Nitrate storage by phytoplankton in a coastal upwelling environment

The storage of nitrate by phytoplankton cells during the early phases of upwelling was studied in coastal stations off northern Spain (southern Bay of Biscay) between 1990 and 1994. In this region, a persistent upwelling during summer is characterised by intermittent pulses of variable intensity, and increased nutrient concentrations in the surface layer. The main effect of an upwelling pulse on phytoplankton distribution is the shifting of the chlorophyll a and primary production maxima to near the surface. When the upwelling relaxes, thermal stratification of the water column occurs, and a distinct subsurface chlorophyll maximum develops below the production maximum. An accumulation of intracellular nitrate characterized the early phases of upwelling (mean = 2.73 μmol N m⁻³), maximum concentrations being attained at depths where biomass and production values were moderate. In contrast, phytoplankton cells from non-upwelling situations contained significantly lower concentrations of intracellular nitrate (mean = 0.17 μmol N m⁻³). The variations in the intracellular pool of nitrate may result from the differential allocation of resources within the cell as a result of variations in the energy available, since the uptake and assimilation of nitrate is a relatively expensive process involving several enzymatic systems. We hypothesize that nitrate storage by phytoplankton cells is characteristic of early phases of upwelling and is linked to patterns of carbon fixation. Average nitrogen budgets for upwelling and non-upwelling situations indicate that intracellular nitrate reserves are not responsible for maintaining high phytoplankton growth rates, since they only account for <2% of daily primary production during upwelling events. Received: 28 August 1996 / Accepted 3 December 1996     

Vertical and horizontal structure in the picoplankton communities of a coastal upwelling system

The distribution of prokaryotic and eukaryotic picoplankton in the west coast upwelling-region off the South Island of New Zealand was investigated during midwinter (1988) the time of year when several commercially important fish species migrate into the region to breed. Picoplanktonic cells were major contributors to the autotrophic biomass, with > 80% of the particulate nitrogen and 39 to 55% of the total chlorophylla contained in the <2µm size-fraction. The prokaryotic picoplankton concentrations ranged from 6.3 × 10⁵ to 2.1 × 10⁷ cell l^–1, and the eukaryotic picoplankton between 3.9 × 10⁵ to 1.2 × 10⁷ cells l^–1. Picoplankton numbers increased with distance offshore to a maximum of ~ 3.0 × 10⁷ cells l^–1 at ~ 35 km from the coast, and then diminished towards the outer shelf and open ocean. The ratio of prokaryotic to eukaryotic cells varied between 1.01 and 4.71 in the mixed layer. Both groups declined substantially beneath the pycnocline, with no evidence of deep maxima. Prokaryotic cells dominated the planktonic cell concentrations at all but two stations, but eukaryotic cells dominated picoplankton biovolume as a result of their larger average cell size. The prokaryotic to eukaryotic picoplankton cell-number ratios in this system were considerably lower than often recorded elsewhere, and were inversely correlated with nitrate concentration. These observations show that a eukaryoticdominated picoplankton community makes a substantial contribution to autotrophic biomass in this nutrient-rich upwelling system, and may thereby play a major role in the food-web dynamics of this coastal fishery.     

Grazing of phytoplankton by copepods in eastern Antarctic coastal waters

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

In-situ measurements of nitrogenous nutrient uptake by kelp (Ecklonia maxima) and phytoplankton in a nitrate-rich upwelling environment

Nitrogen uptake by the kelp Ecklonia maxima Osbeck and phytoplankton was examined under different conditions of nutrient availability in a kelp bed off the Cape of Good Hope by measuring nutrient depletion in large plastic bags by the kelp and ¹⁵N uptake by phytoplankton. E. maxima took up nitrate and ammonia, but not urea, and showed only a weak preference for reduced nitrogen. Phytoplankton absorbed all three forms of nitrogen available, with a preference for ammonia and urea. Ambient nitrate concentration exhibited a marked and rapid decrease with northerly winds and an increase in response to offshore southerly winds. Nitrogen uptake by E. maxima was linearly related to ambient concentration and did not saturate even at nitrate concentrations >20g-at N l^-1, resulting in a significantly higher tissue nitrogen content under upwelling conditions. Nitrate imported by upwelling was the chief source of nitrogen utilised within the kelp bed. Locally regenerated nitrogen (ammonia and urea) was calculated to contribute only ca 4% of total nitrogen uptake during upwelling and 30% during the relaxation or downwelling phase.     

