Nutrient recycling and shifts in N:P ratio by different zooplankton structures in a South Andes lake

In South Andes lakes, zooplankton succession is characterizedby a change in the dominance from the calanoid copepod Boeckellagracilipes in winter and spring, to the cladoceran Bosmina longirostrisin midsummer, and the rotifer Polyarthra vulgaris in late summer.We performed three series of field experiments at differenttimes (late spring, summer and late summer) to examine the roleof zooplankton constitution in the released nitrogen:phosphorus(N:P) ratio. We observed that changes in the zooplankton constitutionover the annual cycle may change the nutrient supply ratio.In South Andes lakes, Boeckella gracilipes would decrease theP limitation, lowering the N:P ratio, whereas Bosmina longirostristends to increase the N:P ratio and therefore increase the Plimitation during summer.     

Survivorship of Cyclops abyssorum tatricus (Cyclopoida, Copepoda) and Boeckella gracilipes (Calanoida, Copepoda) under ambient levels of solar UVB radiation in two high-mountain lakes

We performed in situ experiments during the summer of 1995 and 1996 to assess the potential effect of solar ultraviolet B (UVB) radiation (290-320 nm) on the survival of Cyclops abyssorum tatricus Kozminski and Boeckella gracilipes Daday. These species are numerically dominant within the crustacean zooplankton living in two high-mountain lakes, one located in the Austrian Alps [Gossenkollesee (GKS), 2417 m above sea level, maximum depth 9.9 m] and another in the Chilean Andes (Laguna Negra, 2700 m above sea level, maximum depth 320 m). The copepods were incubated in quartz tubes (1 1) or in quartz tubes wrapped with Mylar D® to exclude most of the UVB radiation. The organisms were exposed at 0.5 m depth for 10-72 h on cloudless days. Both lakes were very transparent to UVB and 10% of the surface radiation at the nominal wavelength of 305 nm was still present at 9.6 m in GKS and at 12.8 in Laguna Negra. These species migrate vertically and have a maximum daytime distribution close to the bottom (C.abyssorum tatricus) or below 15 m depth (B.gracilipes). Both species were red, but the carotenoid concentration was higher in C.abyssorum tatricus than in B.gracilipes (6.5 and 2.3 g mg^-1 dry weight, respectively). UV-absorbing compounds with a maximum absorption at 334 nm were also detected. Cyclops abyssorum tatricus was highly resistant to UVB radiation and no significant lethal effect was observed. Boeckella gracilipes had a mortality 5 times higher in the treatment receiving full sunlight than in the Mylar treatment (3.2%) only when exposed for 70 h. The resistance of B.gracilipes was higher than that reported in the literature for the same species, suggesting the existence of intraspecific differences in UV sensitivity.      

Effects of the ice-edge bloom and season on the metabolism of copepods in the Weddell Sea,Antarctica

The metabolic responses of several species of Antarctic copepods to primary productivity and changes between seasons were investigated. To examine the influence of the spring ice-edge bloom on the metabolism of copepods, oxygen consumption rates were determined on specimens from three zones of widely different ice coverage and chlorophyll biomass: pack ice (pre-bloom), ice edge (bloom) and open water (post-bloom). Summer metabolic rates were compared with published winter rates. Field work was done in the Weddell Sea in the region of 60 °S, 36°W in late November and December 1993. Oxygen consumption rates were determined by placing individuals in syringe respirometers and monitoring the oxygen partial pressure for 10–20 hours. Higher metabolic rates were observed in the primarily herbivorous copepods, Calanoides acutus, Rhincalanus gigas and Calanus propinquus in regions of higher primary production: ice edge and open water. The carnivorous Paraeuchaeta antarctica showed a similar pattern. The omnivorous copepods Metridia gerlachei and Gaetanus tenuispinus showed no changes in metabolism between zones. Data on routine rates of copepods from the winter were available for C. propinquus and P. antarctica. In P. antarctica, rates were higher in the summer. Calanus propinquus showed a higher metabolic rate in the summer than in the winter, but the difference was not significant at the 0.05 level. It was concluded that copepods near the ice zone in the ice zone in the Antarctic rely on the spring ice-edge bloom for growth and completion of their life cycle.     

