Inverse vertical migration and feeding in glacier lanternfish (<Emphasis Type="Italic">Benthosema glaciale</Emphasis>)

A bottom-mounted upward-facing 38-kHz echo sounder was deployed at ~400 m and cabled to shore in Masfjorden (~60^°52′N, ~5^°24′E), Norway. The scattering layers seen during autumn (September–October) 2008 were identified by trawling. Glacier lanternfish (Benthosema glaciale) were mainly distributed below ~200 m and displayed three different diel behavioral strategies: normal diel vertical migration (NDVM), inverse DVM (IDVM) and no DVM (NoDVM). The IDVM group was the focus of this study. It consisted of 2-year and older individuals migrating to ~200–270 m during the daytime, while descending back to deeper than ~270 m during the night. Stomach content analysis revealed increased feeding during the daytime on overwintering Calanus sp. We conclude that visually searching glacier lanternfish performing IDVM benefit from the faint daytime light in mid-waters when preying on overwintering Calanus sp.     

Vertical distribution and nocturnal migration of zooplankton in relation to the development of the seasonal thermocline in Patraikos Gulf

Vertical distribution and nocturnal migration of zooplankton species in relation to the development of the seasonal thermocline in the shallow waters (90 m) of Patraikos Gulf (Ionian Sea, Greece) were investigated using a WP-2 closing net. Juvenile and adult copepods accounted for a mean of 91% of the total collected in three sampling periods, i.e. May, July and September 1985.Ctenocalananus vanus, Paracalanus parvus andOithona plumifera were the dominant copepods. The majority of the zooplankton tend to aggregate at the thermocline layer. Among copepods the two congeneric speciesClausocalanus pergens andC. furcatus exhibited different migratory responses to the development of the thermocline.C. pergens occurred in the lower part of the thermocline andC. furcatus in the upper region or above. The diel vertical migration of all species could be divided into four types: (1) no vertical migration; (2) upward migration at night; (3) occasional migration; and (4) reverse migration (down at night). In July when the strongest thermocline developed, most zooplankters rose close to the surface at night. For most species, temperature discontinuity did not limit their diel migration.Please address all correspondence and requests for reprints to Dr J.J. Lykakis     

Movements and environmental preferences of the shortfin mako, <Emphasis Type="Italic">Isurus oxyrinchus</Emphasis>, in the southeastern Pacific Ocean

Nine individuals of shortfin mako, Isurus oxyrinchus, were tracked in the southeastern Pacific Ocean, off northern Chile, by means of pop-up satellite archival tags. No common pattern was observed in their trajectories, apart from a movement onshore of all the fish tracked during June–August. The average estimated rate of movement was of c. 27 km day⁻¹. Data were collected and processed for a total of 341 days, including 33 days for one recaptured fish specimen, allowing high-resolution archived data to be downloaded. The fish spent most of their time in the mixed layer but undertook dives down to 888 m. Ambient temperatures ranged between 4.6 and 24.1°C, and the sea surface temperatures recorded ranged from 13.4 to 24.1°C during the study period. No clear diel pattern in depth behavior was observed, but mean vertical distribution was deeper during the daytime. Moreover, a foraging pattern, consisting of rapid descents below the thermocline followed by slower ascents, was generally observed during daylight hours. Dissolved oxygen concentration and water temperature seem to be the main factors affecting the vertical range of the species in the area. This is the first study on electronic tagging of the shortfin mako in the southeastern Pacific Ocean and covers the longest total tracking period reported so far for this species.     

Vertical distribution of <Emphasis Type="Italic">Todarodes pacificus</Emphasis> (Cephalopoda: Ommastrephidae) paralarvae near the Oki Islands,southwestern Sea of Japan

The diel vertical distribution patterns of Japanese common squid, Todarodes pacificus, paralarvae were examined using a Multiple Opening Closing Net and Environmental Sensing System (MOCNESS) in the southwest Sea of Japan near the Oki Islands (Japan) during five late-autumn surveys in 1998–2002. A total of 1,511 paralarvae ranging in mantle length (ML) from 0.7 to 7.3 mm were collected at 63 of the 68 stations surveyed. Most (84%) were collected above 75 m depth and in the mixed layer. The vertical distribution patterns varied little between day and night. Hatchling-sized (<1.0 mm ML) paralarvae were abundant at 0–25 m depth, and paralarval ML increased with increasing sampling depth. Our results suggest that T. pacificus paralarvae do not exhibit large diel vertical migration patterns, but as they increase in size, paralarvae gradually descend in the water column and the variability in depth increases with ontogeny.     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. I. Twilight DVM and its relationship to the diel light cycle

