Historical changes in herbaceous wetland distribution induced by hydrological conditions in Lake Saint-Pierre (St. Lawrence River,Quebec, Canada)

Historical changes (1961–2002) in the distribution of herbaceous wetland plant associations were inferred from the hydrological regime of Lake Saint-Pierre, a 312 km² broadening of the St. Lawrence River (Quebec, Canada), to assess the cumulative effects of human interventions and climatic variability. Relative abundance index (height × percent cover) of wetland plants in 630 field quadrats sampled at 13 sites (1999–2002) were used to derive a model predicting the occurrence of nine herbaceous plant classes with a 71% (24–84%) accuracy. Wetland types included seasonally dry (meadows), mudflats and wet (low marshes and submerged) assemblages. Over the 1961–2002 period, the total surface area of Lake Saint-Pierre herbaceous wetlands ranged between 11 (in 1972) and 128 (in 2001) km² and was negatively correlated (Spearman r = –0.86, p < 0.0001) to average water level during the current growing season. Within-season variability and level conditions over the previous season defined 5 marsh assemblages characterized by different species composition, relative abundance and diversity. Significant hydrological variables included quadrat elevation, water depth, number of days flooded and depth variability experienced over the current and/or previous growth seasons. The hydrological model suggests that for a given level, wetland plant assemblages differed markedly whether the multi-year sequence of water levels was rising or falling. Lake Saint-Pierre alternated between three broad-scale wetland configurations, dominated by meadows and open marsh with floating-leaved vegetation (in the 1960s), scattered tall Scirpus marshes (in the 1970s and early 1980s) and closed marsh with aggressive emergents (since 1996). The strong response of Lake Saint-Pierre wetlands to hydrological conditions in the current and previous growth seasons underlines their vulnerability to future water level variations resulting from regulation and climate variability.     

The influence of macrophyte beds on plankton communities and their export from fluvial lakes in the St Lawrence River

1. To determine the influence of macrophyte beds on plankton abundance within fluvial lakes of the St Lawrence River, planktonic components (macrozooplankton, heterotrophic bacteria, and phytoplankton as chlorophyll- a [Chl- a ]) were sampled in Lake St Francis and Lake St Pierre during summer 1998. We tested the hypothesis that the abundance of planktonic components was higher within macrophyte beds in comparison to the more rapidly flushed open water areas of the fluvial lakes.
2. The large cross channel variation in zooplankton biomass was indeed correlated with the presence of dense beds of submerged macrophytes. Total macrozooplankton biomass was nine-fold greater within the beds (mean=180 μg L−1 dry mass) than in either the open water or areas with only sparse vegetation (mean=20 μg L−1 dry mass).
3. Chl- a and heterotrophic bacterial abundance were also higher in the beds, but only slightly so. There was no difference in total phosphorus or dissolved organic carbon concentrations between areas of dense vegetation, sparse vegetation or open water.
4. Macrophyte beds on the margins of the fluvial lakes allow the development of high planktonic abundance relative to the fast flowing central channel. Macrozooplankton biomass was much higher at the outflows of the lakes (∼50 μg L−1 dry mass) in comparison to the inflows (<20 μg L−1 dry mass). The increase is due to the transfer of organisms from submerged macrophyte beds into the central channel in the downstream quarter of the two lakes where the marginal littoral waters enter central channel waters.
5. Along rivers, shallow fluvial lakes appear to act as sources of plankton which is exported downstream during years of extensive littoral macrophyte development.     

Factors affecting the structure of epiphytic gastropod communities in the St. Lawrence River (Quebec,Canada)

A study of epiphytic gastropods associated with two submerged macrophytes (Myriophyllum spicatum and Vallisneria americana) was conducted at a soft and at a hard water site in the St. Lawrence River during two non-consecutive years in order to compare effects of macrophyte species and biomass, site and year in gastropod community structure. There was no effect of macrophyte species on total gastropod abundance nor on gastropod diversity, and few gastropod species showed a marked preference for either macrophyte species. Inter-site and inter-annual variations in diversity, total gastropod abundance and gastropod community structure were greater than variations among macrophytes. However, analysis of variations of epiphytic gastropod communities in one of the two sites shows that abiotic factors are important in explaining epiphytic gastropod distribution. Our results and results of other studies on gastropod population dynamics in the St. Lawrence River indicate that intra- and interspecific competition between gastropods is important on macrophytes and that they must be carefully considered in order to understand epiphytic community structure and population dynamics.     

