Temperature ecotypes near the southern boundary of the kelp Laminaria saccharina

Effects of temperature on survival, growth, and photosynthesis were compared for two USA populations of Laminaria saccharina Lamour. One population was located in New York State, near the southern latitudinal boundary of the species in the western North Atlantic. This southern boundary population was exposed to ambient temperatures 20°C for about 6 wk each summer. The second population was located in Maine, toward the center of the latitudinal range of the species, and was rarely exposed to temperatures>17°C. sporophytes from the New York (NY) population exhibited greater tolerance of high temperature than plants from the Maine (ME) site. Juvenile sporophytes from the two sites had similar rates of survivorship and growth at temperatures below 20°C, but showed different responses at 20°C in laboratory experiments. NY plants survived and grew for 6 wk at 20°C. ME plants showed negative growth during wk 2 and 100% mortality during wk 3. NY and ME plants held in situ at the NY site during June to September, 1985, also exhibited differential survivorship when ambient temperatures exceeded 20°C. Results of photosynthesis and dark respiration measurements on NY and ME plants grown at various temperatures suggested that the high-temperature tolerance of NY plants was attributable to their ability to maintain positive daily net C-fixation at 20°C. The high-temperature tolerance of the NY plants appeared to be due to genetic adaptation and is probably crucial to the persistence of the species near its southern boundary.     

High-temperature tolerance of photosynthesis in the red alga Chondrus crispus

Chondrus crispus (Stackhouse) is a perennial red seaweed, common in intertidal and shallow sublittoral communities throughout the North Atlantic Ocean. In the intertidal zone, C. crispus may experience rapid temperature changes of 10 to 20C° during a single immerison-emerision cycle, and may be exposed to temperatures that exceed the thermal limits for long-term survival. C. crispus collected year-round at Long Cove Point, Chamberlain, Maine, USA, during 1989 and 1990, underwent phenotypic acclimation to growth temperature in the laboratory. This phenotypic acclimation enhanced its ability to withstand brief exposure to extreme temperature. Plants grown at summer seawater temperature (20°C) were able to maintain constant rates of lightsaturated photosynthesis at 30°C for 9 h. In contrast, light-saturated photosynthetic rates of plants grown at winter seawater temperature (5°C) declined rapidly following exposure to 30°C, reached 20 to 25% of initial values within 10 min, and then remained constant at this level for 9 h. The degree of inhibition of photosynthesis at 30°C was also dependent upon light intensity. Inhibition was greatest in plants exposed to 30°C in darkness or high light (600 mol photons m^-2s^-1) than in plants maintained under moderate light levels (70 to 100 mol photons m^-2s^-1). Photosynthesis of 20°C-acclimated plants was inhibited by exposure to 30°C in darkness or high light, but the degree of inhibition was less than that exhibited by 5°C-grown plants. Not only was light-saturated photosynthesis of 20°C plants less severely inhibited by exposure to 30°C than that of 5°C plants, but the former also recovered faster when they were returned to growth conditions. The mechanistic basis of this acclimation to growth temperature is not clear. Our results indicate that there were no differences between 5 and 20°C-grown plants in the thermal stability of respiration, electron transport associated with Photosystems I or II, Rubisco or energy transfer between the phycobilisomes and Photosystem II. Overall, our results suggest that phenotypic acclimation to seawater temperature allows plants to tolerate higher temperatures, and may play an important role in the success of C. crispus in the intertidal environment.     

Physiological ecology of four Polysiphonia species (Rhodophyta,Ceramiales)