Marine copepods commonly exhibit vertical movements in the water column over the diel cycle, termed diel vertical migration (DVM), with the most common pattern being an ascent in the water column to minimum depth around sunset and descent to maximum depth around sunrise. The present study characterized the DVM pattern of the pontellid copepod Calanopia americana Dahl in the Newport River estuary (North Carolina, USA, in July 2003). The estuary is shallow and well-mixed, and the study site (34°43N; 76°40W), 1.5 km inside the estuary entrance, is unusual in lying within a gyre where tidal currents are always in the seaward direction. Changes in C. americana vertical abundance were related to spectrally relevant changes in light throughout the diel cycle. Simultaneous measurements of light and zooplankton abundance near the surface (0.5 m depth) and near the bottom (0.5 m above bottom) were made over one 4-h period and two 3-day periods during different phases of the tide. These observations suggest that C. americana undertook twilight DVM in the Newport River estuary; an ascent to the surface occurred at sunset, followed by a descent to near the bottom around midnight, with a second ascent to the surface and then descent to near bottom at sunrise. DVM in C. americana was independent of the tidal cycle, with the initial ascent in the water column at sunset possibly associated with relative rates of irradiance change. Copepod vertical movements were consistent with a night-active endogenous rhythm, and appeared independent of the abundance of predatory chaetognaths, Sagitta spp. In DVM studies with migrators like C. americana that are broadly sensitive to visible wavelengths of light, measuring photosynthetically active radiation may be a reasonable alternative to measuring light in a spectrally relevant photometric unit.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-005-1569-x.Communicated by J.P. Grassle, New Brunswick     

Mechanisms to initiate a Heterosigma akashiwo red tide in Osaka bay

The ecological role of diel vertical migration of Heterosigma akashiwo Hada to initiate red tide was investigated in Tanigawa Fishing Port and Sano Harbor, Osaka Bay, Japan during red tide seasons in 1979 and 1980. This species migrated toward the surface early in the morning at a velocity of 1.0 to 1.3 m h^-1. Downward migration was found in the afternoon, and more than 2.0×10³ cells ml^-1 aggregated in the bottom layer at night. The upward migration started before sunrise and downward shifting occurred prior to sunset. The movement is presumably associated with circadian rhythm, which is known as one of the biological periodicities. H. akashiwo crossed steep temperature and salinity gradients (6.5°C and 5.7 S) during the diel vertical migration. High values of particulate organic carbon and nitrogen concentrations were obtained in dialysis bags suspended in situ at identical layers with high cell concentration, while the values for surface and bottom bags were comparatively low. The results reveal that H. akashiwo migrates toward the sea surface to carry out photosynthesis effectively, and to the bottom to utilize nutrients efficiently.     

Diel vertical movements of adult male dolphinfish (Coryphaena hippurus) in the western central Atlantic as determined by use of pop-up satellite archival transmitters

The vertical movements of six adult male dolphinfish (Coryphaena hippurus) (95–120 cm estimated fork length), caught using standard sportfishing methods, were investigated using high-rate single-point pop-up satellite archival transmitters from 2005 to 2011 in the western central Atlantic. Data revealed a diel activity pattern within the mixed surface layer with dives below the thermocline suggesting temperature is not a barrier to vertical movements for short periods of time. Dolphinfish were tracked for periods of 4.96–30.24 day (Σ = 83.37 day), reaching depths >200 m, and in temperatures ranging from 16.20 to 30.87 °C. The six tags allowed comprehensive vertical movement analyses by time of day, duration at depth, and based on vertical movement patterns. The longest (>60 min), deepest (>30 m), and most extensive vertical movement patterns occurred during night rather than day, with the most time spent near the surface during the day. Dolphinfish spent 66 % of their time in the surface layer (0–9.9 m) and only one individual spent 8 % of the monitoring period diving >8 °C from the maximum surface temperatures recorded while tracked. Two tags were analyzed based on lunar phase and revealed contrasting relationships between vertical movements during new and full phases. Our results suggest dolphinfish vertically shift between surface and at-depth feeding strategies to exploit aggregating epipelagic and mesopelagic prey items leading to predictable diel vertical movements.     