Delayed gametogenesis and progesterone levels in soft-shell clams (Mya arenaria) in relation to in situ contamination to organotins and heavy metals in the St. Lawrence River (Canada)

Wild carp, Cyprinus carpio, were sampled in January and March 2000 in a section of the Anoia River (NE Spain) known to be polluted by estrogenic compounds. At each sampling time, three groups were distinguished: (1) apparently normal males; (2) apparently normal females; and (3) affected fish. The latter were characterized by the simultaneous development of male and female tissue in their gonads at a macroscopical level (six out of 31 fish sampled at this particular point), or testicular atrophy (three out of 31). Plasmatic and hepatic vitellogenin (VTG) levels and plasma testosterone (T) and estradiol (E2) were measured to observe the particular estrogenic response of the affected fish. Moreover, the response in the xenobiotic metabolizing capacity in liver was tested. This involved the analysis of mixed function oxygenase (MFO) system such as: total cytochrome P450 content, NAD(P)H cytochrome c reductases and the associated CYP1A1, EROD activity. Also, glutathione S-transferase (GST) and UDP-glucuronosyltransferase (UDPGT) as detoxifying enzymes were measured. Our results showed: (1) a highly variable VTG content in all fish groups; (2) an increase in sex hormones content in March for the female group; and (3) an enhanced xenobiotics metabolism in the affected fish group, measured as total cytochrome P450, EROD activity in the January survey and cytosolic GST in March. The observed increase in VTG, sex hormones and in most of the enzymatic activities from January to March that could also be attributed to higher water temperature.     

Wind-driven river plume dynamics and toxic Alexandrium tamarense blooms in the St. Lawrence estuary (Canada): A modeling study

In the lower St. Lawrence estuary (LSLE, eastern Canada), blooms of the toxic dinoflagellate Alexandrium tamarense are a recurrent phenomenon, resulting in paralytic shellfish poisoning outbreaks every summer. A first coupled physical–biological model of A. tamarense blooms was developed for this system in order to explore the interactions between cyst germination, cellular growth and water circulation and to identify the effect of physical processes on bloom development and transport across the estuary. The simulated summer (1998) was characterized by an A. tamarense red tide with concentrations reaching 2.3 × 10⁶ cells L⁻¹ along the south shore of the LSLE. The biological model was built with previously observed A. tamarense cyst distribution, cyst germination rate and timing, and A. tamarense growth limitation by temperature and salinity. The coupled model successfully reproduced the timing of the A. tamarense bloom in 1998, its coincidence with the combined plumes from the Manicouagan and Aux-Outardes (M-O) rivers on the north shore of the estuary, and the temporal variations in the north-south gradients in cell concentrations. The simulation results reveal that the interaction between cyst germination and the estuarine circulation generates a preferential inoculation of the surface waters of the M-O river plume with newly germinated cells which could partly explain the coincidence of the blooms with the freshwater plume. Furthermore, the results suggest that the spatio-temporal evolution of the bloom is dominated by alternating periods of retention and advection of the M-O plume: east or north-east winds favor the retention of the plume close to the north shore while west or north-west winds result in its advection toward the south shore. The response of the simulated freshwater plume to fluctuating wind forcing controls the delivery of the A. tamarense bloom from the northern part of the estuary to the south shore. In addition, our results suggest that a long residence time of the M-O plume and associated A. tamarense population in the LSLE during the summer 1998 contributed to the development of the red tide. We thus hypothesize that the wind-driven dynamics of the M-O plume could partly determine the success of A. tamarense blooms in the LSLE by influencing the residence time of the blooms and water column stability, which in turn affects A. tamarense vertical migrations and growth.     

Exotic plant species of the St Lawrence River wetlands: a spatial and historical analysis

