LIFE HISTORY VARIATIONS IN A PREDOMINANTLY GAMETOPHYTIC POPULATION OV IRIDAEA CORDATA (GIGARTINACEAE, RHODOPHYTA)1

An Iridaea cordata ( Turner) Bory population made up of 83% gametophytes and 17% tetrasporophytes demonstrated little variation in these proportions over three years. These data contrast with an expectation that alternations of generations, as demonstrated by culture studies, should result in equal numbers of isomorphic generations. Analyses of reproductive and demographic attributes of the population were made to better understand the role of alternation of generations in this population .
Growth and mortality rates in the summer could not account for the success of gametophytes. however, size class frequencies demonstrated that tetrasporophytes were smaller individuals. Sporelings recruited into this intertidal region and made up 20% of the spring population numbers. Even with this level of recruitment, gametophyte-to-tetrasporophyte ratios did not change. Perennating holdfasts regenerated 80% of the spring number and 90% of the total blade area. Consequently, perennation and differential survival of gametophyte spores or sporelings are proposed as mechanisms maintaining gatnetophyte dominance in this San Juan Island, Washington population .     

Population modelling of Gelidium sesquipedale (Rhodophyta, Gelidiales)

A matrix population model of Gelidium sesquipedale, a commercial agarophyte from the Northeast Atlantic, was developed based on demographic data obtained during two years in a commercial stand of Cape Espichel, Portugal. G. sesquipedale individuals were classified into categories such as life cycle phase, spores, juveniles and adult frond size, because the species vital rates, fecundity, fertility, survival, growth and breakage depend on them. We also exemplify the use of a user-friendly modelling software, Stella, to develop a structured-population model. This is the first time this software has been used to model the demography of seaweed populations. The Stella model developed here behaved very similarly to the matrix model, because of its particular construction, which causes the forcing functions to be discrete rather than continuous. The relative importance of spore recruitment and vegetative growth of new fronds in both population growth and population structure was investigated. Elasticity analysis suggests that vegetative recruitment is the most important demographic parameter controlling population growth together with survival and transitions between juveniles (1–6 cm fronds) and class 1 fronds (6–9 cm fronds). On the other hand, sexual reproduction may, by itself, efficiently control the relative proportion of gametophytes and tetrasporophytes in the population, even though its contribution to recruitment is extremely small. A 40% difference in the growth rates of gametophyte and tetrasporophyte submatrices resulted from natural differences in spore recruitment rates. This revised version was published online in June 2006 with corrections to the Cover Date.     

Size structure and dynamics of an invasive population of lineage 2 of Asparagopsis taxiformis (Florideophyceae) in the Alboran Sea

In this study, we present basic population data of the red macroalga Asparagopsis taxiformis, widely recognized as invasive in the Mediterranean Sea. A 13‐month field study was carried out on a population located in southern Spain, addressing its phenology, population dynamics and demography. We further tested whether biomass variations were related to environmental variables at the study site. Gametophytes were present year‐round while tetrasporophytes were only found in spring and summer. Recruitment capacity and vegetative growth of the gametophytes are discussed as important modulators for the population structure and enhancers of its persistence. Thallus size‐time histograms revealed a high prevalence of small shoots that showed high mortality that was not related to self‐thinning. Biomass of A. taxiformis was higher from March to July. Vegetative growth was the main way the gametophyte population was maintained, although the presence of tetrasporophytes and fertile gametophytes in the field confirms that sexual reproduction also occurs. Its continuous and high recruitment, in terms of the number of smallest shoots, makes this southern population of A. taxiformis a source of future invasive populations due to the intensive maritime traffic in the region.     

