Breeding seasonality among long-tailed macaques,Macaca fascicularis,in Peninsular Malaysia

The females among more than 7000 long-tailed macaques, Macaca fascicularis,that had recently been trapped for biomedical research were examined for pregnancy. The animals were taken from the wild in the north of Peninsular Malaysia during the period 1978 to 1979 and in the south from 1978 to 1982. Both samples showed some pregnancies in all months of the year and a peak during the months of May to July. Additional peaks in other months in some years showed that, while the midyear pregnancy peak was significant, it was not as marked a phenomenon as is the breeding peak in many other species. This might be due to the complexity and unpredictability of both the climate and the macaques’ food supply in Peninsular Malaysia. The additional peaks also illustrated the pitfalls of generalizing about breeding seasonality on the basis of small samples.     

Abundance dynamics and functional role of predaceous <Emphasis Type="Italic">Leptodora kindtii</Emphasis> in the Curonian Lagoon

The abundance and distribution of predatory cladoceran Leptodora kindtii was investigated in the estuarine lagoon (Curonian Lagoon, SE Baltic Sea). Three hydrodynamically different parts of the lagoon were selected, representing transitory oligohaline, intermediate freshwater and stagnant freshwater sites. L. kindtii was least abundant at the oligohaline site, never occurring at salinities greater than 4 psu. At the two freshwater sites, the abundance of L. kindtii varied from a low of <0.1 up to 2.2 indv. L⁻¹ during peak abundance. Two peaks of L. kindtii abundance were observed with timing differences between stations: at the stagnant site the population of L. kindtii peaked two weeks earlier relative to the more hydrodynamically active sites, likely due to a 2°C higher May temperature. The small body size of L. kindtii in the lagoon (seasonal mean 2.68±0.6 mm) shows high fish predation pressure and predicts small cladocerans, juvenile copepods and rotifers being in the preferred prey size range. The calculated L. kindtii daily consumption during the population peak was as high as 100% of the daily zooplankton production, which implies high potential of this predator to shape the grazing zooplankton community in the lagoon.     

Composition,size, and biomass of zooplankton in large productive Florida lakes

Crustacean zooplankton data were compiled from long-term observational studies at seven large shallow Florida lakes, to determine whether there are general characteristics in regard to species composition, body size, and biomass. In particular, we examined whether patterns in body size and species richness fit empirical models developed by Stanley Dodson. The lakes included range in size from 125 to 1730 km² and encompass mesotrophic to hyper-eutrophic conditions. We found that zooplankton biomass was strongly dominated by one species of calanoid copepod—Arctodiaptomus dorsalis. Large daphnids were absent, and Cladocera assemblages were dominated by small taxa such as Ceriodaphnia, Chydorus, and Eubosmina. The total number of species of pelagic cladocerans (8–12) was consistent with Dodson’s predictions based on lake area. The average size of crustacean zooplankton in Florida lakes is small in comparison with temperate communities. A. dorsalis is the smallest calanoid copepod in North America, and the mean length of Cladocera (0.6 mm) is consistent with Dodson’s results that size decreases from temperate to tropical zones. Total biomass of crustacean zooplankton was very low, ratios of zooplankton to phytoplankton biomass (0.01–0.1) are among the lowest reported in the literature, and the zooplankton displayed short-lasting early spring peaks in biomass. Cladocera were almost entirely absent in spring and summer. Factors known to occur in Florida lakes, which appear to explain these characteristics of biomass, include intense fish predation and high summer water temperature.     

Climate change in Norway: warm summers limit grouse reproduction

Grouse and vole numbers may peak after peaks in the seed crop of bilberry (Vaccinium myrtillus) because of reduced levels of feeding deterrents in bilberry plants. We predicted that grouse reproduction depends also on summer (June–September) temperatures in the 2 previous years, because bilberry plants will be less exhausted after a high seed crop in or after warm summers, and thus rebuild their chemical defence more quickly. After berry peak years, population indices of capercaillie (Tetrao urogallus) and bank vole (Myodes glareolus) in southern Norway were negatively related to summer temperatures in the previous year or previous 2 years. Willow grouse (Lagopus lagopus) chick production in five areas in Norway was negatively related to summer temperatures in the 2 previous years when controlling for vole density. A similar pattern was found for the bilberry-feeding moth (Eulithis populata), an important prey for grouse chicks. In eastern Norway, autumn densities of capercaillie and black grouse (Tetrao tetrix) were more likely to peak in vole peak years at high altitudes, where summer temperatures are low. We conclude that high summer temperatures may limit grouse reproduction through the effect on bilberry plants and that a warm climate thus adversely affects population levels of grouse.     

