Calanoid copepod escape behavior in response to a visual predator

Calanoid copepods typically exhibit escape reactions to hydrodynamic stimuli such as those generated by the approach of a predator. During the summers of 2000, 2001 and 2004, two small calanoid species, Temora turbinata Dana, 1849 and Paracalanus parvus Claus, 1863 were exposed to a visual predatory fish, the blenny Acanthemblemaria spinosa Metzelaar, 1919, and their predator–prey interactions were recorded using both high-speed and standard videographic techniques. Copepod escape reaction components, including swimming pattern, reactive distance, turning rate, and jump kinetics, were quantified from individual predation events using motion analysis techniques. Among the observed escape reaction components, differences were noted between the species’ swimming patterns prior to attack and their response latencies. Temora turbinata was a continuous cruiser and P. parvus exhibited a hop-and-sink swimming pattern. During periods of sinking, P. parvus stopped beating its appendages, which presumably reduced any self-generated hydrodynamic signals and increased perceptual abilities to detect an approaching predator. Response latency was determined for each copepod species using a hydrodynamic stimulus produced by a 1 ms acoustic signal. Response latencies of T. turbinata were significantly longer than those of P. parvus. Despite some apparent perceptual advantages of P. parvus, the blenny successfully captured both species by modifying its attack behavior for the targeted prey.     

Photosynthate partitioning by phytoplankton in a New Zealand coastal upwelling system

A stimulation model of copepod population dynamics (development rate, fecundity, and mortality) was used to compute the predatory consumption necessary to control population growth in three dominant copepod species (Pseudocalanus sp., Paracalanus parvus, and Calanus finmarchicus) on Georges Bank, given observed seasonal cycles of copepod and predator populations. The model also calculated secondary production of each species. Copepod development rate and fecundity were functions of temperature while mortality was a function of predator abundance and consumption rate. Daily inputs of temperature and predator abundance (chaetognaths, ctenophores, and Centropages spp.) were derived from equations fit to field data. Model runs were made with various consumption rates until the model output matched observed copepod seasonal cycles. Computed consumption rates were low compared with published values from field and laboratory studies indicating that, even at conservative estimates of consumption, predators are able to control these copepod populations. Combined annual secondary production by the small copepod species, Pseudocalanus sp. and P. parvus, was nearly twice that of the larger C. finmarchicus with P. parvus having the highest total annual production.     

Fine-scale observations of the predatory behaviour of the carnivorous copepod <Emphasis Type="Italic">Paraeuchaeta norvegica</Emphasis> and the escape responses of their ichthyoplankton prey,Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>)

Paraeuchaeta norvegica (8.5 mm total length) and yolk-sac stage Atlantic cod larvae (4 mm total length) (Gadus morhua) larvae were observed in aquaria (3 l of water) using silhouette video photography. This allowed direct observations (and quantitative measurement) of predator–prey interactions between these two species in 3-dimensions. Tail beats, used by cod larvae to propel themselves through the viscous fluid environment, also generate signals detectable by mechanoreceptive copepod predators. When the prey is close enough for detection and successful capture (approximately half a body-length), the copepod launches an extremely rapid high Reynolds number attack, grabbing the larva around its midsection. While capture itself takes place in milliseconds, minutes are required to subdue and completely ingest a cod larva. The behavioural observations are used to estimate the hydrodynamic signal strength of the cod larva’s tail beats and the copepod’s perceptive field for larval fish prey. Cod larvae are more sensitive to fluid velocity than P. norvegica and also appear capable of distinguishing between the signal generated by a swimming and an attacking copepod. However, the copepod can lunge at much faster velocities than a yolk-sac cod larva can escape, leading to the larva’s capture. These observations can serve as input to the predator–prey component of ecosystem models intended to assess the impact of P. norvegica on cod larvae.     

