Long-distance transport of 14C-labelled assimilates in the Fucales: directionality,pathway and velocity

A directed translocation of ¹⁴C-labelled photoassimilates was shown to occur in the thalli of six species of brown seaweeds belonging to the Fucales. The apical growing regions of these algae act as strong sinks in the source-tosink relationship of the long-distance transport. Physical interruptions of the thalli showed that the midrib is the main pathway of the assimilates in Fucus vesiculosus and F. serratus. ¹⁴C-histoautoradiography has shown that translocation takes place through the filamentous cells of the medulla. The translocation velocity of the labelled assimilates, calculated from time source profiles of the tracer, was estimated to be 2 to 4 cm · h^-1. It is suggested that the sieve elements of the Fucales act in a way similar to the conducting sieve elements of the Laminariales.     

Autecology of Fucus distichus ssp. distichus (Phaeophyceae: Fucales) in Nova Scotia,Canada

The autecology of Fucus distichus L. ssp. distichus was investigated at several sites on the Atlantic coast of Nova Scotia, Canada, throughout a 2-year period. This species is confined to high-littoral rock pools at exposed sites, where there appears to be little competition from other organisms. Water temperature, salinity, dissolved oxygen, and hydrogen-ion concentrations underwent both considerable seasonal and short-term variation. Pools with extremes of salinity did not contain F. distichus ssp. distichus. Receptacles are formed during winter, and by late spring they are cast. Sporelings were first visible during September; their development was slow, and these plants did not become reproductive until the second year. During the period February to May, rapid growth of new fronds occurred; an increase in both length and bulk of the plants resulted from this growth. Fronds of mature plants continued to elongate slowly throughout the summer and the early autumn with the formation of receptacles in winter. Plants were frequently damaged, which resulted in regeneration of new fronds from the wounded surface.Issued as NRCC No. 14510     

Arsenic accumulation by ferns: a field survey in southern China

The objective of this study reported here was to characterize arsenic (As) accumulation by Pteris ferns by comparing 3 of the ferns of this genus with each other as well as with four non-Pteris ferns growing on seven sites in southern China with different As levels. A total of 112 samples, including 78 Pteris vittata, 13 P. cretica, 3 P. multifida and 18 ferns from other non-Pteris genera, with the soils in which they grew were collected for As and other elemental analyses. P. vittata was found to be the most dominant species and the most efficient As-accumulator, whereas P. multifida was the lowest As-accumulator among the Pteris ferns, with 4.54–3599, 28.7–757 and 11.2–341 mg kg^–1 As recorded in the fronds of P. vittata, P. cretica and P. multifida, respectively. Arsenic concentrations in non-Pteris ferns were generally much lower than those in Pteris ferns, with 0.81–1.32, 3.59, 10.7, 6.17–24.3 mg kg^–1 in the fronds of Blechumum orientale, Dicranopteris dichotoma, Pteridium aquilinum and Cyclosorus acuminatus, respectively. For P. vittata, the As bioaccumulation factor (ratio of As in fronds to that in soils) changed, whereas the As translocation factor (ratio of As in fronds to that in roots) remained unchanged among the different sites. The concentrations of Fe were very high in all of the collected fern sample, with the exception of B.␣orientale, with 207–6865, 637–3369, 375–1856, 1876, 493-6865 and 492 mg kg^–1 in the fronds of P. vittata, P. cretica, P. multifida, C. acuminatus, P. aquilinum and D. dichotoma, respectively. The association between Fe accumulation and As accumulation and tolerance in these ferns indicates the unique role of Fe in As-hyperaccumulation.     

Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera in a low-nitrogen environment

