Mitochondrial and nuclear DNA analysis of population subdivision among young-of-the-year Spanish mackerel (Scomberomorus maculatus) from the western Atlantic and Gulf of Mexico

The degree of population subdivision among collections of young-of-the-year (YOY) Spanish mackerel Scomberomorus maculatus sampled from along the Atlantic coast of the United States (Atlantic) and Gulf of Mexico (Gulf) was examined. Young-of-the-year were collected from spawning areas to minimize possible mixing among putative spawning stocks that may occur at later life history stages along migratory pathways or at over-wintering grounds. Restriction fragment length polymorphism (RFLP) analysis of the mitochondrial DNA (mtDNA) ND4 region and a nuclear actin intron was conducted. Collections in each of two years were analyzed to minimize bias associated with sampling at a single point in space or time. Substantial variation was detected at ND4 (haplotype diversity=0.81) and at the actin intron locus (gene diversity=0.21). Significant differences in gene diversity or allele frequencies were not detected among temporal samples at any of three locations. A homogeneous distribution of genetic variance among samples from widely spaced geographic regions was consistent with the hypothesis that Spanish mackerel comprise a single intermingling genetic stock. Power analysis showed that the mitochondrial analysis from this study was much more likely to detect population subdivision than analysis of the nuclear actin locus, despite lower mtDNA sample size. Received: 3 January 2000 / Accepted: 28 August 2000     

Mitochondrial DNA variation in king mackerel (Scomberomorus cavalla) from the western Atlantic Ocean and Gulf of Mexico

King mackerel (Scomberomorus cavalla Cuvier) collected in 1992 and 1993 from 13 localities along the Atlantic coast of the southeastern USA and in the northern Gulf of Mexico were surveyed for variation in mitochondrial (mt)DNA and a nuclear-encoded dipeptidase locus (PEPA-2). Both polymorphic and fixed mtDNA restriction sites were identified and mapped using conventional and polymerase chain-reaction (PCR)-based methods. Heterogeneity in mtDNA haplotype frequencies was found only in comparisons of pooled haplotypes from Atlantic localities versus pooled haplotypes from Gulf localities. This finding indicates weak genetic divergence between king mackerel from the Atlantic and those from the Gulf. Frequencies of two PEPA-2 alleles essentially paralleled previous findings: one allele (PEPA-2a) was common among samples from western Gulf localities, whereas the other allele (PEPA-2b) was common among samples from Atlantic and eastern Gulf localities. There was considerable variation in PEPA-2 allele frequencies within broadly-defined regions. Variation in mtDNA haplotypes and PEPA-2 genotypes was independent, as was variation in mtDNA haplotypes with sex or age of individuals. Variation in PEPA-2 genotypes was not independent of sex or age of individuals. The latter result suggests that frequencies of PEPA-2 alleles in samples of king mackerel may stem, in part, from sex and age distributions of individuals within samples, and indicates that caution should be exercised in using allelic variation at PEPA-2 as a measure of population (stock) structure in king mackerel. The discordance in spatial patterning of mtDNA haplotypes versus PEPA-2 alleles across the Gulf (i.e. homogeneity in mtDNA haplotype frequencies versus heterogeneity in PEPA-2 allele frequencies) may be due to either female excess at several localities, sex-biased migration, or both. Observed patterns of genetic variation also are consistent with the hypothesis that king mackerel in the western Atlantic may have been subdivided during Pleistocene glaciation, and that the current distribution of PEPA-2 alleles may be a historical artefact. Received: 17 December 1996 / Accepted: 2 April 1997     

Application of the Red List Index for conservation assessment of Spanish vascular plants

