Interaction between solar orientation and landscape visibility in Talitrus saltator (Crustacea: Amphipoda)

A series of experiments on the littoral amphipod Talitrus saltator (Montagu) was carried out between April and September, 1978–1981, both under natural conditions and inland with artificial landscapes of different heights on the horizon, in order to assess the visual importance of the landscape in zonal orientation in populations from the Mediterranean coast, and to determine interactions between solar orientation and orientation based on the landscape. Inland, orientation in controls (permitting only vision of the sky and the sun) was compared to that of the experimental individuals, who could see a simulated landscape positioned landwards to them and seawards. In nature, sandhoppers released in a level arena with the landscape screened from view were compared with others released in an unscreened arena and on the sand in absolutely natural conditions. Situations where solar orientation contradicted local cues were produced both by releasing the sandhoppers on a shore diversely orientated to their own, and by shifting their internal clock by nine hours. Results show that T. saltator uses the landscape as a cue in its orientation towards the sea, in conjunction with solar orientation: the latter being the principal factor involved even when the sandhoppers are separated from the local optical factor in the sky (in trials at a distance from the sea). In fact, with the artificial landscape set seawards, none of the populations we studied showed any orientation based on the landscape comparable in accuracy to orientation based exclusively on the sun. From the experiments carried out in natural conditions, it is possible to deduce that conflicting conditions gave rise to two types of results: deviation from the mean direction and an increase in dispersion, up to total dispersion when the local factors were in total contrast to solar orientation (clock-shifted sandhoppers released on the sand).     

Moon orientation in sandhoppers: effects of lighting treatments on the persistence of orientation ability

The moon orientation rhythm persists in sandhoppers removed from environmental entrainments relative to moon periodicity. In order to investigate the underlying mechanism, the crustacean amphipods Talorchestia capensis and Talitrus saltator were collected at new moon and exposed in the laboratory to an artificial light during the day, but to a variety of lighting treatments during the night: (1) continuous dark, (2) dim light in phase with the natural moon, (3) continuous dim light. The animals were tested under the moon 9–21 days later by using a new type of arena. The ability to orientate in a direction perpendicular to the shore persisted in sandhoppers under treatments 1 and 2, but not under treatment 3. A disturbance due to a phototactic tendency in Talorchestia was also observed in animals captured on the day of the test when exposed to unnatural lighting, but not when they were kept under natural light. The present findings show that the timing mechanism allowing compensation for changes in the moon’s position also persists in animals that have been long removed from entraining factors. The dependence of this orientation ability on nocturnal lighting and the disturbing effect of sudden changes in light intensity support the idea that the phasal lighting of the moon, and perhaps sunrise and sunset act as resetting factors for the moon’s orientation rhythm.     

Sexual reproduction of the reef coral Montastreacavernosa (Scleractinia: Faviidae) in the Santa Marta area, Caribbean coast of Colombia

Sexual reproduction of the reef-building coral Montastreacavernosa was studied in the Santa Marta area, Caribbean coast of Colombia, from October 1990 to October 1991. The area is subjected to a seasonal upwelling–outwelling regime. From microscopical analysis of tissue sections sampled each lunar month around the full moon, it was confirmed that this is a gonochoric, broadcasting species, with a single gametogenic cycle per year, and a 1:1 sex ratio. Oogenesis began a few weeks after spawning, and extended for 11 mo. The onset of spermatogenesis occurred just before the full moon of June, later than at other localities where sea-water temperatures during the first half of the year are not as low. The amount of reproductive tissue strongly increased in both sexes after July, in association with an increase in sea-water temperature and a decrease in photoperiod. The gonad index of fully mature female colonies was about four times lower than at other localities, perhaps due to the stressful seasonal regime. The male gonad index was lower than that of the female, indicating sex-related differences in the rates of biomass allocation to reproduction. As in other Caribbean localities, spawning occurred after the full moons of August and September. However, there was some evidence of a third spawning episode after the full moon of October, possibly associated with a delay in the occurrence of maximum sea-water temperatures in near-equatorial localities compared to higher latitudes. The probability of cross-fertilization in this species with a gonochoric breeding system and a broadcasting mode of reproduction is favored by its balanced sex ratio, its usually high within-reef zone abundance, and by localized and repeated spawning episodes, synchronized by lunar phase. Received: 19 April 1996 / Accepted: 24 September 1996     

