Some aspects of the life history of the subarctic copepodNeocalanus cristatus (Calanoida) in Sagami Bay,central Japan

A typical subarctic copepod,Neocalanus cristatus, occurred in the mesopelagic layer (500 to 1000 m) in Sagami Bay, central Japan, throughout the year. Specimens were collected from 1982 to 1986. A small number of adult females were distributed from 800 to 900 m only, but no adult males were collected. This species appeared to be abundant in April and August, when intermediate Oyashio water flowed strongly into Sagami Bay. Mean prosome lengths of copepodite stages IV and V and adults were 4.33, 6.87 and 6.87 mm, respectively. The condition factor [wet wt/(prosome length)³ × 100] of copepodite stage V did not vary remarkably, and mean values ranged from 4.7 to 5.0. Prosome length, body weight and condition factor ofN. cristatus collected from Sagami Bay were smaller than those of copepods in the northern North Pacific.N. cristatus transported from the north cannot molt to adult stages (except for those originating in mesopelagic waters) due to the adverse environmental conditions in Sagami Bay; instead, they die in the mesopelagic layer and sink to the bathypelagic layer (1 000 to 1 400 m), close to the bottom. Since nauplii and early copepodite stages did not occur in any season,N. cristatus probably do not reproduce in Sagami Bay.     

Grazing rates of the copepods Calanus glacialis and C. finmarchicus in arctic waters of the Barents Sea

Natural feeding rates of Copepodite Stages IV and V, and adult female Calanus glacialis (Jaschnov) and Copepodite Stage V and adult female C. finmarchicus (Gunnerus) were estimated using fluorescence analysis of gut contents. Measurements were made on copepods sampled from arctic waters east of Svalbard (Barents Sea) during the spring phytoplankton increase, in the period from 27 May to 13 June, 1983. Observations on Copepodite Stages IV and V and adult female C. glacialis suggest that the gastric evacuation rate is independent of developmental stage, whereas C. finmarchicus Copepodite Stage V showed a lower gastric evacuation rate than adult females. Gut fullness displayed a low correlation with the ambient chlorophyll concentrations. Ingestion rates calculated for C. glacialis were 0.3, 2.3, and 11.0 g C h^-1 for Copepodite Stages IV and V and adult females, respectively. Copepodite Stage V and adult female C. finmarchicus ingested 0.9 and 1.1 g C h^-1 at a temperature of ca.-1.0°C. The maximum ingestion rate in terms of percent body carbon d^-1 was higher for adult female C. glacialis and C. finmarchicus than for the respective Copepodite Stage V's. The results are discussed both in relation to the physiological state of the species and to the environmental conditions.     

Vertical distribution and nocturnal migration of Nyctiphanes couchi (Crustacea: Euphausiacea) in relation to the summer thermocline in the Celtic Sea

Vertical distributions and nocturnal migrations of the developmental stages of Nyctiphanes couchi (Bell) in relation to the summer thermocline in the Celtic Sea, 25 to 26 August 1982, have been investigated using the Longhurst-Hardy Plankton Recorder (LHPR). The vertical distributions of the metanauplii and adult females suggest that N. couchi liberates its young within the euphotic zone as mature metanauplii which, in a matter of hours, moult into the first feeding stage (Calyptopis I). The ascent migration by adult females took a maximum of 3 h (17.10 to 20.05 hrs) and had an amplitude of 50m (54 to 4 m) from below to above the thermocline. A 7C° thermocline occurred between 20 to 30 m in these profiles. The nocturnal migrations by the females were for the purpose of breeding as well as feeding within the euphotic zone and were not influenced by the presence of the thermocline. The majority of the calyptopes and furciliae remained above the thermocline over the sampling period. The post-larval males and females migrated; their vertical distributions showed a pattern similar to those of the adult females. The larger the developmental stage, the deeper was the mode of its vertical distribution. The zooplankton dry weight in the profiles ranged from 3.74 to 6.91 g per haul (=1.85 to 3.45 g C m^-2, 0 to 100 m). The euphausiids represented 35% of total zooplankton dry weight and their migrations removed a large percentage of the total zooplankton biomass from the euphotic zone for 18 h d^-1. Such a large displacement of biomass would have a major impact on the biological interactions within the ecosystem.     

