A commensal relationship between the scyphozoan medusae<Emphasis Type="Italic"> Catostylus mosaicus</Emphasis> and the copepod<Emphasis Type="Italic"> Paramacrochiron maximum</Emphasis>

The copepod Paramacrochiron maximum was found in high numbers (up to 5,675 copepods/medusa) on the oral arms of the scyphozoan Catostylus mosaicus. This association was considered to be commensalism for the following reasons: P. maximum (Lichomolgidae) was abundant on the medusae (approximately 805 copepods/kg of medusae) and very rare in the water column (approximately 5.99×10^-4 copepods/kg of water); copepodites and adults of the symbiont were present on the host; the copepods were on the medusae both day and night, at different times (nine occasions between March 1999 and May 2000) and different locations (Botany Bay and Lake Illawarra, NSW, Australia). Over 40 taxa of plankton were found on the oral arms of C. mosaicus (including protists, cnidarians, polychaetes, molluscs, a wide range of holoplanktonic and meroplanktonic crustaceans, chaetognaths and fish eggs). These taxa were abundant in the water column and we concluded that they were prey. Symbiotic amphipods and carangid fishes were found with medusae. We conclude that there is a symbiotic association between P. maximum and C. mosaicus and care should be taken not to confound these copepods with the prey of C. mosaicus. Poecilostomid copepods are well known for consuming mucus and feeding is likely to be a major reason for the association.Communicated by G.F. Humphrey, Sydney     

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     

Purification and sequencing of a mate-recognition protein from the copepod<Emphasis Type="Italic"> Tigriopus japonicus</Emphasis>

Male copepods in the genus Tigriopus use surface proteins of female copepods to recognize appropriate mates. In a previous study of pre-copulatory mate-guarding in T. japonicus Mori, a monoclonal antibody was developed against one protein that contributed to the attractiveness of female T. japonicus as mate-guarding partners. In the present study, which was performed between December 1998 and November 2000, the antibody was used to purify two proteins from homogenates of pooled T. japonicus. A 70 kD protein was partially sequenced using mass spectrometry, revealing sequence similarity to α₂-macroglobulin. It was shown that the 70 kD protein was highly expressed in stage CV females, the most attractive stage for mate-guarding. A purified 36 kD protein was also shown to be a cognate of α₂-macroglobulin. Because the structure of both the 70 and 36 kD proteins suggested that protease activity and its inhibition might play a role in male mate choice, three protease inhibitors were tested in bioassays of male discrimination of conspecifics. The male's ability to discriminate among females was not completely blocked by the protease inhibitor, but their behavior was affected in more subtle ways. This suggests that the 70 kD protein, designated CSP70, is a variant of α₂-macroglobulin, but with more specific activity.     

Rapid response to changing light environments of the calanoid copepod<Emphasis Type="Italic"> Calanus sinicus</Emphasis>

Calanus sinicus is a large calanoid copepod and a dominant species in the coastal waters of Japan. During a research cruise in Sagami Bay on 18 June 1996, we found C. sinicus performing an unusual diel vertical migration (DVM), a behavior that has not been reported in previous studies on this species. This study examined the DVM of C. sinicus under different light environments and revealed the copepods characteristic response to light. Field and laboratory results show that the DVM of C. sinicus is flexible and also confirmed its sensitivity and its rapid response to changing light environments. It is suggested that C. sinicus reacts to changes in absolute light intensity. This feature may be common in oceanic copepod species. The copepods quick reaction to light variation provides decreased predation risks and increased feeding opportunities, which make them a dominant survivor in coastal water habitats.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.     

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.     

