Larval development of Euphausia nana (Crustacea: Euphausiacea)

The larvae of Euphausia nana Brinton in Sagami Bay and Suruga Bay, Central Japan, are described and illustrated. The furcilia stage is separated into six stages on the basis of the form of the pleopods and the number of terminal telson spines. Furcilia I, no pleopod or non-setose pleopods and seven terminal telson spines. Furcilia II, nonsetose and setose pleopods and seven terminal telson spines. Furcilia III, five pairs of setose pleopods and seven terminal telson spines. Furcilia IV, five pairs of setose pleopods and five terminal telson spines. Furcilia V, five pairs of setose pleopods and three terminal telson spines. Furcilia VI, five pairs of setose pleopods and one terminal telson spine. Segmentation of the antennal endopod and mandible palp occurred at the furcilia V stage. The dominant forms of early furcilia larvae of E. nana differed during different seasons of the year. The total lengths of calyptopis III and furcilia I to VI larvae also showed seasonal changes; they were largest in May and smallest from September to December.     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

Influence of variability in larval development on recruitment success in the euphausiid Euphausia pacifica : elasticity and sensitivity analyses

Simple life-history models were employed to identify those larval stages of Euphausia pacifica (Hansen) with the greatest potential contribution to variability in recruitment success. Using stage-specific vital parameters (i.e. stage duration and specific mortality) the models explored the influence of developmental variability without ascribing the variability to any particular cause or forcing. Through such a non-mechanistic modelling approach, the critical periods in larval development were determined. The results led to inferences about larval population responses to temperature/food heterogeneity in the natural environment. Peaks in recruitment-success elasticity were evident for variability imposed during the egg and Furcilia I and II stages. The high elasticity of the egg stage suggested the importance of female nutrition and the timing and location of egg release in determining their viability and subsequent recruitment success. Peak elasticity during the Furcilia I and II stages supported the notion that indirect developmental pathways of pleopod development during these stages may be associated with diminished recruitment success. Sensitivity analyses further highlighted the influence of these early furcilia stages in determining recruitment success, and also suggested the importance of the Furcilia III and VII stages. The robustness of elasticity and sensitivity results with respect to the life-history structure utilized is discussed. A criterion is presented for the identification of life-history structures which might obscure the true elasticity relationships of a developmental sequence. Received: 27 April 1998 / Accepted: 14 September 1998     

Energetics of Euphausia pacifica. II. Complete carbon and nitrogen budgets at 8° and 12° C throughout the life span

Complete carbon and nitrogen budgets at 8° and 12° C over the life span of Euphausia pacifica were constructed from data published in the foregoing publication. The relative magnitudes of physiological functions such as metabolism and growth for the different life history stages were compared. The carbon net growth efficiency (NGE) for E. pacifica increased to a maximum of 60 to 74% at Calyptopis 3 (a larval stage) and then decreased rapidly to a level of 10 to 12% for adults. The cumulative amount of assimilated carbon or nitrogen in reproductive products was equal to that in growth. The cumulative net production efficiency (tissue plus molts plus reproductive products) is 24 to 29%, and is slightly higher for nitrogen than for carbon. For juveniles and adults, the largest proportion (40 to 65%) of assimilated material is used in metabolism, and is about 10% higher at 8° than 12° C. The difference between measured and predicted ingestion (the sum of metabolism, leakage, defecation, growth, molting and reproduction) probably results from poorly understood aspects of zooplankton physiology (e.g. the effect of body weight on leakage). Nitrogen budgets at 8° C for furcilia bulanced the best, with deviations of less than 10%. In general, predicted ingestion was increasingly less than measured ingestion as body weight and temperature increased. Possible omissions or errors in assumptions or methods that may cause the imbalances are discussed.     

