Leading role of the sea urchin <Emphasis Type="Italic">Arbacia lixula</Emphasis> in maintaining the barren state in southwestern Mediterranean

Sympatric sea urchin species are usually considered to belong to the same grazer guild. Nevertheless, their role in community dynamics may vary due to species-specific morphological traits, feeding preferences and foraging behavior. In the Mediterranean Sea, the two species Paracentrotus lividus and Arbacia lixula co-occur in barren areas. Whereas P. lividus is usually considered responsible for creating a barren ground, the roles of the two sea urchin species in its maintenance are currently unclear. The relative and combined effects of P. lividus and A. lixula on maintaining the benthic community in the barren state were tested experimentally, using orthogonal exclusion of the two species. Results show that exclusion of A. lixula, regardless of the presence of P. lividus, led to a significant decrease in the surface of bare rock and a significant change of the algal assemblages, thus demonstrating the major role of this species in maintaining the barren state.     

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     

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     

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.     

Time and space: genetic structure of the cohorts of the common sea urchin <Emphasis Type="Italic">Paracentrotus</Emphasis><Emphasis Type="Italic">lividus</Emphasis> in Western Mediterranean

Spatio-temporal variability in settlement and recruitment, high mortality during the first life-history stages, and selection may determine the genetic structure of cohorts of long-lived marine invertebrates at small scales. We conducted a spatial and temporal analysis of the common Mediterranean Sea urchin Paracentrotus lividus to determine the genetic structure of cohorts at different scales. In Tossa de Mar (NW Mediterranean), recruitment was followed over 5 consecutive springs (2006–2010). In spring 2008, recruits and two-year-old individuals were collected at 6 locations along East and South Iberian coasts separated from 200 to over 1,100 km. All cohorts presented a high genetic diversity based on a fragment of mtCOI. Our results showed a marked genetic homogeneity in the temporal monitoring and a low degree of spatial structure in 2006. In 2008, coupled with an abnormality in the usual circulation patterns in the area, the genetic structure of the southern populations studied changed markedly, with arrival of many private haplotypes. This fact highlights the importance of point events in renewing the genetic makeup of populations, which can only be detected through analysis of the cohort structure coupling temporal and spatial perspectives.     

Geographic,population, and seasonal patterns in the reproductive parameters of the sea urchin <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>

Generalized additive models were used to synthesize the data from 52 publications on the gonad growth and spawning seasonality of Paracentrotus lividus and identify spatial and temporal patterns in the reproductive processes. According to our results, Atlantic populations spawn in spring and develop large gonads (6.4 ± 0.2% of fresh body weight (mean ± SE)), which increase in size toward higher latitudes. While in the Mediterranean, the gonads are smaller (3.1 ± 0.1%), possibly as the result of successive spawnings which impede nutrients accumulating in the gonads. Differences were also observed between habitats: gonad production was higher in seagrass meadows in the Mediterranean and in subtidal rocky substrates in the Atlantic. Food availability might cause these variations as well as the negative correlation observed between depth and the gonad index for rocky substrates. Sex and body size do not seem to influence greatly gonad growth.     

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.     

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     

Acetylcholinesterase activity is affected by stress conditions in<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> coelomocytes

The presence and function of acetylcholinesterase (AChE) in blood cells and plasma is well known. The enzyme activity is associated to red blood cell membranes, as well as to the perinuclear region of leucocytes, suggesting a role in the function of the latter. Recently coelomocytes, acknowledged progenitors of immune cells in vertebrate systems, have been shown to respond to stress conditions, both in laboratory experiments and field studies. Bearing in mind the similarity between coelomocytes and vertebrate blood cells, we investigated the presence of AChE activity in coelomocytes of the sea urchin Paracentrotus lividus characterized by specific inhibitors, in relation to cold stress induced in the laboratory. Exposure to stress was followed by electrophoretic analysis of AChE activity and Western blot analysis of AChE-like immunoreactivity. Moreover, quantitative analysis of the different molecular forms of AChE was carried out with the aid of a spectrophotometric method. The effects of stress on AChE activity were time dependent. In fact, the enzyme activity was detected by the catalytic activity in cleaving the substrate acetyl thiocholine iodide, which appears to be formed by different isoforms of the enzyme, compared to that of Electrophorus, which was used as a control. Western blot analyses showed the presence of different immunoreactive molecules, with molecular mass ranging from 265 kDa (corresponding to Electrophorus molecules) to about 180 kDa. According to quantitative analysis, the isoforms that were differentiated corresponded to an AChE with the capacity to cleave acetyl--methyl thiocholine iodide (the so-called true AChE, E.C. 3.1.1.7, corresponding to the 265 kDa band) and to a ChE with the capacity to cleave propionyl thiocholine iodide as well. The latter is an ancestral form of AChE, mainly present in invertebrates. Each of these forms was affected differently by varying time exposures.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.     