The International Union for Conservation of Nature (IUCN) Red List Index (RLI) is used to measure trends in extinction risk of species over time. The development of 2 red lists for Spanish vascular flora during the past decade allowed us to apply the IUCN RLI to vascular plants in an area belonging to a global biodiversity hotspot. We used the Spanish Red Lists from 2000 and 2010 to assess changes in level of threat at a national scale and at the subnational scales of Canary Islands, Balearic Islands, and peninsular Spain. We assigned retrospective IUCN categories of threat to 98 species included in the Spanish Red List of 2010 but absent in the Spanish Red List of 2000. In addition, we tested the effect of different random and taxonomic and spatial Spanish samples on the overall RLI value. From 2000 to 2010, the IUCN categories of 768 species changed (10% of Spanish flora), mainly due to improved knowledge (63%), modifications in IUCN criteria (14%), and changes in threat status (12%). All measured national and subnational RLI values decreased during this period, indicating a general decline in the conservation status of the Spanish vascular flora. The Canarian RLI value (0.84) was the lowest, although the fastest deterioration in conservation status occurred on peninsular Spain (from 0.93 in 2000 to 0.92 in 2010). The RLI values based on subsamples of the Spanish Red List were not representative of RLI values for the entire country, which would discourage the use of small areas or small taxonomic samples to assess general trends in the endangerment of national biotas. The role of the RLI in monitoring of changes in biodiversity at the global and regional scales needs further reassessment because additional areas and taxa are necessary to determine whether the index is sufficiently sensitive for use in assessing temporal changes in species’ risk of extinction.     

Assessment of the nutritional condition of larval and early juvenile tuna and Spanish mackerel (Pisces: Scombridae) in the Panamá Bight

The nutritional condition of first-feeding and late larval/early juvenile scombrids was investigated in waters of the northwestern Panamá Bight from May through early November 1988. Wild-caught larvae and juveniles of three taxa, black skipjack tuna (Euthynnus lineatus), bullet and/or frigate tuna (Auxis spp.) and sierra (Scomberomorus sierra), were examined histologically to determine nutritional condition. The incidence of malnourishment in wild-caught preflexion (first feeding—prior to notochord flexion) larvae of all taxa was high. Starvation rates for E. lineatus and Auxis spp. preflexion larvae ranged from 62 to 63% d^-1, while the percentage of larvae actually dying of starvation was estimated at 41 to 43% d^-1. The nutritional point-of-no-return for preflexion larvae was estimated at 1 to 2 d maximum. The cellular condition of liver hepatocytes, particularly the relative amount of vacuolation related to storage of glycogen and lipid, proved to be a sensitive indicator of nutritional condition. In laboratory trials, late larval (postflexion) and early juvenile black skipjack exhibited a nutritional point-of-no-return of 2 to 3 d. Although postflexion larvae were moderately vulnerable to malnourishment in laboratory trials, <13% of wild-caught postflexion larvae exhibited even mild nutritional stress, and no postflexion larvae or juveniles showed signs of severe malnourishment. This pattern of starvation incidence suggests that tropical scombrids undergo stagespecific starvation mortality. Preflexion larvae can suffer significant daily losses due to starvation, while postflexion larvae and early juveniles seem to experience a rapid improvement in feeding ability and/or food availability.     

Branchial Na+−K+-ATPase activity during osmotic adjustments in two freshwater euryhaline teleosts,tilapia (Sarotherodon mossambicus) and orange chromid (Etroplus maculatus)

Effects of osmotic stress on the branchial Na⁺–K⁺-ATPase activity in two freshwater euryhaline teleosts (the tilapia Sarotherodon mossambicus Peters and the orange chromid Etroplus maculatus Bleeker) were studied. Direct transfer from fresh water to salt water with a salinity of 35 S caused extensive mortality in tilapia. Of the remaining four saltwater concentrations (4.375, 8.75, 17.5, and 26.25 S), the higher two increased the enzyme activity significantly in the first week without affecting it further during the remaining seven weeks of acclimation. The lower two concentrations failed to elicit any significant change. In the case of orange chromid, all three salt-water concentrations (4.375, 8.75, and 17.5 S) which the fish survived produced a more significant and extensive change in the enzyme activity, which continued to rise during the entire acclimation period. Employing a procedure of gradual transfer, tilapia and orange chromid were successfully adapted to saltwater concentrations of up to 61.25 and 35 S, respectively. In this experiment, almost each successive increase in salinity elevated the enzyme activity further, but the total change in both species was considerably less extensive than might be expected from the direct transfer study. When the freshwater fish were transferred to identical concentrations of pure NaCl or whole ocean salt, the former produced a relatively greater change in the enzyme activity in tilapia. No such difference was seen in orange chromid. During reacclimation of the saltwater-adapted fish to fresh water, the enzyme activity began to decrease immediately in both species, but remained above the freshwater control levels at the end of eight weeks.     