Dominance of larval forms in euphausiid (Crustacea:Eucarida) ontogenesis

Variability and dominance of Euphausia superba Dana larval forms (Furcilia-I and Furcilia-II stages) in the plankton of the Scotia and Weddell Seas have been investigated. New forms (1, 2, 3 and 14) have been recorded. The dominant forms of Furcilia-I stage are proposed to represent the initial dominants. In accordance with the initial dominance, five types of larval form combinations have been established. During the initial dominance change — from 5 to 2, the larval form number increases. This phenomenon is due to parallel development of euphauiid larvae; it results from a developmental pathway change in the main larval mass. Such deviations are reflected in an initial dominance change. In each pathway, the larvae pass through an unequal number of moults before reaching Form 5 (in accordance with the proposed moult scheme of Furcilia-I stage larvae). In the case of initial dominance of less-developed forms, the presence of different parallel pathways increases. As a result, the total number of larval forms present in the plankton changes, causing increased variability of the total number of larval forms. Each type of larval form combination is the result of summation of all larval forms, which derive from different developmental pathways of the larvae in different localities. The most usual developmental pathway for Euphausia superba is through Form 5, which may, thus, be regarded as the main developmental pathway for this species. This pathway probably corresponds to optimal environment conditions for larval growth and morphogenesis with maximal synchronisation of both processes. Consequently, it may be suggested that changes in developmental pathways of larvae are connected with suboptimal environmental conditions.     

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.     

Diet of calyptopes of the euphausiidEuphausia pacifica in the Yellow Sea

Foregut contents of calyptopes (the first feeding stage) ofEuphausia pacifica Hansen, collected from the Yellow Sea in June 1989, were examined using scanning electron microscopy with the dry-fracturing method. There was no clear evidence of phytoplankton ingestion by calyptopes. The foregut contents consisted entirely of amorphous organic detritus, crustacean remains and inorganic particles. The inorganic particles adhered to amorphous organic detritus, and bacterial populations were found on peritrophic membranes of fecal pellets which enveloped crustacean remains. These materials seemed to have originated from aggregates of organic and inorganic detritus. It is quite possible that the detrital aggregates are utilized extensively as food by calyptopes ofE. pacifica in the Yellow Sea.     

Recruitment,growth, and diel vertical migration of Euphausia pacifica in a temperate fjord

The pelagic crustacean Euphausia pacifica Hansen was sampled with a multiple-sample 1.0 m² Tucker trawl and a multiple-sample 1.0 m² vertical net in Dabob Bay, Washington on 17 dates between May 1985 and October 1987. Size (stage) structure and abundance of the population were determined for each date, while vertical distribution and diel migration were determined for 13 dates. Although internannual variability in both timing and magnitude of events occurred, consistent patterns were discernable. The population produced a large pulse of larvae (2 to 5 mm) in late spring of each year, apparently in response to the vernal phytoplankton bloom. Much lower abundances of larvae occurred during summer and autumn of each year, and larvae were completely absent during winter. Recruitment to the juvenile (6 to 9 mm) and adult (10 mm) stages was strongest during the summer, with abundances of these individuals peaking in summer and autumn. Individual growth rates, determined by modal progression analysis, were calculated for E. pacifica. Rates ranged from zero for some adult cohorts during the winter to 0.12 mm d^-1 for larvae during spring. The latter are among the highest ever reported for this species in the field. The vertical distributions and diel vertical migrations (DVM) of E. pacifica varied seasonally and between size (stage) classes. At night, all size classes were distributed in the surface layer (upper 25 m) irrespective of season or year. During the day, the larger/older stages were always distributed at middepths (50 to 125 m). In contrast, the daytime distribution of the larvae was more variable, being concentrated at the surface during spring and early summer of 1985, and at increasing depths later in the summer and autumn of 1985 and again in spring of 1986. This resulted in invariant DVM in the juveniles and adults, but variable DVM in the larvae, the latter of which is hypothesized to be a response to variable abundances of zooplanktivorous fish.     

Effects of water-body characteristics other than temperature and salinity on respiration in Euphausia pacifica

Ample evidence exists that many planktonic species are limited in their distribution to a given water mass. It has been suggested that the temperature-salinity values which define a given water mass are in themselves limiting these species' distribution. Other workers have suggested that other unique properties of a water mass may be involved. Euphausia pacifica is distributed throughout the Pacific subarctic water mass; its distribution also extends into British Columbia coastal waters. Experiments have shown that E. pacifica can withstand temperature changes of up to 15 C°; it can also withstand salinity changes of up to 10. Coastal E. pacifica appear to be able to better withstand stress from changes in temperature and salinity in water from some sources than in water from others. These differences are thought to result from differences in the past history of the water in question.Contribution No. 75008 from the Bigelow Laboratory for Ocean Sciences.     