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

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

Strain-related physiological and behavioral effects of <Emphasis Type="Italic">Skeletonema marinoi</Emphasis> on three common planktonic copepods

Three strains of the chain-forming diatom Skeletonema marinoi, differing in their production of polyunsaturated aldehydes (PUA) and nutritional food components, were used in experiments on feeding, egg production, hatching success, pellet production, and behavior of three common planktonic copepods: Acartia tonsa, Pseudocalanus elongatus, and Temora longicornis. The three different diatom strains (9B, 1G, and 7J) induced widely different effects on Acartia tonsa physiology, and the 9B strain induced different effects for the three copepods. In contrast, different strains induced no or small alterations in the distribution, swimming behavior, and turning frequency of the copepods. 22:6(n-3) fatty acid (DHA) and sterol content of the diet typically showed a positive effect on either egg production (A. tonsa) or hatching success (P. elongatus), while other measured compounds (PUA, other long-chain polyunsaturated fatty acids) of the algae had no obvious effects. Our results demonstrate that differences between strains of a given diatom species can generate effects on copepod physiology, which are as large as those induced by different algae species or groups. This emphasizes the need to identify the specific characteristics of local diatoms together with the interacting effects of different mineral, biochemical, and toxic compounds and their potential implications on different copepod species.     

Propulsion efficiency and cost of transport for copepods: a hydromechanical model of crustacean swimming

In the absence of direct measurement, costs of locomotion to small swimming Crustacea (<10 mm) have been derived exclusively through application of the fluid dynamic theory. Results indicate very low swimming costs, and contradict experimental data on larger Crustacea (15 to 100 mm) that suggest a three-fold increase in metabolic rate with increasing swimming speed. This paper introduces a swimming model that analyzes the hydrodynamic forces acting on a crustacean swimming at non-steady velocity. The model treats separately the hydrodynamic forces acting on the body and the swimming appendages, approximating the simultaneous solution of equations quantifying the drag and added-mass forces on each by stepwise integration. Input to the model is a time-series of instantaneous swimming-appendage velocities. The model output predicts a corresponding time-series of body velocities as well as the mechanical energy required to move the swimming appendages, dissipated kinetic energy, and metabolic cost of swimming. Swimming of the calanoid copepod Pleuromamma xiphias (Calanoida) was analyzed by extrapolating model parameters from data available in the literature. The model predictions agree well with empirical observations reported for larger crustaceans, in that swimming for copepods is relatively costly. The ratio of active to standard metabolism for P. xiphias was >3. Net cost of transport was intermediate to the values found experimentally for fish and larger crustaceans. This was a consequence of the predicted mechanical efficiency (34%) of the copepod's paddle propulsion, and of increased parasitic resistance resulting from non-steady velocity swimming.     

Demographic and temporal structure of an allele frequency cline in the mussel Mytilus edulis

Feeding rates, patterns of prey selection, and starvation tolerance were investigated for adult males and females of the cyclopoid copepod Corycaeus anglicus collected from the waters of Friday Harbor, Washington, USA. Selection by C. anglicus was determined largely by prey body-size, but was also affected by species and developmental stage. Small developmental stages of all prey species were fed upon at relatively low rates. The small calanoid species Acartia clausii was increasingly vulnerable to predation by C. anglicus as it progressed through successive developmental stages. Larger prey species, Pseudocalanus sp. and Calanus pacificus, were more vulnerable in intermediate stages, the C3 and N6 stages, respectively. Larger and smaller prey were characteristically attacked at different sites on their bodies; however, attack sites fell within a similar range of body widths, 130 to 170 m. Males of Corycaeus anglicus killed a maximum of 1.4 prey d^-1 when feeding on the optimally-sized adult females of Acartia clausii, which are approximately equivalent to its own body length. Males fed at approximately double the rates of females. Despite its small size and apparent lack of metabolic stores, this cyclopoid is highly tolerant of starvation conditions. Median survival time without food is at least 2 wk for both males and females. In its predatory behavior, C. anglicus employs an ambush-type strategy and seems to be adapted for infrequent encounters with relatively large prey.Contribution No. 1412 from the School of Oceanography, University of Washington, Seattle     

Vertical distribution and mortality of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during overwintering in oceanic waters southwest of Iceland

