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.     

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 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.     

Vertical habitat-partitioning by copepodites and adults of subtropical oceanic copepods

Vertical distributions of the abundant larger copepods, both adults and late copepodites, were observed day and night in the upper 500 m of the North Pacific central gyre in early November, 1971. Densities of the copepodites usually equalled or exceeded those of the adults. Copepod species with maximum densities at or above 100 m (Calanus spp., Nannocalanus minor f. major, Undinula darwini, and Euchaeta rimana) usually had no ontogenetic or diel migration. Neocalanus spp. and Haloptilus longicornis exhibited ontogenetic but not diel migrations. Nannocalanus minor f. minor, Aetideus acutus, Euchaeta media, Scolecithrix spp. and Pleuromamma spp., had both ontogenetic and diel migrations. Adults and copepodites of E. media and Pleuromamma spp. usually had their night modes at the same depth, but the daytime modes were at progressively deeper depths for progressively older stages. Daytime modes for adults and copepodites of A. acutus and Scolecithrix bradyi were at the same depth, but the nighttime modes were at shallower depths for progressively older stages. Night modes of all these migrators were usually in the mixed layer (75 m), where primary production rates were maximal. Congeners usually had similar migratory behavior, but competition probably has been a significant determinant of vertical distribution, since congeners, particularly sibling species, consistently had different depths of maximum occurrence during both day and night.     

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.     

Community structure and vertical distribution of cyclopoid copepods in the Red Sea

The species abundance, vertical distribution and diurnal vertical migration of cyclopoid copepods was analyzed in the central Red Sea in October–November 1980. Samples were taken to a depth of 450 m with a multiple opening — closing plankton net with 0.1-mm mesh-size. Selected important species were allocated to five different groups according to their depth distributions during daytime. The greatest number of species (9) was found in the lower epipelagic zone (40 to 100 m), below the strong seasonal thermocline. The lowest number of species (1) occurred in the upper part of the upper mesopelagic zone (100 to 250 m), which is characterized by a strong dissolved oxygen gradient. Five species had a bimodal vertical distribution, with dual peak abundances in the epipelagic and upper mesopelagic zones. Distinct differences in distribution patterns were noted between sexes and/or developmental stages. The vertical range of diurnally migrating species was small, usually less than 50 to 100 m. Characteristic diurnal changes in the vertical succession of dominant species occur within the epipelagic zone (0 to 100 m). Species-specific vertical distribution patterns are compared with published data from other areas. A conspicuous difference in the proportion of carcasses was noted between species: small species (<0.5 mm in length) had a much higher proportion of carcasses, usually between 20 and 40% of the total standing stock, than larger ones (<5%). The potential causes of this phenomenon, which may be due to (1) methodological bias, (2) a lower sinking velocity of small carcasses, or (3) a higher mortality rate of small species, are discussed.     

Movement patterns of large bigeye tuna (Thunnus obesus) in the open ocean, determined using ultrasonic telemetry

 The horizontal and vertical movements of large bigeye tuna (Thunnus obesus Lowe, 1839; 25 to 50 kg) captured in the south Pacific Ocean (French Polynesia) were determined using pressure-sensitive ultrasonic transmitters. Bigeye tuna swam within the first 100 m below the surface during the night-time and at depths between 400 and 500 m during the daytime. The fish exhibited clear relationships with the sound scattering layer (SSL). They followed its vertical movements at dawn and dusk, and were probably foraging on the organisms of the SSL. Bigeye tuna did, however, make regular rapid upward vertical excursions into the warm surface layer, most probably in order to regulate body temperature and, perhaps, to compensate for an accumulated oxygen debt (i.e. to metabolize lactate). The characteristics of these dives differ from those reported from previous studies on smaller bigeye tuna (∼12 kg) near the main Hawaiian Islands. During the daytime, the large fish in French Polynesia made upward excursions approximately only every 2.5 h, whereas smaller fish in Hawaiian waters made upward excursions approximately every hour. Our data are the first observations on the role of body size in the vertical behavior of bigeye tuna. Received: 9 September 1998 / Accepted: 25 November 1999     

Diel vertical migration and feeding of copepods at an oceanic site near South Georgia