Annual biomass and production of the oceanic copepod community off Discovery Bay,Jamaica

Monthly samples were collected in oceanic waters off Discovery Bay, Jamaica, in 60- and 200-m vertical hauls, using 200- and 64-m mesh plankton nets, from June 1989 to July 1991. Length-weight regressions were derived for twelve genera of copepods (R²=0.79 to 0.97). For eight occasions spanning the study period, biomass estimates generated from these length-weight regressions differed by only 3% from direct weight determinations. The mean ash content of copepods was 7.1%, and the energy density was 20.8 kJ g^-1 ash-free dry weight (AFDW). Mean annual biomass of the total copepod community in the upper 60 m was 1.83 mg AFDW m^-3 (range 1.14 to 2.89 mg AFDW m^-3), and for the 200-m water column was 0.96 mg AFDW m^-3 (range 0.12 to 1.99 mg AFDW m^-3). Estimates of generation times for five common taxa ranged from 16.1 to 33.4 d. None of the taxa investigated displayed isochronal development; in general, stage duration increased in later copepodite stages. Weight increments showed a significant decrease in later copepodite stages, but with strong reversal of the trend from stage 5 to adult female in most species. Daily specific growth rates also declined in later copepodite stages, and ranged from 1.49 d^-1 in stage 1–2 Paracalanus/Clausocalanus spp. to 0.04 in stage 5-female of Oithona plumifera. Progressive food limitation of somatic copepodite growth and egg production is postulated. Naupliar production was 50.4 to 59.5% of copepodite production, and egg production was 35.1 to 27.7% of copepodite production in the 60-and 200-m water columns, respectively. Total annual copepod production, including copepodites, nauplii, eggs and exuviae, was 160 kJ m^-2 yr^-1 for the upper 60 m and 304 kJ m^-2 yr^-1 for the upper 200 m. Secondary production of the copepod community in oceanic waters off Discovery Bay approaches 50% of the corresponding value in tropical neritic waters.     

Vertical distribution, life cycle, and developmental characteristics of the mesopelagic calanoid copepod Gaidius variabilis (Aetideidae) in the Oyashio region, western North Pacific Ocean

Vertical distribution, life cycle, and developmental characteristics of the mesopelagic copepod Gaidius variabilis Brodsky in the Oyashio region were investigated by combining analyses of field copepodite populations with laboratory-rearing data of egg hatching and naupliar development. Field samplings from five discrete depths between the surface and ≤2000 m were made approximately every month for 1 year. Most populations of G. variabilis occurred between 600 and 1000 m depth. A modest degree of reversed diel vertical migration behavior and some stage-specific depth-distribution patterns were noted. All copepodite stages were observed throughout the year, suggesting a year-round spawning of G. variabilis. From a prominent abundance peak of Copepodite Stage 1 (C1) seen in June to August, together with development times of eggs and nauplii obtained in laboratory-rearing experiments, the major spawning season was extrapolated to be April to June, the phytoplankton bloom season. Tracing the peak abundance of each copepodite stage (distinguishing males and females for C4 to C6), the generation times of males and females were deduced as 2 and 1 year, respectively. All between-stage increments in terms of wet-, dry-, and ash-free dry weights were greatest in C3/C4, and least in C5/C6 for both males and females. The increments in C3/C4 and C4/C5 were greater for males than for females, reflecting a longer stage duration of the males. These weights did not increase in C5/C6 males, possibly because feeding ceased in C6 males. These results for G. variabilis are compared with those for some mesopelagic copepods previously reported from other regions. Received: 25 October 1999 / Accepted: 20 March 2000     

Life cycle of Pseudocalanus acuspes Giesbrecht (Copepoda, Calanoida) in the Central Baltic Sea: I. Seasonal and spatial distribution

The seasonal and spatial distribution of Pseudocalanus acuspes in the Bornholm Basin (Central Baltic Sea) was studied on 16 cruises between March 2002 and May 2003 from stratified (10 m) multinet samples. The highest abundances were reached in May 2002 and April 2003 (618×10³ and 869×10³ ind. m⁻², respectively). Ontogenetic vertical distribution was stage specific with differences of mean annual weighted mean depth >30 m between nauplii and males; it followed closely the hydrography which was characterized by a permanent halocline and a summer thermocline. The vertical distribution showed a positive correlation with salinity especially in the older developmental stages; the relationship to temperature was negative in the nauplii and copepodite stage I (CI). Most of the stages performed a seasonal migration. The consequences of the vertical distribution patterns in relation to the effects of climate and predation are discussed. A stage shift from nauplii in April/May to CIV and CV as overwintering stages indicated slow seasonal development. However, nauplii were observed all the year round, and the resulting stage structure did not allow to distinguish generations. Changes in the prosome length of females seemed to be related to the advection of water masses with different temperatures rather than to different generations. It could not be clarified whether the strong increase of nauplii and adults after an inflow event of cold, saline North Sea water in the beginning of 2003 was a result of advection or improvement in habitat conditions.     