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.     

Fouling organisms in the port of Mar del Plata (Argentina). I. Siphonaria lessoni: ecological and biometric aspects

The present paper deals with ecological aspects of the pulmonate gastropod Siphonaria lessoni (Blainville, 1824), a common member of fouling communities in Mar del Plata's port (38°0817 S, 57°3118 W). Populations on colonized panels of an experimental raft were studied during three annual cycles, to determine feeding habits, growth and various factors which restrict their distribution on floating substrata. Although S. lessoni is harmless as a fouling organism, its grazing on the algal belt clears a part of the substratum's surface, which is then quickly colonized by truly aggressive organisms such as Balanus amphitrite.     

Diurnal migration and vertical distribution of phyllosoma larvae of the western rock lobster Panulirus cygnus

Studies off the west coast of Australia showed that the phyllosoma larvae of Panulirus cygnus George undergo a diurnal vertical migration, with light as an important factor influencing the depth distribution of all 9 phyllosoma stages. The early stages (I to III) occurred at the surface at night regardless of moonlight intensity, whereas late stages (VI to IX) concentrated at the surface only on nights with less than 5% of full moonlight. Midday peak densities of early-stage larvae occurred in the 30 to 60 m depth range while those of mid (IV to VI) and late stages were in the 50 to 120 m range. Depths of peak densities of larvae increased with distance offshore. The limits of vertical distribution of the phyllosoma remained within ranges of illuminance which were estimated to be in the order of 50 to 250 E m^-2 sec^-1 for early stages, 20 to 200 E m^-2 sec^-1 for mid stages and 5 to 50 E m^-2 sec^-1 for late stages. Minimal rates of net vertical movement were estimated for the larvae. Early stages exhibited mean net rates of ascent and descent of 13.7 and 13.0 m h^-1, respectively, while the rates for mid stages were 16.0 and 16.6 m h^-1 and for late stages 19.4 and 20.1 m h^-1. Diurnal migrations and vertical distribution are shown to have a vital role in the relationships between circulation in the south-eastern Indian Ocean and the transport and dispersal of the phyllosoma larvae. The diurnal migrations of early stages place them at the surface at night, when offshore vectors of wind-driven ocean-surface transport dominate, and below the depth of wind-induced transport during the day, when offshore vectors are small or negative, thus accounting for their offshore displacement. Mid and late stages, because of their deeper daytime distribution and absence from the surface on moonlight nights, are predominantly subject to circulation features underlying the immediate surface layer. This is hypothesized to account for the return of the phyllosoma to areas near the continental shelf edge by subjecting them to a coastward mass transport of water which underlies the immediate surface layer.The western rock lobster is referred to as P. longipes or P. longipes cygnus in some of the literature quoted; these are synonymous with P. cygnus.     

Growth and spatial distribution of Cassidulus mitis (Echinodermata: Echinoidea) on a sandy beach in Southeastern Brazil

Using a hand dredge, a population of Cassidulus mitis Krau, 1954 was sampled monthly from February 1986 to January 1987 at 15 stations offshore from Vermelha Beach (22°5718 S; 43°948 W), in water depths ranging from 0 to 6 m. The sex ratio indicated a slight numerical dominance of females for adults and a larger numerical female dominance for juveniles. A von Bertalanffy growth curve was estimated as L=71.00 (1-e^(-0.390t)), where L=length (mm) and t=time (yr). The instantaneous natural mortality rate (M) was estimated as 0.812. The population occupies a zone parallel to the beach, and displays a spatially aggregated pattern, with higher densities in areas characterized by coarse and very well sorted sand containing low amounts of organic matter.     