An adult giant kelp plant (Macrocystis pyrifera), moved from an inshore kelp forest to an offshore, low-nitrogen environment near Santa Catalina Island, California (USA), maintained growth for 2 wk on internal nitrogen reserves. Frond elongation rates decreased significantly during the third week, and plant growth rate (wet wt) dropped from an initial inshore rate of 3.6 to 0.9% d^-1. During this 3 wk period, nitrogen contents and free amino acid concentrations decreased, while mannitol and dry contents increased in frond tissues. After depletion of internal nitrogen reserves, the nitrogen content of lamina and stipe tissues averaged 1.1 and 0.7% dry wt, respectively. The experimental plant was exposed to higher ambient nitrogen concentrations during the last 2 wk. Rates of frond elongation and plant growth increased, but nitrogen content and amino acids in frond tissues remained low. Of the total nitrogen contained in frond tissue of the plant before transplantation, 58% was used to support growth in the absence of significant external nitrogen supply. Amino acids constituted a small proportion of these internal nitrogen reserves. Net movement of nitrogen occurred within large fronds, but not between different frond size classes. The nitrogen content of holdfast tissue remained relatively constant at 2.4% dry wt and accounted for 18 to 29% of the total nitrogen. Holdfast nitrogen was not used to support growth of nitrogen-depleted fronds. In comparison to Laminaria longicruris, which is adapted to long seasonal periods of low nitrogen availability, M. pyrifera has small nitrogen-storage capacity. However, internal reserves of M. pyrifera appear adequate to make nitrogen starvation uncommon in southern California kelp forests.     

Nitrogen supply,tissue composition and frond growth rates for Macrocystis pyrifera off the coast of southern California

Seasonal changes in ambient NO ₃ ^– and NH ₄ ⁺ , tissue composition (N, C, and C/N ratio), and frond growth rates for Macrocystis pyrifera (L.) Agardh were examined. Ambient NO ₃ ^– showed distinct seasonal variations. Frond growth rates were variable, but showed no clear correlation to ambient NO ₃ ^– . The average N content of plant tissue did, however, show the same seasonal variations as ambient NO ₃ ^– . The longitudinal distribution of total tissue N and various components of tissue N along fronds were also analyzed. Several distinct patterns were found: high levels of protein N at growing tips and elevated levels of soluble N in lower parts of the frond. Free amino acids accounted for a major portion of the soluble N, but neither NO ₃ ^– nor NH ₄ ⁺ accumulated in the plant tissue. The longitudinal distribution of N along the fronds is compared to reported variations in C metabolism, and it is concluded that C and N sourcesink relations do not always coincide and bidirectional translocation may occur.     

Morphology and growth of the giant kelp Macrocystis pyrifera in New Zealand and California

At three sites (Papanui Inlet, Aquarium Point and Seacliff) on the Otago coast of New Zealand and at one site (Point Loma) off southern California, morphometric and shortterm growth measurements were made of single fronds of plants of Macrocystis pyrifera (L.) C. Ag. near the time of the autumn equinox, in 1977. Near the apex, internodes and laminae were shorter in wave-exposed Seacliff and California populations than in those of Papanui Inlet and Aquarium Point in strong currents, although lamina thickness was similar. In the immature region near the apex, linear relationships were apparent from double logarithmic plots of stipe distance from apex vs node number, internode fresh weight vs length (different in the two areas), lamina width vs length, fresh weight vs lamina area and immature lamina, pneumatocyst and internode fresh weight vs node number. Morphologically, Seacliff plants resembled those in California more closely than those in the other New Zealand populations. The relative growth rate in length of stipe internodes decreased linearly with the logarithm of the distance from the apex, but was less closely related to node number. Extrapolation gave zero growth at 1.6 m at Aquarium Point and 2.9 m in California, where the internode lengths coincided with a change in their weight/length relationship. Slow relative growth proximal to these points indicated a different type (mainly elongation). It is suggested that the first, apical, type of growth, being apparently unrelated to frond length, would be a useful measure of activity over short time-periods. The relative growth rate in lamina area (and probably volume), while also related to distance from the apex, showed a very different pattern, increasing slightly to a peak at 100 to 200 mm and declining to nil by 1 m at Aquarium Point and 2 m in California. Maximal lamina size was maintained for only 10 to 15 nodes at Aquarium Point. Calculated node initiation frequencies were 2.7 d at Papanui Inlet, 2 d at Aquarium Point, and 1 d in California. At the latter sites it took about 2 mo for laminae and 2^1/2 mo for stipe internodes to mature. Deduced relative growth rates in stipe tissue and pneumatocyst fresh weight decreased from 0.24 (log_e units per day) at the apex to 0.11 at 200 mm, where basal lamina tissue had a rate of 0.29, a doubling in just over 2 d. The production of tissue per frond was estimated as 25 and 36 g fresh wt d^-1 at Aquarium Point and in California, respectively.     