Abundances,growth rates,and production of tropical neritic copepods off Kingston,Jamaica

Weekly samples were collected near Kingston, Jamaica in 27 m vertical hauls, using 200 and 64µm mesh plankton nets, from July 1985 to January 1987. Thirtytwo copepod species were identified; nauplii and all copepodite stages were enumerated. Total copepod abundance ranged from 2.56 to 87.3 × 10⁴ m^–2. The annual abundance cycle was bimodal with peaks in October–November and May–June corresponding to the rainy seasons. Mean annual copepodite biomass was 0.15 g AFDW m^–2 ranging from 0.03 to 0.41 g AFDW m^–2. Mean generation time (from egg to adult) at 28°C was 19.5 d for the common speciesCentropages velificatus, Paracalanus aculeatus, andTemora turbinata. Isochronal development was demonstrated for copepodites ofP. aculeatus andT. turbinata, but not forC. velificatus. Mean daily specific growth rates (G) were 0.63, 0.63, and 0.48 d^–1 forC. velificatus, P. aculeatus, andT. turbinata, respectively. In general, daily specific growth rates decreased in the later copepodite stages. Thus, it is postulated that growth of later stages and egg production may be food limited. Annual copepodite production was estimated as 419 kJ m^–2 yr^–1, while annual exuvial production and naupliar production were 35 and 50 kJ m^–2 yr^–1, respectively. Egg production was estimated as 44% (184 kJ m^–2 yr^–1) of the total copepodite production. Thus, mean total annual copepod production was 688 kJ m^–2 yr^–1. This estimate is within the range of copepod production estimates in coastal temperate regions.     

Respiration and excretion rates of Calanus glacialis in arctic waters of the Barents Sea

The respiration and excretion rates of Calanus glacialis (Jaschnov) Copepodite Stages III, IV, V, and adult females from the drift-ice area east of Svalbard (Barents Sea) were measured in shipboard experiments during the period from 27 May to 13 June, 1983. The phytoplankton biomass and abundance varied considerably between localities, but these variations were not generally reflected in the respiration and excretion rates of the copepod. The respiration and excretion rates of C. glacialis at the ambient temperature of-1.8°C (average respiration rates of 0.95, 0.73, 0.57, and 0.60 l O₂ mg^-1 dry wt h^-1 for Copepodite Stage III, IV, V, and adult females, respectively) were similar to those previously reported for other large-sized copepods from cold or temperate areas. Average respiration and excretion rates tended to decrease with incubation time or time after capture. Measurements on ten occasions within a period of 27 h after capture revealed excretion rates of ammonium ranging between 2.9 and 16.8 for C III, 3.7 and 21.1 for C IV, 1.3 and 28.4 for C V, and 1.6 and 18.7 for adult females, all expressed as nmol mg^-1 dry wt h^-1. In all experiments, excretion rates of inorganic phosphate varied between 0.7 and 1.5 (C III), 0.5 and 1.1 (C IV), 0.2 and 0.8 (C V), and 0.3 and 1.0 (adult females) nmol mg^-1 dry wt h^-1. Ratios of O:N, O:P, and N:P indicated that much of the metabolic energy was derived from catabolism of proteins. Comparison of the turnover rate of carbon and nitrogen showed, however, that nitrogen turnover was between 2.6 and 8.9 times higher than that of carbon. This may indicate that the copepods deaminate ingested protein, with the carbon skeleton of the amino acids subsequently being used in the synthesis of lipid compounds, possibly wax esters.     