Ontogenetic diel vertical migration of the planktonic copepodCalanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepodCalanus sinicus was investigated in the Inland Sea of Japan in summer 1988, when the water was thermally stratified with a thermocline of ca 5 °C between 35 and 45 m. Stage-specific differences in the diel vertical migration behavior ofC. sinicus were found. Eggs were spawned primarily within the surface-waters between midnight and dawn by ascending females, and sank gradually to deeper waters until they hatched into nauplii. Non-feeding nauplius stages (NI and II) were distributed throughout the water column, but the first feeding stage (NIII) performed an ontogenetic upward migration. NIV to VI and copepodite (C) stages I to III continuously aggregated in the phytoplankton-rich euphotic layer. However, the depth of the median CI to III populations descended as stage progressed. The onset of prominent diel vertical migration took place in CIV, and the amplitude of vertical migration increased with age, being maximal in adult females (CVI). Adult males (CVI), however, remained in the layer below 20 m, and did not migrate dielly. The ecological significance of ontogenetic diel vertical migration is discussed.     

Physiological and behavioral aspects of Calanus euxinus females (Copepoda: Calanoida) during vertical migration across temperature and oxygen gradients

 In the Black Sea, during summer stratification, Calanus euxinus (Hulsemann) undertakes diel vertical migrations with an amplitude of about 117 m from oxygenated, warm (18 °C) surface layers to hypoxic (∼0.8 mg O₂ l⁻¹) zones with lower temperature (7.9 °C). When such changes in temperature and oxygen concentration are reproduced in the laboratory, total metabolism, basal metabolism and scope of activity of copepods decrease 7.2, 7.8 and 6.7 times, respectively, while the frequency of locomotory acts and mechanical power decline 3.4- and 9.5-fold, respectively. These changes allowed the copepods to conserve a significant portion of food consumed near the surface for transformation to lipid reserves. Diel respiratory oxygen consumption of migrating individuals, calculated so as to include actual duration of residence in layers with different temperature and oxygen concentrations, is estimated at 17.87 μg O₂ ind⁻¹. The net energy cost of vertical migration made up only 11.6% of the total. Copepods expend 78.6% of diel energy losses during approximately 10 h in the surface layers, while about 5.4% is required during about 9 h at depth. Hypoxia is shown to have a significant metabolic advantage during diel vertical migrations of C. euxinus in the Black Sea. Received: 1 October 1999 / Accepted: 11 July 2000     

Vertical distributions of eggs,nauplii and copepodites of Calanus helgolandicus (Copepoda: Crustacea) in the Celtic Sea

The vertical distributions of eggs, nauplii, copepodites and adults of Calanus helgolandicus (Claus) from five oblique plankton-net hauls taken in May (1980), March and September (1981) and January (1982) at a site in the shelf sea to the south-west of the United Kingdom are described. The water depth is approximately 95 m and becomes thermally stratified during the summer months when a thermocline of ∼6 C° develops. In early spring when the water column was isothermal (∼8 °C), the development of the eggs and nauplii took place below 60 m and a single ontogenetic migration was observed between Nauplius VI (NVI) and Copepodite I (CI). As the temperature of the water increased, this migration occurred in progressively earlier naupliar stages. The eggs were distributed throughout the water column in the profile taken in early May when a 1 C° thermocline occurred between 30 to 40 m. The majority of the NI to NIV stages occurred below 40 m, with the ontogenetic migration taking place in the NIV stage; the NV and NVI stages were found above the thermocline. In September, the eggs were again distributed throughout the water column (101 490 m^-2), with a maximum number of >4 500 m^-3 occurring in the surface to 5 m depth interval. Nauplius I and II were found at all depths, demonstrating that hatching occurred throughout the water column. The ontogenetic migration in these late-summer profiles took place between the NII and NIII stages, the remainder of the nauplii being found above the thermocline in the top 20 m. This is the first time that an ontogenetic migration, similar to the developmental ascent observed in the naupliar stages of the euphausiid Euphausia superba in the deep ocean, has been shown for a copepod nauplius.     