Multiple regression analysis of the sources of variation in orientation of two sympatric sandhoppers, <Emphasis Type="Italic">Talitrus saltator</Emphasis> and <Emphasis Type="Italic">Talorchestia brito</Emphasis>, from an exposed Mediterranean beach

Two sympatric species from an exposed sandy beach in north-western Tunisia, the sandhoppers Talitrus saltator (Montagu) and Talorchestia brito Stebbing, were compared as regards orientation under natural conditions. Sets of experiments were carried out on the beach using two experimental arenas, one permitting the view of both the sky and landscape and the other of the sky only. Replicates were made at different hours of the day, in different days and period of the year and with various climatic conditions. Multiple regression models were fitted to the angular data obtained. These models allow an estimate of the trends of variation depending on a combination of factors. The simultaneous effects of environmental factors influencing orientation were quantified for each species. The sun was confirmed as the major orientation cue in these populations, but the concentration of directional choices was enhanced by the landscape vision. The climatic factors affected orientation in different ways depending on the time of day and year. When the two species were compared, Talitrus saltator showed a higher flexibility of response than Talorchestia brito under the same environmental conditions, supporting the hypothesis of a higher level of terrestrialization of the former species.     

Sexual reproduction in the coral Pocillopora verrucosa (Cnidaria: Scleractinia) in KwaZulu-Natal, South Africa

The mode of reproduction in Pocillopora verrucosa Ellis and Solander, 1786 varies between geographically isolated regions. This scleractinian coral is common along the KwaZulu-Natal coast, and its reproductive mode and period of reproduction were assessed using histological preparations. The study was undertaken from 1992 to 1994 and showed that P. verrucosa is a simultaneous hermaphrodite and broadcast spawner in KwaZulu-Natal. Gametogenesis occurs from October to January, with the gametes maturing simultaneously so that spawning can be synchronised at new moon in January (mid-summer). Zooxanthellae are present in the mature oocytes. Received: 10 September 1997 / Accepted: 6 July 1998     

Home range, activity and distribution patterns of a temperate rocky-reef fish, Cheilodactylus fuscus

Red morwong (Cheilodactylus fuscus; Cheilodactylidae: Perciformes), are large temperate rocky reef fish that are patchily distributed in local aggregations of 3 to >100 fish. The home ranges and aggregating behaviour of red morwong were investigated using external tags (n = 114, over 35 mo) and ultrasonic transmitters (n = 9 over 20 d) at coastal and estuarine sites in south-east Australia. Adult red morwong had a mean home range during the day of 1865 m² (SE = 268). Night movement, determined by ultrasonic telemetry, indicated a significantly greater mean home range of 3639 m² (SE = 416). Home range did not differ significantly with size or sex. Fish movements were greatest and aggregation sizes varied most during the non-reproductive mid-summer period. Tagged fish (N = 20) displaced 200 to 900 m returned in 1 to 3 d to the point of capture, often traversing open sand habitat and other aggregations. During the day, fish were commonly found on bouldered habitat. During the night, these site-specific aggregations fragmented as fish dispersed over a variety of substrata, with crepuscular peaks in activity. Therefore, estimates of habitat-specificity and feeding patterns collected only by day may give misleading results. Received: 28 June 1996 / Accepted: 8 July 1998     

Grazing of phytoplankton by copepods in eastern Antarctic coastal waters

Chlorophyll a, primary productivity and grazing by copepods on phytoplankton were measured in the upper water column during the summer of 1994/1995 at a coastal site near Davis Station, East Antarctica. Chlorophyll a was at a maximum in mid-December, then dropped markedly as the coastal fast ice melted and broke‐out. Phytoplankton biomass increased again from mid‐ to late‐February. Copepods accounted for at least 65% of zooplankton biomass in the water column before sea ice break‐out, whereas larval polychaetes and ctenophores dominated after ice break‐out. Oncaeacurvata was the numerically dominant species throughout the study. The highest grazing rate (8.7 mg C␣m⁻³␣d⁻¹) was recorded on 21 December when O.␣curvata accounted for 64% of the total. Grazing had decreased markedly by 28 December (0.9 mg C m⁻³ d⁻¹); again O. curvata accounted for over 50% of the total ingested. Copepod grazing increased after ice break-out until the last experiment on 20 February (⋍5 mg C␣m⁻³␣d⁻¹). The main species responsible for grazing during this period were O. curvata, Oithonasimilis, Calanoidesacutus and unidentified copepod nauplii. It was estimated that copepods removed between 1 and 5% of primary productivity. Received: 11 October 1996 / Accepted: 22 October 1996     