Spatial and temporal variations of Chondrus crispus (Gigartinaceae, Rhodophyta) carrageenan content in natural populations from Galicia (NW Spain)

Qualitative and quantitative differences in carrageenan composition of gametophytes and tetrasporophytes of Chondrus crispus were observed in this study. Carrageenans in gametophytes belong to the kappa family (κ-, ι-, ν- and μ-carrageenan). The dominant fractions were κ- and ι-carrageenan (more than 50% of the total carrageenans). In tetrasporophytes, the presence of λ-carrageenan was confirmed. Carrageenan content in gametophytes (37.4?±?1.68% DW) was higher than in tetrasporophytes (29.13?±?0.76% DW). Spatial and temporal variation in carrageenan content in both life cycle phases appears to be related mainly to seawater and air temperatures, insolation, water movement and desiccation. The highest values of carrageenan content were recorded in those localities where higher values of precipitation, wind speed or water movement occurred. A bimodal temporal pattern on carrageenan content was observed. Fronds showed a high carrageenan content in spring and autumn. During these seasons, the content was over 40% in gametophytes and 30% in the tetrasporophytes. In summer and winter, these values down in both life cycle phases below 30%. In general the highest carrageenan contents were related to highest seawater temperatures. On the contrary, high air temperature and high insolation appeared to be unfavourable for carrageenan production. GLM models were obtained to predict carrageenan production from natural C. crispus populations, along Galician coast.     

Biomass variation and reproductive phenology of Gracilaria gracilis in a Patagonian natural bed (Chubut, Argentina)

The biomass variation and the reproduction of the natural Gracilaria gracilis bed in Bahía Bustamante (Patagonia, Argentina) were analyzed for 2 years, with the aim of determining the present situation of the population for an updated status overview; establishing the relevant features of the temporal variation in both biomass and reproductive states in relation to environmental factors, epiphytes and associated algae; and assessing carpospore availability for future spore-culture development. Field measurements and sampling were performed monthly between March 2006 and February 2008. In both years, G. gracilis biomass presented marked seasonal variations, with a minimum in winter and a maximum in late spring and in summer. During both years, coexistence of the three life-cycle phases was found, with dominance of tetrasporophytes. Two data sets from individuals originated from sexual reproduction (tetraspores and carpospores) and from asexual reproduction by thallus fragmentation were analyzed separately. In the fragmentation fraction, tetrasporophyte frequencies remained higher than those for gametophytes. However, in the spore-originated fraction, a generation ratio close to 0.5 was observed. Female gametophytes bearing cystocarps were always present, with a maximum in summer and autumn. Biological data were related to environmental factors by means of canonical correspondence analysis (CCA). The first year was characterized by higher biomass values of G. gracilis and Undaria pinnatifida, lower epiphytism, larger Gracilaria thalli and greater proportion of mature tetrasporophytes and gametophytes. The second year was characterized by a high proportion of Gracilaria vegetative thalli and high epiphyte density. The best time to obtain spores from cystocarpic thalli would be in summer and early autumn.     

Seasonal variation of carrageenan yield,gel strength and viscosity in Sarcopeltis (ex Gigartina) skottsbergii from Southern Chile

Chile is one of the top carrageenan producers worldwide, and Sarcopeltis (ex Gigartina) skottsbergii one of the topmost exploited carrageenophytes from the wild in the world. Total yield, gel strength and viscosity from two contrasting environments Calbuco and Ancud (Inner and Outer Sea, Chile) were estimated monthly in approximately 2 years for this species. While carrageenan yields did not show differences between localities, gametophytes in spring–summer had 15% higher, compared to tetrasporophytes. Sizes (frond surface) normally did not affect carrageenan yields. Gametophytes showed clear differences in gel strength between seasons, but not between localities, with maximum peaks during winter–spring months in Calbuco and autumn-winter months in Ancud. Seasonal variations in viscosity were also significant. While gametophyte viscosity did not exceed 120 cPs, tetrasporophytes reached 1400 cPs in Calbuco and 1000 cPs in Ancud. More remarkably, a positive correlation between viscosity and gel strength was found in S. skottsbergii gametophytes, which is significantly different between both localities. These results suggest that selective harvesting in spring–summer should be preferred to optimize cost–benefit of harvesting activities and subsequent carrageenan productivity.     