The seasonal vertical distribution and the predatory regime encountered by Calanus finmarchicus were studied along a transect across the Reykjanes Ridge in the oceanic area southwest of Iceland from data collected during four cruises between November 1996 and June 1997. The mortality for the overwintering period was estimated using linear regressions of density estimates from November 1996 to April 1997. In addition, we also estimated the mortality of the oldest population stages (C4 and older) in April and June by applying the vertical table method. During winter (November/December–January/February), the animals mainly resided at a depth of ∼300–1,500 m in the water of Atlantic origin. Ascent to upper layers took place mainly during March and April, and continued until May. During all cruises, continuous deep-scattering layers were observed, mainly within the range of 400–500 m to 700–800 m depth. Based on sampling with a Harstad pelagic trawl in April, the scattering was mainly ascribed to jellyfish (mainly Periphylla periphylla), small mesopelagic oceanic fishes (several species but Benthosema glaciale and Maurolicus muelleri were most abundant), euphausiids (mainly Meganyctiphanes norvegica) and shrimps (mainly Sergestes arcticus). These species may represent a predatory threat to overwintering C. finmarchicus. From November to April, daily per capita mortality rates were estimated to be (mean ± 95% CL) 0.004 ± 0.0028 for the total data set, and 0.004 ± 0.0033 and 0.004 ± 0.0023 (day^-1) for the Iceland Basin and Irminger Basin, respectively. Mortality rates were higher later in life (mean ± 95% CL) for C5/females (0.13 ± 0.044) and C5/males (0.19 ± 0.051) than for C4/C5 (0.00 ± 0.035) when averaged over all samples taken in April and June 1997. We discuss how the observed distribution and mortality rates of overwintering C. finmarchicus might be related to predatory regime.     

Foraging behavior in early Atlantic cod larvae (Gadus morhua) feeding on a protozoan (Balanionsp.) and a copepod nauplius (Pseudodiaptomussp.)

Food limitation is likely to be a source of mortality for fish larvae in the first few weeks after hatching. In the laboratory, we analyzed all aspects of foraging in cod larvae (Gadus morhua Linnaeus) from 5 to 20 d post-hatching using protozoa (Balanion sp.) and copepod nauplii (Pseudodiaptomus sp.) as prey. A camera acquisition system with two orthogonal cameras and a digital image analysis program was used to observe patterns of foraging. Digitization provided three-dimensional speeds, distances, and angles for each foraging event, and determined prey and fish larval head and tail positions. Larval cod swimming speeds, perception distances, angles, and volumes increased with larval fish size. Larval cod swam in a series of short intense bursts interspersed with slower gliding sequences. In 94% of all foraging events prey items were perceived during glides. Larval cod foraging has three possible outcomes: unsuccessful attacks, aborted attacks, and successful attacks. The percentage of successful attacks increased with fish size. In all larval fish size classes, successful attacks had smaller attack distances and faster attack speeds than unsuccessful attacks. Among prey items slowly swimming protozoans were the preferred food of first-feeding cod larvae; larger larvae had higher swimming speeds and captured larger, faster copepod nauplii. Protozoans may be an important prey item for first-feeding larvae providing essential resources for growth to a size at which copepod nauplii are captured. Received: 20 April 1999 / Accepted: 12 January 2000     

Impact of copepod grazing on the red-tide flagellate Chattonella antiqua

Although planktonic copepods are major suspension feeders in the sea, the impact of their grazing pressure upon red-tide flagellates has not been fully investigated. In the present study, the grazing of adult females of several copepod species is examined using three food types: viz. natural suspended particles, natural suspended particles mixed with cultured Chattonella antiqua, and cultured C. antiqua. The functional response on C. antiqua was investigated for five species of copepods (Acartia erythraea, Calanus sinicus, Centropages yamadai, Paracalanus parvus and Pseudodiaptomus marinus). Ingestion rates increased linearly with increasing cell concentrations until a maximum level was reached, beyond which the rates were constant. This cell concentration was higher for larger copepods. The weight-specific maximum ingestion rates were higher in the small species. In general, copepods tended to feed selectively on larger particles when feeding on natural particles. This tendency was strongest in a simulated red-tide environment. Thus, it can be surmised that copepods may selectively graze on C. antiqua during the outbreak of a red tide. Grazing pressure by the natural copepod community in Harima Nada, the Inland Sea of Japan, was calculated by integration of the laboratory determined feeding rates and field measurements of zooplankton biomass. The daily removal rate was 3.4 to 30.8% (mean: 12.3%) of C. antiqua biomass at 20 cells ml^-1 and decreased to 0.6–4.3% (mean: 1.8%) at 500 cells ml^-1. Therefore, the grazing pressure by the copepod community is important at the initial stage of the red tide.     

