Creation of artificial upwelling areas for brown trout, Salmo trutta, spawning in still water bodies

Abstract  Brown trout, Salmo trutta L., spawning sites were constructed by creating areas of artificial upwelling water, 252 ± 37 mL m⁻² min⁻¹ (95% CL), through appropriately sized spawning gravel substrate in 3 m² vessels buried in the bottom of a 150-m² pond. Natural spawning occurred in the vessels during autumn 2001–2004, with hatching and alevin swim up the following spring. In areas of upwelling, egg survival was 85–95%, while no live eggs were observed in areas without upwelling. In areas with upwelling, the maximum density of live eggs at the eyed stage was 570–1510 eggs m⁻². In spring 2004 and 2005, the density of alevins was estimated at 322 (±187) m⁻² and 567 (±217) m⁻², respectively, in areas with upwelling water, compared with 35.2 ± 25.4 m⁻² in areas without upwelling water in 2004.     

Variations in spawning ground area and egg density of the Japanese sardine in Pacific coastal and oceanic waters

The spawning ground of the Japanese sardine, Sardinops melanostictus (Schlegel), was distributed over the oceanic waters as well as the coastal waters along the Pacific coasts of western and eastern Japan during 1978–1992. The area of the spawning ground in the coastal waters on the continental shelf has ranged from 95 000 km² in 1992 to 143 000 km² in 1988, constituting 44–77% of the total area of the spawning ground. The area of the coastal spawning ground was relatively constant in spite of the large fluctuations in egg abundance, i.e. size of the spawning population, from 88 trillion (1987) to 668 trillion (1989) in the waters. Spawning adults seemed to extend over the coastal waters irrespective of the size of the spawning population. In contrast to the coastal waters, the spawning area in the oceanic waters offshore of the continental shelf increased from 31 000 km² in 1978 to 183 000 km² in 1988 and then shrank to 83 000 km² in 1992, as a function of the spawning population size. The egg distribution density in the coastal waters stayed less than 6000 m⁻² mo⁻¹, but it reached as high as 27 400 m⁻² mo⁻¹ in the expanded spawning ground in the oceanic waters. The oceanic waters seemed to function as a reserve spawning ground for the sardine in years of extremely high spawning population.     

Remotely sensed spawning habitat of Pacific sardine (Sardinops sagax) and Northern anchovy (Engraulis mordax) within the California Current

We use trivariate kernel density estimation to define spawning habitat of northern anchovy ( Engraulis mordax ) and Pacific sardine ( Sardinops sagax ) in the California Current using satellite data and in situ egg samples from the Continuous Underway Fish Egg Sampler (CUFES) deployed during surveys in April by the California Cooperative Oceanic Fisheries Investigations (CalCOFI). Observed egg distributions were compared with monthly composite satellite sea surface temperature (SST) and surface chlorophyll a (chl a ) data. Based on the preferred spawning habitat, as defined in SST and chl a space, the satellite data were used to predict potential spawning habitat along two areas of the west coast of North America. Data from the southern area (21.5 to 39°N) were compared to observations from the CUFES data for the period 1998–2005. Northern anchovy and Pacific sardine exhibited distinctly different spawning habitat distributions. A significant relationship was found between satellite-based spawning area and that measured during surveys for sardine. CUFES area estimated for sardine was similar in magnitude to that estimated from satellite data (∼60 000 km²). In contrast, spawning habitat of anchovy averaged between 1000 and 200 000 km² for the period 1998–2005, for CUFES and satellite estimates, respectively. Interannual variability in the area (km²) and duration (months) of estimates of suitable habitat varied between species and between the northern (39 to 50.5°N) and southern portions of the California Current. Long-term monitoring of habitat variability using remote sensing data is possible in the southern portion of the California Current, and could be improved upon in the northern area with the addition of surveys better timed to describe relationships between observed and estimated spawning habitats.     