Spring bloom succession,grazing impact and herbivore selectivity of ciliate communities in response to winter warming

This study aimed at simulating different degrees of winter warming and at assessing its potential effects on ciliate succession and grazing-related patterns. By using indoor mesocosms filled with unfiltered water from Kiel Bight, natural light and four different temperature regimes, phytoplankton spring blooms were induced and the thermal responses of ciliates were quantified. Two distinct ciliate assemblages, a pre-spring and a spring bloom assemblage, could be detected, while their formation was strongly temperature-dependent. Both assemblages were dominated by Strobilidiids; the pre-spring bloom phase was dominated by the small Strobilidiids Lohmaniella oviformis, and the spring bloom was mainly dominated by large Strobilidiids of the genus Strobilidium. The numerical response of ciliates to increasing food concentrations showed a strong acceleration by temperature. Grazing rates of ciliates and copepods were low during the pre-spring bloom period and high during the bloom ranging from 0.06 (Δ0°C) to 0.23 day⁻¹ (Δ4°C) for ciliates and 0.09 (Δ0°C) to 1.62 day⁻¹ (Δ4°C) for copepods. During the spring bloom ciliates and copepods showed a strong dietary overlap characterized by a wide food spectrum consisting mainly of Chrysochromulina sp., diatom chains and large, single-celled diatoms. Priority programme of the German Research Foundation—contribution 4.     

Life history and production of Agapetus quadratus (Trichoptera: Glossosomatidae) in a temporary, spring-fed stream

1. The life history and trophic basis of production of the caddisfly grazer Agapetus quadratus were studied in the torrent Gorg Blau, a spring‐fed stream on the island of Majorca that dries annually during summer. 2. Quantitative random samples were taken every 2–3 weeks during an annual surficial flow period, from November 2000 to mid‐July 2001. Instars of field‐collected larvae were determined by measurements of head width and pronotum length, and the sex of all pupae was determined to study sexual dimorphism and sex ratio. 3. Stage‐frequency histograms suggested a trivoltine population, with an average cohort time of 4 months. Larval development was asynchronous, with continuous growth and overlapping generations. Recruitment peaks were identified in mid‐November, early March and late June, indicative of winter, spring and summer generations. On average, females were larger than males and the mean sex ratio was 2 : 3 (females : males). Population densities and biomasses derived from the field data were used to calculate production and turnover rate. 4. Annual production of A. quadratus in the torrent Gorg Blau (4.80 g dry mass m^?2 year^?1) was the highest ever reported for the genus, being comparable with that estimated for some insects with rapid development and multiple cohorts. 5. Estimates of production of A. quadratus were combined with foregut content analysis to estimate the fraction of total production derived from the principal food sources: algae and organic detritus. Algae supported a major proportion of the production of this grazer. 6. The low density of predators characteristic of many temporary streams, and the small amplitudes in discharge and temperature during most of the wet period that characterise the spring habitats might allow high levels of grazer production in this particular Mediterranean stream.     

Patterns of energy storage in Pseudoboeckella poppei (Crustacea,copepoda) from two contrasting lakes on Signy Island,Antarctica

The copepod Pseudoboeckella poppei (Daday) (Calanoida, Centropagidae) was sampled from Sombre and Heywood Lakes on Signy Island, Antarctica (60° S, 45° W) between January 1984 and March 1985. Sombre Lake is clear and oligotrophic with little phytoplankton and a bottom sediment low in organic content. By contrast Heywood Lake is turbid and mesotrophic; a substantial phytoplankton develops in summer and the bottom sediments are comparatively rich in organics. Both lakes freeze over for much of the year, forcing the copepods to adopt a benthic feeding strategy over winter. Adult Pseudoboeckella feed on phytoplankton when this is available, but also on detritus, diatoms and short algal filaments stirred up from the sediment. In Heywood Lake, male copepods show a smooth seasonal trend in lipid content with lipid being synthesised in early summer and utilised in late summer and winter. The summer increase in lipid content is associated with an increase in dry weight. Female lipid contents show evidence of two peaks of egg production. In Sombre Lake both male and female copepods increase in size during summer and show a wider range of lipid contents than in Heywood Lake; it is likely that this is due to the poorer winter feeding conditions which necessitate the synthesis of a much larger store of reserves during the summer. In contrast to marine calanoid copepods, lipid stores are exclusively triacylglycerol with no trace of wax ester.     