Determination of the swimming trajectory and speed of chain-forming dinoflagellate <Emphasis Type="Italic">Cochlodinium polykrikoides</Emphasis> with digital holographic particle tracking velocimetry

The marine dinoflagellate Cochlodinium polykrikoides is a harmful and highly motile algal species. To distinguish between the motility characteristics of solitary and chain-forming cells, the swimming trajectories and speeds of solitary cells and 2- to 8-cell chains of C. polykrikoides were measured using a digital holographic particle tracking velocimetry (PTV) technique. C. polykrikoides cells exhibited helical swimming trajectories similar to other dinoflagellate species. The swimming speed increased as the number of cells in the chain increased, from an average of 391 μm s⁻¹ (solitary cells) to 856 μm s⁻¹ (8-cell chain). The helix radius R and pitch P also increased as the number of cells in the chain increased. R increased from 9.24 μm (solitary cell) to 20.3 μm (8-cell chain) and P increased from 107 μm (solitary cell) to 164 μm (8-cell chain). The free thrust-generating motion of the transverse flagella and large drag reduction in the chain-forming cells seemed to increase the swimming speed compared to solitary cells. The measured swimming speeds agreed with those from field observations. The superior motility of chain-forming C. polykrikoides cells may be an important factor for its bloom, in addition to the factors reported previously.     

Population structure of the planktonic copepod <Emphasis Type="Italic">Calanus pacificus</Emphasis> in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Embryogenesis and larval biology of the ahermatypic scleractinian<Emphasis Type="Italic"> Oculina varicosa</Emphasis>

The ivory tree coral Oculina varicosa (Leseur, 1820) is an ahermatypic branching scleractinian that colonizes limestone ledges at depths of 6–100 m along the Atlantic coast of Florida. This paper describes the development of embryos and larvae from shallow-water O. varicosa, collected at 6–8 m depth in July 1999 off Fort Pierce, Florida (27°32.542 N; 79°58.732 W). The effect of temperature on embryogenesis, larval survival, and larval swimming speed were examined in the laboratory. Ontogenetic changes in geotaxis and phototaxis were also investigated. Embryos developed via spiral cleavage from small (100 µm), negatively buoyant eggs. Ciliated larvae developed after 6–9 h at 25°C. Embryogenesis ceased at 10°C, was inhibited at 17°C, and progressed normally at 25°C and 30°C. Larval survival, however, was high across the full range of experimental temperatures (11–31°C), although mortality increased in the warmest treatments (26°C and 31°C). Larval swimming speed was highest at 25°C, and lower at the temperature extremes (5°C and 35°C). An ontogenetic change in geotaxis was observed; newly ciliated larvae swam to the water surface and remained there for approximately 18 h, after which they swam briefly throughout the water column, then became demersal. Early larvae showed no response to light stimulation, but at 14 and 23 days larvae appeared to exhibit negatively phototactic behavior. Although low temperatures inhibited the development of O. varicosa embryos, the larvae survived temperature extremes for extended periods of time. Ontogenetic changes in larval behavior may ensure that competent larvae are close to the benthos to facilitate settlement. Previous experiments on survival, swimming speeds, and observations on behavior of O. varicosa larvae from deep-water adults indicate that there is no difference between larvae of the deep and shallow populations.Communicated by J.P. Grassle, New Brunswick     

Mating behaviour of <Emphasis Type="Italic">Pseudodiaptomus annandalei</Emphasis> (Copepoda,Calanoida) at calm and hydrodynamically disturbed waters