Photosynthesis and respiration of 4 species of the marine red algal genus Polysiphonia were evaluated under a variety of light, temperature and salinity conditions. The manometric results were compared with the local distribution and abundance of each species. The species can be separated into two distinct categories based on their overall distribution and temperature optima: (1) cold water plants [P. lanosa (L.) Tandy and P. elongata (Hudson) Sprengel], with peak photosynthesis at 21° to 24°C, but with active photosynthesis as low as 5°C; (2) plants with warm-water affinities [P. nigrescens (Hudson) Greville and P. subtilissima Montagne], having photosynthetic optima at 27° to 30°C, and exhibiting little or no photosynthesis below 10°C. The plants from the first group exhibit thermal injury at temperatures of 25°C and show a narrow tolerance to low salinities during periods of high temperatures. The plants from the second group show thermal injury at 30°C and have a wider tolerance to low salinities. The horizontal distribution of the 4 Polysiphonia species within the Great Bay Estuary System of New Hampshire, USA, is primarily governed by their tolerances to high temperatures and low salinities. The temperature optimum for each of the species corresponds to its particular estuarine distribution. Thus, P. subtilissima, having the highest temperature optimum, penetrated furthest into the Estuary, while P. lanosa, having the lowest temperature optimum, was restricted to the more coastal stations. There was a good correspondence between the natural distribution patterns and the manometric results.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 731.Scientific Contribution No. 4 of the Jackson Estuarine Laboratory.     

Plasmatic resistance and rate of respiration and photosynthesis of Zostera marina at different salinities and temperatures

Zostera marina L. was studied at the Izembek Lagoon, Alaska Peninsula. Two morphologically different forms, tidepool and subtidal, can be distinguished. Both show a high tolerance to different salinities and temperatures. The plasmatic resistance was found in a range of distilled H₂O up to 3.0 seawater (24 h) and between-6° and 34°C (12 h). Within these resistance limits, the photosynthesis, which has its maximum in normal (1.0) seawater, decreases nearly to zero not only in distilled H₂O but even in 2.0 seawater, and increases with the temperature in the tidepool form up to 35°C, but in the subtidal form up to 30°C only. At higher temperatures photosynthesis declines sharply in both forms. Respiration has its minimum in distilled H₂O and at 0°C and increases with increasing salinity and temperature.Dedicated to Prof. Dr. Dr. h. c. mult. K. Mothes, Halle, Germany (DDR) on the occasion of his 70th birthday.     

Temperature tolerance and latitudinal range of brown algae from temperate Pacific South America

Lower and upper survival temperatures of microthalli of 25 species of South American Phaeophyceae isolated from central Peru (14°S) to the Canal Beagle (55°S) were determined using 2-wk exposure for the upper and 4-wk exposure for the lower limit. All species survive 4 wks at -2°C. With respect to the upper limit, species reported only from southernmost South America tolerate 19.9 to 24.5°C (n=8), and species occurring from Cape Horn to central Chile 24.6 to 27.4°C (n=7). Three species that occurred as far north as northern Chile and Peru before the 1982–1983 El Niño event, and whose northern limit was dramatically shifted southwards in 1983, tolerate 20.8 to 25.3°C, whereas five species that have survived in Peru tolerate 25.6 to 28.5°C. Tinocladia falklandica which tolerates 27.8 to 28.1°C but lives only in southernmost South America and Striaria attenuata, which tolerates 31.6 to 31.9°C but occurs at ca. 42°S, are exceptional. Their high temperature tolerance may have no adaptive value in South America. They are restricted to the cold-temperate region due to low temperature requirements for reproduction or for reasons yet unknown. In general, the northern distributional limits of the Phaeophyceae studied along the temperate Pacific coast of South America are reproduction boundaries, except in El Niño years when they are redefined according to the species' upper suvival limits. Temperature tolerance of isolates from northern Chile and Peru agrees well with maximum temperatures reached during the 1983 El Niño.     

Growth patterns of Codium fragile ssp. tomentosoides in response to temperature,irradiance, salinity,and nitrogen source

Seasonal patterns of growth, reproduction, and productivity of Codium fragile spp. tomentosoides (van Goor) Silva were monitored at 3 locations in Rhode Island. Maximal growth occurred during the summer and was more significantly correlated with temperature than any other factor measured in this study. Multiple correlation models suggested an interaction between temperature, irradiance, and available nitrogen. Maximal reproduction occurred in late summer and early fall. The maximal productivity, based on harvested quadrats, was 2. 10 g dry weight m^-2 day^-1. A large amount (up to 87.3%) of the annual production entered the detrital food chain during the winter by fragmentation of the thallus. Culture studies examined the effects of temperature (6° to 30°C), irradiance (7 to 140 E m^-2 sec^-1), daylength (8 h light: 16 h dark to 24 h light: O h dark) and salinity (6 to 48) on growth. Differentiated thalli grew over a broad range of experimental conditions, with maximal growth at 24°C, 24 to 30 S, a minimal irradiance of 28 E m^-2 sec^-1, and 16 h daylength. The effect of increasing daylength was due to increased total daily irradiance rather than to a true photoperiodic effect. Undifferentiated sporelings survived and grew in a narrower range of environmental conditions than thalli. c. fragile spp. tomentosoides grew equally well with nitrate, nitrite, ammonium, and urea as a nitrogen source. The addition of NaHCO₃ stimulated growth at levels of 2.4 to 4.8 mM, suggesting an inorganic carbon limitation in static cultures. This study supports the hypothesis that the in situ seasonal growth pattern of c. fragile spp. tomentosoides is primarily due to the interaction of temperature and irradiance.     