Simultaneous nitrogen and silicate deficiency of a phytoplankton community in a coastal jet-front

The nutritional status of a phytoplankton community was investigated in a coastal jet-front located in the Gulf of St. Lawrence, Canada, in 1987. During the sampling period, the frontal community was mainly composed of the diatomsChaetoceros debilis, Skeletonema costatum, Thalassiosira gravida andC. pelagicus. As previously reported for the St. Lawrence, some frontal stations were depleted both in nitrate and silicate. At stations impoverished in nitrate, internal nitrate pool concentrations were low or undetectable, suggesting that cells had not, recently, been exposed to a nitrate flux which exceeded the nitrate assimilation rate. At these impoverished stations, however, ambient and intracellular concentrations of ammonium and urea were high, suggesting that the community was not nitrogen-deficient. The comparison between the ambient silicate concentrations and the silicate requirement (K _s ) of the dominant diatoms suggests thatC. debilis andS. costatum were Si-deficient. This is further supported by the low silicate uptake rates and intracellular concentrations measured at the silicate impoverished stations. The silicate deficiency also resulted in a decrease in the seston and phytoplankton N:C ratios.Please address all correspondence and requests for reprints to Dr Levasseur at his new address: Maurice Lamontagne Institute, 850 Route de la Mer, Mont-Joli, Québec G5H 3Z4, Canada     

Fuzzy modelling of chlorophyll production in a Brazilian upwelling system

In the context of coastal management the aim of this paper is to present the development of a fuzzy model through the application of a Genetic algorithm in order to select the most appropriate set of variables and improve our understanding with a set of rules. The case studied is the chlorophyll response as bioindicator of ecological status in the Northeast coastal upwelling system of Rio de Janeiro state, Southeastern of Brazil. The prediction of the fuzzy model has shown an improved performance when compared to the traditional approaches as Multiple Regression modelling. The results show that the set of inferred rules can assess three different water masses. Despite the increased occurrence of upwelling is observed in spring–summer period and some instability of the model, it is able to forecast some chlorophyll peaks. We conclude that the sampling frequency is crucial to reach a better performance.     

Microzooplankton grazing and selectivity of phytoplankton in coastal waters

Microzooplankton grazing activity in the Celtic Sea and Carmarthen Bay in summer 1983 and autumn 1984 was investigated by applying a dilution technique to high-performance liquid chromatographic (HPLC) analysis of photosynthetic pigments in phytoplankton present within natural microplankton communities. Specific grazing rates on phytoplankton, as measured by the utilisation of chlorophyll a, were high and varied seasonally. In surface waters during the autumn, grazing varied between 0.4 d^-1 in the bay and 1.0 d^-1 in the Celtic Sea, indicating that 30 and 65% of the algal standing stocks, respectively, were grazed daily. Grazing rates by microzooplankton within the thermocline in summer suggest that 13 to 42% of the crop was grazed each day. Microzooplankton showed selection for algae containing chlorophyll b, in spite of a predominance of chlorophyll c within the phytoplankton community. Changes in taxon-specific carotenoids indicated strong selection for peridinin, lutein and alloxanthin and selection against fucoxanthin and diadinoxanthin. This indicates a trophic preference by microzooplankton for dinoflagellates, cryptophytes, chlorophytes and prasinophytes and selection against diatoms, even when the latter group forms the largest crop within the phytoplankton. Interestingly, those algal taxa preferentially grazed also showed the highest specific growth-rates, suggesting a dynamic feed-back between microzooplankton and phytoplankton. Conversion of grazing rates on each pigment into chlorophyll a equivalents suggests firstly, that in only one experiment could all the grazed chlorophyll a be accounted for by the attrition of other chlorophylls and carotenoids, and secondly that in spite of negative selection, a greater mass of diatoms could be grazed by microzooplankton than any other algal taxon. The former may be due either to a fundamental difference in the break-down rates of chlorophyll a compared to other pigments, or to cyanobacteria forming a significant food source for microzooplankton. In either case, chlorophyll a is considered to be a good measure of grazing activity by microzooplankton.     