Aim To evaluate the importance (number of species, plant cover) of the exotic flora in seven well‐defined sectors of one of the most important transportation waterways in North America. To determine the impact of exotic species on wetland plant diversity and reconstruct the spread of some invasive species. Location St Lawrence River, southern Québec. Methods The exotic flora (vascular plants) of wetlands bordering the St Lawrence River was studied using 713 sampling stations (25 m²) along a 560‐km long corridor. Results Exotic species represent 13.7% of the vascular flora of the St Lawrence wetlands. The relative plant cover occupied by exotic species is high in some of the fluvial sectors (42–44%), but low (6–10%) in the estuarine sectors. Wetlands (marshes) surrounding islands were particularly susceptible to invasion by exotic plants. Historical, abiotic and landscape factors may explain the differences observed between sites. Purple loosestrife (Lythrum salicaria L.) is the most common exotic species of the St Lawrence wetlands, but other species, namely flowering‐rush (Butomus umbellatus L.) and reed canary grass (Phalaris arundinacea L.) are much more invasive. There is no linear relationship between the exotic species cover and the diversity of wetland plants; low diversity sites can be dominated by either exotic or native plant species. In the other sites, exotic species generally have little impact on plant communities and can contribute to increase diversity. Common reed (Phragmites australis (Cav.) Trin. ex Steudel) and reed canary grass, both considered as exotic species in this study, clearly have a stronger impact on plant diversity than flowering‐rush and purple loosestrife. Main conclusions This study shows that the global impact of an invader cannot be adequately evaluated with only a few highly invaded sites. While nationwide strategies have been developed to control exotic species, large surveys are essential to adapt them to regional particularities.     

Factors influencing biomass and nutrient content of the submersed macrophyte Egeria densa Planch. in a pampasic stream

We identified factors influencing biomass and nutrient content in E. densa in an enriched pampean stream of Argentina. Physical (current velocity, temperature), chemical (pH, conductivity, dissolved oxygen, nutrient content in water and sediments), and biological variables (biomass and nutrient content of E. densa, biomass of periphyton and other macrophytes) were estimated at each sampling occasion, and mean monthly values estimated. Biomass and nutrient content in E. densa were correlated with these physical-chemical and biological variables. Biomass was positively correlated with ammonium in stream water (P<0.05) and sediment total nitrogen (P<0.01). Nitrogen showed a positive relationship with ammonium (P<0.01), and a negative one with nitrate and periphyton biomass (P<0.05). Phosphorus was positively correlated with soluble reactive phosphorus (P<0.01). The growth of other macrophyte species in the stream seemed to influence E. densa biomass, probably through competition for light. Current velocity was low and not significantly related with E. densa biomass, however, a flood at the beginning of the study washed the macrophyte stand downstream.     

Importance of geographic origin for invasion success: A case study of the North and Baltic Seas versus the Great Lakes–St. Lawrence River region

Recently, several studies indicated that species from the Ponto‐Caspian region may be evolutionarily predisposed to become nonindigenous species (NIS); however, origin of NIS established in different regions has rarely been compared to confirm these statements. More importantly, if species from certain area/s are proven to be better colonizers, management strategies to control transport vectors coming from those areas must be more stringent, as prevention of new introductions is a cheaper and more effective strategy than eradication or control of established NIS populations. To determine whether species evolved in certain areas have inherent advantages over other species in colonizing new habitats, we explored NIS established in the North and Baltic Seas and Great Lakes–St. Lawrence River regions—two areas intensively studied in concern to NIS, highly invaded by Ponto‐Caspian species and with different salinity patterns (marine vs. freshwater). We compared observed numbers of NIS in these two regions to expected numbers of NIS from major donor regions. The expected numbers were calculated based on the available species pool from donor regions, frequency of shipping transit, and an environmental match between donor and recipient regions. A total of 281 NIS established in the North and Baltic Seas and 188 in the Great Lakes–St. Lawrence River. Ponto‐Caspian taxa colonized both types of habitats, saltwater areas of the North and Baltic Seas and freshwater of the Great Lakes–St. Lawrence River, in much higher numbers than expected. Propagule pressure (i.e., number of introduced individuals or introduction effort) is of great importance for establishment success of NIS; however in our study, either shipping vector or environmental match between regions did not clarify the high numbers of Ponto‐Caspian taxa in our study areas. Although we cannot exclude the influence of other transport vectors, our findings suggest that the origin of the species plays an important role for the predisposition of successful invaders.     

Hydrological and biological processes modulate carbon,nitrogen and phosphorus flux from the St. Lawrence River to its estuary (Quebec,Canada)