Twelve Longhurst Hardy Plankton Recorder (LHPR) profiles were taken over a 16 h period in January 1990, in order to study feeding of four copepod species at an Antarctic oceanic site near South Georgia. Vertical distributions of their life stages, as well as those of dominant competitors and predators, are described in relation to the feeding cycles of Calanoides acutus CV, Calanus simillimus CV, Calanus propinquus CV and Rhincalanus gigas CIII, CV and CVI. Comparisons with vertical ring-net catches, which were used for concomitant gutevacuation experiments, demonstrated the suitability of the LHPR for these fine-scale studies. Planktonic predators, with the exception of the diel migrant Themisto gaudichaudii, resided deeper than the herbivores. During the day and around midnight, when feeding rates were low, species and stages reached their maximum vertical separation. At these times, new generation copepodites of the four species lived progressively deeper and the overwintered generation (i.e., R. gigas Stages CIV, CV, CVI) were progressively shallower. During the afternoon or evening (depending on species), all stages older than CII, as well as Euphausia frigida and T. gaudichaudii, migrated upwards, to amass in the surface mixed layer. Feeding was restricted to darkness, although R. gigas commenced several hours before dusk. In detail their migration and feeding differed widely, with combinations of unimodal and apparent bimodal cycles. As a whole, the results suggest that (1) feeding could occur during sinking as well as during upward migrations, (2) upward migrations were not always associated with feeding increases, and (3) individuals appeared to descend after filling their guts.     

Comparison of the feeding habits of migratory and non-migratory Stenobrachius leucopsarus (Myctophidae)

Stenobrachius leucopsarus, the most abundant species of myctophid fishes off Oregon, USA, has a bimodal distribution at night, with a peak of abundance in the upper 100 m composed of diel vertical migrants, and another peak at 300 to 500 m composed of fish that did not migrate the night they were caught. We compared the feeding habits of these two groups of fish in an attempt to learn if deep fish migrated to surface waters. Low similarity of diets, differences in the rank order of common prey, and similar states of stomach fullness and digestion of prey suggest that fish captured in deep water at night probably did not feed exclusively in shallow water on previous nights. They probably fed in deep water. The similarity in food habits between deep and shallow fish is most readily explained by daytime feeding by fish in deep water and by broad vertical distributions of prey.     

Species composition, biomass and vertical distribution of micronekton over the mid-slope region off southern Tasmania, Australia

Seasonal sampling was carried out based on day/night, vertically stratified tows (100 or 125 m strata) in the upper 900 m of the water column over the mid-slope commercial fishing grounds south of Tasmania. A large midwater trawl (105 m² mouth area) was used with an opening/closing cod-end. Subtropical convergence and subtropical species dominated the fauna, but many less abundant, more widely-distributed species were also present. Fishes, which contributed 89% of micronekton biomass and 135 of 178 species, were dominated by the Myctophidae (48% biomass and 48 species). Twenty micronekton species made up 80% of the total biomass. Overall, the micronekton fish biomass in this region was 2.2 g m⁻² wet weight. A pronounced day/night shift in the distribution of biomass was attributable to diel migratory species. During the day, <0.2% of the total micronekton biomass was found in 0 to 300 m; most biomass was below 400 m, with peaks at 400 to 525 m and 775 to 900 m. At night, 53% of the biomass was found in 0 to 300 m, with progressively less in each deeper stratum. The vertical ranges of individual species typically exceeded 400 to 500 m during the day and night and were non-coincident, although nyctoepipelagic migrators were concentrated in the surface 200 m at night. Distinct epipelagic, lower and upper mesopelagic assemblages were identified, and patterns of epipelagic migration, limited migration and non-migration were categorised for species from each of the lower and upper mesopelagic assemblages. The vertical distribution of these assemblages was coincident with the primary water masses: subantarctic mode water (∼250 to 600 m) and antarctic intermediate water (below ∼700 m). The flux of migrating micronekton, estimated at 0.94 to 3.36 g C m⁻² yr⁻¹ to the lower mesopelagic and 1.14 to 4.06 g C m⁻² yr⁻¹ to the upper mesopelagic, appeared to be considerably outweighed by the consumption needs of aggregated mid-slope benthopelagic predators. We suggest that advection of mesopelagic prey in antarctic intermediate water may sustain aggregated populations of orange roughy (Hoplostethus atlanticus) and other predators on the micronekton in mid-slope depths at this site. Received: 2 April 1997 / Accepted: 21 August 1997     

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.     