Behavioural observations of male copepods revealed that they commonly follow female footprints to find their mates. Copepods can perceive signals generated by females either hydromechanically or chemically. Signal intensity is affected by hydrodynamic conditions which clear chemical and mechanical cues and modulate copepod’s ability to sense signals of their biotic environment, such as in their search for mates. We studied the patterns and efficiency of the copepod Pseudodiaptomus annandalei to mate in calm and hydrodynamically disturbed waters, in illuminated and dark conditions in experimental containers of different shapes and volumes. Courtship in P. annandalei was a negative function of hydromechanical disturbance, since successful mating events were observed in calm water only. In weakly turbulent conditions (air-bubbling of 100 ml/min), males were not able to pursue females properly; swimming speed decreased about three times in comparison with that in calm water. In calm water conditions, sequential and simultaneous taxis mechanisms were used by P. annandalei males to pursue females. The ability of P. annandalei males to track a three-dimensional trail probably depended on the persistence of fluid-borne signals.     

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     

Mechanics of prey selection by ephyrae of the scyphomedusa Aurelia aurita

In situ feeding patterns of ephyrae of the jellyfish Aurelia aurita (Linnaeus) revealed the importance of relatively large (>1 mm) prey in the diet of these scyphozoan predators. These studies were carried out in Narragansett Bay, Rhode Island, USA in March and April, from 1993 through 1996. Rotifers were the only small prey ingested in quantity, and then only when they were unusually abundant in the plankton. Copepod nauplii, similar in size to rotifers and equally abundant, were rarely consumed. Since copepods evince rapid escape responses, this observation suggested a role for prey escape in determining prey vulnerability, while the predominance of large prey in the diet suggested a role for prey size. Using two dimensional video observations of free-swimming ephyrae and their prey in the laboratory we tested hypotheses about the mechanisms underlying these dietary patterns, comparing mechanisms for capture of large versus small prey and for prey of equal size but differing escape behaviors. Capture efficiencies of ephyrae feeding on large prey were 4 to 12 times greater than for small prey taxa. Capture efficiencies for prey of equal size also differed significantly, indicating that other factors influence the outcome of predator–prey interactions. Most prey captures occurred while the ephyrae were swimming and creating fluid flows that entrained prey into the subumbrellar region. Even copepod nauplii were frequently drawn into the subumbrella of swimming ephyrae despite average potential escape velocities (25.7 mm s⁻¹) that exceeded mean maximum velocity of fluid flows around the ephyrae (13.1 mm s⁻¹). Large prey were more likely than small prey to contact nematocyst-bearing surfaces both before and after entrainment in flow fields. With regard to behavior, prey escape speeds were not the only predictor of prey vulnerability. Prey that continued swimming after entrainment (rotifers and brine shrimp) were captured more often than prey of equal size that ceased normal swimming (copepod nauplii and barnacle nauplii). Copepod nauplii were the prey least likely to be captured because they either “played dead” and were expelled from the subumbrella of the ephyrae before contacting a surface, or they eventually escaped at high velocity. These observations indicate that size-selective predation by ephyrae of A. aurita can be influenced by a variety of behavioral responses of the prey. Received: 9 April 1997 / Accepted: 5 September 1997     

Swimming ability of eels (<Emphasis Type="Italic">Anguilla rostrata,Conger oceanicus</Emphasis>) at estuarine ingress: contrasting patterns of cross-shelf transport?