Human impact on <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>: the result of harvest restrictions and accessibility of locations

The sea urchin Paracentrotus lividus is common in the Mediterranean Sea in shallow subtidal rocky habitats, and it is intensely harvested for commercial and recreational purposes. This study is aimed at investigating whether the effects of harvest restrictions of P. lividus in rocky reef habitats interact with the accessibility of locations in structuring sea urchin population (total and commercial-sized individuals). These results are important for generating hypotheses about the influence of human harvesting on P. lividus and for addressing suitable measures of conservation. Paracentrotus lividus was sampled after the end of the sea urchin harvesting period (May–July 2007) within the Gulf of Alghero (North West Sardinia), where the Capo Caccia–Isola Piana Marine Protected Area (MPA) was established since 2002. Paracentrotus lividus was sampled at sixteen locations and attributed in groups of four to 4 combinations of harvest restrictions (Restricted Harvest, RH, vs. Unlimited Harvest, UH) and accessibility (Boat vs. Car), which correspond to a gradient of potential human activity on P. lividus in the ranked order of very low (RHBoatfar), low (RHBoatclose), moderate (RHCar) and high (UHCar). At each location, two depth ranges of 3–7 and 8–12 m were considered. At each of these depths, two areas of about 100-m² size were chosen. The density of P. lividus was assessed in ten quadrats of 1 × 1 m, and the size of 100 individuals (test diameter) was considered. Human activity has been found to significantly affect population structure of P. lividus influencing the proportion of individuals larger than 50 mm. Although harvest was restricted by MPA regulations, a significantly lower abundance of large individuals was found at sites accessible by car. This result highlights that there is an effect of harvest restrictions in relation to accessibility and emphasizes the need to carefully address the enforcement of the MPA toward easily accessible sites. Thus, surveillance and investment in enforcement of marine reserves seem crucial points that may provide the greatest return on maintaining the ecological benefits to the fishery activities.     

Classification of juvenile rockfish,<Emphasis Type="Italic"> Sebastes inermis</Emphasis>, to<Emphasis Type="Italic"> Zostera</Emphasis> and<Emphasis Type="Italic"> Sargassum</Emphasis> beds,using the macrostructure and chemistry of otoliths

Young-of-the-year (YOY) Sebastes inermis use Zostera and Sargassum beds as nursery grounds, although it is not known which habitat YOY prefer. In this study, YOY S. inermis were accurately assigned to Zostera or Sargassum beds by two approaches: the width and length of the otolith nucleus and the composition of trace elements in otoliths. The otolith nucleus was initially opaque and then showed a marked shift to hyaline deposition once YOY settled in the nursery grounds. The first hyaline zone (FHZ) was deposited earlier in Zostera beds (from mid-May to early June) than in Sargassum beds (around mid-summer). Likewise, irrespective of settlement year, the FHZ was formed at both significantly younger ages and shorter back-calculated sizes (total length, TL) in the Zostera bed (overall mean: 131±3 days; 2.5±1.7 mm TL) than in the Sargassum bed (overall mean: 158±12 days; 61.3±1.00 mm TL). YOY collected in the Zostera bed were born earlier (mainly in January) than YOY from the Sargassum bed (mainly in February). In addition, a significant linear relationship was found between the age at formation of the FHZ and nucleus dimensions, suggesting that nucleus dimensions were a reliable macroscopic indicator of the time of formation of the FHZ and, consequently, also an indicator of the nursery where YOY grew. Linear discriminant function analysis (LDFA) based on the width and length of the otolith nucleus could distinguish juveniles from Zostera (88–96%) and Sargassum (96–97%) beds with a high degree of accuracy. In the other approach, six detectable trace elements (Li, Mn, Ni, Cu, Zn, and Ba) in otoliths of YOY collected in the nursery grounds were measured by high-resolution, inductively coupled mass spectrometry. LDFA based on the trace elemental composition separated YOY from three nurseries with 100% of accuracy. Findings suggest that both the trace elemental composition and nucleus dimensions of otoliths can be used as natural tags of the nursery grounds of S. inermis, offering a considerable potential for answering questions on habitat use and the contribution of nursery grounds to the adult stock.Communicated by T. Ikeda, Hakodate     

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.     