Some mechanisms of salinity acclimation in the euryhaline teleost,Etroplus maculatus

Mechanisms of salinity acclimation in the euryhaline freshwater living teleost, Etroplus maculatus (Bleeker), were studied. Brain water content decreased steadily with increasing salinity of the medium. Liver and muscle water content increased initially in 30% sea-water but in higher salinities decreased markedly, approximating control levels. Osmotic pressure of tissue fluid increased by 33% from 0.45% NaCl equivalent in freshwater to 0.60% NaCl equivalent in 100% sea-water. K and Mg decreased in dilute saline media but in 100% sea-water increased markedly approximating control levels; this is especially so in regard to Mg. Na decreased steadily and Ca increased with increasing salinity. Cl increased in 60% sea-water but decreased in 100% sea-water; however, its concentration in 100% sea-water exceeded that in freshwater. Free amíno acids increased by 11% in 30% sea-water but decreased in 60% and 100% sea-water, such that their concentration was lower by 28% in 60% sea-water and by 34% in 100% sea-water relative to the control level. Ascorbic acid decreased by 37% in 30% sea-water but in 60% and 100% sea-water increased steadily approximating the control level. The significance of these changes in the physiology of salinity acclimation is discussed and a correlation suggested between acclimation to salinity and to high temperature.     

Aspects of osmotic and lonic regulation in two Baltic teleosts: Effects of salinity on blood and urine composition

Hydromineral regulation was studied by examining the response to different environmental salinities in two Baltic brackish-water (BW) teleosts—a species of marine ancestry, Myoxocephalus scorpius (L.), and a glacial relict, M. quadricornis (L.). M. scorpius tolerated fresh water (FW) and M. quadricornis sea water (SW) for only about 24 h, but the survival time of M. scorpius in SW and M. quadricornis in FW was one to several weeks. M. scorpius seems able to balance plasma ionic concentrations in salinities down to about 2 to 3. Death of M. scorpius in FW was associated with partial haemolysis, increased volume of red blood cells (RBC), increased plasma K⁺ concentration, and decreased concentrations of Na⁺, Cl^- and Mg²⁺ in plasma and, to a lesser extent, in urine. Death of M. quadricornis in SW was associated with increased plasma osmolality, and Na⁺, Cl^- and Mg²⁺ concentrations, but the renal excretion of ions approached that generally found in marine teleosts. In most cases, RBC volume followed the changes in plasma osmolality or Na⁺ and Cl^- concentrations. Both species showed an ability to increase tubular Mg²⁺ secretion much over that needed in BW, and increased secretion was associated with high urine Cl^- concentration. M. quadricornis, but not M. scorpius, reabsorbed Na⁺ effectively in SW also. Differences between Oceanic and Baltic specimens of M. scorpius are discussed.     

Effect of low dissolved oxygen concentrations on behavior and predation rates on red sea bream Pagrus major larvae by the jellyfish Aurelia aurita and by juvenile Spanish mackerel Scomberomorus niphonius

A shift in outcomes of predator-prey interactions in plankton community may occur at sublethal dissolved oxygen concentrations that commonly occur in coastal waters. Laboratory experiments were conducted to investigate how a decline in dissolved oxygen concentration alters the predation rate on fish larvae by two estuarine predators. Behavior and consumption of larval fish by moon jellyfish Aurelia aurita (103.1±12.4 mm in bell diameter) and by a juvenile piscivore, Spanish mackerel Scomberomorus niphonius (30.1±2.1 mm in standard length: SL), were observed under four oxygen concentration treatments (1, 2 and 4 mg l^–1 and air-saturated: 5.8 mg l^–1). Larvae of a coastal marine fish species, red sea bream Pagrus major (7.21±0.52 mm SL), were used as prey for the experiment. Bell contraction rate of the jellyfish did not vary among the oxygen concentrations tested, indicating a tolerance to low oxygen concentration. Gill ventilation rate of the Spanish mackerel increased and swimming speed decreased as the oxygen concentration decreased, indicating that oxygen concentrations 4 mg l^–1 are physiologically stressful for this species. The number of larvae consumed in 15 min. by jellyfish increased whereas those consumed by Spanish mackerel decreased with the decrease in oxygen concentration. Low oxygen concentrations that are commonly observed in coastal waters of Japan during summer have the potential to increase the relative importance of jellyfish as predator of fish larvae and to change the importance of alternative trophic pathways in estuarine ecosystems.Communicated by T. Ikeda, Hakodate     