Differences in somatic and gonadic growth of sea urchins ( Stronglyocentrotus droebachiensis ) fed kelp ( Laminaria longicruris ) or the invasive alga Codium fragile ssp. tomentosoides are related to energy acquisition

The rocky subtidal community off the Atlantic coast of Nova Scotia has historically undergone a cyclical transition between Laminaria-dominated kelp beds and sea urchin-dominated barrens. Since the introduction of the invasive alga Codium fragile ssp. tomentosoides, a third community state has emerged: Codium-dominated algal beds. We conducted a 42-week feeding experiment in the laboratory, which mimicked the quantity and quality of food available to urchins (Strongylocentrotus droebachiensis) in each of these community states. Feeding rate, growth, reproduction, and survival of urchins fed either Laminaria longicruris or C. fragile ad libidum, or L. longicruris 2 days per month, were measured. Although the ad libidum feeding rate on C. fragile was higher than that on kelp, energy intake was lower. Urchins in the ad libidum kelp treatment were larger and had larger gonads than those in the C. fragile treatment. Urchins fed kelp infrequently exhibited little somatic and gonadic growth over the course of the experiment. Regression analysis revealed that urchin performance on these diets was strongly related to energy intake. Diet treatment had no effect on survival or gonad maturation. Although urchins can consume substantial amounts of C. fragile, it appears that they cannot, or do not, feed quickly enough to compensate for its lower nutritional value. Our results suggest that, although urchins feeding on C. fragile are capable of surviving, growing, and reproducing, the replacement of kelp by C. fragile in some areas might negatively affect urchin populations as they continue to repopulate the shallow subtidal zone.     

Temperature acclimation of respiration and photosynthesis in the brown algaLaminaria saccharina

Sporophytes of the brown algaLaminaria saccharina (L.) Lamour grown at 15°C contained significantly more chlorophylla (chla) than did similar plants grown at 5°C. The increase in chla in 15°C plants was due to increased numbers of photosystem II reaction centes, and possibly to increased photosynthetic unit size, compared with 5°C plants. These changes were associated with increased values (photosynthetic efficiencies) in 15°C-grownL. saccharina relative to 5°C-grown plants. The changes in together with reduced respiration rates allowed 15°C-grownL. saccharina to achieve net photosynthesis and light-saturated photosynthesis at a lower photon fluence rate (PFR) than 5°C plants when both groups were assayed at the same temperature (15°C). The photon fluence rates necessary to reach the compensation point and achieve light-saturated photosynthesis (I _c andI _k , respectively) increased with increasing incubation temperature inL. saccharina grown at both 5 and 15°C. However, acclimation responses to growth temperature compensated for the short-term effect of temperature onI _c andI _k . Consequently, plants grown at 5 and 15°C were able to achieve similar rates of light-limited photosynthesis, and similarI _c andI _k values at their respective growth temperatures. These responses are undoubtedly important for perennial seaweeds such asL. saccharina, which frequently grow in light-limited habitats and experience pronounced seasonal changes in water temperature.Please address all correspondence and requests for reprints to I.R. Davison     

Nitrogen-fixation by cyanobacteria associated withCodium fragile (Chlorophyta): Environmental effects and transfer of fixed nitrogen