Changes in atmospheric deposition, stream water chemistry, and solute fluxes were assessed across 15 small forested catchments. Dramatic changes in atmospheric deposition have occurred over the last three decades, including a 70% reduction in sulphur (S) deposition. These changes in atmospheric inputs have been associated with expected changes in levels of acidity, sulphate and base cations in streams. Soil retention of S appeared to partially explain rates of chemical recovery. In addition to these changes in acid–base chemistry we also observed unexpected changes in nitrogen (N) biogeochemistry and nutrient stoichiometry of stream water, including decreased stream N concentrations. Among all catchments the average flux of dissolved inorganic nitrogen (DIN) was best predicted by average runoff, soil chemistry (forest floor C/N) and levels of acid deposition (both S and N). The rate of change in stream DIN flux, however, was much more closely correlated with reductions in rates of S deposition rather than those of DIN. Unlike DIN fluxes, the average concentrations as well as the rates of decline in streamwater nitrate (NO₃) concentration over time were tightly linked to stream dissolved organic carbon/dissolved organic nitrogen ratios DOC/DON and DON/TP rather than catchment characteristics. Declines in phosphorus adsorption with increasing soil pH appear to contribute to the relationship between C, N, and P in our study catchments. Our observations suggest that catchment P availability and its alteration due to environmental changes (e.g. acidification) might have profound effects on N cycling and catchment N retention that have been largely unrecognized.     

Postglacial climate in the St. Lawrence lowlands, southern Québec: pollen and lake-level evidence

Pollen and lake-level data are used to reconstruct past climate changes in the St. Lawrence lowlands, southern Québec. Past lake-level changes are assessed from sedimentological, pollen and macrofossil records from a single shallow-water core from Lac Hertel, which lies in the central part of the studied area. Three low lake-level phases are recognised: prior to 8000, 7600-6600 and 4800-3400 cal. BP. The modern analogue method is applied to pollen data from seven well-dated sites from the St. Lawrence lowlands and adjacent mountain areas, constrained and unconstrained by lake-level changes. The reconstructed climate changes are congruent with the pattern of climate changes known from eastern North America: a dry and cold late-glacial episode due to the presence of pro-glacial lakes and seas; a rapid warming between 12?500 and 11?000 cal. BP possibly caused by increasing summer insolation; a dry period from 10?000 to 6500 cal. BP; a brief cooling between 9000 and 8000 cal. BP, possibly related to a summer cooling of Arctic airmasses; a temperature maximum around 8000 cal. BP; and finally, a progressive decrease in summer temperature and an increase in (winter?) precipitation over the 4500 last years. These results show that it is possible to reveal seasonal patterns in climate by combining pollen and lake-level data.     

Distribution and abundance of young fish in the St. Clair River and associated waters,Ontario

Fish larvae were sampled in 1986 in the St. Clair River, and adjacent waters. Species richness (9 taxa as larvae; 4 others as juveniles) and abundance was lowest in the river, where many larvae (e.g., burbot, rainbow smelt, and yellow perch) were in transit from Lake Huron. The most abundant, and localized, species was gizzard shad, which reached a peak mean density of 4600 larvae 100 m^-3 in an agricultural canal. Adjacent waters contribute greatly to the fish communities of the river and adjoining Lakes Huron and Erie, especially in terms of the number and quantity of forage species.     

Effects of pollution on benthic invertebrate communities of the St. Marys River, 1985

A survey was undertaken in 1985 to assess spatial and temporal trends in the benthic community structure in relation to sediment contamination and wastewater sources at 70 stations between Whitefish Bay and lower Lake George in the St. Marys River. Cluster analysis identified seven benthic communities. Three were identified as pollution impacted, based on a preponderance of tubificids and nematodes, usually at high densities (up to 259 000 m^-2), but sometimes at low densities (< 100 m^-2) at individual stations. Impacted communities occurred downstream of industrial and municipal sources and in depositional areas, and were confined mainly to Canadian waters. Unimpacted communities had greater numbers of taxa, and occurred upstream of point sources, along the U.S. shoreline, and in most areas of downstream lakes. Impacted and unimpacted communities were separated along particle size and contaminant gradients in river sediments. Despite recent reductions in pollutant loadings and improvements in sediment quality, no major changes were apparent in the status of the benthic community from earlier surveys.     