Adaptations to osmotic stress in the fresh-water euryhaline teleost Tilapia mossambica. IV. Changes in blood glucose,liver glycogen and muscle glycogen levels

Some aspects of osmoregulation energetics have been studied in the euryhaline teleost Tilapia mossambica (Peters) acclimated to media of different salinities. In stress media (75 and 100% sea water) the blood glucose of the fish increases significantly, accompanied by a corresponding increase in oxygen consumption and cytochrome-oxidase activity, suggesting that oxidative degradation of blood glucose is the predominant energy source for osmotic work in these stress media. It is likely that the variations in the blood-glucose level as a function of acclimation to the heterosmotic media —except the natural fresh-water medium — are governed by the combined effects of salinity of the medium and blood-medium osmotic gradient, rather than by the effect of any one of them separately. Perhaps, metabolic homoeostasis is in operation in the natural fresh-water medium. Depletion of muscle glycogen at significant levels is noticed only in stress media. Presumably, there is an augmentation of oxidative metabolism with glycolysis to meet the exacting energy demands for heavy osmotic work in high-stress media. Prior acclimation to 75% sea water (24.375 S) facilitates subsequent acclimation to 100% sea water (32.50 S) with less energy cost —an instance of facilitation acclimation. Smaller individuals of T. mossambica osmoregulate with less energy expenditure than larger ones. Thus, smaller individuals are osmotically more efficient.     

Effects of fluctuating salinity on the behaviour of the West African blood clam Anadara senilis and on the osmotic pressure and lonic concentrations of the haemolymph

Specimens of the West African blood clam Anadara senilis were exposed to both sinusoidal and abrupt salinity regimes. Measurements showed that haemolymph osmolality and concentrations of Na, K⁺, Ca²⁺ and Mg²⁺ followed the concentrations of the external medium up to the time of shell valve closure. Shell valves closed when the seawater concentration had fallen to about 15.4% S and reopened at a similar salinity. The closure mechanism was effective in preventing excessive haemolymph dilution.     

Changes in the spatial distribution of spawning activity by north-east Atlantic mackerel in warming seas: 1977–2010

Migratory species with a broad geographic range, such as north-east Atlantic mackerel, may be amongst the fauna most able to respond to warming seas, typically with a poleward shift in range. Habitat heterogeneity could, however, produce more complex patterns than a simple polewards translation in distribution. We tested for changes in the central location and spatial spread of mackerel spawning over a 33-year period. Spatial statistics [centre of gravity (CoG) of egg production, spatial variance, and degree of anisotropy] were used to summarise interannual changes in the spawning locations of the western spawning stock of north-east Atlantic mackerel (NEA-WSC) using data from the ICES triennial egg survey. A northwards shift in CoG of egg production estimates was observed, related to both an expansion in the distribution in survey effort and warming waters of the north-east Atlantic. Sea surface temperature (SST) had a significant positive association with the observed northward movement of NEA-WSC mackerel, equivalent to a displacement of 37.7 km °C^?1 (based on spring mean SST for the region). The spatial distribution of spawning around the CoG also changed significantly with SST, with a less elongated spatial spread in warm years. An increase in the proportion of spawning over the Porcupine Bank demonstrated how habitat interacts with positional shifts to affect how north-east Atlantic mackerel are distributed around the centre of their spawning range.     