Multiple feeding stimulants in <Emphasis Type="Italic">Sinapis alba</Emphasis> for the oligophagous leaf beetle <Emphasis Type="Italic">Phaedon cochleariae</Emphasis>

Summary. In earlier investigations on host plant discrimination of leaf beetles glucosinolates were described as feeding stimulants for the Brassicaceae specialist Phaedon cochleariae F. (Coleoptera: Chrysomelidae). However, since these findings could not be confirmed in later studies offering 2-propenylglucosinolate in concentrations corresponding to those detected in host plant leaf material, the identification of feeding stimulants of this leaf beetle species remained unclear. In order to investigate which compounds of the host plant Sinapis alba (Brassicaceae) are involved in feeding stimulation, leaf extracts of different polarities were tested in bioassays with adults of P. cochleariae. Number of feeding beetles and net consumption rates were highest on pea leaves painted with methanol extracts of S. alba, whereas weak feeding responses were also detectable for hexane extracts. In subsequent bioassay-guided fractionations of methanol extracts with semi-preparative high performance liquid chromatography, two distinct fractions, one containing glucosinolates and another containing flavonoids, were found to stimulate beetles to feed to variable degrees. Other collected fractions had zero activity. The combination of both active fractions evoked significantly higher consumption rates and stimulated more beetles to feed than fractions tested individually. At least one compound of each fraction, among these the main glucosinolate of S. alba, 4-hydroxybenzylglucosinolate, act additively. Effects of two different naturally-occurring ratios of glucosinolates and flavonoids on the strength of feeding responses were investigated by use of extracts of two sets of host plants differently exposed to radiation. One set was outdoors-exposed, whereas the second set was kept in the greenhouse. However, the feeding behaviour of P. cochleariae was not affected by the significantly different relative compositions of both compound classes in the host material. In conclusion, mustard leaf beetles need a combination of distinct plant metabolites acting in concert for feeding stimulation, whereby the mere presence of these stimulants, but probably not the ratio of involved compounds, determines their feeding response.     

The effect of feeding behavior of the gastropods<Emphasis Type="Italic"> Batillaria zonalis</Emphasis> and<Emphasis Type="Italic"> Cerithideopsilla cingulata</Emphasis> on their ambient environment

Feeding behaviors of the gastropods Batillaria zonalis, a suspension and deposit feeder, and Cerithideopsilla cingulata, an obligate deposit feeder, were studied to examine their effect on dynamics of suspended materials, total nitrogen (TN) and total organic carbon (TOC) in sediments. Suspension feeding in B. zonalis was observed in detail visually, as it had been previously unreported. An experimental system where B. zonalis and C. cingulata were cultured for 10 weeks, using previously frozen microalgae Nannochloropsis oculata as food, was then constructed. During feeding observations, the suspension-feeding B. zonalis formed a mucus food cord to entangle particulate materials, which were subsequently ingested. The feeding mode of B. zonalis is hence categorized as ctenidial filter feeding. For the culture experiments, decreases in suspended materials were seen only in the B. zonalis cultures, while the control (no gastropods) and C. cingulata cultures remained nearly unchanged. Sediment TN and TOC showed no significant differences between B. zonalis (with mean TN at 0.0345% and mean TOC at 0.261%) and control cultures (with TN at 0.0389% and TOC at 0.331%), but the sediments in C. cingulata cultures had lower levels (with TN at 0.0204% and TOC at 0.156%). The C/N ratios were similar for both B. zonalis (7.55) and C. cingulata (7.68) cultures, and both were lower than the control cultures (8.55). The filtration rate for B. zonalis was lower than that previously observed in bivalves inhabiting the same intertidal flat (e.g. Cyclina sinensis, Grafrarium tumidum and Barbatia virescens). However, Batillaria zonalis occurs at higher abundances than these bivalves. Therefore, it is expected that this species has a large affect upon the transport of suspended materials to the sediments. The addition of TN and TOC to sediments in B. zonalis cultures was probably caused by biodeposition, but deposit feeding by B. zonalis may have restrained the accumulation of those components. The impact of deposit feeding in Cerithideopsilla cingulata cultures was most probably stronger than sedimentation and biodeposition, because of the lower sediment TN and TOC. Bioturbation by both B. zonalis and C. cingulata yields the same effect on sediment quality, as indicated by the low C/N in the culture sediment of both treatments, despite difference in feeding modes. This paper demonstrates, for the first time, the importance of gastropods in bioturbation and removal of suspended materials in subtropical tidal flat habitats.Communicated by T. Ikeda, Hakodate     

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     

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     

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.     