Diel vertical migrations and nocturnal feeding of a dense coastal krill scattering layer (Thysanoessa raschi and Meganyctiphanes norvegica) in stratified surface waters

To study the nocturnal feeding of euphausiids during vertical migrations and its impact on the phytoplankton, a phytoplankton-rich water mass (drogue marked) drifting over a dense krill scattering layer (acoustic 104kHz) in the lower St. Lawrence estuary was monitored over 46 h in July 1982. Phytoplankton >20 m was abundant and mostly concentrated at the bottom of the photic layer above the pycnocline. Less than 42% of the particulate carbon was due to phytoplankton. The krill scattering layer was about 2 to 3 km in width, elongated along the 100-m bathymetric contour, and absent when the bottom was shallower than 50 m. Its upper day depth was 50 m. At deeper depths, its vertical distribution frequently changed from unimodal to polymodal shapes and viceversa, often with large concentrations of zooplankton just above the bottom. Typical vertical migrations were observed on both days. At night the scattering layer had a lower scattering strength. Most of it was below the thermocline but net catches showed that large concentrations of euphausiids (up to 57 individuals m^-3) crossed it. Stomach pigment content of Thysanoessa raschi was generally low, but mean stomach fullness was always high. They were more opportunistic than herbivorous. From stomach fullness and the presence of a food bolus in mouth parts, feeding in surface waters appeared to be intensive, but gut content indicated that food was not processed there. It is therefore suggested that individuals underwent vertical interchanges across the thermocline while feeding during the night. Meganyctiphanes norvegica had significant herbivorous activity during the night. The grazing pressure impact of the scattering layer on phytoplankton was negligible.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. II. Proximate role of exogenous light cues and endogenous rhythms

The marine copepod Calanopia americana Dahl undergoes twilight diel vertical migration (DVM) in the Newport River estuary, North Carolina, USA, in synchrony with the light:dark cycle. Copepods ascend to the surface at sunset, descend to the bottom around midnight, and make a second ascent and descent before sunrise. Behavioral assays with C. americana in the laboratory during fall 2002/2003 and summer 2004 investigated aspects of three hypotheses for the proximate role of light in DVM: (1) preferendum hypothesis (absolute irradiance), (2) rate of change hypothesis (relative rates of irradiance change), and (3) endogenous rhythm hypothesis. Results suggest that C. americana responds to exogenous light cues consistent with its DVM pattern; changes in absolute irradiance evoked swimming responses that would result in an ascent at sunset and descent at sunrise, while relative rates of irradiance decrease at sunset (–0.0046 s^–1) evoked an ascent response, and relative rates of irradiance increase at sunrise (0.0042 s^–1) evoked a descent response. Furthermore, C. americana expressed an endogenous rhythm in vertical migration that was positively correlated with field observations of twilight DVM. Collectively, these results indicate that both exogenous light cues and endogenous rhythms play a proximate role in twilight DVM of C. americana, providing redundancy in the causes of its vertical migration.Communicated by J.P. Grassle, New Brunswick     

Combining fishing and acoustic monitoring data to evaluate the distribution and movements of spotted ratfish <Emphasis Type="Italic">Hydrolagus colliei</Emphasis>

Direct and indirect methods have been used to describe patterns of movement of fishes, but few studies have compared these methods simultaneously. We used 20 years of trawl survey data and 1 year of acoustic telemetry data to evaluate the vertical and horizontal movement patterns of spotted ratfish Hydrolagus colliei in Puget Sound, WA, USA. Densities of large ratfish (≥30 cm) were higher at the deepest depths trawled (70 m) during daylight hours, whereas densities were similar across depth zones (to 10 m) at night. Acoustic tracking of ratfish showed distinct diel patterns of movement and activity level; ratfish moved into shallow, nearshore habitats at night from deeper, offshore habitats during the day and made ~3 times more moves at night than day in shallow habitats. Broader spatial patterns depended on where ratfish were tagged: one tag group remained in one general location with few excursions, whereas a second tag group moved within a 20-km band with some individuals moving >90 km. These data will help inform food web models’ abilities to quantify interspecific interactions between ratfish and other components of their community.     