Demographic models to simulate the stable ratio between ecologically similar gametophytes and tetrasporophytes in populations of the Gigartinaceae (Rhodophyta)

In populations of the Gigartinaceae (Rhodophyta), gametophytes often predominate numerically over tetrasporophytes. Several hypotheses have been proposed to explain this dominance, based on the usually implicit assumption that the stable ratio between gametophytes and tetrasporophytes (G:T ratio) should be 1 if both reproductive phases are ecologically similar. We developed demographic models to test this assumption, for which we considered that both phases are ecologically similar. Defining ecologic similarity for most demographic rates is relatively straightforward, except for rates of spore output. The first set of models considered the same spore output per thallus of both phases as representing ecologic similarity. Model iterations led to stable G:T ratios of 1 for triennial and for perennial thalli, regardless of the initial G:T ratio, but not for annual thalli with initial G:T ratios different from 1. However, equal spore output may not represent ecologic similarity, due to size differences between carpospores and tetraspores. The second set of models considered the lowest possible spore output for each phase, according to the life history of this family: only one carposporangium, with one carpospore, is produced from every two gametophytes and only one tetrasporangium, with four tetraspores, is produced by every tetrasporophyte. Model iterations led to stable G:T ratios of 2.8 for most cases, a ratio of 1 being obtained only every 2 years for annual thalli with an initial G:T ratio of 1. Increasing absolute spore output, without altering the relative output between phases and incorporating density-independent mortality through a matrix model, given the same mortality rate for both phases, did not modify results. We suggest that the combination of both modeling and field research may uncover more rapidly than otherwise the most relevant ecologic differences between phases, if any, that underlie the G:T ratio observed for a given population.     

Intraspecific variation in Gracilaria caudata (Gracilariales,Rhodophyta): growth,pigment content,and photosynthesis

In Brazil, one of the probable reasons for failure in attempts at macroalgal mariculture is the lack of previous studies under controlled conditions. Gracilaria caudata is an important marine red alga which is locally exploited for the production of agar. In this study the aim was to compare in vitro growth rates, pigment content, and photosynthesis in gametophytes and tetrasporophytes of G. caudata from two distinct geographical areas located 2,500 km apart on the Brazilian coast, one in a warmer area closer to the equator (northeastern population), and the other in a colder area closer to the Tropic of Capricorn (southeastern population). Additionally, the artificial ultraviolet B (UVB) radiation effects on strains were evaluated. Under UVB, deleterious effects were observed in all strains. Although the strains from the southeastern population had higher growth rates than those from the northeastern under control condition, the opposite was observed under UVB condition. Under controlled conditions and regardless of the population, growth rates, net photosynthesis, P _max, I _k, and pigment contents were higher in tetrasporophytes than in gametophytes. Consequently, when determining the real potential of a certain phase in cultivation, the tetrasporophyte appears to be the more promising for future experiments along the Brazilian coast. Furthermore, although the growth rate of southeastern strains under control condition was higher, their higher sensitivity to UVB radiation emphasizes the importance of careful selection of the most suitable sites prior to experimental cultivation. The differences in performance between the southeastern and northeastern strains provide support for the hypothesis of their ecotypic differentiation.     

Differential ecological responses to environmental stress in the life history phases of the isomorphic red alga Gracilaria chilensis (Rhodophyta)

In order to better understand the alternation of generations that characterizes haploid–diploid life cycles, we assessed the existence of ecological differences between the two phases (haploid gametophyte and diploid tetrasporophyte) in Gracilaria chilensis, a rhodophyte with a typical Polysiphonia-type life cycle. We investigated the effect of light intensity and salinity on viability and growth of both phases at different ontogenetic stages: juveniles and adults. In our study, the survival of juvenile gametophytes (n) was higher than the survival of juvenile tetrasporophytes (2n) despite culture conditions; however, low salinity had greater effect on carpospores (2n) than on tetraspores (n). On the other hand, a complex interaction between salinity and light intensity within each life history phase generated observed differences between juvenile growth rates. Low light was shown to trigger early onset of alteration of the holdfast growing pattern. In addition, adult tetrasporophytes showed, despite the conditions, a faster vegetative growth than female and male gametophytes. These differences between phases could have led to the complete dominance of tetrasporophyte fragments of fronds observed in G. chilensis farms. We hypothesize that Chilean fishers could have unknowingly selected for tetrasporophyte thalli during domestication of the species, thus enhancing the natural trend of tetrasporophytes dominance already present in estuarine natural populations of free-floating plants.     

