibiotics during film culture. Furthermore, the nature of the cue of the cultured diatom-based film was also investigated. Results showed that P. depressus and A. crassispina metamorphosed both on natural microbial films and diatom-based films in a tank. In the sea, the metamorphosis (%) of P. depressus increased gradually in accordance with the immersion period of film formed on glass slides, whereas the larval metamorphosis of A. crassispina had a bell-shaped response curve. In the tank, although the diatom-based films showed a low inducing activity for larval metamorphosis of A. crassispina, the metamorphosis of P. depressus larvae increased linearly in accordance with the diatom density. These results suggest that diatom-based films could promote the larval metamorphosis of P. depressus, but are less important in A. crassispina. In a simultaneous larval assay (May), P. depressus showed a higher percentage of metamorphosis than A. crassispina. We concluded that the former is more sensitive to diatom-based film than the latter and that this is due to differences in their natural habitats. For laboratory-cultured diatom-based film, both species of sea urchins showed a similar response, in which reduction in diatom and bacteria density resulted in a decrease in the original inducing activity. There seems to be a synergistic effect between diatom and bacteria in inducing larval metamorphosis. Films subjected to treatment with 0.1 N HCl were no longer inductive for either sea urchin, while those films treated with 40^°C heat or EtOH (5% and 10% EtOH) showed a significant reduction in the inducing activity. Thus the surface-associated cues may be highly susceptible to the above treatments.Communicated by T. Ikeda, Hakodate     

Ground gleaning in horseshoe bats: comparative evidence from<Emphasis Type="Italic"> Rhinolophus blasii</Emphasis>,<Emphasis Type="Italic"> R. euryale</Emphasis> and<Emphasis Type="Italic"> R. mehelyi</Emphasis>

The 71 species of horseshoe bat (genus Rhinolophus) use echolocation calls with long constant-frequency (CF) components to detect and localize fluttering insects which they seize in aerial captures or glean from foliage. Here we describe ground-gleaning as an additional prey-capture strategy for horseshoe bats. This study presents the first record and experimental evidence for ground-gleaning in the little-studied Blasius horseshoe bat (Rhinolophus blasii). The gleaning bouts in a flight tent included landing, quadrupedal walking and take-off from the ground. The bats emitted echolocation calls continuously during all phases of prey capture. Both spontaneously and in a choice experiment, all six individuals attacked only fluttering insects and never motionless prey. These data suggest that R. blasii performs ground-gleaning largely by relying on the same prey-detection strategy and echolocation behaviour that it and other horseshoe bats use for aerial hawking.We also studied the Mediterranean horseshoe bat (R. euryale) in the flight tent. All four individuals never gleaned prey from the ground, though they appeared to be well able to detect fluttering moths on the ground. It is not known yet whether ground-gleaning plays a role in Mehelys horseshoe bat (R. mehelyi). In a performance test, we measured the ability of these three European species of middle-sized horseshoe bats (R. euryale, R. mehelyi and R. blasii) to take-off from the ground. All were able to take flight even in a confined space; i.e. the willingness to ground-glean in R. blasii is not related to a superior take-off performance. In contrast to ground-gleaning bats of other phylogenetic lineages, R. blasii appears not to be a specialist, but rather shows a remarkable behavioural flexibility in prey-capture strategies and abilities. We suggest that the key innovation of CF echolocation paired with behavioural flexibility in foraging strategies might explain the evolutionary success of Rhinolophus as the second largest genus of bat.Communicated by T. Czeschlik     

Heart rate reflects osmostic stress levels in two introduced colonial ascidians <Emphasis Type="Italic">Botryllus schlosseri</Emphasis> and <Emphasis Type="Italic">Botrylloides</Emphasis><Emphasis Type="Italic">violaceus</Emphasis>