Relationship of oxygen consumption to swimming speed in Euphausia pacifica

Swimming efficiency (the ratio of thrust power required to overcome hydrodynamic drag to net metabolic energy expenditure) was calculated for the vertically migrating euphausiid Euphausia pacifica swimming at speeds of 1–20 cm s^–1 and at temperatures of 8° and 12°C. Efficiencies ranged from 0.014 to 2.8% at 8°C and 0.009 to 1.69% at 12°C. A comparison with efficiency in fishes 2–3 orders of magnitude larger in weight (efficiency range 10–25%) indicates that locomotion in E. pacifica is far less efficient, a probable result of the organism's small size (x=33.5 mg WW) and multiple-paddle mode of propulsion. Net cost of transport of E. pacifica is three to six times the cost of a hypothetical value for sockeye salmon. Low swimming efficiencies in zooplankton such as E. pacifica are responsible for the underestimation of zooplankton swimming costs. Multiple-paddle propulsion is less efficient than the undulatory mode of fishes.     

Comparison of three techniques for administering radiolabeled substrates to sediments for trophic studies: uptake of label by harpacticoid copepods

Microbial grazing by two species of meiofaunal harpacticoid copepods (Heteropsyllus nunni and Thompsonula hyaenae) was determined by uptake of radioactive labels following their introduction into natural sediments from a low-energy intertidal site in Florida (29°5440N; 84°3130W) in May 1986. Grazing was related to three methods of radioactive-label introduction: injection, porewater replacement, and slurry. Uptake of label by harpacticoids was examined using two dual-label combinations, ³H-thymidine/¹⁴C-bicarbonate and ³H-thymidine/¹⁴C-acetate. The injection and porewater-replacement methods yielded statistically indistinguishable results. Results obtained by the slurry method differed significantly from the other two methods. We suggest that the unique results obtained in the slurry method were a consequence of the disruption of microbial-meiofaunal spatial relationships.     

Population structure and life cycle of the mesopelagic ostracod Conchoecia pseudodiscophora in Toyama Bay,southern Japan Sea

The population structure and life cycle of the mesopelagic ostracod Conchoecia pseudodiscophora Rudjakov in Toyama Bay, southern Japan Sea, were investigated using a time-series of samples collected during 0 to 500 m vertical hausls with twin-type Norpac nets (0.35 and 0.10 mm mesh) over one full year (1 February 1990 to 30 January 1991). Additional samples were also collected with a single-type closing Norpac net (0.06 mm mesh) to examine the vertical distribution patterns of eggs and all instars of this species. The proportion of gravid females present indicated that reproduction of C. pseudodiscophora continues throughout the year, but peaks in April–July. Eggs and Instars I and II were distributed below 500 m, while the more advanced instars were most abundant in the 350 to 500 m stratum by both day and night. Based on the abundance peaks of each instar in the time-series samples, development times were estimated to be 2.5, 4, 3, 7 and 11 mo for Instars III, IV, V, VI, and VII, respectively. Thus, a total of 30 mo is required for newly spawned eggs to hatch and reach adulthood. Stomach-fullness indexes revealed no seasonality in the feeding activity of any instar stage, but that feeding activity was low in older instars, particularly in adult males. The present results are compared with those for a few other ostracod species, in an attempt to characterize the life cycle of C. pseudodiscophora inhabiting waters of subzero temperature in the mesopelagic zone of the Japan Sea.     

Population genetics of the northern Pacific seastar Asterias amurensis (Echinodermata: Asteriidae): allozyme differentiation among Japanese,Russian, and recently introduced Tasmanian populations

The northern Pacific seastar Asterias amurensis Lütken was recently introduced to Tasmanian waters, possibly through ballast water discharged from ocean-going vessels. Three Tasmanian populations and seven native populations from Japan and eastern Russia were examined in 1994 for variation at 22 allozyme loci. There was significant spatial heterogeneity in gene frequencies among the native populations, especially for the ocus APK ^*, where G _ST was a very high 0.47. The Tasmanian populations could not be separated genetically from one another, suggesting they have a common origin from a single introduction. The average heterozygosity per locus of the Tasmanian populations (0.116 to 0.127, mean 0.123) was 30 to 40% less than that of the native populations (0.177 to 0.216, mean 0.192), suggesting that colonisation of Tasmanian waters was accompanied by a population-size bottle-neck. The Tasmanian seastars were genetically more closely related to the two populations from central Japan (Suruga and Tokyo Bays) than to populations from Vladivostok, northern Japan (Yoichi, Nemuro and Mutsu Bays) or southern Japan (Ariake Sea). However, there were significant differences between the populations from Tasmania and central Japan, which may be attributable to founder effects or may mean that the true source of the Tasmanian introduction has yet to be identified.     