Influence of inter- and intra-daily light-field variability on photosynthesis by coastal phytoplankton

An investigation was carried out to determine the relationships between the principal photosynthetic parameters of natural marine phytoplankton and the properties of the irradiance of the preceding five days and the intra-daily ambient irradiance. Samples used in the study were collected from Strangford Lough, Northern Ireland during both winter and summer between 1979 and 1981. Photosynthetic parameters were determined for both constant and ambient incubation light fields. The fluctuation properties of the irradiance field exerted little influence on the photosynthetic parameters. In summer populations, the parameters obtained from morning samples were strongly influenced by the total irradiance recorded on the fourth day previous to experimentation, irrespective of the incubation irradiance. In contrast, the corresponding parameters from afternoon samples were most influenced by the total irradiance received during incubation. The influence of these two separate factors over a day may contribute to variability in the magnitude of diel changes in marine phytoplankton photosynthesis.     

Trophic role of small cyclopoid copepod nauplii in the microbial food web: a case study in the coastal upwelling system off central Chile

Copepod grazing impact on planktonic communities has commonly been underestimated due to the lack of information on naupliar feeding behaviour and ingestion rates. That is particularly true for small cyclopoid copepods, whose nauplii are mainly in the microzooplankton size range (<200 μm). The trophic role of Oithona spp. nauplii was investigated off Concepción (central Chile, ~36°S) during the highly productive upwelling season, when maximum abundances of these nauplii were expected. Diet composition, ingestion rates, and food-type preferences were assessed through grazing experiments with different size fractions of natural planktonic assemblages (<3, <20, <100, and <125 μm) and cultures of the nanoflagellate Isochrysis galbana. When the Oithona spp. nauplii were offered a wide range of size fractions as food (pico- to microplankton), they mostly ingested small (2–5 μm) nanoflagellates (5–63 × 10³ cells nauplius⁻¹ day⁻¹). No ingestion on microplankton was detected, and picoplankton was mainly ingested when it was the only food available. Daily carbon (C) uptake by the nauplii ranged between 28 and 775 ng C nauplius⁻¹, representing an overall mean of 378% of their body C. Our relatively high ingestion rate estimates can be explained by methodological constraints in previous studies on naupliar feeding, including those dealing with “over-crowding” and “edge” effects. Overall, the grazing impact of the Oithona spp. nauplii on the prey C standing stocks amounts up to 21% (average = 13%) for picoplankton and 54% (average = 28%) for nanoplankton. These estimates imply that the nauplii of the most dominant cyclopoid copepods exert a significant control on the abundances of nanoplankton assemblages and, thereby, represent an important trophic link between the classical and microbial food webs in this coastal upwelling system.     

Killing of marine phytoplankton by a gliding bacterium Cytophaga sp., isolated from the coastal sea of Japan

A marine gliding bacterium Cytophaga sp. (strain J18/M01) was isolated from Harima-Nada, eastern Seto Inland Sea, Japan in 1990. This bacterium preys upon various species of marine phytoplankton. All of the five raphidophycean flagellates, all of the four diatoms, and one of the two dinoflagellates examined were killed within a few days when cultured with the bacterium. The bacterium presumably achieves this by direct attack, because the culture filtrate in which host organisms were totally destroyed had no significant effects on the growth of the same host organism (Chattonella antiqua). If one or a few bacterial cells were inoculated into C. antiqua culture, all of the host organisms were killed. The bacterium proliferated in filter-sterilized seawater, suggesting its ubiquitous existence in the coastal sea. The killing of phytoplankton by bacteria such as Cytophaga sp. J18/M01 may be a significant factor influencing the population dynamics of phytoplankton in nature and may contribute to the sudden disappearance of red tides in the coastal sea. Bacterial destruction of phytoplankton may also be a factor that regulates primary productivity in marine ecosystems.     