Parasitic regulation in a population of Boeckella hamata Brehm (Copepoda: Calanoida)

SUMMARY. 1. The impact of parasitism by a fungus Aphanomyces sp. (Phycomycetes: Saprotegniaceae) on a population of Boeckella hamata in Lake Mahinerangi in New Zealand was studied over 20 months. The incidence of infection among adult female copepods exceeded 12% throughout winter (May-October) and was highest in autumn (46%) when population birth rates were depressed by 48.8%. During most of the study, parasitism lowered the rates of population recruitment and growth by more than 10%. These effects on the B. hamata population exceeded those of Aphanomyces on Boeckella dilatata Sars in another study.
2. It is hypothesized that the high incidence of parasitism by Aphanomyces and its importance at times in regulating populations of Boeckella in New Zealand lakes may be an ecological consequence of life in predator-poor ecosystems.     

Population structure and reproductive biology of Pygospio elegans (Polychaeta: Spionidae) on an intertidal sandflat, Firth of Forth, Scotland

Abstract. The population structure and reproductive biology of a population of the spionid polychaete Pygospio elegans was studied at Drum Sands, Firth of Forth, Scotland, between March 1997 and February 1998. P. elegans was numerically dominant at the site and exhibited a population density increase through the sampling period. The population showed a seasonal pattern with two acute periods of recruitment during the year, one in spring and one in winter. Planktotrophic larvae were the only larval type produced throughout the sampling period. Larvae were released at the 3–5 setiger stage and settled at a size of 18–22 setigers. The spring recruits grew at a rate of 23 setigers per month, reaching sexual maturation size during September of the same year. The adult sex ratio was female-biased throughout the study period (mean = 2.99:1 ± 0.51). When female:ripe male sex ratios were highest (summer and fall), the proportion of females brooding larvae decreased suggesting that sexual reproduction is seasonal. Asexual reproduction was not observed in the population of P. elegans on Drum Sands. These findings suggest that a high degree of variability is a trait of the reproductive biology of P. elegans. .     

Seasonality of birth and sex ratio in Tehran, Iran

234 218 births (1975-1983) from a hospital in Tehran have been analysed for seasonality of birth and sex ratio. The results show more births in winter and less births in fall. The sex ratio is higher in summer and lower in fall. The occurrence of multibirths shows higher twinning and triplet rates in spring and fall, respectively, whereas lower twinning and triplet rates occur in winter and summer, respectively. It could be demonstrated that geographical factors influence the rates of births and multibirths as well as the sex ratio.     

Morphological and molecular analysis of centropagids from the high Andean plateau (Copepoda: Calanoidea)

In small metazoan invertebrates classical taxonomic analyses can be ambiguous due to the limited number of morphological characters available. This difficulty can yield incorrect estimates of species richness or taxa distribution. The Boeckella genus has been described as the dominant taxon of zooplankton assemblages in the Andean biogeographical region. In this genus, taxonomic classification and delimitation of boundaries between species has long been problematic and controversial. Among South American centropagids Boeckella gracilipes has been regarded as one of the most broadly distributed species, its presence having been reported from Ecuador to Tierra del Fuego. However, in the high Andean plateau some centropagid populations identified as Boeckella gracilipes have also been considered as B. gracilipes titicacae or even identified as a different species, namely Boeckella titicacae. In an attempt to resolve the taxonomic status of the Centropagidae family from the high Andean plateau, we combined traditional and multivariate morphological analyses (integral approach) with the molecular phylogenetic approach. The results obtained allow us to conclude that centropagids collected from the high Andean plateau actually represent a different species, B. titicacae, not B. gracilipes. The phylogenetic reconstruction of the South American Centropagidae family indicated that B. gracilipes represents a sister taxon to B. titicacae. The present study stresses the usefulness of integrating alfa-taxonomy with morphometric and molecular approaches in order to resolve species boundaries, to determine geographical distributions and to investigate evolutionary processes.     