Transversal profiles of carbon assimilation in the fronds of three Laminaria species

The brown macroalgae Laminaria digitata (Huds.) Lamour., L. hyperborea (Gunn.) Foslie, and L. saccharina (L.) Lamour. (Laminariales, Phaeophyceae) have been investigated for carbon assimilation and assimilate biosynthesis in different tissues (meristoderm, cortex, medulla) of stipes and blades. Carboxylation via ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) and phosphoenolpyruvate carboxykinase (EC 4.1.32) follows a transversal gradient from the outer to the inner tissues with maximum activity of enzymatic CO₂ fixation in the meristoderm. Such transversal profiles are also recorded for photosynthetic and dark carbon assimilation as well as for kinetics of assimilate ¹⁴C-labelling in isolated tissues. Differential chemical constitution of the blade and stipe tissues examined and results from assimilate biosynthesis are discussed with regard to a transversal translocation of photosynthates from meristodermal to cortical and medullar cells.     

Differences between native and invasive<Emphasis Type="Italic"> Caulerpa taxifolia</Emphasis>: a link between asexual fragmentation and abundance in invasive populations

The tropical and subtropical marine green alga Caulerpa taxifolia has invaded several temperate regions throughout the world, including southeastern Australia. In this study, I examined how invasive C. taxifolia from temperate southeastern Australia differed from native C. taxifolia from subtropical Moreton Bay, Australia, in the traits proposed as being important to its invasion success: thallus size and density, levels of asexual reproduction (fragmentation) and total biomass. Against the prediction of a large size for invasive C. taxifolia, native populations from Moreton Bay had larger stolons and fronds than invasive populations. However, invasive populations consistently had much higher densities of stolons, fronds and fragmented fronds; and a greater biomass compared to native populations. Average densities at invasive sites exceeded 4,700 stolons and 9,000 fronds/m² and were as high as 27,000 stolons and 95,000 fronds/m², which are the highest reported for C. taxifolia anywhere. Average densities of fragmented fronds at invasive sites were as high as 6,000/m² and up to 45% of all stolons at invasive sites could be directly linked to asexual recruitment via fragmented fronds. Importantly, at invasive locations there was a strong association between asexual reproduction and abundance demonstrated by positive correlations between the density of fragmented fronds and total biomass. These findings are the first to describe quantitative differences between native and invasive C. taxifolia and to demonstrate a link between the high levels of asexual reproduction and high abundance in invasive populations. Although the causes and consequences of high levels of asexual reproduction remain to be explored, this study suggests that changes in demographic and life-history traits during the invasion by C. taxifolia into temperate habitats may contribute to its success there.Communicated by M. S. Johnson, Crawley     

Morpho-functional patterns of photosynthesis and UV sensitivity in the kelp <Emphasis Type="Italic">Lessonia nigrescens</Emphasis> (Laminariales,Phaeophyta)

The photosynthetic responses of the south Pacific kelp Lessonia nigrescens of the coast of Valdivia, Chile (40°S), were investigated by exposing its different thallus parts, fronds, stipes and holdfasts, to UV radiation in the laboratory. Biologically effective doses (BED_{photoinhibition300}) between 400 and 800 kJ m⁻² were required for a 40% inhibition in photosynthesis under UVA+UVB radiation. At BED_{photoinhibition300} close to 250 kJ m⁻² (in treatments without UVB), the inhibition of photosynthesis did not exceed 20%. These UV doses were in the range of current daily doses measured in Valdivia on cloudless summer days. In general, exposure to UVB for periods longer than 12 h reduced photosynthesis, measured as maximal quantum yield (F _v/F _m) and electron transport. The fronds were the most UV-sensitive section of this alga, coinciding with the highest pigments contents and carbon fixation. Evidence of a photodamage was also seen. After a 48 h exposure to PAR+UVA+UVB, a decrease of F _v/F _m in the fronds was close to 41%, while in the stipes and holdfasts it was 12 and 18%, respectively. Although the thalli from the different size classes showed marked differences in their morphology and morphometry, no obvious differences in the UV tolerance of the fronds were detected. The results indicated that the UV-related responses are integrated in the suite of morpho-functional adaptations of the alga. Although the fronds are spatially more exposed to solar radiation than basal structures (stipes and holdfast), due their high turnover rate they may compensate better detrimental effects of UV. In contrast, stipes and the holdfast are key support structures characterized by low replacement rates and designed to confer hydrodynamic resistance to drag forces.     