Metabolism and body composition of a copepod (<Emphasis Type="Italic">Neocalanus cristatus</Emphasis>: Crustacea) from the bathypelagic zone of the Oyashio region,western subarctic Pacific

Metabolism [respiratory oxygen consumption, electron-transfer-system (ETS) activity] and body composition [water, ash, carbon (C), nitrogen (N), carbon/nitrogen (C/N) ratio] of stage C5/C6 Neocalanus cristatus from 1000 to 2000 m depth of the Oyashio region, western subarctic Pacific, were determined during the period of July 2000 through June 2003. Compared with the C5 specimens from shallow depths (<250 m), those from 1000 to 2000 m were characterized by quiescent behavior, reduced respiration rates (30% of the rates at active feeding), very low water content (61–70% of wet weight), but high C content (56–64% of dry weight) and C/N ratios (7.2–10.6, by weight). Artifacts due to the recovery of live specimens from the bathypelagic zone appeared to be unlikely in this study, as judged by the consistent results between re-compression (100 atm) and non-compression (1 atm) respiration experiments, and between ETS activities and respiration rates directly measured. In addition, the respiration rates of C6 males and females of N. cristatus from the same 1000–2000 m depth were two to three times higher than the rates of C5 individuals, but were similar to the rates of a bathypelagic copepod, Paraeuchaeta rubra. Combining these results with literature data, C budgets of: (1) diapausing C5 specimens, weighing 6–10 mg dry weight; (2) molt to C6 females; and (3) the complete the life span were established, taking into account assorted losses in respiration during diapause at stages C5 and C6, molt production and egg production. Respiratory C losses by C5 and C6 specimens were estimated on the basis of body N as adjusted metabolic rates [AMR; µl O₂ (mg body N)^–0.843 h^–1], then N budgets were also computed subtracting N lost in the form of cast molts and eggs from the initial stock. Calculations revealed that allocation of the C stock was greatest to egg production (34–57%), followed by respiration (27%) and cast molts (3%), leaving residual C of 13–36% in spent C6 females. The present results for N. cristatus from the North Pacific are compared with those of Calanus spp. in the North Atlantic.Communicated by O. Kinne, Oldendorf/Luhe     

Ecological investigations of the zooplankton community of balsfjorden,northern norway: The genital system in Calanus finmarchicus and the role of gonad development in overwintering strategy

The structure of the reproductive system in Copepodite Stages IV and V of the marine copepod Calanus finmarchicus (Gunnerus) is described, together with seasonal variations in gonad development and sex ratio, from Balsfjorden (69°21N; 19°06E), a subarctic fjord in northern Norway. The genital system in Copepodite Stage IV consists of a small gonad with two genital ducts. The gonad of Copepodite Stage V in its most immature condition resembles that seen in Copepodite Stage IV, whereas the sex of Stage V copepodites with mature gonads can be easily and reliably determined. The genital ducts are the key characters for separating males and females in Copepodite Stage V; potential males have one and potential females two genital ducts connecting the gonad to the genital opening on the first urosome segment. Adults males were rare compared to females, and were only in the majority at the end of January and beginning of February. The sex-ratio also favoured females in Copepodite Stage V. Seasonal peaks in the proportion of adult males reflected peaks of males in Copepodite Stage V, suggesting that seasonal variations in the adult sex-ratio are a direct result of a situation which has already been determined in Copepodite Stage V. The seasonal variation in gonad development in Copepodite Stages IV and V of C. finmarchicus reveals that the size of maturity of the gonad varies throughout the year. This study indicates that visible sex-differentiation in Stage V and moulting into adults occurs in Balsfjorden at least 2 months before spawning in April. Phytoplankton levels are immeasurable before the spring diatom increase starts at the end of March, implying that development and maturation of gonads are dependent upon internal energy resources, whereas the final act of spawning seems to require energy input from phytoplankton.     

Seasonal development of scyphozoan medusae and the predatory impact of Aurelia aurita on the zooplankton community in the Bornholm Basin (central Baltic Sea)

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita and Cyanea capillata was studied in the Bornholm Basin (central Baltic Sea) in 2002. Seasonal changes in prey composition and predatory impact were investigated by analyzing stomach contents. A. aurita occurred from July to November, with a maximum mean abundance of 2.3 ind. per 100 m³ in August, whereas C. capillata was caught in much smaller numbers from July to September. No ephyrae of either species were found; therefore, advection of medusae from the western Baltic Sea is assumed. From July to October, ~80% of A. aurita medusae was distributed in the upper 20 m above the thermocline, whereas C. capillata occurred only in the halocline below 45 m. A. aurita did not migrate vertically and fed mainly on the most abundant cladoceran species Bosmina coregoni maritima. Further prey organisms were the cladocerans Evadne nordmanni and Podon spp., mollusk larvae and copepods. Copepod nauplii and copepodite stages I–III were not eaten by the medusae, neither were fish eggs and larvae used as prey. Based on mean medusa and zooplankton abundance from the upper 20 m, the predatory impact was very low. In August, when mean abundance of A. aurita was highest, only 0.1% of the copepod and 0.5% of the cladoceran standing stock were eaten per day. However, in regions with higher medusa or lower zooplankton abundance, up to 7.9% of the cladoceran standing stock was consumed per day. Hence, A. aurita did not regulate the zooplankton community in the Bornholm Basin, and fish larvae did not suffer from competition with and predation by the medusae.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Bimodal vertical distribution and diel migration of the copepods Metridia pacifica,M. okhotensis and Pleuromamma scutullata in the western North Pacific Ocean