Abundance of small cetaceans in waters of the central Spanish Mediterranean

Seasonal aerial surveys were conducted in the waters of the central Spanish Mediterranean from 2001 to 2003 using the line transect sampling methodology to estimate cetacean abundance. The density of the three most abundant species, striped dolphin (Stenella coeruleoalba), bottlenose dolphin (Tursiops truncatus) and Risso’s dolphins (Grampus griseus), was estimated. In the case of the first two species, the density was estimated accounting for the proportion of submerged animals, while for Risso’s dolphin only the surface density could be estimated. The striped dolphin was the most abundant species in the study area with a mean density of 0.489 dolphins km⁻² (95% CI = 0.339–0.705) and a mean abundance of 15,778 dolphins (95% CI = 10,940–22,756). This density is comparable to that obtained in the International Ligurian Sea Cetacean Sanctuary. Striped dolphins were observed throughout the whole year and no seasonal changes in the density were detected. The mean density of bottlenose dolphins was an order of magnitude lower than that of striped dolphins (0.041 dolphins km⁻²; 95% CI = 0.023–0.075) with a mean abundance of 1,333 dolphins (95% CI = 739–2,407). The Risso’s dolphin had a surface estimated density of 0.015 dolphins km⁻² (95% CI = 0.005–0.046) and a mean abundance of 493 dolphins (95% CI = 162–1,498). These results provide valuable biological information useful to develop conservation plans and establish a baseline for future population trend studies.     

Day/night differences in determination of growth rate of mackerel Scomber scombrus larvae during a patch-study in the Celtic Sea

The growth rate of mackerel, Scomber scombrus, larvae was calculated by analysing mean cohort length over time. The study took place on the central spawning ground of mackcrel in the Celtic Sea (Great Sole Bank) in April 1986. In order to minimize horizontal catch variability, repeated small-scale sampling with the Mocness plankton net was undertaken close to a satellite-tracked drifting buoy during a period of 78 h. Eighteen samples from nine hauls distributed over the whole sampling period were considered. One hundred larvae per sample were measured using a semi-automatic method. The proportion of larger larvae was significantly higher in night catches than in day catches. Independently calculated growth rates were 0.18 mm d^-1 for day catches and 0.14 mm d^-1 for night catches. This difference was not statistically significant. However, confidence in the calculation of growth rate, computed from night sampling, was much higher than that for day sampling. This may result as an effect of avoidance behaviour by larvae in daylight. Thus, night sampling seems to be a more reliable tool in the study of larval growth and should be used more extensively than daylight sampling.This paper was presented at the Early life history of fish symposium, Bergen, October 3 to 5, 1988     

Northward shifts of the distributions of Spanish reptiles in association with climate change

It is predicted that climate change will drive extinctions of some reptiles and that the number of these extinctions will depend on whether reptiles are able to change their distribution. Whether the latitudinal distribution of reptiles may change in response to increases in temperature is unknown. We used data on reptile distributions collected during the 20th century to analyze whether changes in the distributions of reptiles in Spain are associated with increases in temperature. We controlled for biases in sampling effort and found a mean, statistically significant, northward shift of the northern extent of reptile distributions of about 15.2 km from 1940-1975 to 1991-2005. The southern extent of the distributions did not change significantly. Thus, our results suggest that the latitudinal distributions of reptiles may be changing in response to climate change.     

Growth-selective survival of young jack mackerel Trachurus japonicus during transition from pelagic to demersal habitats in the East China Sea

To examine the growth-selective survival in early life stages of jack mackerel from the East China Sea, we compared growth rates of young fish collected from the surface layer in April with those of survivors collected from the near-bottom layer in May?CJune. We performed this analysis from 3?year classes with different levels of relative abundance: low in 2005, intermediate in 2008 and high in 2009. Growth rates during the larval stage of the original population were slower than the survivors in all survey years, indicating that faster-growing individuals in the surface layer had a higher probability of survival to the near-bottom habitat than slower-growing individuals. Selection intensity, defined as differences in growth rates between the two groups, was higher in year classes with a lower abundance. Hence, growth rates during the early life stages of jack mackerel are considered to be a critical determinant of successful survival to the demersal habitat.     