Seasonal dynamics of phytoplankton in the Gulf of Aqaba,Red Sea

Seawater samples were collected biweekly from the northern Gulf of Aqaba, Red Sea, for Phytoplankton analysis during the period May 1998 to October 1999. Microscopic counts and HPLC methods were employed. Procaryotic and eucaryotic ultraplankton dominated throughout most of the year, with larger nano- and microplankton making up only 5% of the photosynthetic biomass. Moderate seasonal variations in the 0–125 m integrated Chl a contrasted with a pronounced seasonal succession of the major taxonomic groups, reflecting the changes in the density stratification of the water column: Prochlorococcus dominated during the stratified summer period and were almost absent in winter. Chlorophyceae and Cryptophyceae were dominant during winter mixing but scarce or absent during summer. Diatoms and Synechococcus showed sharp and moderate biomass peaks in late winter and spring respectively, but remained at only low Chl a levels for the rest of the year. Chrysophyceae, Prymnesiophyceae and the scarce Dinophyceae showed no clear seasonal distribution pattern. The implications of alternating procaryotic and eucaryote dominated algal communities for the Red Sea pelagic food web are discussed. Electronic Supplementary Material Electronic supplementary material is available for this article at and accessible for authorised users.     

Regulation of rotifer densities by crustacean zooplankton in an oligotrophic montane lake in British Columbia

Summary To examine the relative demographic effects of predation and competition for food in rotifers during spring and summer in an oligotrophic lake, predator and competitor densities and food supplies were experimentally altered inside large enclosures. Abundances of rotifer species were positively correlated with experimental densities of fourth instar Chaoborus trivittatus larvae, a major crustacean predator in 1976. Experimental alteration of the densities of Daphnia rosea and Diaptomus leptopus and D. kenai in 1978 produced highly significant increases in rotifer biomass only under Daphnia removal, but not under copepod removal. Inorganic fertilizer additions to enclosures in 1978 and 1979 revealed minimal rotifer increases unless pulsed additions were large or Daphnia were also excluded. Large demographic reponses of rotifers to low fertilizer loadings in the absence of Daphnia confirmed the pre-eminence of competitive food limitation in producing rotifer scarcity in summer.     

Zooplankton variability related to environmental changes in a eutrophic coastal lagoon in the Po Delta

Zooplankton variability in a lagoon of the Po Delta, the Sacca di Goro, was studied in relation to changes in its hydrodynamic and hydrological features. From 1987 to 1992 the lagoon was affected each summer by severe anoxia due to the decomposition of dense and widespread beds of Ulva rigida. In August 1992 a canal was opened through the sand bank closing off the lagoon from the sea in order to improve the water circulation. This hydraulic intervention led to significant environmental changes in the lagoon: during the two subsequent years the summer dystrophic crises were brief and less severe, due to a sharp decrease in the Ulva cover. At the same time a clear increase in phytoplankton biomass values was observed. Following the opening of the channel, the most remarkable change in zooplankton was a significant density increase of calanoids, particularly of Acartia tonsa.     

Ecology of the zooplankton of a meromictic antarctic lagoon with special reference to Drepanopus bispinosus (Copepoda: Calanoida)