Decomposition of urea associated with photosynthesis of phytoplankton in coastal waters

Decomposition of urea in seawater was studied in Mikawa Bay, a shallow eutrophic bay on the southern coast of central Japan. The urea concentration in seawater ranged from 1.3 to 5.9 μg-at. N/1 and comprised 12 to 40% of the dissolved organic nitrogen. Using ¹⁴C labelled urea, the rate of CO₂ liberation from urea and the incorporation rate of urea carbon into the particulate organic matter were determined. For the surface samples, high rates of CO₂ liberation from urea as well as the incorporation of urea carbon into the particulate organic matter were observed in the light, while much lower rates were obtained in the dark. Incubation experiments with exposure to different light intensities revealed that the rate of CO₂ liberation from urea and the incorporation of urea carbon into particulate organic matter changed with light intensity, showing a pattern similar to that of photosynthesis. The highest liberation and incorporation rates were observed at 12,000 lux. Incubation in light and in dark produced marked decreases and increases, respectively, in urea and ammonia, while no appreciable changes were observed for nitrate and nitrite. It is suggested that urea decomposition associated with photosynthetic activity of phytoplankton is one of the major processes of urea decomposition, and that it plays a significant role in the nitrogen supply for phytoplankton in coastal waters.     

Stimulation of phytoplankton production in coastal waters by natural rainfall inputs: Nutritional and trophic implications

Recent evaluations of estuarine and coastal nutrient budgets implicate atmospheric deposition as a potentially significant (20 to 30%) source of biologically available nitrogen. We examined the potential growth stimulating impact of atmospheric nitrogen loading (ANL), as local rainfall, in representative shallow, nitrogen limited North Carolina mesohaline estuarine and euhaline coastal Atlantic Ocean habitats. From July 1988 to December 1989, using in situ bioassays, we examined natural phytoplankton growth responses, as¹⁴CO₂ assimilation and chlorophylla production, to rain additions over a range of dilutions mimicking actual input levels. Rainfall at naturally occurring dilutions (0.5 to 5%) stimulated both¹⁴CO₂ assimilation and chlorophylla production, in most cases in a highly significant manner. Parallel nutrient enrichments consistently pointed to nitrogen as the growth stimulating nutrient source. Generally, more acidic rainfall led to greater magnitudes of growth stimulation, especially at lower dilutions. Nutrient analyses of local rainfall from May 1988 to January 1990 indicated an inverse relationship between pH and NO ₃ ^- content. There have been growing concerns regarding increasing coastal and estuarine eutrophication, including ecologically and economically devastating phytoplankton blooms bordering urban and industrial regions of North America, Europe, Japan, and Korea. It appears timely, if not essential, to consider atmospheric nutrient loading in the formulation and implementation of nutrient management strategies aimed at mitigating coastal eutrophication.     

Differences in photosynthetic pigment signatures between phytoneuston and phytoplankton communities in a coastal lagoon of Baja California

In order to understand the relationships between the dynamics of phytoplankton populations in the surface microlayer (MIL) and in the water column below (SSW), this study used high-performance liquid chromatography-derived pigment markers in samples from a coastal lagoon of Baja California (Estero de Punta Banda, EPB) under summer (October 2003) and winter (December 2003) conditions. Photosynthetic pigment signatures of phytoplankton at the air–sea interface (phytoneuston) and subsurface measurements were related to bottom-up (temperature, salinity, nutrient concentrations) and top-down factors (zooplankton abundance). Slicks and scum layers were observed in the inner part of the lagoon and coincided with greater stratification of layers just below the sea surface and lower wind intensities. In general, spatial variability in pigment markers and ancillary data was very high and resulted in non-significant differences between MIL and subsurface samples when different regions of EPB or sampling dates were compared. However, different patterns were found between pigments and environmental factors of MIL and SSW samples when the relative numbers of stations with positive and negative differences (ΔX = X _MIL−X _SSW) were computed. For each survey, pigment markers of phytoneuston and phytoplankton samples were not necessarily correlated. Further analysis revealed that those markers (19′-butanoyloxyfucoxanthin, prasinoxanthin, divinil-chlorophyll a) corresponded to picophytoplankton groups (haptophyte, prasinophyte, and prochlorophyte). On both dates, the MIL was enriched in 19′-hexanoyloxyfucoxanthin (a marker for a type 4 haptophyte) and fucoxanthin (marker for bacillariophytes, haptophytes, and crysophytes) and depleted in peridinin (marker for dinophytes). Different zooplankton grazers accumulated in the MIL (loricate tintinnids) and in SSW (copepod nauplii).     