N₂-fixation associated with the green macroalgaCodium fragile subsp.tomentosoides (van Goor) Silva from Long Island, New York, USA, was attributable to several species of endophytic cyanobacteria. Rates of N₂-fixation ranged from 0.03 to 3.2µg N g^–1 dry wt h^–1 in freshly collected plants from several sites. Growth of the cyanobacteria appeared to be light-limited, due to the transmission of only 5 to 10% of incident light through the pigmented surface-layer of the macroalga. Daily irradiance was the most important factor determining both abundance of cyanobacterial cells and rate of N₂-fixation. The rate was also affected by instantaneous irradiance, and increased twofold from dark to ambient surface irradiance. Rates were reduced at low temperature (8°C) but showed no temperature effect between 12° and 26°C. External concentrations of dissolved inorganic nitrogen (DIN) up to 20µM did not influence N₂-fixation rate, but long-term exposure to 60µmol l^–1 d^–1 of NH ₄ ⁺ caused a reduction in the rate. InC. fragile grown under high daily irradiance and low external DIN concentration, ~50% of the assimilated-N was attributable to N₂-fixation. However, chlorophyllb extracted from plants grown with¹⁵N₂ showed an atom % excess¹⁵N of less than 0.1, suggesting that only a small proportion of the bacterially fixed-N was transferred to the seaweed. The association betweenC. fragile and its endophytic cyanobacteria appears to be based primarily on microhabitat suitability, rather than mutual metabolic dependence. It is doubtful that N₂-fixation by cyanobacteria is important to the ecological success of this seaweed species.     

Effect of temperature,salinity and food level on sexual and asexual reproduction in Brachionus plicatilis (Rotifera)

The reproductive response of sexual and asexual female Brachionus plicatilis (Muller) was examined over temperatures ranging from 20° to 40°C, salinities from 5 to 40 S, and food levels from 0.25 to 20 g Chlorella vulgaris dry-weight per ml. Reduced food levels, as well as temperature and salinity extremes, reduced reproduction of both sexual and asexual females, but did so differentially. Reproduction by sexual females was reduced to a greater extent at environmental extremes than asexual females. The broad, flat reproductive response curve of asexual females extended beyond the limits of the narrower, more sharply peaked curve of sexual females. Thus zones of exclusively asexual reproduction exist at environmental extremes where sexual reproduction is physiologically restricted. These results are corroborated by a comparison of the lifetime fecundity of individual sexual and asexual females over a 20°C temperature range. No differences in lifetime fecundity occurred between sexual and asexual females at 18° and 28°C. At 38°C, however, asexual female fecundity reached its highest level, while sexual female fecundity declined 15%. The appearance of sexual females in rotifer populations in the result of both inducible and repressible factors.     

Autecological investigation on marine diatoms 3. Thalassiosira nordenskioeldii and Chaetoceros diadema

The temperature range for the best competitive position by growth of Thalassiosira nordenskioeldii Cleve has been determined by comparing generation times. It ranges from-1.5° to 6°C. At these temperatures, especially at lower light intensities, it was one of the fastest growing species, whereas above 6°C many other species grew faster. High light intensities at increasing temperatures became damaging. A flowering of the cold oligo-eurytherm diatom T. nordenskioeldii occurs not only in the upper layers, but can also occur at greater depth simultaneously, because decreasing daylengths at 6°C had the least influence on its growth. Continuous light at 6°C had a positive influence on its growth. The start of the T. nordenskioeldii spring flowering under the Arctic Sea Ice is discussed in connection with the occurrence of enclosed marine diatoms in Polar Sea Ice. The influence of the winter temperature on the spring flowering of the North Sea, the southern border for flowerings of T. nordenskioeldii, is discussed. For Chaetoceros diadema (Ehrenberg) Gran the best competitive position by growth is reached at-1.5° to about 6°C. It has the best opportunity of reaching high cell numbers at the lowest temperatures of the range. The occurrence of the cold oligo-eurytherm diatom Ch. diadema in plankton samples at temperatures above 10°C need not be incorret, for the species did grow in cultures at 12° and 16°C. The wrong interpretation of the experimentally determined optimum temperature of e.g. T. nordenskioeldii caused a discrepancy between experimental results and field data that does not exist. The question is discussed whether ecologically it is relevant to talk about a temperature optimum. On account of the results of T. nordenskioeldii the question of the adaptation of diatom cultures for the start of the real experiments is discussed.     