Spawning migrations and local movements of a tropical sparid fish

Between June 1994 and September 1996, a mark–recapture study was conducted in the Hinchinbrook Channel area of tropical northeastern Australia to investigate the movements of the sparid fish Acanthopagrus berda. A total of 962 A. berda were tagged, 922 within a single estuary system, Deluge Inlet. No movements among estuaries were detected. Of the fish tagged in Deluge Inlet, 12.9% were recaptured. Two movement regimes were detected: local movements (mostly <500 m) during non-spawning times, and longer migrations (up to 3.12 km) during spawning seasons, to and from a spawning site at the estuary mouth. Although there appeared to be a major spawning aggregation in the mouth of Deluge Inlet, ripe female A. berda were present in upstream sites, indicating that spawning probably occurs there also. This contrasts to the situation in South Africa, where A. berda is reported to spawn only close to estuary mouths. Differences such as this highlight our limited knowledge of variations in the reproductive biology of widespread species such as A. berda. Received: 25 November 1997 / Accepted: 10 September 1998     

Diurnal and lunar patterns of larval recruitment of Brachyura into a mangrove estuary system in Ranong Province, Thailand

The present study followed the temporal recruitment pattern of brachyuran larvae in a mangrove tidal creek on the Andaman Sea coast of Ranong Province, Thailand, based on the assumption that the processes governing recruitment are important for the overall population dynamics of mangrove brachyuran crabs. Plankton net samples were taken on five occasions: on two new moon spring tides, one waxing moon neap tide, one full moon spring tide and one waning moon neap tide during October and November 1997. In addition collectors for larval crab megalopae were employed every 3 days through one dry season and one wet season (March–October 1998). Both the plankton net samples and collector samples revealed four major brachyuran groups in three families: Ocypodidae, Grapsidae and Portunidae. The grapsid group was further separated into two morphotypes which were identified as Metaplax and sesarmid species. Identified group mean numbers per cubic metre were ocypodids 3.0, sesarmids 0.8 and Metaplax 0.5, while portunid megalopae were very scarce (≪0.1 m⁻³). Further analysis of plankton net samples showed that when considering the parameters date, depth, current direction and the diel cycle, Metaplax and ocypodids distribute differently in the tidal and lunar cycle. Metaplax recruitment dominates on flood tides and on bottom layers, followed by middle and surface layers. Conversely, ocypodid abundance varied significantly with date only. Notably recruitment was not dependent on the diel cycle for either group. The collector samples of megalopae showed that recruitment of ocypodids, Metaplax and sesarmids occurred on full and new moon spring tides, while portunid megalopae preferred to settle on full moon spring tides. Since tidal currents were related to the lunar cycle megalopa groups are also cross-correlated with tidal amplitude, except for the portunid group. It is concluded that megalopae recruit in a similar manner to what has been found in other regions of the world, except that the abundance of ocypodids and Metaplax is not influenced by the diel cycle. Received: 14 February 2000 / Accepted: 24 November 2000     

Introduction of the northern Pacific asteroid Asterias amurensis to Tasmania: reproduction and current distribution