The zooplankton of Burton Lake, a meromictic lagoon with a tenuous connection to Crooked Fjord near Davis, Antarctica, was studied during the summer of 1981, when the lake was still ice-covered. Sampling occurred through a 10 cm diameter ice-hole. A plankton pump was used to investigate vertical changes in population density. A newly designed collapsible net was hauled vertically through the entire oxylimnion (6.5 m thick) to examine temporal changes in population structure and in the species composition of the zooplanktonic community. Four metazoan species occurred in the zooplankton: Drepanopus bispinosus and Paralabidocera antarctica (Copepoda), Rathkea lizzioides (Anthomedusae) and a cydippid ctenophore. Except for R. lizzioides, the lower limit of the vertical distributions occurred precisely at the oxycline at a depth of 8.15 m. The densities of both R. lizzioides and the ctenophore were low throughout the depth range 5.0–7.25 m and considerably higher above and below this stratum. The density of R. lizzioides increased greatly in a downward direction starting about 35 cm above the oxycline, and many individuals were capable of making excursions of about 10 cm downward past the chemocline into anoxic water. D. bispinosus had a more even vertical distribution than the medusa and ctenophore, but densities were higher in that portion of the oxylimnion lying below 6.0 m than above. The vertical distributions of R. lizzioides and D. bispinosus were stable over 24 h, but vertical migration occurred in that portion of the ctenophore population located above the low-density stratum. P. antarctica occurred in very small numbers. The ctenophore was uniformly distributed horizontally, but significant horizontal differences (among four stations) in mean vertical density occurred in adult D. bispinosus and R. lizzioides. The lowest density of D. bispinosus occurred at the station with the highest density of R. lizzioides. In early summer (11 December 1981), the mean densities of zooplankters within that portion of the oxylimnion overlying the anoxylimnion were D. bispinosus 550 m⁻³, ctenophore 410 m⁻³ and R. lizzioides 230 m⁻³. Throughout summer the absolute density and relative community proportion of R. lizzioides fell continuously, and these measures showed a comparable decline for adult D. bispinosus (almost entirely females). At the same time there was a consistent increase in the numbers of immature D. bispinosus. The density and community percentage of the ctenophore remained high (c. 480 m⁻³ and 44%, respectively) throughout December and early January, but declined thereafter. D. bispinosus is univoltine. Mature males appear in late autumn and mating occurs mainly in early winter, when significant numbers of mature females first appear. The species overwinters as adults, and ovigerous females first appear in midwinter. Males are short-lived and are already rare by the end of winter. Stage I nauplii are present in early spring, and naupliar development continues throughout spring and into early summer in company with an almost entirely female adult population. Stage I copepodites first appear in December and reach peak numbers in January. The last of the adult females from the previous winter disappear by late summer, but development of the immature copepodites continues into autumn. Females may store sperm for up to six months. Highly significant dry weight-length regressions were obtained for D. bispinosus. Mean clutch size for this species remained constant at about 24–25 eggs. The mean summer standing crop of total zooplankton was almost 80 mg (dry weight) m⁻³. This value is high compared with those of most other marine waters. The structure and composition of the zooplanktonic community of the lagoon were quite distinct from those of the fjord with which water was exchanged during January and February. The summer standing crop of the lagoon was substantially greater than that of the fjord in terms of both unit surface area and unit volume of water. A tentative food web for the lagoon is presented.     

Mesozooplankton beneath the summer sea ice in McMurdo Sound,Antarctica: abundance,species composition,and DMSP content

The summer Phaeocystis antarctica bloom increases under-ice phytoplankton biomass in McMurdo Sound, Antarctica. The magnitude of mesozooplankton grazing on this bloom is unknown, and determines whether this production is available to the pelagic food web. We measured mesozooplankton abundance and body content of dimethylsulfoniopropionate (DMSP) during the McMurdo Sound austral summer (2006 and 2006–2007). Abundance varied from 20 to 4,500 ind. m⁻³ (biomass 0.02–274.0 mg C m⁻³), with peaks in mid-December and late-January/February. Abundance was higher but total zooplankton biomass lower in our study compared to previous reports. Copepods and the pteropod Limacina helicina dominated the zooplankton in both abundance and biomass. DMSP was detected in all zooplankton groups, with highest concentrations in copepod nauplii and L. helicina (95 and 54 nmol mg⁻¹ body C, respectively). Experiments suggested that L. helicina obtains DMSP by directly grazing on P. antarctica, which often accumulates to high biomass under the summer sea ice in McMurdo Sound.     

Life cycle,population dynamics,growth and production of <Emphasis Type="Italic">Abra segmentum</Emphasis> (Mollusca,Bivalvia) at low salinities in a Mediterranean lagoon