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     

Intraspecific competition in<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> F</Emphasis>.<Emphasis Type="Italic"> evanescens</Emphasis> (Phaeophyceae: Fucales) germlings: effects of settlement density,nutrient concentration,and temperature

Survival and growth of early post-settlement stages are critical for the development of seaweed populations. Fucoid germlings commonly settle in dense monospecific aggregates, where intraspecific competition and environmental variables (e.g. nutrient concentration and temperature) may affect survival and growth. Using factorial experiments, we determined the effects of settlement density (~10, ~50 and ~250 germlings cm^–2), nutrient enrichment (from ~10 to ~40 µM N and from ~0.5 to ~2.5 µM P), and temperature (7°C and 17°C) on Fucus serratus and F. evanescens germlings in laboratory cultures over 3 months. Settlement density, nutrient concentration and temperature interactively affected growth of germlings, and the magnitude of this interaction varied between the two species. This represents the first record of such factorial interactions in Fucus spp. germlings. Intraspecific competition, estimated as the relative reduction in germling growth and survival from low to high densities, increased with decreasing nutrient concentration and increasing temperature in both species. While temperature and nutrient concentration had little effect on germling size distributions, size inequality and skewness generally increased with germling density, indicating that a few large individuals gained dominance and suppressed many smaller ones at high density. Self-thinning increased with settlement density and depended on nutrient concentration and species at high density. At high density, self-thinning increased with decreasing nutrient levels in F. evanescens, but not in F. serratus. At low density, nutrient enrichment increased germling growth in F. evanescens, but not in F. serratus, whereas growth in both species was stimulated by nutrient enrichment at higher densities. These results suggest that germling growth and self-thinning are more sensitive to variation in nutrient concentration in F. evanescens than in F. serratus. The potential implications of our findings for the understanding of eutrophication-related abundance changes in both species in southern Norway are discussed.Communicated by L. Hagerman, Helsingør     

Competitive interactions between two fucoid algae with different growth forms,<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> Himanthalia elongata</Emphasis>

Competitive interactions between two fucoid algae with different growth forms, Fucus serratus L. and Himanthalia elongata (L.) S.F. Gray were examined both in the laboratory and on a shore of the Isle of Man, Irish Sea. The growth of germlings of both species declined with increasing density, irrespective of whether they were with cohorts or rival species, indicating that intra- and interspecific competition occurred between germlings. H. elongata suppressed the performance of F. serratus at the germling stage by virtue of its larger initial size, and at the mushroom stage by forming a miniature canopy with the caps of the adjacent plants. In a field experiment, the mortality of H. elongata juveniles generally increased in mixtures with F. serratus and was highest when F. serratus were 50% of the plants. At the juvenile stage, the negative effect of F. serratus on H. elongata was more severe than the other way round. This was because F. serratus grows predominantly upwards, whereas H. elongata had already begun to expand laterally at the distal end. If F. serratus survives in sparse mixed stands with H. elongata juveniles, it can overgrow them and inhibit their subsequent survivorship and growth, probably by both shading and physical sweeping. H. elongata and F. serratus maintain their discrete monospecific stands because of the varying outcomes of mutual competitive exclusion resulting from their differing growth patterns. Thus it is possible for them to co-occur at a similar shore height.Communicated by O. Kinne, Oldendorf/Luhe     

Competition and facilitation between germlings of<Emphasis Type="Italic"> Ascophyllum nodosum</Emphasis> and<Emphasis Type="Italic"> Fucus vesiculosus</Emphasis>