Seasonal changes in composition and biomass of epiphytic algae on the macrophyte Ranunculus penicillatus in a chalk stream,with estimates of production,and observations on the epiphytes of Cladophora glomerata

The epiphytic algae on surfaces of the macrophyte Ranunculus penicillatus (Dumort.) Bab. var. calcareus (R. W. Butcher) C. D. K. Cook and on Cladophora glomerata (L.) Kütz growing in the River Itchen at Otterbourne near Southampton were studied between February 1984 and June 1985. The river at this site has a mean flow rate of 0.33 m s^–1, and is about 16 m wide and on average 20 cm deep, with a discharge ranging through the year between 0.34 and 2.46 m³ s^–1. The pH of the river varies little around 8.2, with a mean alkalinity of 236 mg HCO _inf3 ^sup1 l^–1, because of its origin from chalk springs. Ranunculus grows throughout the year, with peaks of biomass in spring and autumn. It forms a very large surface for attachment of epiphytes, and covers on average 40% of the stream bed. Numbers of epiphyte cells removed from Ranunculus ranged through the year between 52 × 10³ and 271 × 10³ cells mm^–2 stream floor, with maximum numbers in April, and a secondary peak in October. This pattern partly reflects fluctuations in the biomass of Ranunculus; the number of cells per unit area of plant surface showed a broader spring peak and lower fluctuations in other seasons. Diatoms formed 65 to 98% of these epiphyte cells, with chlorophytes reaching their peak (10%) in summer and cyanophytes (25%) in autumn. Estimates of biomass of these epiphytic forms, derived from measurements of chlorophyll c, indicate a range between 30 and 100 g dry weight m^–2 of weed bed. Colonisation studies showed that the algae grow and reproduce throughout the year, with a mean generation time of about 5 days, suggesting an annual production of about 3 kg dry weight m^–2 of weed bed, which makes epiphytic algae the principal primary producers in the stream. The numbers and biomass of epiphytic algae on Cladophora are considerably less.The species of epiphytic algae found on Ranunculus were generally similar to those growing on Cladophora, and to epilithic algae on pebbles of the stream bed, but different species were dominant on the different substrata. Algal cells in the water column were all derived from benthic habitats, although their relative abundance was very different.     

Spatial distribution of fishes and decapods in eelgrass (Zostera marina L.) and sandy habitats of a New Brunswick estuary, eastern Canada

We investigated spatial distribution of fishes and decapods in eelgrass (Zostera marina L.) and non-vegetated sandy habitats in Kouchibouguac Estuary, New Brunswick, Canada. During the ice-free season in 1999 and 2000, mobile fauna were sampled using fyke nets, minnow traps and an epibenthic sled. In general, fishes and decapods were more abundant and diverse (species richness) in eelgrass beds than in nearby sandy habitats, although the trend in abundance of decapods was not statistically significant. Abundance was greater at night than during daylight hours for most species. Atlantic tomcod (Microgadus tomcod), winter flounder (Pleuronectes americanus) and sand shrimp (Crangon septemspinosa) dominated night/crepuscular catches, whereas threespine stickleback (Gasterosteus aculeatus) and cunner (Tautogolabrus adspersus) predominated in diurnal periods. The nursery function of eelgrass habitat was most evident for juvenile white hake (Urophycis tenuis) and small cunners (<3 cm), which were found only in such habitat. This study is one of the few to have investigated the ecological importance of eelgrass habitats for mobile faunal communities in eastern Canada.     

Mitochondrial DNA analysis confirms the existence of two glacial races of rainbow smelt Osmerus mordax and their reproductive isolation in the St Lawrence River estuary (Quebec, Canada)

A phylogeographic analysis of mitochondrial DNA variation was performed in order to test the hypothesis that north-eastern North America has been postglacially recolonized by two races of rainbow smelt Osmerus mordax . This was accomplished by documenting the geographical distribution of two major mtDNA phylogenetic clades among 1290 smelt from 49 lacustrine and anadromous populations covering most of the species' native range. The data set was built by combining previously published results with those generated in this study. The two mtDNA clades showed a geographical dichotomy, independent of life-history types, whereby the more eastern populations were either fixed or largely dominated by one clade and western populations for the other. Such geographical pattern implying a phylogenetic discontinuity provided strong evidence for the persistence of smelt in two distinct glacial refugia as well as their differential postglacial dispersal. The most likely refugium for the so-called Atlantic race was the Atlantic coastal plains, whereas that of the so-called Acadian race was the exundated Grand Banks area. Patterns of postglacial dispersal interpreted from palaeogeographic events suggested that the Atlantic race recolonized northern regions about 5000 years prior the Acadian race. Both races came into contact in the St Lawrence River estuary. While gene flow has been possible, the sympatric occurrence in the estuary of anadromous populations alternatively dominated by one mtDNA clade or the other indicated that reproductive isolation mechanisms between the two races developed within this contact zone. This represents the first evidence of secondary intergradation among distinct races of aquatic organisms in an estuarine environment.     