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.     

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     

Morphology and growth of the deep-water gorgonians <Emphasis Type="Italic">Primnoa resedaeformis</Emphasis> and <Emphasis Type="Italic">Paragorgia arborea</Emphasis>

The morphology of the gorgonian corals Paragorgia arborea and Primnoa resedaeformis was studied from video records and colonies collected from different locations in Atlantic Canada, at depths between 200 and 600 m. Growth was studied by relating colony height to age (number of growth rings) in P. resedaeformis, and from a photographic time-series of a P. arborea colony in a Norwegian fjord. The highest P. resedaeformis and P. arborea colonies were 86 and 180 cm, respectively. The height of P. arborea seemed to be restricted by the size of the boulder it was attached to. When the coral exceeds a critical height (approximately twice the stone size), the drag of strong currents can turn the coral and its substrate over. No limiting factors for the height of P. resedaeformis colonies were identified. P. arborea occurred in three colour varieties: red, salmon red, and white. The red and white contributed 41% to the population each, while 18% of the colonies were salmon red. On average the salmon red P. arborea were taller than the red and white. P. arborea colonies >50 cm were mainly concave fan shaped. The orientation of these indicated a near-bottom current pattern similar to what is known from previous current measurements in the area. P. resedaeformis occurred mainly on the up-current side of boulders, but its bushy morphology does not indicate influence by unidirectional current to the same degree as P. arborea. The different height, morphology, and position on boulders of the two species indicate that they utilize different food sources. P. resedaeformis seems to be adapted to a near-bottom environment with turbulent currents, whereas P. arborea utilize uni- or bidirectional currents higher above bottom by developing planar colonies perpendicular to the current. The oldest P. resedaeformis colony was 61 years. The relationship between height and age indicated an average growth of 1.7 cm year^–1 for P. resedaeformis. X-ray images of skeletal sections of P. arborea showed clear growth bands with a maximum band width of 1.3 cm. It is not clear what time scales these bands represent, and they could therefore not be used for indicating age. The limited previously reported data on age and growth of P. arborea indicate an average growth rate of 1 cm year^–1. This gives an age of about 180 years for the largest colony in this study. The time-series photographs, however, indicated a much higher growth rate (varying between 2 and 6 cm year^–1 within the colony), which may be more representative for colonies of an intermediate size.Communicated by R.J. Thompson, St. Johns     

Effects of dietary fatty acids on the reproductive success of the calanoid copepod<Emphasis Type="Italic"> Temora longicornis</Emphasis>

Egg production and hatching success of the copepod Temora longicornis were measured in laboratory experiments and in the field (North Sea). In the laboratory, ingestion of four algal species (Thalassiosira weissflogii, Phaeocystis globosa, Isochrysis sp. and Dunaliella tertiolecta) was followed and the content of fatty acids in the algae was determined. The two food types (T. weissflogii and Isochrysis sp.) that provided the highest ingestion of carbon and long chain fatty acids also resulted in the highest egg production rate (E_r) and hatching success (H%). In contrast, D. tertiolecta led to both low ingestion of carbon and long chain fatty acids, resulting in low reproductive success. There was a positive relationship between the amount of eicosapentaenoic fatty acid [20:5(n-3), EPA] ingested and E_r and H%, and of the ratio between docosahexaenoic and eicosapentaenoic fatty acid [22:6(n-3)/20:5(n-3), DHA/EPA] in the ingested food and H%. In the field, chlorophyll a and specific fatty acids were measured and protists were enumerated, in order to investigate the link between these factors and the reproductive success of T. longicornis. Hatching time was found to be related to temperature and exceeded 120 h at 6°C. No relationship was found between chlorophyll a and reproductive success (E_r or H%). E_r correlated with the concentration of diatoms and ciliates, which were the dominating protists in early spring, indicating that food quantity was the limiting factor for E_r. As in the laboratory experiments, H% was dependent on the fatty acid DHA and the ratio of DHA/EPA, which indicates that the quality of eggs (H%) is linked to the quality of food.Communicated by M. Kühl, Helsingør     

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.     

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