Diel changes in fine-scale vertical distributions of three calanoid copepods Metridia pacifica, M. okhotensis and Pleuromamma scutullata in the subarctic waters of the western North Pacific were examined. Sampling was carried out in June and August 1983, at two stations in Oyashio water using a Longhurst-Hardy Plankton Recorder (LHPR). Sampling, down to about 1 000 m, was repeated four to five times at intervals of several hours. Vertical resolution was 5 to 40 m. Copepods were concentrated in two strata, the surface (0 to 60 m) and the mesopelagic (200 to 300 m) layers, throughout the day at both stations. Younger M. pacifica (C III and C IV) were dominant in both strata. Although most female C V and adult females demonstrated diel vertical migration at 20 to 30 m h^-1, a significant number of females did not migrate upward but remained in the deep stratum at night. The same trend was evident in M. ohkotensis and P. scutullata. Foregut content observations indicated that feeding activities of the deep mode populations were as high as those of the surface mode, though food of deep individuals was different. Such a bimodal distribution may increase intraspecific diversity of copepod populations and is possibly why metridiid copepods dominate during late summer to winter in the relatively simple ecosystems of high latitudes.     

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

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

Male preference for large females and assortative mating for body size in the sand lizard (Lacerta agilis)

Summary Male sand lizards (Lacerta agilis) are polygynous and guard individual females for several hours to days after copulation. Even though the copulation itself only lasts 2–4 min, the total time that a male invests per female is considerably more and may constitute a substantial investment during a mating season. In such situations, when male copulation frequency is constrained, or when variation in female fecundity is high, mate choice by males may be adaptive. Large body size in female sand lizards is correlated with higher fecundity. In choice experiments performed in the laboratory, male sand lizards preferred to court large females rather than small females. In addition, when there was little difference in size between the females in the experiment, the males visited the two females more often before they started to court the preferred female. The results from a field study during 1984 and 1987–1990 showed that females are non-aggressive, have small neighboring home ranges (c. 100 m²) and may share burrows and sites for thermoregulation. This means that females can be found close together and thus gives males the opportunity to choose a mate. Assortative mating with respect to size was observed in a natural population, as well as a limited number of direct choices of females by males. These results support the results of the choice experiment.     

Photosynthetic characteristics of nanoplankton and picoplankton from the surface mixed layer

Nanoplankton and picoplankton primary production has been studied at two oceanic stations in the Porcupine Sea-bight and at one shelf station in the Celtic Sea. At both sites, low wind conditions in June and July 1985 resulted in greatly reduced vertical turbulent mixing and a secondary, temporary thermocline developed in what is usually a well-mixed surface layer; as a result, there was physical separation of the phytoplankton within two zones of the surface mixed layer. The photosynthetic characteristics of three size fractions (>5 m, <5 to >1 m and <1 to >0.2 m) of phytoplankton populations from the two zones have been measured. Phytoplankton was more abundant at the oceanic stations and chlorophyll a values were between 1.3 and 2.2 mg chlorophyll a m^-3, compared with 0.3 to 0.6 mg chlorophyll a m^-3 at the shelf station; at both stations, numbers of cyanobacteria were slightly higher in the lower zone of the surface mixed layer. There was no effect of the temporary thermocline on the vertical profiles of primary production and most phtosynthesis occurred in the surface 10 m. Photosynthetic parameters of the three size fractions of phytoplankton have been determined; there was considerable day-to-day variation in the measured photosynthetic parameters. Assimilation number (P _m ^B ) of all >5 m phytoplankton was lower for the deeper than for the surface populations, but there was little change in initial slope (a ^B ). The small oceanic nanoplankton (<5 to >1 m) showed changes similar to the >5 m phytoplankton, but the same size fraction from the shelf station showed changes that were more like those shown by the picoplankton (<1 m) viz, little change in P _m ^B but an increase in a ^B with depth. Values of a ^B were generally greater for the picoplankton fraction than for the larger phytoplankton, but values of adaptation parameter (I _k )(=P _m ^B /) were not always less. There was little evidence to support the hypothesis that these populations of picoplankton were significantly more adapted to low light conditions than the larger phytoplankton cells. When photosynthetic parameters of the picoplankton were normalised to cell number (P _m ^C /a ^C ) rather than chlorophyll a, P _m ^C was comparable to other published data for picoplankton, but a ^C was much lower. The maximum doubling time of the picoplankton at saturating irradiance is calculated to be ca. 8.5 h for the oceanic population and ca. 6.2 h for the shelf population.     