Spawning habitat of the Atlantic menhaden in Onslow Bay, North Carolina

The Continuous, Underway Fish Egg Sampler (CUFES) was used to sample pelagic eggs of the Atlantic menhaden ( Brevoortia tyrannus ) from 3-m depth off North Carolina in winter 1993–94 and 1994–95. Simultaneous measurements were made of temperature, salinity, and the concentration of chlorophyll a . The maximal concentration of eggs was 346 eggs m⁻³. Eggs were highly aggregated in patches which occurred between the Gulf Stream and mid-shelf fronts (17–23°C, 36.0–36.4‰). Unexpectedly, eggs were found almost exclusively in water of 20–60 m (mode 20 m) bottom depth. Thus, spawning appears related to bathymetry as well as hydrography. Variograms for egg concentration indicated a mean (± SE) patch scale of 3.6 ± 1.7 km and a high degree of spatial variance explained by CUFES sampling. Lagrangian modelling of particles moving in response to tides, winds, and a prescribed flow from the north indicated that the region of observed, maximal occurrence of eggs is favourable for the retention of eggs and larvae on the shelf adjacent to inlets used to enter nursery areas.     

A continuous, underway fish egg sampler

We describe a method to sample the highly contagious distribution of pelagic fish eggs. CUFES, the continuous, underway fish egg sampler, consists of a submersible pump, concentrator, electronics and sample collector. This system operates continuously and under nearly all sea conditions, providing a real-time estimate of the volumetric abundance of pelagic fish eggs at pump depth, usually 3 m. CUFES-derived estimates of volumetric abundance agree well with those from nets towed at pump depth and with areal abundance estimated from vertically integrated plankton tows. CUFES has been used successfully to sample the eggs of menhaden, pinfish, sardine, and anchovy off the coasts of the eastern and western United States and South Africa. Two large patches of eggs of the Atlantic menhaden were sampled off North Carolina in winter 1993–94, had a linear scale of 5–10 km, and were found in waters between the Gulf Stream and mid-shelf front. Spawning location may be related to bathymetry. CUFES is now being used to estimate spawner biomass by the daily egg production method. An optical plankton counter provided accurate estimates of the number of Atlantic menhaden eggs sample by CUFES.     

Variability in the spawning habitat of Pacific sardine (Sardinops sagax) off southern and central California

The spatial pattern of sardine spawning as revealed by the presence of sardine eggs is examined in relation to sea surface temperature (SST) and mean volume backscatter strength (MVBS) measured by a 150 kHz acoustic Doppler current profiler (ADCP) during four spring surveys off central and southern California in 1996–99. Studies in other regions have shown that MVBS provides an excellent measure of zooplankton distribution and density. Zooplankton biomass as measured by survey net tows correlates well with concurrently measured MVBS. The high along‐track resolution of egg counts provided by the Continuous Underway Fish Egg Sampler (CUFES) is a good match to the ADCP‐based data. Large interannual differences in the pattern and density of sardine eggs are clearly related to the concurrently observed patterns of surface temperature and MVBS. The strong spatial relationship between sardine eggs and MVBS is particularly evident because of the large contrast in zooplankton biomass between the 1998 El Niño and 1999 La Niña. The inshore distribution of sardine spawning appears to be limited by the low temperatures of freshly upwelled waters, although the value of the limiting temperature varies between years. Often there is an abrupt offshore decrease in MVBS that is coincident with the offshore boundary of sardine eggs. Possible reasons for this association of sardine eggs and high zooplankton biomass include an evolved strategy that promotes improved opportunity of an adequate food supply for subsequent larval development, and/or adult nutrient requirements for serial spawning. Hence, the distribution of these parameters can be used as an aid for delineating the boundaries of sardine spawning habitat.     

Modeling the transport and survival of Japanese sardine larvae in and around the Kuroshio Current

We have numerically modeled the advection and diffusion of sardine eggs and larvae to investigate the larval transport processes of Japanese sardine from the spawning grounds by the Kuroshio.
The results indicated that the offshore drift current induced by the winter monsoon and the location of the spawning ground have significant effects on the survival of the Japanese sardine. The contribution of the drift current, the distance of the spawning ground from the Kuroshio axis, and the eddy diffusivity to the larval retention in the coastal area is approximately expressed by the following equation: where R is the retention rate in the coastal area, a the variance of initial distribution of eggs, T the time after the eggs were spawned, – V₀ the velocity of the wind-induced offshore current, y₀ the distance of the center of the spawning area from the Kuroshio axis, and K the coefficient of horizontal eddy diffusivity.
The year-to-year variation in larval survival rates stimulated by the two-dimensional model are consistent with those estimated previously by using field data of egg and larval abundance during 1978–1988.     