Competitive interactions between germlings of Ascophyllum nodosum (L) Le Jolis and Fucus vesiculosus L. were studied both in the laboratory and on a shore of the Isle of Man, in the Irish Sea. Both intra- and interspecific competition were investigated by comparing the performance of algal germlings both in monocultures and mixed populations of the two species. The growth of germlings of both species reduced with increasing density. F. vesiculosus always grew faster than Ascophyllum and did best in mixed cultures, whereas Ascophyllum did least well when mixed with Fucus germlings. Clearly the adverse effects of F. vesiculosus on A. nodosum were greater than those of Ascophyllum cohorts. At the same total density, the survival and growth of Ascophyllum declined with an increasing proportion of Fucus germlings, implying that poor recruitment of A. nodosum results from strong competition with F. vesiculosus. However, under desiccation stress on the shore, F. vesiculosus enhanced the survival of A. nodosum at the early germling stage even though competition may occur again at the late stage. Thus, whether interactions between germlings take the form of competition or facilitation depends on the environmental conditions.Communicated by O. Kinne, Oldendorf/Luhe     

<Emphasis Type="Italic">Patella aspera</Emphasis> and<Emphasis Type="Italic"> P. ulyssiponensis</Emphasis>: genetic evidence of speciation in the North-east Atlantic

The group of subspecies of Patella ulyssiponensis, described by Christiaens, was widely known as Patella aspera until recently. The group extends throughout the Mediterranean, on all Macaronesian islands, along the North African coast, and in Europe, as far north as southern Norway. Throughout its range it displays great variation in shell sculpture and colour. The aim of this work was to re-examine the various subspecies proposed by Christiaens and, genetically, to test the hypothesis that European continental populations belong to a different species than the one composed by Macaronesian (north-west African) populations. In the present work, this group was studied by allozyme electrophoresis of 21 loci and by six morphological variables. The monomorphic locus of malate dehydrogenase (Mdh-1) was found to be diagnostic for distinguishing European continental populations from those of north-west African archipelagos, confirming the specific status of both groups. The allele observed at this locus in the Macaronesian populations was novel, while European continental populations showed a plesiomorphic one, shared with all other north-west Atlantic patellids with the exception of Patina pellucida, suggesting a more recent origin of the Macaronesian species from the continental forms. Both species showed a genetic identity of 0.730±0.061, which allowed rough estimations of 6.5–3 Mya since speciation. The subspecies Patella ulyssiponensis deserta described by Christiaens was not confirmed by our genetic data. Other loci (glyceraldehyde-3-phosphate dehydrogenase and the second locus of lactate dehydrogenase) were partially diagnostic, in which both species showed different most common alleles. Morphologically, both species are easily recognised by shell characters and the results agree with previous findings, that continental populations are more homogeneous in shell morphology and radula characters than populations from the Macaronesia. This work supports retention of the earliest valid name, Patella ulyssiponensis Gmelin, with Lisbon, Portugal as type locality, for the European continental species, and Patella aspera Röding for the Macaronesian populations. Population subdivision within species was measured by theta, the estimator of F_st, showing in both P. aspera and P. ulyssiponensis a high degree of genetic structuring (=0.226 and 0.182, respectively) mostly explained by the large distances separating the populations within species.Communicated by J.P.Thorpe, Port Erin     

Auditory defence in the peacock butterfly (<Emphasis Type="Italic">Inachis io</Emphasis>) against mice (<Emphasis Type="Italic">Apodemus flavicollis</Emphasis> and <Emphasis Type="Italic">A. sylvaticus</Emphasis>)

Morphological and behavioural traits can serve as anti-predator defence either by reducing detection or recognition risks, or by thwarting initiated attacks. The latter defence is secondary and often involves a ‘startle display’ comprising a sudden release of signals targeting more than one sensory modality. A suggested candidate for employing a multimodal defence is the peacock butterfly, Inachis io, which, by wing-flicking suddenly, produces sonic and ultrasonic sounds and displays four large eyespots when attacked. The eyespots make small birds retreat, but whether the sounds produced thwart predator attacks is largely unknown. Peacocks hibernate as adults in dark wintering sites and employ their secondary defence upon encounter with small rodent predators during this period. In this study, we staged predator–prey encounters in complete darkness in the laboratory between wild mice, Apodemus flavicollis and Apodemus sylvaticus, and peacocks which had their sound production intact or disabled. Results show that mice were more likely to flee from sound-producing butterflies than from butterflies which had their sound production disabled. Our study presents experimental evidence that the peacock butterfly truly employs a multimodal defence with different traits targeting different predator groups; the eyespots target birds and the sound production targets small rodent predators.