The northern Pacific asteroid Asterias amurensis (Lütken) has been recently introduced to Tasmania and is now well-established in east and southeast Tasmania. This sea star is conspicuous throughout the Derwent River estuary and is particularly abundant in the Hobart port area. Reproduction of two populations of A. amurensis at Sullivans Cove and at Sandy Bay was investigated from August 1993 to October 1994 by gonadal histology and measurement of the gonad index (GI). An intense period of vitellogenic and spermatogenic growth started in April, with the breeding condition reached by June. In 1994, peak GI was recorded at Sullivans Cove in July and at Sandy Bay in August. Although these data indicated that the Sullivans Cove population spawned before the Sandy Bay population, histological examination revealed that major spawning activity occurred in both populations from July onwards. In the early part of the breeding season, gametogenesis proceeded in parallel with spawning, with released gametes being replaced by continual gametogenesis. Maintenance of a higher GI during the early part of the breeding season at Sandy Bay was due to prolonged gametogenic replacement at this site. Spawning continued to October 1994, resulting in a sharp decline in the GI. The cyclic expansion and regression of the genital haemal sinus coincident with germinal proliferation and growth, respectively, support the contention that haemal fluid provides nutrients for gametogenesis. In Tasmania, A. amurensis experiences temperature and photoperiod regimes similar to those experienced by endemic populations of this species in the north Pacific. Comparison of reproduction of A. amurensis in Japan and Tasmania at similar latitudes shows that gametogenesis of the northern and southern populations is 6 mo out of phase. This phase shift provides evidence for photoperiodic regulation of gametogenesis in A. amurensis. The similar trend in sea-temperature regimes associated with the stages of gametogenesis in northern and southern populations suggests that temperature also plays a modulatory role. Based on the periodicity of reproduction and the ontogeny of A. amurensis, the planktotrophic larvae of this species would be expected to be in the plankton for several months from the end of winter through summer. Considering the highly fecund, dispersive life-history of A. amurensis, the southeast Tasmanian populations have considerable potential to serve as a seed source for establishment of new populations of this asteroid elsewhere in Tasmania and in mainland Australia. Received: 18 October 1996 / Accepted: 22 October 1996     

Spawning and larval development of the black turban snail Tegula funebralis (Prosobranchia: Trochidae)

An understanding of spawning and larval development can be fundamental to interpreting the abundance, distribution, and population structure of marine invertebrate taxa. Tegula funebralis (A. Adams, 1855), the black turban snail, has been the focus of numerous ecological studies on the Pacific coast of North America. To date, there are only conflicting and anecdotal reports of spawning, and there is no information on larval or juvenile development for this conspicuous and abundant species. On 19 September 1995, two individuals of T. funebralis were observed free-spawning gametes into seawater in tanks at the Oregon Institute of Marine Biology. Embryos and larvae were subsequently reared to metamorphosis and beyond. Development was pelagic and similar to development described for other trochids, and larvae were observed not to feed at any stage. Larvae began to metamorphose at 5.7 to 6.7 d and settled at 260 μm shell length. Juveniles grew ≃ 10 μm in shell length per day and appeared to feed on detritus. Juveniles lacked some adult diagnostic shell characters, including two columellar nodes and a closed umbilicus. In the field, small (<3 mm) juveniles occurred in the adult habitat on all sampling dates between October and March. Small juveniles were found only under rocks and were most abundant under rocks partially buried in coarse sand, suggesting that juveniles may utilize a specific microhabitat within the adult T. funebralis habitat. Received: 7 October 1996 / Accepted: 17 October 1996     

Seasonal abundance and toxicity of the dinoflagellate Gambierdiscus spp. (Dinophyceae), the causative agent of ciguatera in Tahiti, French Polynesia

The benthic dinoflagellate Gambierdiscus spp. is the primary causative agent of ciguatera fish poisoning in French Polynesia. However, the initiation of ciguatera outbreaks and the factors that control ciguatoxin production in this dinoflagellate are still poorly understood. In this paper, we present some original data concerning the seasonal abundance and toxicity of natural populations of Gambierdiscus spp. based on a long-term survey in a ciguateric site of the island of Tahiti. From February 1993 to December 1997, Gambierdiscus spp. population densities were monitored weekly in the Atimaono barrier reef of Papara in relation to temperature and salinity. Densities peaked at 4992 cells g⁻¹ in October 1994 and constituted ≥1000 cells g⁻¹ on several occasions. A total of 58 blooms were recorded in the area, of which 65% occurred in 1995 and 1996 alone. Our data confirmed the high endemicity of Gambierdiscus spp. in the Papara area. Refined analysis of our raw data by a principal-component analysis of processes (PCAP) revealed a seasonality in cell densities from February 1993 to May 1995. During this period, Gambierdiscus spp. populations tended to reach maximum abundance at the beginning and end of the hot season. In contrast, salinity did not appear to be a determining factor in the seasonal abundance of this dinoflagellate. The noticeable increase in both peak densities and frequency of blooms further noticed in 1995 and 1996 was preceded by unusually high water temperatures in January to April 1994, concomitant with a severe coral-bleaching episode. During the course of the study, a total of 303 × 10⁶ cells obtained from 48 successive blooms was harvested. Toxicity screening revealed that toxin production was maximum from October 1994 through December 1996. No correlation was found between toxicity of these blooms and their biomass nor the seasonal pattern of temperatures. It is suggested that the toxicity of naturally-occurring blooms of Gambierdiscus spp. and, consequently, the severity of ciguatera incidents in a given area, is mainly dependent on the clonal nature of cells which coexist within local populations of this dinoflagellate. Received: 12 October 1998 / Accepted: 15 June 1999     