Influence of high temperature and residual chlorine on marine phytoplankton

Marine phytoplankton forms are frequently exposed to sudden biological changes such as rapid rise in water temperature and chlorine content of their environment, resulting from the use of sea water for cooling purposes by electric generators. The direct influence of these effluents, i.e. inhibitory effects of high temperature and residual chlorine on growth and photosynthesis of Chlamydomonas sp. and Skeletonema costatum, were investigated experimentally. Chlamydomonas sp. and S. costatum exposed to high temperatures were affected in their growth from 43° and 35°C, respectively, by immersion of the respective cultures in a warm bath for 10 min. Treatment at high temperatures of 40 °C and 30° 35°C for 10 min, influenced their photosynthetic activities, which were completely inhibited immediately after 10 min exposure at 42° and 37 °C, respectively. S. costatum was killed by chlorine at a concentration of 1.5 2.3 ppm when exposed for exactly 5 or 10 min, while Chlamydomonas sp. was not irreversibly damaged even at 20 ppm chlorine or more with the same exposure period. These results lead to the conclusion that the high temperature of, and residual chlorine in, effuents from a power plant discharging into the open sea, should not cause great damage to marine phytoplankton in that area.     

Response of gametophytes of Ecklonia radiata (Laminariales) to temperature in saturating light

Gametophytes of Ecklonia radiata (C.Ag.)J.Ag. from two New Zealand locations with different field temperature ranges were exposed to temperatures of 5° to 26°C in saturating light. Plants from Goat Island Bay (Lat. 36° 16S, Long. 174°48E) grew in 9.3° to 25°C and reproduced in 9.3° to 24°C. There was no growth at 8°C and plants died at 26°C. Plants from the cooler location, Houghton Bay (Lat. 41°20S, Long. 174°40E), grew from 8° to 24°C and reproduced up to 15°C but not at 21.5°C. The plants did not grow at 6°C and died at 26°C. The timing of the first cell division and subsequent growth rate were retarded close to the upper and lower tolerance limits. Reproduction was a broad optimum of roughly 12° to 20°C. Within this range, fertile female gametophytes grown at lower temperatures had fewer, larger cells and thus fewer potential ova than those grown at higher temperatures.     

Effect of temperature on life cycles of nematodes associated with the mangrove (Rhizophora mangle) detrital system

The effect of temperature on the life history of various representatives of the meiofauna associated with decaying mangrove (Rhizophora mangle) leaves was investigated. Life cycles, recorded in days, for 6 species of marine nematodes cultured at a temperature of 24°C are as follows: Rhabditis marina 2 1/4; Diplolaimelloides sp., 7; Diplolaimella ocellata, 11 1/2; Enoplus paralittoralis, 22; Oncholaimus sp., 29; Haliplectus dorsalis, 34. In general, life cycles become shorter with increased temperatures; however, as temperatures approach the upper limits which support reproduction, life cycles become slightly lengthened. For most species, the ability to complete a life cycle was inconsistent within the temperature range of 33° to 35°C. Studies with two harpacticoid copepod and two foraminifera species tend to support the 33° to 35°C range as being the thermal stress zone.Contribution No. 1697 from the University of Miami, Rosentiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

Interactions between light and temperature on the physiological ecology ofGracilaria tikvahiae (Gigartinales: Rhodophyta)

Main effects and interactions of light and temperature on rates of growth (), net photosynthesis (P_s), and dark respiration (R) of the red seaweedGracilaria tikvahiae were investigated in outdoor, nutrient-replete continuous-flow seawater culture chambers. Below 15°C,G. tikvahiae did not grow and between 15° and 30°C, both main effects and interactions of light and temperature on and P_s were significant, which explains the occurrence of this alga as a summer annual in its northern range. Temperature interacted with light (I) through its influence on the vs I and P_s vs I curves. The initial slope of the vs I curve, , the light saturation intensity, I_s, and maximum growth rate, _max, were all significantly lowerat 15°C compared to 20°, 25°, or 30°C. Maximum values of _max, the P_s:R ratio and the net photosynthesis:gross photosynthesis ratio (P_s:P_g) all occurred at 25°C, suggesting that this is the best temperature for growth ofG. tikvahiae. Values for P_max increased up to 30°C, indicating that the temperature for maximum growth and net photosynthesis are not the same forG. tikvahiae. Significant photoinhibition of growth and photosynthesis at full incident sunlight (I₀) occurred at 15°C but not at 20°, 25°, or 30°C. Steele's equation fit the 15°C vs I data best, whereas the hyperbolic tangent function fit the 20°, 25°, and 30°C data best. Main effects and interactionof light intensity and temperature on rates of R were also significant (P<0.001). R was highly intercorrelated with and P_s (0.86r0.94), indicating that R inG. tikvahiae is primarily regulated by growth rate and not temperatureper se. Environmental factors that regulate growth, such as light intensity, exert a great influence on R inG. tikvahiae.     