Exchange and flux of larval fishes across the western Gulf Stream front south of Cape Hatteras, USA, in winter

To describe exchange and to frame the magnitude of the flux of larval fishes across the Gulf Stream front in winter, collections were taken on the inshore and offshore sides of the frontal zone in 1991, 1993, 1994 and 1995, off Onslow Bay, North Carolina. Collections yielded larval fishes representing 75 families. Of these, the larvae of the families Gonostomatidae or Phosichthyidae consistently occurred on both sides of the frontal zone, yet are exclusively of mesopelagic, deep-oceanic origin; hence, they indicate Gulf Stream-to-shelf water-exchange. The larvae of the clupeid, Brevoortia tyrannus , and the sciaenid, Leiostomus xanthurus , occurred on both sides of the frontal zone, yet spawn on the outer continental shelf in or near the Gulf Stream front; hence these taxa indicate shelf-to-Gulf Stream water-exchange. The mechanisms of exchange, as indicated by the distribution of heat and salt across the front, along with satellite images of sea-surface temperature, were not completely resolved, largely because of the two-dimensional ( x , z ) rather than three-dimensional ( x , y , z ) coordination employed. The observed distributions of indicator taxa did not always conform with expected, given the distribution of heat and salt across the front. Parcels of water exchanged across the front upstream (to the south) and encountered during observation along the cross-shelf transects explains the lack of conformity. Where the expected and observed distributions agreed, the flux was 0.0009 larvae m⁻¹ s⁻¹ for the indicators of Gulf Stream-to-shelf exchange, and 0.0007–0.0030 larvae m⁻¹ s⁻¹ for the indicators of shelf-to-Gulf Stream exchange.     

Cage culture of the Pacific white shrimp Litopenaeus vannamei (Boone, 1931) at different stocking densities in a shallow eutrophic lake

Postlarvae of Litopenaeus vannamei were acclimated and stocked in lake-based cages at the following stocking densities: 10, 20, 30 and 40 shrimp m⁻². Another set of shrimp was stocked in concrete tanks as reference samples at 30 shrimp m⁻². Significant differences were observed among stocking densities throughout the 95-day culture. The final weight at harvest decreased with increasing stocking density: mean weights of 23.3, 15.8, 13.0, 10.9 and 14.6 g for the 10, 20, 30, 40 shrimp m⁻² and reference tanks were observed respectively. There were no significant differences in survival throughout the culture period, ranging between 69% and 77%. Daily growth rates (range: 0.11–0.24 g day⁻¹) and specific growth rates (range: 3.54–4.34%) also differed significantly among stocking densities, both increasing with decreasing stocking density. The feed conversion ratio in the cages did not differ among the stocking densities, ranging from 1.53 to 1.65. The relationship between stocking density and mean individual weight at harvest followed the equation y =81.06 x ^−0.54 ( R ²=0.938) and that of stocking density and production (in g m⁻²) is y =58.01 x ^−0.46 ( R ²=0.834).     

Accuracy and precision of the continuous underway fish egg sampler (CUFES) and bongo nets: a comparison of three species of temperate fish

We examine the accuracy and precision of the continuous underway fish egg sampler (CUFES) relative to bongo nets based on the catch ratio of Atlantic cod (Gadus morhua), American plaice (Hippoglossoides platessoides), and cunner (Tautogolabrus adspersus). We derived expectation of catch ratios based on the application of a one-dimensional model of the vertical distribution of fish eggs applied to cod and on prior data on egg vertical distribution. Samples were collected in May and August 2001, two periods when the vertical structure of the water column differed substantially. Stationary CUFES collections did not yield significant differences in accuracy or precision relative to the underway CUFES. In May, when there was relatively little stratification, the CUFES-to-bongo catch ratio of cod and plaice eggs was well within expectations based on model predictions. In August, the CUFES-to-bongo catch ratios of cod and cunner were higher than expected. Generally, there was a greater proportion of early stage eggs in bongo than in CUFES samples, with the strongest differences in American plaice. The replicate variance of the CUFES was ∼25 times greater than that of the bongo nets, probably because of the large volumes sampled by bongo nets relative to the CUFES. Given that the CUFES provides greater accuracy in mapping but lower precision than bongo nets, multiple sampling gears may be the most effective method for surveying fish eggs of pelagic and demersal species.     