Mass spawning by the sea urchin Evechinus chloroticus (Echinodermata: Echinoidea) in a New Zealand fiord

A mass synchronous spawning of the sea urchin Evechinus chloroticus (Valenciennes) was observed in situ in Doubtful Sound, a large New Zealand fiord. Spawning occurred between 17:30 hrs and 18:30 hrs on 27 January 1994 and coincided with a full moon, spring tides and a period of decreasing sea temperatures. During spawning, the sea urchins formed a dense spawning aggregation of both sexes, with >90% of the urchins observed spawning at the time. Spawned gametes clouded the water column, and some were eaten by small labrid fish species. The spawning, which may have been as widespread as 40 km, marked a 42 to 50% decrease in gonad indices and resulted in a widespread, dense cohort of  E. chloroticus larvae within the fiord. Received: 25 September 1997 / Accepted: 6 March 1998     

The role of daytime cues in the development of magnetic orientation in a night-migrating bird

To assess the role of celestial rotation during daytime in the development of the magnetic compass course, pied flycatchers (Ficedula hypoleuca Pallas, Muscicapidae) were handraised in Latvia under various celestial and magnetic conditions. Tests were performed during autumn migration in the local geomagnetic field (50 000 nT, 73° inclination) in the absence of celestial cues. A group of birds that had never seen the sky showed a bimodal preference for the migratory southwest-northeast axis, whereas a second group that had been exposed to the natural sky from sunrise to sunset in the local geomagnetic field showed a unimodal preference for the seasonally appropriate southwesterly direction. A third group that had also been exposed to the daytime sky, but in the absence of magnetic compass information, also oriented bimodally along a southwest-northeast axis. These findings demonstrate that observing celestial rotation during daytime enables birds to choose the right end of the migratory axis for autumn migration at the Latvian test location. This transformation of axial behavior into appropriate migratory orientation, however, requires the birds to have simultaneous access to information on both celestial rotation and the geomagnetic field. Received: 19 September 1997 / Accepted after revision: 22 November 1997     

Interception and dispersal of artificial food falls by scavenging fishes in the abyssal Northeast Atlantic: early-season observations prior to annual deposition of phytodetritus

The hypothesis that the behaviour of deep-sea scavenging fishes is influenced by seasonal input of organic matter from the ocean surface was investigated by observing responses to baits placed on the sea floor at 4800 m depth in the NE Atlantic (48°50′N; 16°30′W) during spring (April 1994). Data from the present study are compared with those from previous studies of the same location made in summer 1989. The first fishes to arrive at baits were the grenadier Coryphaenoides (Nematonurus) armatus and the eel Histiobranchus bathybius, after delays of 28 and 29 min, respectively; these results are not significantly different from those of summer 1989. Similarly, other indices of activity (staying time and swimming speed) showed no evidence of differences between years/seasons. However, the rate of radial dispersal of bait (0.009 m s⁻¹) by C. (N.) armatus was much slower than in all previous studies. A change in the size distribution of C. (N.) armatus to smaller individuals in spring 1994 was also evident. It is suggested that the fish on the abyssal plain may not comprise a steady-state population and that major episodic or seasonal migrations may occur. Received: 18 October 1996 / Accepted: 20 December 1996     

Tracking adult North Atlantic bluefin tuna (Thunnus thynnus) in the northwestern Atlantic using ultrasonic telemetry