Anti-grazing activity and seasonal variation of dimethylsulfoniopropionate-associated compounds in the invasive alga <Emphasis Type="Italic">Codium fragile</Emphasis> ssp. <Emphasis Type="Italic">tomentosoides</Emphasis>

In this study, we present evidence that the invasive alga Codium fragile ssp. tomentosoides is chemically defended against grazing by a wound-activated chemical defense involving dimethylsulfoniopropionate (DMSP) and the products of its cleavage, dimethylsulfide (DMS), and acrylic acid (AA). DMSP in C. fragile ssp. tomentosoides was present throughout the year, but concentrations varied seasonally and were highest in the winter. Intra-thallus variation was neither large, nor consistent over time. High DMSP concentrations were uncommon among northwestern Atlantic macrophytes. Of 26 other species tested, only two, Ulva lactuca and Polysiphonia harveyi contained concentrations comparable to, or higher than, those of C. fragile ssp. tomentosoides. DMS and AA, both individually and together, deterred grazing by the green sea urchin Strongylocentrotus droebachiensis at “natural” concentrations. These results suggest that DMS and AA contribute to the avoidance of C. fragile ssp. tomentosoides by S. droebachiensis. As a result, the production of DMSP and its subsequent cleavage, upon injury, may reduce herbivory on C. fragile ssp. tomentosoides and contribute to its success.     

Effect of temperature on the proliferation of Gymnodinium breve and Gomphosphaeria aponina

The effect of temperature on the growth and proliferation of two marine microorganisms, the toxigenic dinoflagellate Gymnodinium breve, and a potential bio-control organism, the blue-green alga Gomphosphaeria aponina, was determined by culturing the organisms in thermal gradients established by heating and cooling the opposite ends of an aluminum bar that had been adapted to hold culture tubes. Gradients were linear and stable for the duration of each trial. There was no relationship between variations in light and growth of the organisms. Gymnodinium breve showed optimum growth at 22°C, and proliferated over a range of temperatures (17° to 30°C). Below 17°C cultures of G. breve declined in growth, and at 4°C the organisms died within 5 h. Above 31°C there was rapid decline in viability of cells, and at 33.5°C the organism died within 24 h. Gomphosphaeria aponina showed optimum growth between 24° and 29°C, with a maximum at 27°C. Growth at temperatures greater than 31°C was minimal, but the organism survived. Limitation may be due to repression of the bio-synthesis of an iron-transport compound.     

Temperature adaptation in strains of the amphi-equatorial green alga Urospora penicilliformis (Acrosiphoniales): biogeographical implications

The temperature requirements for growth and the upper survival temperatures (UST's) of the amiphi-equatorial green alga Urospora penicilliformis collected from several localities within its distribution area between 1986 and 1991 were determined. Ecotypic variation, both with regard to growth ranges and optima and to survival temperatures, was demonstrated. In the polar strains of U. penicilliformis, temperature growth ranges were narrower and the growth optima and UST's were at lower temperatures compared to cold-temperate strains. In particular, the polar strains grew between 0 and 15°C with optimal growth at 0 or 5°C, whereas the cold-temperate isolates grew between 0 and (15) 20°C with almost equal growth rates or a growth optimum between 5 and 15°C. The Arctic strains survived 23 to 24°C, and the Antarctic isolate only 19°C, while the UST's of the cold-temperate isolates were between (24) 25 to 26°C. The data strongly indicate that a cold water history of ca. 3 million yr in the Arctic can be sufficient for changes in the temperature growth ranges and optima as well as for small changes of UST as shown in the Arctic populations of U. penicilliformis. For stronger reduction of upper survival temperatures, longer time periods are necessary as exemplified in the isolate from Antarctica, where low temperatures have existed for at least 14 million yr. The significantly lower UST of the Antarctic strain, points to an early contact of the alga with the cold water of the Antarctic region and may indicate an origin of U. penicilliformis in the Southern Hemisphere. The UST's of the cold-temperate isolates (24 to 26°C) would have allowed a migration across the equator during Pleistocene lowerings of the seawater temperatures in the tropics. Growth, however, would not have been possible during the passage across the equator due to the narrow temperature-growth window. The nature of the geographical boundaries and the control of seasonal development of U. penicilliformis by the temperature conditions in the various geographical regions are discussed in relation to the present local temperature regime.     