Fine scale spatial pattern of Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs

Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs exhibited different spatial structure on the scale of 0.75–2.5 km in two egg patches sampled in the Southern California Bight in April 2000. Plankton samples were collected at 4‐min intervals with a Continuous Underway Fish Egg Sampler (CUFES) on 5 × 5 km grids centered on surface drifters. Variograms were calculated for sardine and anchovy eggs in Lagrangian coordinates, using abundances of individual developmental stages grouped into daily cohorts. Model variograms for sardine eggs have a low nugget effect, about 10% of the total variance, indicating high autocorrelation between adjacent samples. In contrast, model variograms for anchovy eggs have a high nugget effect of 50–100%, indicating that most of the variance at the scales sampled is spatially unstructured. The difference between observed spatial patterns of sardine and anchovy eggs on this scale may reflect the behavior of the spawning adults: larger, faster, more abundant fish may organize into larger schools with greater structure and mobility that create smoother egg distributions. Size and mobility vary with population size in clupeoids. The current high abundance of sardines and low abundance of anchovy off California agree with the greater autocorrelation of sardine egg samples and the observed tendency for locations of anchovy spawning to be more persistent on the temporal scale of days to weeks. Thus the spatial pattern of eggs and the persistence of spawning areas are suggested to depend on species, population size and age structure, spawning intensity and characteristic physical scales of the spawning habitat.     

Ecological variations and El Niño effects off the southern coast of the Korean Peninsula during the last three decades

To explore ecosystem response to environmental changes, we investigated interannual and decadal changes in the marine ecosystem using data collected from the southern coast of the Korean Peninsula during the 1960s–1990s. Water properties such as sea surface temperature (SST) and salinity in April did not show large variation during the 1970s, but a relatively cool water mass appeared from the early to mid 1980s and a warm water mass replaced it after 1989. Chl a concentration in April, which was converted from Secchi disc information, was low (mean=1.18 mg m⁻³) during the period 1968–1980 excluding 1974, but there was a period of greater biomass for 7 years from 1981 (mean=2.78 mg m⁻³). A negative correlation ( r =−0.387, P  < 0.05) between SST and chl a in April matched high chl a during the low-SST period in the early and mid 1980s. Annual zooplankton biomass ranged from 37 to 132 mg m⁻³, but abundance was frequently high after 1984 and low during 1972–1982. The most representative pelagic fishes in the South Sea seemed to respond to the increase in planktonic organisms. Anchovy, mackerel and sardine increased in abundance since the mid 1970s. High catches of anchovy and mackerel lasted until the 1990s, whereas sardine decreased after the early 1990s. From the matrix of simple correlation coefficient, the Southern Oscillation Index (SOI) exhibited a high correlation with SST in December in the study area ( P  < 0.05). Catches of anchovy and mackerel were positively correlated with chl a and zooplankton during their early life periods, as well as with SST in December.     

Changes in the distribution of sardine eggs and larvae off Portugal, 1985–2000

Generalized additive models (GAMs) were fitted to sardine (Sardina pilchardus) egg distribution data from three daily egg production method surveys. The results showed that the area of egg cover off Portugal decreased significantly from 11 800 km² in 1988 to 7000 km² in 1997 and 7400 km² in 1999. This is because of a significant reduction in sardine egg presence off northern Portugal, GAM estimated areas being similar or higher in the late 1990s for southwestern and southern Portugal. The distributional area covered by larvae was not estimated for 1988 (larval distribution extended beyond the survey area), although it was probably higher than the 9600 km² for 1997 and 5500 km² for 1999. In 1997 and 1999, the Gulf of Cadiz was also sampled, indicating extensive areas with sardine eggs and larvae (more than 50% of the total area of distribution off Portugal). Standardized data from 15 ichthyoplankton surveys between 1985 and 2000 show a decline in the mean probability of egg presence within the Portuguese continental shelf from the mid‐1980s to the late‐1990s, because of a marked reduction in egg presence off northern Portugal. Sardine larval data from the same surveys suggest that the reduction in mean probability of presence in the north is less marked than for eggs (although this comparison ignores the presence of sardine larvae beyond the continental shelf in the 1980s). Similar changes off northern Portugal and western Galicia are observed in commercial sardine catches and the acoustically estimated area of fish distribution. It is possible that the observed decline in spawning area off northwestern Iberia during the 1990s is indirectly reflecting the prevalence of environmental conditions detrimental to sardine recruitment (northerly winds during winter that favour coastal upwelling and offshore transport), which have reduced the spawning contribution of young fish in that area.     