Ultrasonic, depth-sensitive transmitters were used to track the horizontal and vertical movements, for up to 48 h, of 11 adult (136 to 340 kg estimated body mass) North Atlantic bluefin tuna (Thunnus thynnus Linnaeus). Fish were tracked in October 1995, September and October 1996, and August and September 1997 in the Gulf of Maine, northwestern Atlantic. The objective was to document the behavior of these fish and their schools in order to provide the spatial, temporal, and environmental information required for direct (i.e. fishery-independent) assessment of adult bluefin tuna abundance using aerial surveys. Transmitters were attached to free-swimming fish using a harpoon attachment technique, and all fish remained within the Gulf of Maine while being followed. Most of the bluefin tuna tagged on Stellwagen Bank or in Cape Cod Bay (and followed for at least 30 h) held a predominately easterly course with net horizontal displacements of up to 76 km d⁻¹. Mean (±SD) swimming depth for all fish was 14 ± 4.7 m and maximum depth for individuals ranged from 22 to 215 m. All but one fish made their deepest excursions, often single descents, at dawn and dusk. In general, adult bluefin tuna spent <8% of their time at the surface (0 to 1 m), <19% in the top 4 m, but >90% in the uppermost 30 m. Mean (±SD) speed over ground was 5.9 km h⁻¹, but for brief periods surpassed 20 to 31 km h⁻¹. Sea surface temperatures during tracking were 11.5 to 22.0 °C, and minimum temperatures encountered by the fish ranged from 6.0 to 9.0 °C. Tagged bluefin tuna and their schools frequented ocean fronts marked by mixed vertebrate feeding assemblages, which included sea birds, baleen whales, basking sharks, and other bluefin schools. Received: 19 July 1999 / Accepted: 25 March 2000     

Occurrence of the Kumamoto oyster Crassostrea sikamea in the Ariake Sea, Japan

The Kumamoto oyster Crassostrea sikamea is distinguished from the closely related Pacific oyster C. gigas by concordant differences in 16S rDNA, allozymes, and a one-way gametic incompatibility. After repeated failures to find this oyster in its native habitat, we speculated in 1994 that “the Kumamoto oyster may be extinct in Japan”. In September 1996, we sampled small, deep-cupped oysters from the Ariake Sea and typed these for 16S rDNA and ITS-1 DNA markers previously shown to be diagnostic for the three most common oysters in the Ariake Sea, C. gigas, C. sikamea and  C. ariakensis. Our earlier suggestion of the demise of  C. sikamea proved incorrect. Of the 256 oysters sampled, 181 (71%) were  C. gigas, 53 (21%) were C. sikamea, and 22 (9%) were  C. ariakensis; no interspecific hybrids were observed. The distributions of C. sikamea and  C. ariakensis are clumped in the Ariake Sea:  C. sikamea occurs on the eastern and northern shores,  C. ariakensis occurs only in the northern part. These results emphasize the value of molecular markers for discriminating these morphologically plastic species both in the field and in aquaculture. Received 6 July 1998 / Accepted 8 October 1998     

Zooplankton responses to hypoxia: behavioral patterns and survival of three species of calanoid copepods

Seasonally recurrent and persistent hypoxic events in semi-enclosed coastal waters are characterized by bottom-water dissolved oxygen (d.o.) concentrations of < 2.0 ml l⁻¹. Shifts in the distribution patterns of zooplankters in association with these events have been documented, but the mechanisms responsible for these shifts have not been investigated. This study assessed interspecific differences in responses to hypoxia by several species of calanoid copepods common off Turkey Point, Florida, USA: Labidocera aestiva (Wheeler) (a summer/fall species), Acartia tonsa (Dana) (a ubiquitous year-round species), and Centropages hamatus (Lilljeborg) (a winter/spring species). Under conditions of moderate to severe hypoxia 24-h survival experiments were conducted for adults and nauplii of these species from August 1994 to October 1995. Experiments on adults used a flow-through system to maintain constant d.o. concentrations. Adults of A. tonsa showed no decline in survival with d.o. as low as 1.0 ml l⁻¹, sharp declines in survival at d.o. = 0.9 to 0.6 ml l⁻¹, and 100% mortality with d.o. = 0.5 ml l⁻¹. Adults of L. aestiva and C. hamatus were more sensitive to oxygen depletion: both species experienced significant decreases in survival for d.o. = 1.0 ml l⁻¹. Nauplii of L. aestiva and A. tonsa showed no significant mortality with d.o. = 1.1 to 1.5 ml␣l⁻¹ and d.o. = 0.24 to 0.5 ml l⁻¹, respectively. In addition, experiments investigating behavioral avoidance of moderate to severe hypoxia were carried out for adults of all three species. None of the three species effectively avoided either severely hypoxic (d.o. < 0.5 ml l⁻¹) or moderately hypoxic (d.o. ≈ 1.0 ml l⁻¹) bottom layers in stratified columns. These results suggest that in␣nearshore areas where development of zones of d.o. < 1.0 ml l⁻¹ may be sudden, widespread, or unpredictable, patterns of reduced copepod abundance in bottom waters may be due primarily to mortality rather than avoidance. Received: 31 August 1996 / Accepted: 24 September 1996     