Positive effects of the introduced green alga, Codium fragile ssp. tomentosoides, on recruitment and survival of mussels

The green macroalga, Codium fragile ssp. tomentosoides, is an important component of sheltered low-shore assemblages on breakwaters along sandy shores in the northern Adriatic Sea. Macroscopic thalli of C. fragile are not perennial, but develop from propagules and/or undifferentiated forms in early spring, when the dominant native space-occupier, the mussel Mytilus galloprovincialis, recruits. By mid-summer, rapid growth of C. fragile leads to the formation of a dense canopy. We investigated the effects of juvenile and adult thalli of C. fragile on recruitment, survival and growth of mussels. Two experiments tested the hypotheses: (1) that recruitment of mussels is greater within patches of juvenile thalli (primordia) of C. fragile than on adjacent bare surfaces; (2) that the presence of a canopy of C. fragile affects the survival and growth of mussel recruits. The number of recruits of mussels was significantly larger within clumps of primordia of C. fragile than on bare surfaces. The removal of the canopy of C. fragile affected negatively the density of mussels after 2 months from the start of the experiment, but there were no effects on the mean size of individuals, nor on the size–frequency distribution. The same trend persisted after 4 months from the start of the experiment. These results show that re-colonisation of space by mussels is enhanced by C. fragile. Given that mussels, in turn, have the potential to reduce recruitment rates of C. fragile, quick recovery of mussel beds after disturbances could be crucial for controlling the abundance of this alga on breakwaters. Results also suggest that the effects of introduced species on native assemblages can be explained only through studies encompassing different life-stages of interacting organisms.     

Energetics of Euphausia pacifica. I. Effects of body carbon and nitrogen and temperature on measured and predicted production

Laboratory production during the life span of Euphausia pacifica was measured directly (as the sum of growth, molting and reproduction) and indirectly (as assimilation minus metabolism and leakage) to test the hypothesis that weight-specific production is a constant for all sizes. Euphausiids were collected in Puget Sound, Washington State, USA, from September 1973 to March 1978. Equations were determined (in terms of carbon and nitrogen at 8° and 12° C) expressing the relationships between body weight and the daily rates of growth, molting, reproduction, ingestion and metabolism. The allometric equation (R=aW ^b ) best related body weight (W) to the rate (R) for growth, molting, ingestion, respiration and excretion for life stages from late larvae through adults. As predicted by the original above hypothesis, the weight-specific coefficient (b) was close to 1.0 for ingestion and excretion; in contrast, b was 0.62 for growth, and 0.77 to 0.85 for molting and respiration. The Q₁₀ s also varied: 3.5 for growth, 2.4 for molting, about 3.0 for ingestion, and 2.0 for respiration and excretion. Assimilation efficiencies, for all weights and at both temperatures, were 81.3% of carbon and 85.9% of nitrogen ingested. The relationships between rate and body weight of early larvae for growth and molting were linear, as was the relationship for reproduction in adults. Weight-specific production was higher by I to 2% at 12° than 8° C for all life stages, and was 2 to 4% for carbon and 2 to 6% for nitrogen in adults, but 13 to 17% for carbon and 14 to 15% for nitrogen in early furcilia larvae. The null hypothesis was rejected for production measured directly, but would have been accepted if only an indirect measurement of nitrogen production had been considered. Clearly, indirect measurement incorporates all errors of measurement and assumption and makes interpretation difficult.