Aspects of the biology of Abra segmentum were investigated at low salinities in a Mediterranean coastal lagoon (Monolimni Lagoon, Northern Aegean Sea). Monthly samples were collected during the period from February 1998 to January 1999. Recruitment occurred from mid-spring to early autumn (0.3–5.7 psu) and recruits grew during summer and autumn (1.2–5.7 psu), while a major part vanished during next autumn, displaying a maximum life span of about 20 months. A positive correlation was found between the percentage of individuals having a shell length of ≤3.5 mm and temperature; age group 0 showed a growth rate of 0.97 mm per month, and the largest individual collected had a 19.76 mm shell length. The population density sharply increased during late spring (0.3–1.2 psu); this increase was followed by a decline during summer and, afterwards, a gradual increase up to late autumn. Secondary production calculated by the size–frequency method gave a mean annual density (n) of 3,357 individuals m⁻², a mean annual biomass (B) of 21.98 g DW m⁻², an annual production (P) of 73.72 g DW m⁻² and a P:B ratio of 3.35. A comparison of the present data with available data of A. segmentum populations from higher salinity habitats revealed that this bivalve in the study area showed a life history pattern similar to that of other populations of the species and a comparatively high growth rate, maximum body size, n, B, P and P:B ratio. Our findings suggest that the studied aspects of A. segmentum biology could not be markedly affected by low salinities.     

Evidence of a planktonic food web response to changes in nutrient input dynamics in the Mar Menor coastal lagoon,Spain

Nutrient input dynamics in the Mar Menor coastal lagoon has recently changed as a consequence of changes in agricultural practises. An interannual comparison of the environmental variables and the planktonic biomass size-spectra was performed between 1988 and 1997. While nitrate concentration was low in 1988, the values in 1997 increased considerably. Since 1995, two alloctonous jellyfish species (Rhyzostoma pulmo and Cotylorhiza tuberculata) occurred in large numbers in summer time and reached peak abundance in summer of 1997. The size-spectra analysis comparison revealed that, in spite of changes in nutrient input that stimulated the growth of larger phytoplankton cells, there were no significant differences in the spectra slope which followed a similar seasonal trend in both years. However, the plankton biovolume considered under the size range compared (between 2 and 1000 m diameter) was, paradoxically, always lower in 1997. Given that there were higher nutrient levels in 1997, this finding suggest a strong top-down control mechanism of size structure. Gut contents of jellyfishes showed their preference for large diatoms, tintinnids, veliger larvae and copepods, corroborating that size structure in these assemblages can be subject to top-down control. The implication of these results is that the feeding activities of large gelatinous zooplankton (jellyfishes) may play an important role controlling the consequences of eutrophication within the Mar Menor coastal lagoon.     

Life history, aspects of reproductive biology and production of Corophium orientale (Crustacea: Amphipoda) in Monolimni lagoon (Evros Delta, North Aegean Sea)

Monthly samples of Corophium orientale were collected during February 1998–February 1999 in both parts of Monolimni lagoon (0.3 < Sal. < 6 psu, 2 < Temp. < 28.5 °C). Corophium orientaleshowed a semiannual life cycle. In the southern part, breeding occurred from early spring to mid autumn having three peaks, in early spring, early summer and early autumn, and three cohorts were produced, a spring cohort, a summer cohort and an overwintering one. In the shallower innermost northern part of Monolimni lagoon, breeding peaked in mid-spring and, also, in early autumn, and two discrete cohorts, a spring and an overwintering one, were produced. A lack of large individuals, attributed to a temporary size – selective predation by migrating shorebirds, was observed during summer. That lack possibly contributed to the hiatus in reproduction. Photoperiod seemed to be a more important cue than temperature in the initiation of the reproductive cycle, while extremely low salinities (<1 psu) did not deter breeding. Females attained maturity at a smaller size in late spring and, especially, in summer than in early spring at lower temperatures. Brood size was a function of females body size. Females, matured during summer at high temperatures, showed the lowest brood size, body length ratio. Mean brood size was small (12–13 early embryos) and embryo loss during development high (54%), possibly due to a negative effect of low salinities. A 1:1 sex ratio existed in the small body length classes, but females preponderated in the large ones. The spring cohort in the southern part, which developed at moderate temperatures, showed the highest growth rate (40 m day^–1). Secondary production of C. orientalecalculated by Hyness method gave a mean annual density of 4562.5 and 9327.6 ind. m^–2, a mean annual crop (B) of 1.03 and 2.67 g DW m^–2, an annual production (P) of 6.91 and 22.54 g DW m^–2 and a P/B ratio of 6.7 and 8.4 in the northern and southern part of the lagoon, respectively.     