Metabolic responses of Azolla pinnata to cadmium stress: photosynthesis,antioxidative system and phytoremediation

This present study deals with the growth, photosynthesis, oxidative stress and phytoremediation character of Azolla pinnata L. exposed to different levels (0.05, 0.1, 0.5, 1.0, 1.5 and 2.0 mg·L^?1) of cadmium (Cd). Significant accumulation of Cd in Azolla fronds was noticed after 24 and 96 h of exposure and the accumulation rate was dose and time dependent. Growth of A. pinnata increased significantly after both exposure times with and without metal. At lower Cd doses (0.05 and 0.1 mg·L^?1), growth and photosynthesis of A. pinnata showed a marginal increase over the respective control, however, at higher Cd doses (0.5, 1.0, 1.5 and 2.0 mg·L^?1), a decreasing trend was noticed. At lower doses, Azolla fronds could counterbalance the negative effect of enhanced levels of superoxide radicals (SOR) and hydrogen peroxide (H₂O₂) through the greater activity of antioxidative enzymes. The decaresing trends in catalase and peroxidase activity at higher Cd doses suggest that Azolla fronds were not able to mitigate the negative effects of H₂O₂, hence an increase in malondialdehyde content was noticed. The study concludes that up to 0.1 ,mg·L^?1 Cd, A. pinnata can flourish and be used as biofertiliser and for phytoremedial purposes in Cd-contaminated fields; beyond this concentration poor growth may restrict its application.     

Growth and production of the red alga Gelidium sesquipedale off the Basque coast (northern Spain)

Several aspects of the growth of the agarophyte Gelidium sesquipedale Clemente (Thuret) were studied in sublittoral populations of the Basque coast (northern Spain) from 1981 to 1986. The standing stock of the populations varied greatly with season. During two consecutive years, 60 to 70% of summer biomass had been lost by the end of the unfavourable period of year to growth (autumn and winter). The decline in biomass was mainly due to partial losses of branches, with more branches being lost from larger fronds. Yearly variations in the standing stock were also recorded. Production estimates obtained by a destructive sampling method were not satisfactory because of the spatial heterogeneity of the populations at the three locations studied. Indirect estimation of production by measuring the growth of marked fronds in a control quadrat gave more reliable estimates. Production values of 622±124, 735±147 and 615±123 g dry wt m^-2 yr^-1 were obtained at the three locations, respectively. Frond elongation rates of 7.5, 6.1, 7.5 and 7.9 cm yr^-1, respectively, were recorded during four consecutive years. In winter elongation rates of fronds were rather low. Elongation did not seem to be affected by frond length, the self-shading effect of the canopy, or changes in depth. Cuts resulting from grazing were frequent on fronds through the year, decreasing significantly with increasing depth. Sheared fronds regenerated promptly, independent of season or height at which the cut was made. For commercial purposes the best season for harvesting is summer, which is the period of maximum standing stock, infertility, and calm warm waters that provide easier working conditions for divers.     

CO2 fixation and translocation in benthic marine algae. III. Rates and ecological significance of translocation in Laminaria hyperborea and L. saccharina

Translocation of ¹⁴C-labelled assimilates in Laminaria hyperborea and L. saccharina is strictly basipetal within the thallus. The lowermost 20 cm of the frond accumulates 71% (L. hyperborea) or 63% (L. saccharina) of the basipetallymoving translocate after a period of 96 h from initial feeding with NaH¹⁴CO₃. Whereas translocation was found to occur at all seasons in L. saccharina, which grows throughout the year, no translocation of newly synthesized photosynthate occurred in L. hyperborea in October, when no frond enlargement takes place, nor in January, when the new fond occurs as a tiny outgrowth. In L. hyperborea, organic material, assimilated by the old frond during the period of rapid growth, is exported at the rate of 0.5 mg C · dm^-2 · h^-1. In L. saccharina, 14 to 18 cm frond length are necessary for an elongation rate of the growing zone, situated in lowermost 10 cm of the frond, of 1 cm per week. The significance of translocation in the two Laminaria species is demonstrated by comparison of export rates with growth rates measured in situ and in the laboratory.Dedicated to Professor Dr. Otto Stocker on his 85th birthday.     