Breeding biology of Tenagomysis tasmaniae Fenton,Anisomysis mixta australis (Zimmer) and Paramesopodopsis rufa Fenton from south-eastern Tasmania (Crustacea: Mysidacea)

The number, size and developmental stage of young in the brood pouch of female Tenagomysis tasmaniae, Anisomysis mixta australis and Paramesopodopsis rufa was recorded throughout the year. Breeding was intensive from spring till the end of autumn for the three species. Calculation of the egg ratio for each species showed that their major reproductive peaks occurred during spring and summer. A winter depression in the breeding cycle was observed for T. tasmaniae and P. rufa, but A. mixta australis ceased breeding during winter. Seasonal variation in the length of gravid females and number of young carried was evident for these three species. Females were longer in spring and summer and carried more young than in autumn and winter. A linear relationship between female length and brood size was demonstrated for each species; annual and seasonal equations were calculated for females carrying each developmental stage. The seasonal equations showed that for a female of given length fecundity was greater during spring than any other season. Natality was estimated to be highest during late spring, summer and early autumn for the three species. No seasonal variation in the size of eggs was evident for the three species. The reproduction pattern of T. tasmaniae, A. mixta australis and P. rufa appears to be very similar to that reported for the majority of iteroparous coastal temperate mysids throughout the world.     

Sex, sex-ratios, and the dynamics of pelagic copepod populations

I examine how the population biology of pelagic copepods depends on their mating biology using field data and a simple demographic model. Among calanoid copepods, two distinct patterns emerge. Firstly, copepods that lack seminal receptacle and require repeated mating to stay fertilized have near equal adult sex ratios in field populations. Winter population densities are orders of magnitude less than the critical population density required for population persistence, but populations survive winter seasons as resting eggs in the sediment. Population growth in these species is potentially high because they have on average a factor of 2 higher egg production rates than other pelagic copepods. Secondly, other copepods require only one mating to stay fertile, and populations of these species have strongly female-skewed adult sex-ratios in field populations. Resting eggs have not been described within this group. Winter population sizes are well predicted by the critical density required for population persistence which, in turn, is closely related to the body-size-dependent mate-search capacity. Thus, the different requirements for mating lead in the first case to a more opportunistic reproductive strategy that implies rapid colonization of the pelagic during productive seasons, and in the second case to a strategy that allows maintenance of a pelagic populations during unproductive seasons. Positive density dependent population growth during periods of low population density (‘Allee effect’) amplifies population density variation during winter into the subsequent summer, thus explaining why summer densities appear to depend more on winter densities than on current growth opportunities in pelagic copepods.     

Calanoid copepod grazing on phytoplankton: seasonal experiments on natural communities

Calanoid copepods are major components of most lacustrine ecosystems and their grazing activities may influence both phytoplankton biomass and species composition. To assess this we conducted four seasonal, in situ, grazing experiments in eutrophic Lake Rotomanuka, New Zealand. Ambient concentrations of late stage copepodites and adults of calanoid copepods (predominantly Calamoecia lucasi, but with small numbers of Boeckella delicata) were allowed to feed for nine days on natural phytoplankton assemblages suspended in the lake within 1160 litre polyethylene enclosures. The copepods reduced the total phytoplankton biomass of the dominant species in all experiments but were most effective in summer (the time of highest grazer biomass) followed by spring and autumn. In response to grazing pressure the density of individual algal species showed either no change or a decline. There were no taxa which increased in density in the presence of the copepods. The calanoid copepods suppressed the smallest phytoplankton species (especially those with GALD (Greatest Axial Linear Dimension) < µm) and there appeared to be no selection of algae on the basis of biovolume. Algal taxa which showed strong declines in abundance in the presence of the copepods include Cyclotella stelligera, Coelastrum spp., Trachelomonas spp., Cryptomonas spp., and Mallomonas akrokomos. Calanoid copepods are considered important grazers of phytoplankton biomass in this lake. The study supports the view that high phytoplankton:zooplankton biomass ratios and large average algal sizes characteristic of New Zealand lake plankton may, at least partly, be caused by year round grazing pressure on small algae shifting the competitive balance in favour of larger algal species.     