Monitoring acoustically tagged king prawns <Emphasis Type="Italic">Penaeus</Emphasis> (<Emphasis Type="Italic">Melicertus</Emphasis>) <Emphasis Type="Italic">plebejus</Emphasis> in an estuarine lagoon

Fine-scale movement patterns in penaeid prawns are rarely observed in situ, but are essential in understanding habitat use, foraging, and anti-predator behaviour. Acoustic telemetry was applied to examine the activity, space utilization, and habitat use of the eastern king prawn Penaeus (Melicertus) plebejus, at small temporal and spatial scales. Tracking of sub-adult P. plebejus (n = 9) in Wallagoot Lake (36.789°S, 149.959°E; 23 April–12 May 2009) and calculation of a minimum activity index (MAI) revealed high variation in activity rates across diel periods and in different habitats. Elevated activity rates and movement indicated foraging in unvegetated habitats during the night. Areas within the 95 and 50% space utilization contours averaged 2,654.1 ± 502.0 and 379.9 ± 103.9 m², respectively, and there was a significant negative relationship between these areas and prawn activity rates in unvegetated habitats. This study provides the first estimates of prawn activity rates and space utilization in the field. Application of acoustic telemetry can increase knowledge of prawn movements and their interactions with other marine species in different habitats.     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in June 1989, when the water column was thermally weakly stratified. Because of fewer eggs and less variation in their abundance, nocturnal spawning was not apparent. A pronounced upward migration occurred in NIII. NIII to CIII resided in the upper 20 m layer throughout the day, and from CIV on their median depths descended. CV and adult females underwent significant diel vertical migration, whereas adult males did not migrate. By integrating the results from the present study and those from our previous investigations (in August–September 1988, November 1988 and March 1989), we review seasonal variation in the ontogenetic diel vertical migration of C. sinicus. Spawning was largely nocturnal, reaching its maximum level around dawn, but spawning depth and fecundity changed seasonally. The distribution of pre-feeding stages, NI and NII, was similar to that of eggs. A pronounced upward migration always occurred in the first feeding stage, NIII, and late nauplii and early copepodites always resided in the food-rich upper layer, indicating that upward migration by NIII is feeding migration. As the stages progressed, they extended their vertical distribution range, and CV and adult females usually underwent diel vertical migration. However, the pattern and strength of this migration differed seasonally. Their day depths increased with the increase of relative biomass of planktivorous fish, indicating that predator avoidance induces their diurnal downward migration. High chlorophyll a concentrations in the upper layer (<15 m deep) relative to the lower layer (>20 m deep) amplified their diel vertical migrations. Diel vertical migration of C. sinicus is a phenotypic behavior.     

Vertical distribution,and seasonal and diurnal migration of Calanus helgolandicus in the Celtic Sea

The vertical distribution and migration (seasonal, diel and ontogenetic) of Calanus helgolandicus are described from the shallow (100 m) shelf-seas to the south-west of the British Isles. In 1978 and 1979, the overwintering population of C. helgolandicus consisted primarily of Stage V copepodites and adults. By late winter/early spring the copepodites had moulted to adult females (>90%), which matured and bred the first cohorts of the year, prior to onset of the spring phytoplankton bloom in April/May. C. helgolandicus reached a peak of numerical abundance in August of 20x10³ copepodites m^-2 (over the depth range sampled -0 to 70 m), which was 200 times the population in winter. The seasonal peak of abundance occurred 4 mo after the peak of the bloom of phytoplankton in spring. The yearly development of the copepod was not always out of phase with the diatom bloom, as seen when the data from 1978 was placed in the context of a longer time-series collected at 10 m over 22 yr (1960–1981, inclusive). Large vertical migrations were observed in the younger copepodites (CI and II) in May from below to above the thermocline. In the remainder of the year, the CI and CII stages behaved differently and were located above the thermocline within the euphotic zone. The largest vertical displacements of biomass were seen in the summer months due to the migrations of the CV stages and adults, which had developed from the spring cohorts. It was contended that the seasonal and vertical migrations of C. helgolandicus are part of a more complex pattern of inherent behavior than has been reported previously and that, however difficult this is to discern in the natural populations, it always expresses itself.     