Aspects of a zooplankton,phytoplankton, phosphorus system

In this paper a three-dimensional dynamical system which models the three-species system made up of phytoplankton, zooplankton and organic phosphorus nutrient in a lake environment is studied. The system is part of a more general limnological model for eutrophic lakes and impoundments which has been developed by Battelle Northwest Laboratories. It is shown that this system, henceforth referred to as Z-P-P, has a phase portrait comprised of a plane portrait embedded in the three-dimensional space R³ as an “attractor”.¹ Under a small perturbation of the nutrient equation it is shown that the system is essentially a classical Volterra-Lotka system embedded in a three-dimensional phase space R³.The system derived from Z-P-P by the addition of a term to the nutrient equation which represents the organic phosphorus contribution of dying phytoplankton is also considered. The equilibria of this system are studied and what can be deduced of the phase portrait is compared with that of the above systems. It is found that these phase portraits are qualitatively indifferent to the form of the growth rate functions for zooplankton and phytoplankton provided they are monotone increasing. Some discussion about the stability of these systems is included. Throughout this paper results are interpreted in limnological terms.     

Phosphate uptake behavior of natural phytoplankton during exposure to solar ultraviolet radiation in a shallow coastal lagoon

The effect of solar UV radiation (UVR: 290–400 nm) on the ³²P-phosphate uptake rates of natural phytoplankton from a southern Atlantic Ocean coastal lagoon was studied during two consecutive summers at one station located in the marine-influenced area. Due to the shallowness of this lagoon and also to the generally high UV water transparency in this area, phytoplankton are exposed to high UV irradiances. The ³²P-phosphate uptake rates measured at several phosphate concentrations were inhibited up to 59.2% by UVR, although uptake stimulation was also observed in four of nine experiments (up to 28%). The effect of UVR on the apparent maximum velocity of ³²P-phosphate uptake (V _uptake) ranged from an inhibition of 49% to a stimulation of 31%. Although the highest inhibition values were associated with the maximum registered incident UV irradiance, a significant correlation between these two parameters was not observed. Changes in microalgal community structure were not related to the observed UV effect; however, a significant relationship was found between the inhibition of ³²P-phosphate uptake rates and V _uptake used as a proxy for phosphate deficiency. This relationship suggests that the phytoplankton phosphorus nutritional status modulates their sensitivity to UV exposure. Overall, our results suggest that solar UVR has the potential to affect phosphorus cycling.Communicated by O. Kinne, Oldendorf/Luhe     

Orthophosphate uptake by phytoplankton and bacterioplankton from the Los Angeles Harbor and Southern California coastal waters

Orthophosphate (P) uptake on a seasonal basis in surface waters and in vertical profiles was directly proportional to the standing stocks of phytoplankton and bacterioplankton in the outer Los Angeles Harbor and in southern California coastal waters during 1978–1979. A phytoplankton-enriched size fraction (PEF) which was retained on a 1 m pore-size filter contained 83% of the total chlorophyll a but only 18% of the total bacteria. A bacterioplankton-enriched size fraction (BEF) which passed the 1 m filter but was retained on a 0.2 m filter contained 82% of the total bacteria but only 17% of the total chlorophyll a. PEF and BEF accounted for 91 and 9% of the microbial carbon, respectively. The differential uptake of 10 radiolabeled substrates more fully characterized PEF and BEF. ³³P uptake occurred in both PEF and BEF, accounting for 47 and 53%, respectively, of the total uptake. ³³P uptake by both size fractions was inhibited by low concentrations of 2,4-dinitrophenol (DNP), N-ethylmaleimide (NEM) and carbonyl cyanide, m-chlorophenylhydrozone (CCCP). Darkness and low levels of 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) selectively inhibited ³³P uptake by PEF; valinomycin selectively inhibited ³³P uptake by BEF. An experiment measuring ³³P uptake velocity versus P concentration produced sigmoidal saturation kinetics at high levels of exogenous P. Kinetic parameter analyses according to the Hill equation gave a V _max of 7.12 nmol l^–1 h^–1 and aK _t of 0.41 nmol l^–1 for PEF, and a V _max of 5.17 nmol l^–1 h^–1 and aK _t of 112 nmol l^–1 for BEF. Consideration of relative surface areas of phytoplankton and bacterioplankton, their ³³P uptake rates in light and dark, and estimates of the population turnover times emphasizes the potential importance of bacterioplankton in community phosphorus metabolism.     