The influence of abiotic factors on the establishment and success of invasive species is often difficult to determine for most marine ecosystems. However, examining this relationship is critical for predicting the spread of invasive species and predicting which habitats will be most vulnerable to invasion. Here we examine the mortality and physiological sensitivity to salinity of adult colonies of the colonial ascidians Botryllus schlosseri and Botrylloides violaceus. Adult colonies of each species were exposed to abrupt changes in salinity (5, 10, 15, 20, 25, 30 psu) in the laboratory. Salinity ranges used in the laboratory corresponded with those of the field distributions of B. violaceus and B. schlosseri in the Great Bay Estuary, NH. Heart rate was used as a proxy for health to assess the condition of individual colonies. Heart rates were monitored daily for approximately 2 weeks. Results revealed that both species experienced 100% mortality after 1 day at 5 psu and that their heart rates declined with decreasing salinity. Heart rates of B. schlosseri remained consistent between 15 and 30 psu and slowed at 10 psu. Heart rates of B. violaceus remained constant between 20 and 30 psu, but slowed at 15 psu. These laboratory results corresponded to the distribution of these species in estuaries, indicating salinity is a key factor in the distribution and dominance of B. schlosseri and B. violaceus among coastal and estuarine sites. Furthermore, physiological differences to salinity were found between colonies of B. schlosseri in the Venetian Lagoon and colonies in Portsmouth Harbor, suggesting adaptation to environmental variables.     

In vitro reproduction of the turbellarian<Emphasis Type="Italic"> Urastoma cyprinae</Emphasis> isolated from<Emphasis Type="Italic"> Mytilus galloprovincialis</Emphasis>

Urastoma cyprinae is a species infecting the gills of several marine bivalves. Although there is some literature on this turbellarian, its life cycle remains unknown. In our work we have demonstrated that reproduction of U. cyprinae can be completed out of its host. More than 50% of turbellarians isolated from mussels (Mytilus galloprovincialis) secreted and cemented a cocoon to the well bottom during the first 72 h of incubation in seawater (34 salinity) at 14°C. Oviposition started at days 1–18 (average 4.8 days) and occurred in most cases inside the protective cocoon. Each Urastoma laid an average of 2.9 egg capsules (range 1–10) and 3.9 embryos were developed inside each egg capsule (range 1–11). Hatching started at days 20–43 (average 24 days). An average of 12.8 juvenile forms (range 1–64) escaped from the cocoon after hatching. The free-swimming juveniles showed a positive phototactic response and survived about a month after hatching. On the basis of our results, we propose a life cycle for U. cyprinae involving a sexual maturation parasitic period in the bivalve gills and a reproduction period including cocoon secretion, egg laying, and hatching that is entirely completed in the external environment.Communicated by S.A. Poulet, Roscoff     

Geographical genetic structure and phylogeography of the <Emphasis Type="Italic">Sargassum horneri</Emphasis>/<Emphasis Type="Italic">filicinum</Emphasis> complex in Japan,based on the mitochondrial <Emphasis Type="Italic">cox3</Emphasis> haplotype

The genetic structure and phylogeography of the brown seaweed Sargassum horneri/filicinum complex in Japan were studied based on the mitochondrial cox3 haplotype. The cox3 haplotypes found were divided into three clades in a statistical parsimony network, among which there were large numbers of steps. Contrary to the reported large amount of drifting S. horneri along the Japanese coast, the three clades were dividedly distributed on the Japanese coast: the northern Pacific, the central Pacific, and western Japan. The western Japan S. horneri had haplotypes that were phylogenetically closer to those of S. filicinum than to the northern and central Pacific S. horneri populations. The S. filicinum populations were included within the western Japan clade and grouped together with the S. horneri samples from western Japan. Taken together with the unstable morphological diagnosis, this result suggests that S. filicinum should be reduced into a synonymy of S. horneri. The TMRCA analysis suggested that the divergence time of each clade may go back to the last interglacial period and a skyline plot suggested that the last glacial maximum had only a small effect on the population size of S. horneri. The geographic subdivision of the three groups, in spite of a large amount of drifting mats, suggests a limited contribution of drifting mats to gene flow on a large geographic scale. On a small geographic scale, a small number of haplotypes were shared between S. horneri-type and S. filicinum-type populations. This result suggests that populations of these two types are partially, though not completely, isolated from each other, possibly by selfing in S. filicinum-type populations or by a difference in peak reproduction.     