Chitinolytic enzymes in the integument and midgut-gland of the shrimpPalaemon serratus during the moulting cycle

Chitinase andN-acetyl--D-glucosaminidase activity were quantified inPalaemon serratus (Pennant) integument and midgut gland during the moulting cycle. Studies were performed on specimens collected near Concarneau, France, in July 1989. The changes in specific activity are different in the two organs: in the midgut gland enzymatic activity is high throughout the whole moulting cycle with a weak peak at the early premoult Stage D1, whereas in the integument the activity of both enzymes is very low throughout post- and intermoult stages and rises only after premoult Stage D1. The highest specific activity is reached in D1 for chitinase and somewhat earlier (D1) forN-acetyl--D-glucosaminidase. The increase in specific chitinolytic activity coincides with an increase in ecdysteroids.     

Receptor neurones in three heliothine moths responding to floral nd inducible plant volatiles

Summary. Some plant volatiles are produced in response to herbivory of several insect species, including heliothine larvae. In the present study of female heliothine moths, four co-located receptor neurone types were identified, of which three types responded strongest to the inducible compounds E--ocimene, E,E--farnesene and E,E-TMTT, respectively. The fourth type responded strongest to geraniol, which is a common floral volatile. The narrow tuning of each receptor neurone type was demonstrated by responses to a few structurally-related monoterpenes, sesquiterpenes, homo-terpenes and monoterpene alcohols, respectively, out of hundreds of plant constituents tested. The four neurone types showed the same relation of spike amplitudes and ranking of effective compounds in the three heliothine species; the polyphagous Heliothis virescens and Helicoverpa armigera and the oligophagous Helicoverpa assulta. The results indicate the presence of functionally similar types of plant odour-receptor neurones in the three related species, and suggest conservation or reappearance of functionally similar olfactory receptors in related species, independent of the evolution of polyphagy and oligophagy.     

Avoiding predation: alarm responses of Caribbean sea urchins to simulated predation on conspecific and heterospecific sea urchins

Alarm responses to the extracts of conspecifics and hetero-specifics were measured for the Caribbean sea urchins Echinometra viridis, E. lucunter, Lytechinus variegatus, L. williamsi, Tripneustes ventricosus, Diadema antillarum, and Eucidaris tribuloides collected along the Caribbean coast (9°3314N; 78°5523W) during October 1984 and July–December 1985. Responses to seawater and extracts of the gnathostomate echinoid Clypeaster sybdepressus were used as controls. The intensity of the response resulting from exposure to sea-urchin extracts was measured by: (1) the percentage of individuals that responded by moving away from the extract and/or towards shelter, and (2) the mean distance moved. Echinometra viridis, E. lucunter, and L. williamsi responded to sea-urchin extracts by moving towards nearby shelter sites. The distance that individuals of each species moved in the first minute following exposure to conspecific extracts was correlated with the distance that species moved from shelter while foraging. L. variegatus and D. antillarum, living in microhabitats not providing protection from predators, responded to extracts of conspecifics and heterospecifics by moving away from the direction of the extract. Eucidaris tribuloides did not exhibit alarm responses to the extracts of con- or hetero-specifics. E. tribuloides secures itself with its stout spines into protected sites within corals. Similarly, L. variegatus living in long, dense seagrass that provided protection from detection by predators, and D. antillarum occupying crevices, showed no alarm responses to extracts of conspecifics. Presumably, in these situations, sea urchins cannot increase their defenses against predation by moving away from an injured neighbor. T. ventricosus showed a weak response to extracts of L. variegatus, but no response to extracts of other species including conspecifics. The reasons for this lack of an alarm response are unclear. For the five species that demonstrated an alarm response to sea-urchin extracts, the intensity of the response varied depending on the type of extract used. L. variegatus, L. williamsi, and D. antillarum responded most strongly to extracts from conspecifics, while Echinometra viridis and E. lucunter responded strongly to extracts from both conspecifics and congeners. The weakest responses were shown to the extracts of T. ventricosus and Eucidaris tribuloides. Habitat overlap, overlap in predators, and phylogenetic relationships did not consistently explain patterns of alarm responses to the extracts of heterospecific sea urchins.