Reproductive patterns of scleractinian corals in the northern Red Sea

The majority of published accounts on scleractinian coral reproduction are from the tropical Pacific and Caribbean, with very little information known about Red Sea species. This report examines variation in reproductive mode in 24 species of hermatypic corals (belonging to seven families) in the Gulf of Eilat, Red Sea. Eighteen species are hermaphroditic broadcasters, two are hermaphroditic brooders and three are gonochoric broadcasters. In the Pocilloporidae, the gonads project into the body cavity, while in the other six families the gonads reside inside the mesenteries. The number of gonads per polyp in broadcasting species follows family or genus lines. Fecundity (eggs per polyp) increases with polyp size. Brooding species usually exhibit one or two gonads per polyp and each gonad contains only one to three oocytes. Oocyte size varies widely and does not relate to mode of reproduction. The largest oocytes (diameter = 450 μm) occur in the brooding coral Alveopora daedalea and in broadcasting species of the genus Acropora (diameter = 420 μm). Gonad morphology and gonochorism versus hermaphroditism appear to be constrained phylogenetically at the family or genus level. Lastly, this report compares the data presented for Red Sea scleractinian species with the data available on scleractinian corals from other geographical regions. Received: 2 February 1993 / Accepted: 9 March 1998     

Prey selection by the hogchoker, Trinectes maculatus (Pisces: Soleidae), along summer salinity gradients in Chesapeake Bay, USA

We investigated feeding by the hogchoker, Trinectes maculatus (Bloch and Schneider), in freshwater, oligohaline, mesohaline, and polyhaline regions of Chesapeake Bay, USA, and examined prey selection in relation to food availability. Otter trawling for fish and Van Veen grab sampling for benthic macrofauna occurred in July and August 1992 and August and September 1993. Hogchokers exhibited both opportunistic and selective feeding patterns along the estuarine salinity gradient in four tributaries (Potomac, Rappahannock, York, and James Rivers) and in the mainstem Chesapeake Bay. Major prey taxa included annelids, arthropods, and tellinid siphons. In polyhaline habitat, polychaetes dominated both the benthos and gut contents numerically and gravimetrically. On the other hand, oligochaetes were numerically dominant in freshwater/oligohaline areas but were rarely eaten, perhaps because of their burial depth. Arthropods (mostly amphipods) occurred at most salinities, were common in gut contents in low-salinity areas, and were replaced as prey by larger proportions of polychaetes in polyhaline regimes. Although hogchokers ate tellinid siphons, they rarely consumed whole bivalves or gastropods. These diet patterns (and especially the importance of siphon nipping) are similar to those of juvenile or small flatfish elsewhere in Europe, Africa, and North America. A size–salinity relationship for hogchokers occurred along the summer salinity gradient, with smaller fish predominating upstream and larger fish downstream. It was not clear from our data if variation in diet composition reflected changes in prey composition along the salinity gradient rather than changes in fish size. Received: 14 June 1997 / Accepted: 27 June 1997     

Conditional sex allocation in the Red Mason bee, Osmia rufa

Trivers' and Willard's hypothesis that natural selection favors sex allocation in relation to maternal condition assumes iteropary. Though this assumption is not met in most solitary Aculeata, the reproductive life span of semelparous females may be divided into discrete successive cycles by the risk of open-cell parasitism. Females can avoid losing their investment to parasites attacking the open cell only by limiting the provision time for each cell. The restriction of time available for the investment in a single progeny irrespective of the condition of the female leads to de facto iteropary. Moreover, in Hymenoptera, there are no costs for sex allocation due to the haplodiploid mode of sex determination. In sexually size dimorphic species, females in poor condition are predicted to invest in the smaller sex and vice versa. The resulting prediction of a conditional sex allocation in solitary Aculeata was tested in the Red Mason bee, Osmia rufa (Osmia bicornis), a stem or hole-nesting, polylectic, univoltine megachilid bee. Body size is a key component of condition in females of nest-constructing solitary bees. Large females collect the same amount of pollen and nectar in a shorter time than small ones and should suffer less from parasitism. We found that small females dealt with their handicap of a low provisioning performance by shifting the sex ratio toward sons (the smaller sex) and by reducing the body size of daughters. Large females, however, shifted their offspring sex ratio toward daughters, the sex that depends more on body size in its reproductive value. The sex ratio in the population met the expected Fisherian sex ratio. Although females allocated their investment in the sexes according to their body mass, the population-level investment was balanced.     

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.