Feeding rates and productivity of the copepod Acartia bifilosa in a highly turbid estuary; the Gironde (SW France)

Acartia spp. are the dominant copepod species in the Gironde estuary, seaward of the turbidity maximum area. Acartia bifilosa develop a large population in spring and early summer whereas Acartia tonsa appear in late summer. High values and high variability of chlorophyll a/suspended particulate matter ratio are found seaward of the turbidity maximum area. Feeding rates of A. bifilosa were measured by fluorometry. Phytoplankton ingestion was found to be highly variable, between 8 to 80% of copepod carbon body weight. Except for adult females, copepods were heavier in summer than in winter. PB ratios, estimated by the instantaneous growth rate method, varied from 0.03 d^–1 to 0.14 d^–1. The gut contents and P/B ratios of Acartia bifilosa were related to chl a/SPM ratio. From those data, and a few obtained for A. tonsa, it is concluded that only in summer months phytoplankton ingestion is enough to maintain secondary production.     

The Limnology of a New Zealand Reservoir,with Particular Reference to the Life Histories of the Copepods Boeckella propinqua SARS and Mesocylops leuckarti CLAUS

The Auxiliary Nihotupu Reservoir, near Auckland, New Zealand, was studied during 1967-68. The reservoir was emptied for a short time during the study. Changes in physico-chemical factors, phytoplankton and zooplankton were followed throughout the year. Thermal stratification was evident over the summer and the partitioning of oxygen and pH at this time is discussed. An acid heterograde pH curve is reported. From an analysis of oxygen deficits it is suggested that the reservoir may be oligotrophic. An analysis of egg production, length distributions and changes in developmental stages was used in interpreting the life cycle of Boeckella propinqua. Development times from egg to adult, and approximate survival rates are briefly considered. Mesocyclops leuckarti is also discussed and the annual cycles of both copepods are compared with those in other New Zealand lakes.     

Dormancy, dispersal and the survival of cyclopoid copepods (Cyclopoida, Copepoda) in a lowland floodplain

1. The survival of cyclopoid copepods was investigated in a floodplain for 2 years where flooding occurred during the cold season. The cyclopoid community was studied in three waterbodies with distinct hydroperiods: a permanent pond connected to the flooded area during inundation, a temporary pool that is part of the flooded area and an isolated temporary pool.
2. Field studies, including data obtained from samples of water, sediment and soil, showed the overall predominance of species with a summer diapause over those with a winter diapause or without diapause. Emergence of cyclopoid copepods at the onset of flooding, examined using emergence traps and data from recently filled or still isolated temporary pools, showed that some species can survive several months of drying.
3. The ability of the diapausing fourth copepodid stages of Cyclops strenuus and C. insignis , the two cyclopoids most abundant during winter and spring flooding, to survive terrestrial conditions was tested in laboratory experiments. Both species survived for several months, but rates differed among the species. A higher percentage of C. strenuus survived for a longer period, possibly explaining why this species was relatively more abundant in more temporary habitats.
4. Both dormancy and dispersal facilitated survival of cyclopoid copepods in transient habitats connected to each other during flood periods. Dormancy was the most important survival strategy, whereas dispersal could be more important following prolonged periods without flooding.     

Seasonal changes in zooplankton abundance in the lower Rhine during 1987–1991

Zooplankton in the River Rhine was surveyed for five years at the Dutch sampling stations, Lobith (German/Dutch border) and Maassluis (at the point of discharge of the river into the North Sea). The zooplankton abundance showed an apparent seasonal pattern at both stations, characterized by low densities during the winter period, and higher densities during the summer period, with a spring peak. Zooplankton was dominated by rotifers at both stations, although during the winter periods the contribution of copepods was considerable. The rotifers were dominated byBrachionus angularis, B. calyciflorus, Keratella cochlearis andK. quadrata; the copepods by cyclopoid nauplii; the cladocerans by small-sized species mainly belonging toBosmina. At Maassluis the relative contribution of copepods was higher than at Lobith. Furthermore, the zooplankton at Maassluis included the speciesEurytemora affinis, characteristic for estuarine conditions. In spring, the rotifer density and water temperature and rotifer density and chlorophylla concentration were positively correlated. Furthermore, both rotifer density and chlorophylla were inversely correlated with discharge. The possible role of environmental factors (water temperature, chlorophyll content, discharge and biotic factors) controlling the river zooplankton dynamics is indicated.     