Vertical distribution of larvae of Euphausia nana and E. similis (Crustacea: Euphausiacea) in Sagami Bay and Suruga Bay,Central Japan

Vertical distributions of the larval stages of Euphausia nana Brinton and E. similis G. O. Sars in Sagami Bay and Suruga Bay, Central Japan were studied. Most of the metanauplius larvae of E. nana occurred between 25 and 80 m depth, and they were found at greater depths than the eggs and calyptopis larvae. The nauplii and metanauplii of e. similis were mainly found between 50 and 100 m depth, and they also occurred deeper than the eggs and calyptopes. The larvae of the two species from calyptopis I demonstrate diurnal vertical migration. However, this phenomenon was not clear in the season (March) when a seasonal thermocline was absent. Calyptopes and fruciliae of E. similis occurred deeper and migrated over greater vertical distances than those of E. nana. The distance of migration of furcilia I larvae of E. similis was estimated to be about 200 m.     

Offshore–inshore and vertical distributional patterns of heteropod mollusks off leeward Oahu, Hawaii

Replicated zooplankton tows were taken during day and night periods at neritic (1 nmi offshore) and oceanic (5 and 15 nmi offshore) stations off the leeward side of Oahu, Hawaii during fall (September) and spring (April). Plankton samples were obtained with a 1-m² MOCNESS net system towed obliquely through seven depth strata to 80 m (neritic station) and eight depth strata to 200 m (oceanic stations). Five more species were collected in April (17) than in September (12). During both months the number of species decreased from the 15 to the 1 nmi stations; from 11 to 9 in September and 15 to 11 in April. Species diversity, as measured by species accumulation (rarefaction) curves, was higher at the 15 than the 1 nmi stations during both months. Total nighttime water column density in April was about double that in September. In September, the density decreased sevenfold from the 15 to 1 nmi stations, but in April densities were comparable among the three stations; mainly as a result of the shoreward increase in density of Atlanta plana. The most abundant species in both months were A. plana, A. lesueuri, A. inflata, A. peroni, and Protatlanta souleyeti, which in combination accounted for 93% (September) and 91% (April) of the total nighttime water column densities. The vertical ranges of six species were limited to the upper 100 m, and nocturnal migration was suggested for two (A. helicinoides during both months and A. turriculata in April). The ranges of the remaining 11 species extended to 160 or 200 m, and among them nocturnal migration was suggested for five (A. plana, A. meteori and Pterotrachea hippocampus during both months, and A. peroni and Protatlanta soueleyti in April).     

Under-ice feeding and diel migration by the planktonic copepodsCalanus glacialis andPseudocalanus minutus in relation to the ice algal production cycle in southeastern Hudson Bay,Canada

The marine planktonic copepodsCalanus glacialis Jaschnov andPseudocalanus minutus (Kroyer) typically dominate the copepod biomass in spring under the ice in southeastern Hudson Bay, Canada. Females of both species exhibited significant diel feeding cycles, as measured by gut pigment content, throughout a bloom of ice algae at the ice-water interface in 1986. Periods of grazing correlated well with a nighttime vertical migration by females to within 0.2 m of the ice-water interface, suggesting that feeding took place at or just below the thermohaline boundary between seawater and the interfacial layer containing the ice algae. Seasonal melting of the ice bottom in mid-May resulted in freshening of the surface layer and release of the ice algae into the water column. FemaleC. glacialis andP. minutus responded by ceasing migration to the interface. Gut pigment content, and by reasonable assumption, feeding activity in the water column, increased substantially immediately after this event. In mid-May, the water column phytoplankton consisted of flagellates, sedimenting ice algal cells, and diatoms (Navicula pelagica andChaetoceros sp.) previously found at the interface and then growing in the water column. We conclude that algae growing at the ice-water interface, and sedimenting or actively growing algae derived from this interfacial layer, are a regular and principal source of nutrition for these pelagic copepods during and immediately after the ice algal bloom.     

Vertical community structure and ontogenetic distribution of chaetognaths in upper pelagic waters of the Eastern Mediterranean

The vertical distribution of chaetognaths and their developmental stages were investigated in the pelagic waters (0 to 500 m) of the Eastern Mediterranean Sea. Twenty vertical hauls were taken during autumn 1991 in the Ionian, Cretan, Levantine and Rhodes Seas. The chaetognaths Sagitta bipunctata and S. enflata had the same median depth distributions, which differed from those of S. minima, S. serratodentata, S. lyra, Krohnitta subtilis, and S. decipiens, while S. hexaptera occurred only sporadically. S. bipunctata and S. enflata were mainly restricted to <50 m, and while their distributions co-occurred their stage composition differed. The remaining species took up progressively deeper positions in the water column. Ontogenetic vertical distributions, with the older stages occurring at greater depth, were only observed for the mesopelagic species S. lyra and S. decipiens. Diurnal vertical migration was not detected in any of the species nor in their developmental stages.     