Reproductive strategies and life histories in the cheilostome marine bryozoans Chartella papyracea and Bugula flabellata

Seasonal life history phenomena were monitored through 1978 for two sublittoral cheilostome bryozoans from souther Britain: Chartella papyracea (Ellis and Solander), a non-placental brooder, and Bugula flabellata (Thompson in Gray), which is placental. The intracolonial relationships between growth by zooid budding, polypide recycling, and sexual reproduction, were analysed for both species — quantitatively in the case of C. papyracea. Although both species grow only in summer, their life histories differ. C. papyracea is more of a K-strategist, producing perennial fronds that release larvae in several successive years, most being liberated in winter months. The reduced need for maximization of the larval productivity rate is reflected in the intracolonial configuration of polypide and sexual recycling. B. flabellata is more of an r-strategist. The larvae are produced by the ephemeral fronds of two successive colony generations each summer. Colonies are dormant in winter. The need to maximize the larval productivity rate in this species is reflected in the polypide/sexual recycling configuration. It is believed that embryonic placentation contributes to the maximization of the larval productivity rate in this and other placental brooders.     

Population biology ofMacrocystis integrifolia from British Columbia,Canada

Aspects of the population dynamics ofMacrocystis integrifolia Bory were determined from harvested wild populations over a 26-month period and cultivated plants over a 40-month period. Mean wild plant size varied from 0.1 wet kg and two fronds in the winter to 3.1 wet kg and mine fronds in the summer. Plant recruitment probably occurred year-round with recruits making up 20 to 86% of the total population. Plant half-lives, estimated from population data, ranged from 1 to 3 months, with few plants exceedings 2 to 3 years in age. Standing crop ranged from 1.6 plants m^-2 and 0.1 wet kg m^-2 in winter to 8.3 plants m^-2 and 8 wet kg m^-2 in summer.     

Nitrate reductase and glutamine synthetase activity,internal N pools,and growth of<Emphasis Type="Italic"> Ulva lactuca</Emphasis>: responses to long and short-term N supply

Fast-growing macroalgae, including Ulva lactuca Linnaeus, respond rapidly to changes in nutrient conditions, particularly to short-term N supply. This ability to rapidly take up and assimilate N contributes to the increasing occurrence of macroalgal blooms in heavily N loaded coastal ecosystems. To determine whether long-term nutrient histories affect short-term responses in activity of N-assimilating enzymes, including nitrate reductase (NRA) and glutamine synthetase activity (GSA), internal N storage, and macroalgal growth, we conducted an in situ nitrate fertilization experiment between 7 and 22 July 2004, with fronds of U. lactuca collected from estuaries with high and low N loads in Waquoit Bay, Cape Cod, Massachusetts, USA (N 41° and W 70°). Initial NRA, GSA, % N, δ¹⁵N, and growth of U. lactuca fronds were higher in the site where nitrate was in high supply. Differences in NRA persisted even after short-term experimental enrichment. Differences in internal N pools, δ¹⁵N, and growth, in contrast, mirrored the changes of nutrient supply. The rate of turnover of the internal N content of U. lactuca was quite short (<2 d), and turnover of enzyme activity may have been even shorter. N isotopic fractionation by U. lactuca appeared to be of small magnitude, unlike the case of phytoplankton, and similar to that of vascular plants. δ¹⁵N was a better indicator of short-term response to external and internal nutrient supplies in U. lactuca than enzyme activity or N content, and may reliably detect rapid changes in N availability, source, and uptake and assimilation processes.     

Geographic variation in polyphenolic levels of Northeastern Pacific kelps and rockweeds

Brown algal polyphenolic compounds are secondary metabolites whose functions may include protecting plants from pathogens or damage by UV radiation, and deterring feeding by herbivores. We present here the first analysis of spatial variation (at scales from tens of meters to hundreds of kilometers) in concentration of these compounds in two orders of brown algae from the northeastern Pacific Ocean. In kelps (order Laminariales), variation among sites was significant in only 25% of species examined and was consistent within families (high in the Alariaceae and low in the Laminariaceae and Lessoniaceae). In rockweeds (order Fucales, family Fucaceae), site variation was high in three of four species examined. Both the proportion of high polyphenolic kelp species and the magnitude of spatial variation within species from both kelps and rockweeds were much higher than would have been predicted from previous studies in other regions. In one kelp (Laminaria groenlandica), significant differences between sites occurred at scales of only tens of meters. No latitudinal clines were observed. Differences in phenolic concentrations of kelps spanned nearly an order of magnitude in one species, Hedophyllum sessile. Phenolic levels were significantly higher in members of the Fucales than the Laminariales, but showed no significant differences between intertidal and subtidal species. Received: 22 July 1996 / Accepted: 26 October 1998     