Temporal variations in the diet of pearly razorfish Xyrichtys novacula (Osteichthyes: Labridae)

The diet of pearly razorfish Xyrichtys novacula, caught monthly along the shores of the Island of Lampedusa, appeared to be mainly composed of crustaceans, followed by colonial ascidians, molluscs and polychaetes. Among prey, sand dwellers and phanerogam‐associated species were recorded. In winter months, the diet was characterized by a small number of prey items, dominated by colonial ascidians, while in spring and summer a wider prey array was recorded. Dietary indices show that X. novacula do not strictly exploit benthic prey but also pelagic organisms, such as copepods. This feeding behaviour reached its peak in March and October, when the abundance of primary consumers was at its highest after phytoplankton blooms. Furthermore, X. novacula caught prey organisms according to their availability and seasonal patterns during their life cycles, irrespective of fish size.     

Reproductive biology of Philodryas patagoniensis (Snakes: Dipsadidae) in south Brazil: male reproductive cycle

The male reproductive cycle of Philodryas patagoniensis in south Brazil was described through morpho‐anatomical and histological analysis of individuals deposited in zoological collections. Spermatogenesis occurred during late autumn–winter (June–September) and spermiogenesis occurred in spring–summer (October–March). The volume of the testes was smaller (quiescent) in winter, while the tubular diameter and the epithelial height of the seminiferous tubule were larger in summer (January–March). The ductus deferens presented spermatozoa all over the year and had no seasonal variation in diameter. The length of the kidney was larger in winter–spring (July–December), although the tubular diameter and epithelium height of the sexual segment of the kidney (SSK) were larger only in winter (July–September). Total testicular regression was observed in late autumn (May), simultaneously with the peak in SSK. Therefore, at the individual level, males exhibit a discontinuous cyclical reproduction. Considering the population level, the reproductive cycle is seasonal semisynchronous, with most of the individuals showing a reproductive peak in spring–summer (October–March). Here, we present evidence to support the importance of the microscopic approach to reproductive cycle studies. Finally, we discuss the intrinsic and extrinsic factors influencing P. patagoniensis reproductive patterns.     

Sex steroids, growth and condition of Arctic charr broodstock during an annual cycle

Changes in plasma concentrations of sex steroids, growth rate and condition of repeat spawning (3+) male and female Arctic charr were studied throughout an annual reproductive cycle. Individually marked fish (mean weight approx. 500 g) were held under conditions of liberal food supply, constant temperature (4° C) and simulated natural photoperiod (Tromsø, 70° N). Once each month fish were weighed, measured and blood samples taken for steroid analysis. Plasma concentrations of testosterone (T), 11-ketotestosterone (11-KT) and oestradiol-17β (E₂) were determined using radioimmunoassay (RIA). Both male and female fish displayed distinct seasonal changes in plasma concentrations of sex steroids, growth rate and condition. From February (minimal concentrations) to March all sex steroids increased slightly and these elevated concentrations were maintained until May. Thereafter, there was a second, and far more pronounced, increase in plasma steroid concentrations which culminated in peak steroid concentrations in September–October. There was then a rapid decline during the spawning period. In winter, growth rate and condition were generally low, then increased during the spring, reached a peak during the summer, and then declined with the onset of autumn. During spring (March–May), the frequency distributions of plasma testosterone concentrations in both male and female fish were bimodal. The fish of the upper modal group of the distribution had significantly higher growth rates and condition than those in the lower modal group. In summer and early autumn (June–September) the association between T and growth rate changed. Significant negative correlations between T and growth rates were observed in females. There was an increase in endocrine activity, indicated by elevated plasma sex steroid concentrations in March, 7–8 months prior to maturation. It is suggested that this may be one factor influencing the onset of spring growth and energy deposition among maturing charr.