Vertical distribution and diurnal migration of some Cyclopoida (Copepoda) in the tropical region of the Pacific Ocean

The material was collected in the western equatorial part of the Pacific Ocean at a 24 h station on December 21 to 23, 1968. Collections were made by means of 2 Juday nets from 2 winches simultaneously. A series of hauls was made at 2 h intervals. The vertical distribution of 17 Cyclopoida species, belonging to 3 families (Oithonidae, Oncaeidae and Corycaeidae) was studied. From 0 to 300 m, the absolute number of Cyclopoida individuals was practically constant throughout the 24 h period. The bulk of the species performed no diurnal vertical migration, or migrated with low intensity in the usual way (i.e. moved upward at night and downward during the day). Only 2 Oncaeidae species performed significant diuranl vertical migration. No reversed migration in Cyclopoida was discovered. To characterize the vertical distribution of Cyclopoida, the distribution of the cores of populations (abundancies between 25 and 75%) was examined. During the 24 h period, the cores of populations of various Cyclopoida species were found at different depths; this steplike distribution was not disturbed in migrating species. The cores of populations differed both with habitat depth and time of ascent of the migrating species to the surface layer. This peculiarity, probably, tends to lessen the intensity of food competition between Cyclopoida species with similar nutritional habits.     

Laboratory experiments on the diurnal vertical migration of marine dinoflagellates through temperature gradients

The diurnal vertical migrations of 4 marine dinoflagellates (Cachonina niei, Ceratium furca, Gymnodinium splendens, and Prorocentrum micans) were studied in a thermally-stratified Plexiglas column (1.70 m deep, 0.61 m diameter). All species migrated through temperature gradients exceeding 7 C°. Some species exhibited altered migration patterns at different salinities either between experiments or compared to observations by other investigators. P. micans exhibited complex changes in swimming speed that depended both on temperature and on the light cycle. These changes provide insight into the possible time course of a field organism's diurnal exposure to physical variability in the water column. P. micans exhibited compositional changes in response to the light cycle, but not to the cross-thermocline temperature differential. A multi-parameter response matrix is required to model coupling between organisms and biologically-active physical mechanisms realistically.     

西北地区昼夜增温的不对称性对植被动态的影响

全球变暖有很大的差异性,表现在昼夜增温速度和各个季节的温度增加的速率具有不一致性,夜间增温的速度要快于白天增温的速度,夜间气温的增温对植被变化的影响更为明显。文章利用34年逐月最高温、最低温以及降水量数据,用一元线性回归分析法、反距离权重插值以及二阶偏相关分析法,分析了西北地区白天和夜间气温的非对称时空变化和非对称增温趋势对植被动态的影响,结果表明:西北地区昼夜温度都呈现出上升的趋势,且上升趋势比较明显,白天增温的速度和夜间增温的速度呈现出不对称性,夜间温度升高的速率是白天温度升高速率的1.2倍,昼夜的温差呈现出减小的趋势;西北地区昼夜增温趋势存在不一致性,白天气温的增温速率在-0.02-0.11℃?a-1,夜间气温的增温速率在-0.06-0.2℃?a-1。tmax和tmin在绝大部分地区都呈现上升趋势,但在空间变化的区域上具有不对称性;西北地区植被生长对昼夜增温的响应具有明显的空间差异性,大部分地区的植被对昼夜增温表现为积极的响应,夜间增温对植被的影响要比白天增温对植被的影响是比较明显的;白天和夜间增温的不对称性影响着不同类型的植被,白天气温的升高有利于针叶林植被、草地和阔叶林植被的生长,而夜间气温的升高对灌丛和荒漠植被的生长有积极的影响。     

Lunar influence on distribution of a calanoid copepod in the water column of a shallow,temperate estuary

Moonlight had a significant influence on the distribution of the copepodPseudodiaptomus hessei in the water column. Copepod samples were taken between March 1986 and March 1987 in the Sundays River estuary. South Africa. Late copepodid stages and adults showed vertical differences in abundance on moonlit nights, with more animals in bottom waters, but not on dark nights. Numbers of adults present in the water column were markedly lower on full moon nights compared with new moon nights. The strongest decrease in abundance was displayed by ovigerous females. These observed distribution patterns support published predation hypotheses.