Growth of Flustra foliacea (Bryozoa)

The perennial bryozoan Flustra foliacea L. has annual growth-checks which leave lines across the fronds. These growth-checks have been used to determine the age and the pattern of growth of the colonies in terms of height and numbers of zooids. Monthly samples have been used to find the annual growth cycle. Heavy encrustations of epizoites on the F. foliacea colonies reduce growth rate. As the fronds increase in height, frontal budding of zooids thickens, and thereby strengthens, the holdfast.     

砷浓度、形态及碳酸氢盐对蜈蚣草吸收砷的影响

为了探讨超富集植物蜈蚣草在处理高砷地下水方面的可行性,研究了水培条件下砷的浓度、形态和碳酸氢盐(HCO-3)对超富集植物蜈蚣草吸收砷的影响。实验中使用了浓度为0.1～100mg·L-1的As(III)和As(V)溶液。HCO-3处理中,HCO-3浓度范围为0.5～20mmol·L-1,As(III)或As(V)的浓度为5mg·L-1。结果表明,在水培条件下,蜈蚣草具有明显的耐高砷特征。当介质砷含量高达100mg·L-1时,砷的去除率可达到80%,且对As(III)的吸收效率高于As(V)。植物体内砷形态研究表明,蜈蚣草体内2种形态砷的含量与外源砷形态有一定的关系,As(V)处理条件下,植物体中的As(V)比例较As(III)处理高。高浓度的HCO-3(20mmol·L-1)处理对蜈蚣草地上部分生物量没有明显影响,但是抑制了地下部分的生长,并且对砷的吸收表现出明显的抑制作用。     

砷浓度、形态及碳酸氢盐对蜈蚣草吸收砷的影响

大同盆地是典型的高砷地下水分布区。利用从地方性砷中毒严重病区山阴县采集的高砷地下水样品,用稀释培养法实验研究了外加砷源对地下水中微生物数量的影响;同时基于生物学可培养法和16S rDNA序列比对法,选取代表性高砷水样,研究了耐砷菌的种群特征。结果表明,外加砷源对地下水中微生物数量影响显著,高浓度砷会抑制大部分微生物生长,使微生物数量减少;低浓度砷对微生物生长具有一定促进作用。通过多次分离、纯化从3个不同砷含量地下水样中分离到多株砷抗性菌,经鉴定属于主要为Bacillus、Pseudomonas、Paenibacillus、Aeromonas、Enterobacter5个属。从RDP(Ribosomal Database Project)分析显示3个水样可培养微生物组成不同,都有生存能力强能够耐低浓度NaAsO₂的Bacillales,优势耐砷菌是γ-proteobacteria,其中Enterbacter具有耐高浓度NaAsO₂的能力。     

Life history of the red mullet Mullus surmuletus from the bottom-trawl fishery off the Island of Majorca (north-west Mediterranean)

The red mullet Mullus surmuletus is one of the main target species of the trawling fishery along the continental shelf off the Island of Majorca. The size distribution of the catches, and the reproduction, age and growth of this species have been studied based on sampling carried out from 1990 to 1992. The length range of the catches was between 10 and 32 cm, with a main distribution between 15 and 20 cm, but this included small specimens (recruits of 10 to 11 cm) from August. In the >19 cm length-class, females clearly dominated. Monthly variations in the gonadosomatic index (GSI) and in the percentage of mature specimens showed that males spawn from December to June, whereas the reproductive activity of the females centres around spring. Fifty percent of males and females mature at 15 and 16.8 cm, respectively, corresponding to 1 yr of age. Otolith age-readings indicate that the population exploited in the trawl fishery consists of six age-groups, including a very high proportion of individuals between 0 and 4 yr old. Population growth curves revealed that females grow comparatively slowly over a longer period of time and attain greater asymptotic sizes than males. The growth parameters for the whole population are: asymptotic length, L =31.28 cm; growth coefficient, K=0.211 yr^-1; theoretical age when length is zero, t ₀=-2.348 yr.