Escape behaviour of solitary herring (Clupea harengus ) and comparisons with schooling individuals

The escape behaviour of solitary herring (Clupea harengus L.) startled by a sound stimulus was observed by means of high-speed video-filming. The results were compared with data from a previous study on the escape behaviour of schooling herring. Escape responses were divided into “away responses” and “towards responses” according to the orientation of the C-bend of the body relative to the stimulus. The proportion of away responses was smaller for solitary than for schooling herring. In solitary herring, the subsequent escape trajectories of fish making initial away responses showed a bimodal pattern of distribution, with modes at 130 and 180° from the stimulus. Trajectories following towards responses, however, were mainly within the semicircle directed at the stimulus, and their pattern of distribution differed from that of away responses. This result contrasts with observations on schooling herring, whose trajectories following both initial away and towards responses are directed away from the stimulus. In addition, we measured the response latency, defined as the interval of time between stimulus presentation and the first detectable movement of the fish. Solitary herring showed a higher proportion of short-latency responses (latency <50 ms) than schooling herring. Different behaviours appear to be exhibited by herring depending on whether they are solitary or within a school. We hypothesize that schooling may raise the threshold for initiation of fast escape responses, giving longer latencies and slower responses which are more appropriate in their directionality and reduce the possibility of collisions with neighbours. In addition, we suggest that schooling behaviour enhances the directionality and co-ordination of the escape response of the whole school, possibly increasing the probability of surviving a predator attack. Received: 2 October 1996 / Accepted: 17 October 1996     

School fidelity and homing synchronicity of yellowfin tuna, Thunnus albacares

Thirty-eight yellowfin tuna (Thunnus albacares) were tagged with coded ultrasonic beacons between 6 March and 4 December 1996 near two buoys off the western coast of Oahu, Hawaii. Two to four tuna were captured, tagged, and released on the same day in as rapid succession as possible in an effort to tag members of the same school. Automated “listening” monitors attached to the buoys recorded when these marked individuals entered within a radius of ≤1.1 km of the buoys during a 13 mo period. Twenty-seven of the tuna returned to the site of tagging. The mean number of returns was 4.2 per tuna (max. = 17), and visits ranged from 1 to 910 min (median = 2.7 min, mean = 40.1 min). The intervals between successive returns varied from 1 to 257 d (median = 3.0 d, mean = 17.4 d). Seventy-three percent of the tuna returned together with tunas tagged on the same day, exceeding the frequency of returns of tuna tagged on another day or arriving alone. This social cohesion is supported by the pattern of return visits by five tuna tagged on 6 March at Monitoring Station R. Two or more of these tuna arrived together on 24 of 35 d when tagged tuna were detected. All five individuals visited R on 11 April, a month after tagging, three arrived together 5 mo later on 4 August, and three returned 6 mo later on 1 December 1996. Tuna often arrived at the same time of day, e.g. Individuals 1 and 3 visited R at 09:15 hrs on 12 April and at 09:00 hrs 8 mo later. The returns were also site-specific. The 22 tuna tagged at R made 182 return visits to R (92.4%) and only 15 visits to Monitoring Station K (7.6%), 10 km away. An allegiance of tuna to one school, a predilection for returning to the site of tagging, and precise timing when visiting sites, are consistent with tuna having migratory pathways consisting of “way-points” that are visited with temporal regularity. Received: 30 April 1998 / Accepted: 27 October 1998