Space use and social structure in meadow voles,Microtus pennsylvanicus

Summary Free-ranging, sexually mature meadow voles (Microtus pennsylvanicus) were tracked by using radiotelemetry from June through August in Front Royal, Virginia, U.S.A. Estimates of intraspecific spacing were derived from the concurrent movements of up to 16 voles. Positions were recorded hourly for 24 h, twice per week. A total of 16 male and 15 female voles were studied during sixteen 24-h sessions.The daily ranges of males (192.3±109.7 m²) were larger and more variable than those of females (68.6±39.4 m²). Males also changed locations more frequently (Fig. 2).Adult females usually maintained territories free of other females; males overlapped considerably among themselves (Fig. 2). Males temporarily moved into the areas occupied by estrous females, indicating intrasexual competition among males for access to receptive females (Fig. 3). M. pennsylvanicus appears to be promiscuous, is socially organized into territorial, maternal-young units during the breeding season, and fits the female territorial model of population regulation.     

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.     

Pattern of planktonic bioluminescence in the northern Sargasso Sea: Seasonal and vertical distribution

The vertical structure of bioluminescence potential (BPOT) and flash density (FD) were measured on five cruises to the northern Sargasso Sea in 1987 and 1988. Depth-integrated (0 to 150 m) BPOT did not vary seasonally, remaining within the range 9 to 15 × 10¹⁵ photons m^–2 in all months sampled. Conversely, depth-integrated FD was significantly higher (> 2 × 10⁵ flashes m^–2) during winter (November and March) than during summer (< 9 × 10⁴ flashes m^–2 in May and August). The vertical patterns of BPOT and FD were well correlated within a single profile, more highly so in summer than in winter. Despite intracruise variability in the vertical pattern of BPOT and FD, there were clear summer-winter differences in the vertical distribution of BPOT and FD. During winter, BPOT and FD were maximal and relatively uniform throughout the surface mixed layer; for example in November they declined sharply within the thermocline at 130 to 150 m. During summer, BPOT and FD were greatest (12 to 25 × 10¹³ photons m^–3 and 600 to 1 200 flashes m^–3, respectively) at subsurface depths. Commonly in summer, the upper depth limit of high BPOT and FD occurred at the base of the surface mixed layer (10 to 40 m) and the lower depth limit was located at the base of the subsurface fluorescence maximum (usually at 100 to 120 m).     

Production of tropical copepods in Kingston Harbour, Jamaica: the importance of small species

The copepod community observed during an 18-month period at the mouth of eutrophic Kingston Harbour, Jamaica, was dominated by small species of Parvocalanus, Temora, Oithona, and Corycaeus. Mean copepod biomass was 22.1 mg AFDW m⁻³ (331 mg m⁻²). Annual production was 1679 kJ m⁻², partitioned as 174 kJ m⁻² naupliar, 936 kJ m⁻² copepodite, 475 kJ m⁻² egg and 93 kJ m⁻² exuvial production. All nauplii, most copepodites and many adults, equivalent to half of the biomass and production, were missed by a standard 200-μm plankton net, emphasizing the importance of nauplii and small species in secondary production estimates. The evidence suggests that growth rates and production are generally not food limited, and we speculate that size-selective predation shapes the structure of the harbour community. Biomass and production are higher than previous estimates for tropical coastal waters, but comparable to other eutrophic tropical embayments and many productive temperate ecosystems. Far from being regions of low productivity, tropical zooplankton communities may have significant production and deserve greater research attention than they currently receive. Received: 19 September 1997 / Accepted: 21 October 1997