The recovery of a very shallow eutrophic lake, 20 years after the control of effluent derived phosphorus

1. Monitoring at fortnightly to monthly intervals of a very shallow, lowland lake over 24 years has enabled the time course of recovery from nutrient enrichment to be investigated after high external P loading of the lake (>10 g P m^?2 year^?1) was reduced between 1977 and 1980. 2. The lake showed a relatively rapid response during the spring and early summer, with a reduction in phytoplankton biomass occurring after 5 years when soluble reactive phosphorus concentration was <10 μg L^?1. 3. However, during the later summer the response was delayed for 15 years because of sustained remobilisation of phosphorus from the sediment. The greater water clarity in spring and a gradual shift from planktonic to benthic algal growth may be related to the reduction in internal loading after 15 years. 4. Changes in the phytoplankton community composition were also observed. Centric diatoms became less dominant in the spring, and the summer cyanobacteria populations originally dominated by non‐heterocystous species (Limnothrix/Planktothrix spp.) almost disappeared. Heterocystous species (Anabaena spp. and Aphanizomenon flos‐aquae) were slower to decline, but after 20 years the phytoplankton community was no longer dominated by cyanobacteria. 5. There were no substantial changes in food web structure following re‐oligotrophication. Total zooplankton biomass decreased but body size of Daphnia hyalina, the largest zooplankton species in the lake, remained unchanged, suggesting that the fish population remained dominated by planktivorous species. 6. Macrophyte growth was still largely absent after 20 years, although during the spring water clarity may have become sufficient for macrophytes to re‐establish.     

Rare charophytes in Scotland's coastal saline lagoons

Summary

Recently collected data comparing the distribution and performance of the rare lagoonal charophyte Lamprothamnium papulosum across a number of UK coastal lagoon sites, including nine sites in the Western Isles, is presented. The surveys revealed that the species occurred over a wide range of salinities but was absent from sites with high phosphorus concentrations. The sites in the Western Isles held the most extensive populations and were least threatened by nutrient pollution. The discovery of two other endangered and vulnerable charophytes of brackish water, Tolypella nidifica and Chara baltica, at two of the Western Isles sites further highlights the importance of Scotland's saline lagoon resource. The use of the autoecological data on salinity and nutrient conditions for supporting Biodiversity Action Plan implementation for Lamprothamnium papulosum and for saline lagoon conservation in general is considered. Status summaries for all the charophyte species of brackish water recorded in Scotland are also provided.     

Seasonal Variation of the Macrozoobenthic Community Structure at Low Salinities in a Mediterranean Lagoon (Monolimni Lagoon,Northern Aegean)

The macrozoobenthic community structure and dynamics at low salinities (0.3–6 psu) in a Mediterranean lagoon (Monolimni lagoon) were investigated. Samples were collected monthly from February 1998 to February 1999 at two sampling stations. Community structure was analyzed by means of uni‐ and multivariate methods. 21 taxa were collected; the amphipod Corophium orientale and the gastropod Ventrosia maritima dominated the assemblages. Total abundance peaked (50,000–60,000 individuals m^–2) in mid or late autumn. Community structure showed an almost even seasonal periodicity; seasonal changes were mainly derived from the intense variation in abundance of most species and the non–occurrence of a few ones (e.g. Corophium insidiosum, Polydora ciliata) in spring and summer. Non– occurrence, which led to a depression of the most diversity indices, was possibly the only direct impact of the extremely low salinities (~0.3 psu) on community structure. The main structuring factors of the community in the deeper outer part of the lagoon were water temperature and depth, and in the innermost part, where a Ruppia maritima meadow occurred, were water temperature and predation pressure by crabs (Carcinus aestuarii) and gobies (Knipowitchia caucasica). A temporary decline in total abundance in summer followed an increase in abundance of these predators. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Seasonal fluctuations in macrobenthic fauna of theFucus belt in kiel fjord (western baltic sea)

The macrobenthic fauna associated withFucus at a station in the Kiel Fjord was investigated from June 1978 until June 1979. The predominant group in number as well as in biomass were gammarids. They formed, together with the isopodsIdotea spp., approximately 95 % of the total average annual biomass. The total dry weight of all macrobenthic animals (excl.Littorina spp.) increased from 1.9 g per kgFucus in May to about 16 g in June–August, and dropped to 8.3 g in September. Winter average dry weight values were only about 6 % of the summer values.     