The transport of eel early life stages may be critical to their population dynamics. This transport from ocean spawning to freshwater, estuarine and coastal nursery areas is a combination of physical and biological processes (including swimming behavior). In New Jersey, USA, the American eel (Anguilla rostrata) enters estuaries as glass eels (48.7–68.1 mm TL) in contrast to the Conger eel (Conger oceanicus) that enters as larger (metamorphosing) leptocephali (68.3–117.8 mm TL). To begin to understand the mechanisms of cross-shelf transport for these species, we measured the potential swimming capability (critical swimming speed, U _crit) under ambient conditions throughout the ingress season. A. rostrata glass eels were collected over many months (January–June) at a range of temperatures (4–21°C), with relative condition declining over the course of the ingress period as temperatures warmed. C. oceanicus occurred later in the season (April–June) and at warmer temperatures (14–24.5°C). Mean U _crit values for A. rostrata (11.7–13.3 cm s⁻¹) and C. oceanicus (14.7–18.6 cm s⁻¹) were comparable, but variable, with portions of the variability explained by water temperature, relative condition, ontogenetic stage, and fish length. Travel times to Little Egg Inlet, New Jersey, estimated using 50% U _crit values, indicate it would take A. rostrata ~30 and ~60 days to swim from the shelf edge and Gulf Stream, respectively. Travel times for C. oceanicus were shorter, ~20 days from the shelf edge, and ~45 days from the Gulf Stream. Despite differences in life stage, our results indicate both species are competent swimmers, and suggest they are capable of swimming from the Gulf Stream and/or edge of the continental shelf to estuarine inlets.     

Phylogeography of the calanoid copepods <Emphasis Type="Italic">Calanus helgolandicus</Emphasis> and <Emphasis Type="Italic">C. euxinus</Emphasis> suggests Pleistocene divergences between Atlantic,Mediterranean, and Black Sea populations

Molecular systematic analyses of marine taxa are crucial for recording ocean biodiversity, so too are elucidation of the history of population divergence and the dynamics of speciation. In this paper we present the joined phylogeography of the calanoid copepod Calanus helgolandicus (Claus 1863) from the North East (NE) Atlantic and the Adriatic Sea and the closely related C. euxinus (Hulsemann 1991) from the Black Sea based on sequences of a mitochondrial Cytochrome Oxidase subunit I (COI) fragment. Coalescent-based Bayesian methods and minimum spanning networks are used to reconstruct the history of population divergence. Our results reveal that copepod populations from all three basins share a great number of haplotypes and demonstrate a close genetic affinity of C. euxinus with C. helgolandicus. The data do not support significant genetic structuring among samples within seas. Coalescent analyses suggest divergences between NE Atlantic, Mediterranean, and Black Sea populations dating back to the middle Pleistocene, with the NE Atlantic–Mediterranean divergence being the earliest and the Mediterranean–Black Sea divergence the most recent. These middle Pleistocene dates are much older than the estimated dates of colonisation of the Mediterranean and Black Seas based on paleoclimatic scenarios. Our results do not rule out that the assumed colonisations took place but they indicate that the populations colonising the Mediterranean and the Black Sea were already, and have since remained, diverged. The chaetognath Sagitta setosa, which has a comparable distribution pattern and feeds upon the copepods, provides a unique opportunity to compare phylogeographic patterns and distinguish among alternative hypotheses. The dates produced in this paper are in agreement with those estimated elsewhere for S. setosa. We propose that a great deal of the genetic make-up of marine planktonic populations comprises divergences that date back to long before the last glacial maximum. We consider questions on the taxonomic status of C. euxinus to remain open. However, its high genetic affinity to the C. helgolandicus calls for further investigation.     

Effect of the presence of the shore crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, on burrowing behaviour and clearance rate of the common cockle, <Emphasis Type="Italic">Cerastoderma edule</Emphasis>

Bivalves demonstrate various morphological and behavioural adaptations to reduce the risk of being attacked by predators. This paper examines how the presence of the crab Carcinus maenas (L.), a natural predator of the cockle Cerastoderma edule (L.), affects its burrowing depth and clearance or feeding rate. Cockles were placed in experimental tanks and treated with three levels of predatory disturbance: (1) unfed crab loose inside the tank, (2) unfed crab inside a cage suspended in the water column and (3) no crab present. Cockles’ burrowing depth was measured in two sediment types: mud and sand. Cockles burrowed more deeply in treatments with no crabs. Burrowing depth in sand was significantly greater than in mud. Two factors may contribute to the reduction in burial depth of C. edule in the presence of C. maenas: the change in the vertical orientation of the cockle and the ‘cough response’. No significant difference was found in the cockles’ clearance rate among the different levels of predator threat.