Naupliar copepod concentrations in the spawning grounds of Japanese sardine, Sardinops melanostictus, along the Kuroshio Current

Naupliar copepods were distributed at similar concentrations over the waters inshore and offshore of the Kuroshio Current off central Japan in early spring 1993 and 1994, overlapping with the distribution of early feeding larvae of Sardinops melanostictus . Although N, P, Si and chlorophyll a concentrations were higher in the waters inshore of the Kuroshio axis than in the offshore waters, the mean concentrations of nauplii were not statistically different between the two waters. Food availability for larval S. melanostictus did not seem to be different between the two waters in terms of the mean food concentrations. Using the critical food concentration (>9 nauplii L^–1) for 25% survival during 3 days after first feeding derived from a rearing experiment, percentages in number of stations or water samples with nauplii concentrations >9 nauplii L^–1 were higher in the inshore waters than in the offshore waters in both years examined. Considering that fish larvae may depend on small-scale patchiness of food for their survival, the inshore waters seemed to be more favourable for first-feeding larvae than the offshore waters.     

Fertilization efficiency of cryopreserved sperm from striped catfish, Pangasius hypophthalmus (Sauvage)

The fertilization efficiency of cryopreserved sperm was compared with fresh sperm from striped catfish, Pangasius hypophthalmus . Of the two sets of experiments carried out, the first compared four sperm doses using fresh sperm and fresh eggs. The second experiment compared six concentrations of cryopreserved sperm ranging from 6.94 × 10⁷ to 6.94 × 10¹⁰ to fertilize 100 eggs per batch. Fertilization, hatch and survival rates were compared between cryopreserved and fresh sperm. The highest fertilization rate (53.75±1.62%) was achieved with a sperm dose of 6.94 × 10⁸. Increasing the sperm dose to 3.47 × 10⁹ did not increase the fertilization rate, indicating that the optimum sperm:egg ratio lies between 6.94 × 10⁶ and 3.47 × 10⁷ sperm per egg. Both highest (6.94 × 10¹⁰) and the lowest (6.94 × 10⁷) sperm doses resulted in lower fertilization rates (2.04% and 16.90% respectively). No significant differences were found among four fresh sperm doses compared. Mean hatch and survival rates resulting from fresh and cryopreserved sperm were similar. The experiment shows that while only 1.89 × 10⁶ fresh spermatozoa was required to fertilize a fresh egg, 6.94 × 10⁶ (or 3.67 times more) cryopreserved sperm was required to achieve the same level of fertilization. This provides important information for making decision to cryopreserve sperm for commercial and/or conservation purposes.     

Alternating dominance of postlarval sardine and anchovy caught by coastal fishery in relation to the Kuroshio meander in the Enshu-nada Sea

In the mid 1970s, the fishery catch of postlarval Japanese anchovy (Engraulis japonica) in a shelf region of the Enshu‐nada Sea, off the central Pacific coast of Japan, started to decline corresponding to a rapid increase of postlarval sardine (Sardinops melanostictus). In late 1980s, sardine started to decline, and it was replaced by anchovy in the 1990s. This alternating dominance of postlarval sardine and anchovy corresponded to the alternation in egg abundance of these two species in the spawning habitat of this sea. It was also noteworthy that during the period of sardine decline, sardine spawning occurred in April–May, a delay of two months compared with spawning in the late 1970s. The implication of oceanographic changes in the spawning habitat for the alternating dominance of sardine and anchovy eggs was explored using time‐series data obtained in 1975–1998, focusing on the effect of the Kuroshio meander. Large meanders of the Kuroshio may have enhanced the onshore intrusion of the warm water into the shelf region and contributed to an increase in temperature in the spawning habitat. This might favour sardine, because its egg abundance in the shelf region was more dependent on the temperature in early spring than was that of anchovy. In addition, enhanced onshore intrusion could contribute to transport of sardine larvae from upstream spawning grounds of the Kuroshio region. On the other hand, anchovy egg abundance was more closely related to lower transparency at the shelf edge, which may indicate the prevalence and prolonged residence of the coastal water, and therefore higher food availability, frequently accompanying non‐meandering Kuroshio. The expansion/shrinkage of the spawning habitat of sardine and anchovy in the shelf region, apparently responding to the change in the Kuroshio, possibly makes the alternation in dominance of postlarval sardine and anchovy most prominent in the Enshu‐nada Sea, in combination with changes in the abundance of spawning adults, which occurred almost simultaneously in the overall Kuroshio region. The implication of this rather regional feature for the alternating dominance of sardine and anchovy populations on a larger spatial scale is also discussed.     