Metal partitioning in sediment pore water from the Ondo coastal region,Nigeria

Distribution and equilibrium partitioning of metals (Cd, Cu, Cr, Fe, Ni, Pb, Mn, and Zn) between pore water and surface sediments at the Ilaje coast of Ondo State, Nigeria, were studied. The Ilaje River can be one of the interesting research locations because of its economic nature and history of oil pollution. Seasonal variations were observed to investigate possible variations in the availability of metals for organisms throughout the year. The concentrations in both sediments and pore water during the dry and wet seasons were as follows: Cu?>?Fe???Mn?>?Pb?>?Ni?>?Cr?>?Cd?>?Zn. The pore water–sediment partition coefficient (K _p) showed that Zn and Cd were highly mobile while Fe and Cu have restricted mobility. Dissolved organic carbon (DOC) in pore water had a strong influence on mobility and bioavailability of all the metals (p?K _oc), increased levels of Fe and Cu were linked to lithological origin. Concentrations of Pb and Ni were associated with petroleum-related sources. The significance of the field-based techniques for ecotoxicological purposes is discussed.     

Resource partitioning by four species of elasmobranchs (Batoidea: Urolophidae) in coastal waters of temperate Australia

Four species of stingarees (Urolophus lobatus, U. paucimaculatus, Trygonoptera personata and T. mucosa) were abundant in samples collected from over sandy substrates in four regions along ∼200 km of the lower west coast of Australia in each season between summer 1990/1991 and winter 1992. The main aim of this study was to determine whether these four species were segregated by habitat, size, sex and/or diet, thereby reducing the potential for intra- and interspecific competition. The densities of U. lobatus and T. personata were almost invariably greater in deep, offshore waters (20 to 35 m) than in shallow, inshore waters (5 to 15 m), while those of U. paucimaculatus were greatest in the southernmost region, irrespective of depth, and those of T. mucosa were highest at the single inshore site at which the water was deep. None of these species was segregated by either size or sex. The volumetric dietary compositions of the two Urolophus species, which consisted predominantly of various crustaceans, and those of the two species of Trygonoptera, which consisted mainly of different polychaetes, were significantly different from each other. The two Urolophus species initially fed largely on amphipods, mysids and carid decapods but, with increasing size, U. lobatus started to ingest teleosts and U. paucimaculatus commenced feeding on errant polychaetes and penaeid decapods. The contribution of errant polychaetes to the diets of T. personata rose progressively from 0% in the smallest fish to 90% in the largest fish, thereby accounting for a concomitant progressive decline in dietary breadth and for intraspecific dietary overlap typically to occur only between successive size classes. In contrast, the volumetric contribution of sedentary polychaetes to the diets of T. mucosa remained high (65 to 80%) throughout life, which accounts for the relatively low and constant dietary breadth of most size classes and a significant overlap in the diets of all but one pair of size classes. The diets of each of the four stingaree species occasionally differed between sites and sometimes underwent seasonal changes, presumably reflecting spatial and temporal differences in the types of prey in the environment. Since the ratios of benthic to epibenthic prey in the diets of the two Trygonoptera species were greater than in those of the two Urolophus species, the first two species apparently forage deeper into the substrate. The combination of partial segregation of the four stingaree species among habitats and the differences in the dietary compositions and feeding strategies of these species, would presumably facilitate the co-existence of these batoid elasmobranchs when they are abundant in the same general region. Received: 29 October 1997 / Accepted: 25 March 1998     

Numerical behavior of a zooplankton,phytoplankton and phosphorus system

Several numerical features of a three-dimensional dynamical system which models the three species system made up of phytoplankton, zooplankton and organic phosphorus nutrient in a lake environment are presented. Certain properties of this system, such as the existence of limit cycles, are demonstrated numerically. It is shown that in certain time domains the system may be stiff, thus requiring the use of suitable algorithms.