Use of a geographic information system data base to measure and evaluate wetland changes in the St. Marys River,Michigan

A digital data base was constructed by photo interpretation, mapping, and digitizing seven dates of aerial photography on the St. Marys River, Michigan, USA. The data base was used in conjunction with geographic information system software to examine historical changes in wetland area. Total wetland area between 1939 and 1985 ranged from 7 200 to 7 317 ha over a 46-year period of high and low water. There was greatest variation in areas of emergent wetland and scrub-shrub wetland, which appeared to be responding primarily to changes in water level.     

Productivity of the aquatic macrophyte Pontederia lanceolata Nvtt. (Pontederiaceae) on floodplains of the Pantanal Mato-grossense,Brazil

The Pantanal Mato-grossense is an extensive floodplain system located in the center of South America with important macrophytic primary producers. This study describes the dynamics of net primary production and the biomass of the aquatic macrophyte Pontederia lanceolata, the dominant species in some floodplain fields near Poconé. Productivity was determined from a combination of demographic and growth methods. Biomass was determined using squares of known area, periodically distributed at random. The average productivity during the period was 3.11 g dry weight m^-2 day^-1. The minimum and maximum values (0.04 g dry weight m^-2 day^-1 and 8.59 g dry weight m^-2 day^-1, respectively) occurred during the drought and the flood period, respectively. The standing stock varied between 12.7 and 235.9 g dry weight m^-2, determined at the beginning of the flood and at its peak, respectively. The dynamics of the productivity and the standing stock of the species are associated with the flood patterns of the area and with life cycle of the plant.     

Myxozoan communities parasitizing Notropis hudsonius (Cyprinidae) at selected localities on the St. Lawrence River, Quebec: possible effects of urban effluents.

Late summer myxozoan infra- and component communities parasitizing 73 Notropis hudsonius at 5 sites on the St. Lawrence River upstream and downstream from the island of Montreal are described from study of histological sections of individual fish. Community membership included Myxobolus sp. A (intracellular in striated muscle fibers), Myxobolus sp. B (intracellular in striated muscle fibers), Myxobolus sp. C (brain), and Thelohanellus notatus (Mavor, 1916) Kudo, 1929 (loose connective tissue), all of which are histozoic myxobolids displaying strict tissue specificity for trophozoite development. Mean infracommunity richness in fish at the separate localities was estimated to be 0.4 +/- 0.5 to 1.3 +/- 0.7, with a maximum richness of 3 in any 1 fish. Component community richness in host samples was 2 to 4. It is argued that these are relatively high levels of diversity for freshwater fish parasites but that the values are probably conservative because of the study of only portions of individual fish. The percentage of fish infected with myxozoans of any species and infracommunity richness was significantly greater below the island of Montreal compared with above it. It is suspected that increased oligochaete populations at these sites, resulting from sewage-caused organic enrichment of sediments, may have accounted for the observed increased prevalence of infection.     

Energetics of lactation in harp seals (Phoca groenlandica) from the gulf of St. Lawrence,Canada

 This study reports the findings of an integrated, comprehensive analysis of lactation energetics in harp seals conducted using longitudinal measurements of mass, body composition and milk composition from mother-pup pairs in conjunction with water flux measurements in pups. The nursing period of harp seals is a short, intense and relatively efficient period of energy transfer from mothers to pups. The average daily milk intake for pups was 3.65±0.24 kg which is equivalent to 79.5 MJ of energy. Eighty-one per cent of the energy received in the milk was metabolisable and 66% of the energy was stored by the pups as body tissue. The field metabolic rate of pups was 3.9±0.4 time basal metabolic rate. The pups were growing at a rate of 2.2 kg per day during the nursing period. The distribution of this mass gain varied in terms of tissue composition, depending on the age of the pups, but over the whole nursing period approximately half of the tissue was stored as fat. Harp seal mothers lost an average of 3.1 kg per day during lactation which was composed of 37% water, 50% fat, 11% protein and 2% ash. Mothers spent half of their time during the lactation period actively diving and only one-third of their time on the surface of the ice. Milk compositional changes followed the normal phocid pattern with increasing fat content and decreasing water content as lactation progressed. The mean mass transfer efficiency was 73%. However, this value cannot be used without qualification because female harp seals in this study fed to varying degrees, consuming an estimated 0–4.8 kg of fish per day. Feeding does not appear to be required in order to achieve the energy requirements for lactation, given the energy stores possessed by females, and some females do fast through the entire period so feeding may be considered opportunistic in nature. Accepted: 25 April 1996