Physiological responses of juvenile wedge sole Dicologoglossa cuneata (Moreau) to high stocking density

Physiological responses to a high stocking density were tested in juvenile wedge sole Dicologoglossa cuneata (Moreau). Fish were kept at low (1 kg m⁻²), medium (3 kg m⁻²) and high (9 kg m⁻²) stocking densities for 22 days. No differences in the weight, length, survival and hepatosomatic index were observed among treatments. Basal plasma cortisol and osmolality were found to be directly and positively related to stocking density. A mild increase in plasma glucose was seen at medium density, and plasma protein was elevated at medium and high densities. The liver glucose and glycogen content was inversely related to stocking density. The liver triglyceride level was significantly elevated at the highest density, and the α-amino acid content decreased at the highest density. In muscle, glucose levels were significantly higher in fish kept at the lowest density; the α-amino acid content was elevated in fish kept at high density. In conclusion, plasma cortisol levels indicated an increasing stress level depending on the culture density, but significant changes in energy reserves did not occur in tissue (mainly liver and muscle glycogen and glucose reserves were significantly affected).     

Microhabitat use, movements and abundance of gilt darters (Percina evides) in southern Appalachian (USA) streams

Abstract –  We examined microhabitat use by gilt darters ( Percina evides ) in two streams in the south-eastern USA. Darters were over-represented in erosional microhabitats with higher average velocities and more cobble. Male darters tended to show stronger selection than females. Size-based analyses showed that larger (≥60 mm) gilt darters tended to use microhabitats with more heterogeneous substrata and more boulder than smaller (≤59 mm) darters. We also conducted a short-term movement study and calculated population estimates based on mark–recapture data in autumn 2005. Darters moved both long and short distances with 40% of all recaptures occurring within 5 m of the initial capture point. Using Program mark and model-averaged parameter estimates gilt darter density was 0.31 darters·m⁻² (225 darters/730 m²). Conservation of this species will require the preservation of erosional habitats in streams.     

Spatial and temporal distribution and abundance of European hake, Merluccius merluccius, eggs and larvae in the Catalan coast (NW Mediterranean)

Plankton hauls were conducted on five surveys from November 1998 to November 1999 to study the spatial distribution of eggs and larvae of European hake off Catalonia (NW Mediterranean Sea). Also CTD casts to record hydrographic parameters were carried out on a closely spaced station grid. Merluccius merluccius eggs and larvae appeared mainly in late spring, summer and autumn surveys and were very scarce in winter. Strong differences in terms of egg and larval densities were observed between the two November surveys, which could be attributed to the anomalous hydrographic situation during November 1998. M. merluccius egg and larvae were mainly distributed over the continental shelf, with peak abundances between the 100 m isobath and the edge of the shelf. On the evidence of larval size frequency distributions in the different sampling sectors and the closely overlapping distribution patterns for the eggs and the adult spawning stock, drifting of hake eggs and larvae was not a major factor. The larval distribution extended only slightly further offshore than the egg distribution. Using the hydrographic information and the larval distribution data, an attempt was made to relate the different seasonal productivity levels over the spawning period and the distribution of the larvae.     

Longline cultivation of some Laminariaceae in British Columbia, Canada

Abstract. Seedlings of Laminaria groenlandica Rosenvinge. L. saccharina (L.) Lamour. and Cymathere triplicata (Post. & Rupr.) J. Ag. were produced directly from mixed gametophytes and indirectly from cloned gametophytes. When cultivated in the sea, these seedlings produced up to 8 wet kg.m⁻¹ of cultivation rope for L . saccharina , 20 wet kg.m⁻¹ for L. groenlandica and 2·7 wet kg.m⁻¹ for C. triplicata . These values are similar to those reported from other parts of the world. Plant sizes for cultivated L. groenlandica and C. triplicata were greater than observed in wild stands Production by L. groenlandica was positively correlated with nutrient concentrations and negatively correlated with temperature. The time required to achieve harvestable size may he reduced from 16 to 9 months by employing seedlings produced indirectly from cloned gametophytes.