The role of Ulva spp. as a temporary nutrient sink in a coastal lagoon with oyster cultivation and upwelling influence

Bahía San Quintín is a coastal lagoon with large Ulva biomass and upwelling influence. Previous observations suggest that Ulva has increased recently as a result of oyster cultivation. To evaluate the possible role of Ulva as a temporary nutrient sink, biomass and tissue C, N, and P were determined seasonally. Maximum biomass was present during spring and early summer (1,413–1,160 t (dry)) covering about 400 ha. However, the biomass decreased to 35 t (dry) by winter. The mean annual percentage of Ulva C, N, and P was 28%, 2%, and 0.14%, respectively. This study shows that Ulva can store up to 28 t of N and 2 t of P in Bahía San Quintín during spring–summer. Ulva may be displacing the seagrass Zostera marina subtidal beds. A partial removal of the seaweed would reduce the risk of eutrophication and the displacement of eelgrass beds.     

Seasonal dynamics of phytoplankton in a lagoon-type lake Izmenchivoye (Southeast Sakhalin)

The seasonal changes in taxonomic structure, dynamics of number, and biomass of phytoplankton in the Izmenchivoye lagoon lake (southeast Sakhalin) were studied. In all, 266 species and intraspecies taxa of microalgae were revealed. The greatest species diversity (according to the Shannon index) was observed in May, August and October (H = 2.76–2.89), the least (H = 0.5–0.86), in April and January of 2006. The monthly average number varied from 997 up to 84 282 cells/l, and biomass from 18.98 up to 878.62 mg/m³. The average annual number of phytoplankton and its biomass were 32 650 cells/l and biomass 172.13 mg/m³ respectively. The maximum number was registered in August, 2005, and maximum biomass was recorded in January, 2006. Winter, spring and summer peaks of number coinciding with those of biomass were registered. For the first time, winter bloom of phytoplankton was registered in inland waters of the Sakhalin Island. In the winter and spring the basic input to formation of the parameters was composed of diatoms; in summer and autumn it was composed of by flagellates (dinophyta and cryptophyta).     

Spatial and seasonal variation of macroalgal biomass in Laguna Ojo de Liebre, Baja California Sur, Mexico

Laguna Ojo de Liebre is part of El Vizcaíno Biosphere's Reserve, one of the largest protected natural areas in the world. The contribution of seaweeds to the lagoons' total biomass had not been previously quantified. The purpose of this study was to evaluate the spatial and temporal variations of seaweed biomass in the lagoon. Seaweed samples were taken every season during 1995 at six sampling stations distributed throughout the lagoon. Total specific biomass of seaweeds was at its peak in the summer, and minimum in spring. The highest total annual biomass was found at Isla Brosa in the lagoon's central portion, and the lowest in El Dátil at the head. The seasonal and spatial variation of biomass in the lagoon is related with species richness and environmental parameters. Potentially important species in terms of biomass, wide spatial and temporal distribution, and potential use were: Spyridia filamentosa, Entheromorpha clathrata, Dasya baillouviana, Hypnea valentiaeand Sargassum sinicola .Using PCA three groups of stations were defined: one chiefly at the lagoon's mouth, another comprised the islands in the central portion, and the last in the lagoon's head.     

Seasonal growth of the submerged macrophyte Ceratophyllum demersum L. in mesotrophic Lake Vechten in relation to insolation,temperature and reserve carbohydrates

Seasonal growth of a Ceratophyllum demersum stand in Lake Vechten, The Netherlands, varied considerably during 1978, '79 and '81. The biomass was highest in 1981 (max. 115 g ash-free dry weight m^–2). The 1978 growth trend was similar to that of 1981 with slightly lower net annual production (max. 92 g AFDW m^–2). The vegetation almost disappeared in 1979 (max. 24 g AFDW m^–2). These large differences in biomass could not be explained by differences in insolation but were mainly attributed to the severe winter of 1978–'79 which slowed plant development and full maturity was attained only after the diurnal insolation had decreased.Reserve carbohydrates were primarily starch and water-soluble sugars. Starch concentrations varied between 71.5 and 24.7 mg g^–1 AFDW in the winter and summer, and total sugar concentrations between 24.9 and 20.2 mg g^–1 AFDW, respectively. The non-reducing sugars tended to predominate in summer and the reducing sugars in winter. The value of these reserves for maintenance, growth and survival was assessed by calculating plant carbon budgets for several seasons.It was concluded that C. demersum relies heavily on its developmental timing and energy-conserving physiological mechanisms to survive extended periods of stress and in these respects is analogous to a typical terrestrial shade plant.