Temporal changes in feeding habits and daily rations of<Emphasis Type="Italic"> Hoplostethus mediterraneus</Emphasis> in the bathyal Ionian Sea (eastern Mediterranean)

Data on the diet, feeding habits and daily rations of Hoplostethus mediterraneus Cuvier, 1829 in the bathyal eastern Ionian Sea (Mediterranean Sea) are presented. A total of 430 specimens collected by bottom trawls at depths ranging from 473 to 603 m during four 24-h day–night sampling cycles covering the four annual seasons was examined. H. mediterraneus diet consisted of pelagic and vagile epibenthic prey, mainly crustaceans, and was dominated by benthopelagic natantian decapods (83.35% IRI, index of relative importance). Seasonal changes in diet were apparent and related to seasonal fluctuations in suprabenthic and zooplanktonic prey in the environment. Diel patterns in stomach fullness and trends in diel feeding cycles are discussed in relation to the vertical migratory movements of available prey (i.e. suprabenthos and zooplankton). Daily-ration estimates were determined by evacuation-rate models and ranged from 0.143% to 0.397% WW/WW. Overall, daily-ration estimates were within the range of the daily consumption of other deep-sea fish. Deduced from diet contents, we found a constant gross energy intake (305–316 kcal g^–1) during all seasons. As a possible response to the reproductive peak of mature females observed in summer, H. mediterraneus increases its food consumption, which, in turn, is coupled with an increase in food availability.Communicated by S.A. Poulet, Roscoff     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic

Fishes and zooplankton were obtained (March–April 1979 and partly in August 1974) from 45 hauls taken during the day and at night in the central equatorial Atlantic between Latitude 3°N and 2°S from the surface to 1250-m depth, using the RMT 1+8, a combined opening-closing plankton and micronekton trawl. The vertical distribution of 30 myctophid species is described. All species migrate in a diel pattern, Ceratoscopelus warmingii and Lampanyctus photonotus down to at least 1250 m. During daytime most species aggregated at 400-to 700-m depth, therefore only partly occupying the depth of the Deep Scattering Layer (400 to 500 m at 15 kHz). The feeding patterns of seven of the most abundant species were compared, with a total of 1 905 stomach contents being analysed. All seven species are regarded as opportunistic predators, which feed predominantly during the night on calanoid copepods. A total of 66 species of calanoid copepods were identified among the prey items, with smaller species definitely being in the minority. Stomachs of C. warmingii (700 to 1 250 m depth) and Lepidophanes guentheri (500 to 900 m depth) from daytime samples contained copepod species restricted to the upper 150 m of the water column, including Undinula vulgaris, Nannocalanus minor, and Euchaeta marina, thereby confirming an extended vertical migration of predators. Differences in diet and preferences between species in their total food spectrum are described.     

The response of sea scallop (Placopecten magellanicus) veligers to a weak thermocline in 9-m deep mesocosms

We examined the vertical distributions of scallop (Placopecten magellanicus) veligers in deep (0.6 m diameter, 9.5 m deep) polyethylene mesocosms from December 1991 to January 1992. In the mesocosms temperature stratification varied from 0 to 1.5 °C. Profiles of vertical distribution revealed several repeated patterns. Peaks in veliger numbers often appeared at the water surface and just above the thermocline. Higher density patches were seen below the surface peaks, and revealed the presence of bio-convective cells. Distribution away from these discontinuities was usually even. Distribution of veligers was affected by thermoclines above 1.0 °C. Responses to thermoclines varied with larval age and time of day, and 28 to 30 d veligers passed in both directions through a 1.5 °C thermocline. We conclude that larval behaviour is a major determinant of whether veligers pass through a thermocline. Kinematic viscosity may play a role in perception of temperature changes. Two potential consequences of such behaviour are (1) remaining in more productive upper water layers, where feeding opportunities are enhanced, and (2) increased horizontal transport in the region of the thermocline, which may enhance recruitment. Received: 15 May 1996 / Accepted: 11 May 2000