Morpho-functional defences of Mediterranean sea urchins,<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> and<Emphasis Type="Italic"> Arbacia lixula</Emphasis>, against fish predators

Morpho-functional features potentially involved in defence mechanisms against fish predators (i.e. attachment tenacity, spine length, and test robustness and thickness) have been assessed in two Mediterranean sea urchins, Paracentrotus lividus and Arbacia lixula. All four morpho-functional features were significantly and positively related to individual size for both species of sea urchins. Test robustness (i.e. static load needed to break sea urchin tests) was significantly greater for A. lixula (from 3,450 to 15,000 g depending on size) than for P. lividus (1,180–11,180 g). Attachment tenacity (i.e. force needed to dislodge sea urchins from the rocky substrate) was greater in A. lixula (280–3,300 g) than in P. lividus (110–1,450 g), and the difference tended to decrease in relation to smaller sea urchin size. Spine length was greater in A. lixula (1.5–2.9 cm) than in P. lividus (0.5–2.3 cm), but the difference decreased for larger sea urchin size. Test thickness was slightly greater (but not significantly) in A. lixula (0.35–1.10 mm) than in P. lividus (0.12–0.90 mm). These results provide evidence that morpho-functional features of sea urchins could be involved in affecting predation rates by fishes upon P. lividus and A. lixula, with potential implications for the population structure and distribution patterns of the two sea urchins in shallow rocky reefs.Communicated by R. Cattaneo-Vietti, Genova     

Characterization of urease activity in three marine phytoplankton species,<Emphasis Type="Italic"> Aureococcus anophagefferens</Emphasis>,<Emphasis Type="Italic"> Prorocentrum minimum</Emphasis>, and<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis>

The availability of different forms of nitrogen in coastal and estuarine waters may be important in determining the abundance and productivity of different phytoplankton species. Although urea has been shown to contribute as much as 50% of the nitrogen for phytoplankton nutrition, relatively little is known of the activity and expression of urease in phytoplankton. Using an in vitro enzyme assay, urease activities were examined in laboratory cultures of three species: Aureococcus anophagefferens Hargraves et Sieburth, Prorocentrum minimum (Pavillard) Schiller, and Thalassiosira weissflogii (Grunow) Fryxell et Hasle. Cultures of P. minimum and T. weissflogii were grown on three nitrogen sources (NO₃^m, NH₄⁺, and urea), while A. anophagefferens was grown only on NO₃^m and urea. Urease was found to be constitutive in all cultures, but activity varied with growth rate and assay temperature for the different cultures. For A. anophagefferens, urease activity varied positively with growth rate regardless of the N source, while for P. minimum, urease activity varied positively with growth rate only for cultures grown on urea and NH₄⁺. In contrast, for T. weissflogii, activity did not vary with growth rate for any of the N sources. For all species, urease activity increased with assay temperature, but with different apparent temperature optima. For A. anophagefferens, in vitro activity increased from near 0-30°C, and remained stable to 50°C, while for P. minimum, increased in vitro activity was noted from near 0-20°C, but constant activity was observed between 20°C and 50°C. For T. weissfloggii, while activity also increased from 0°C to 20°C, subsequent decreases were noted when temperature was elevated above 20°C. Urease activity had a half-saturation constant of 120-165 wg atom N l^у in all three species. On both an hourly and daily basis, urease activity in A. anophagefferens exceeded nitrogen demand for growth. In P. minimum, urease activity on an hourly basis matched the nitrogen demand, but was less than the demand on a daily basis. For T. weissflogii, urease activity was always less than the nitrogen demand. These patterns in urease activity in three different species demonstrate that while apparently constitutive, the regulation of activity was substantially different in the diatom. These differences in the physiological regulation of urease activity, as well as other enzymes, may play a role in their ecological success in different environments.