IS THERE AN ECOPHYSIOLOGICAL EXPLANATION FOR THE GAMETOPHYTE–TETRASPOROPHYTE RATIO IN GELIDIUM SESQUIPEDALE (RHODOPHYTA)?1

In the fall, when 61% of the fronds of the Gelidium sesquipedale (Clem.) Born. et Thur. population located in Albufeira (southern Portugal) were reproductive, about 90% of these fronds were tetrasporophytes, whereas an equal percentage of female and male gametophytes was found (5%). The comparison of physiological performances of the reproductive phases (males, females and tetrasporophytes) did not reveal a physiological advantage of tetrasporic fronds. There were no significant differences either in the photosynthesis, nitrogen uptake, nitrate reductase activity, or biochemical composition of adult fronds. On the other hand, vegetative recruitment and spore production in the laboratory were significantly different. The re‐attachment to calcareous substrate and the subsequent rhizoidal growth were faster in tetrasporophytes. Particular levels of temperature, rather than irradiance, had an important effect on the phase differences in the spore release, attachment, and germination rates. Significant results were the higher release of carpospores at all irradiances at 17°C, and the higher attachment percentage of carpospores at 13°C versus tetraspores. Under higher temperatures (21°C), tetraspores showed higher attachment rates while carpospores germinated more. G. sesquipedale cystocarps released carpospores for 2 months, while tetrasporangia stopped shedding tetraspores after 1 month, resulting in a 3‐fold higher production of carpospores than tetraspores. Results showed that vegetative and spore recruitment may explain the low gametophyte–tetrasporophyte ratio of the studied population of G. sesquipedale as opposed to the physiological performance of phases.     

Differential responses of tetrasporophytes and gametophytes of Mazzaella laminarioides (Gigartinales,Rhodophyta) under solar UV radiation

The effects of solar UV radiation on mycosporine‐like amino acids (MAAs), growth, photosynthetic pigments (Chl a, phycobiliproteins), soluble proteins (SP), and C and N content of Mazzaella laminarioides tetrasporophytes and gametophytes were investigated. Apical segments of tetrasporophytes and gametophytes were exposed to solar radiation under three treatments (PAR [P], PAR+UVA [PA], and PAR+UVA+UVB [PAB]) during 18 d in spring 2009, Punta Arenas, Chile. Samples were taken after 2, 6, 12, and 18 d of solar radiation exposure. Most of the parameters assessed on M. laminarioides were significantly influenced by the radiation treatment, and both gametophytes and tetrasporophytes seemed to respond differently when exposed to high UV radiation. The two main effects promoted by UV radiation were: (i) higher synthesis of MAAs in gametophytes than tetrasporophytes at 2 d, and (ii) a decrease in phycoerythrin, phycocyanin, and SPs, but an increase in MAA content in tetrasporophytes at 6 and 12 d of culture. Despite some changes that were observed in biochemical parameters in both tetrasporophytes and gametophytes of M. laminarioides when exposed to UVB radiation, these changes did not promote deleterious effects that might interfere with the growth in the long term (18 d). The tolerance and resistance of M. laminarioides to higher UV irradiance were expected, as this intertidal species is exposed to variation in solar radiation, especially during low tide.     

Effects of elevation, wave exposure, and year on the proportion of gametophytes and tetrasporophytes in Mazzaella parksii (Rhodophyta, Gigartinaceae) populations

We investigated the effects of elevation, wave exposure, and year on the proportion of gametophytes and tetrasporophytes in populations of the intertidal red seaweed Mazzaella parksii (= M. cornucopiae, Gigartinaceae) from the Pacific coast of Canada. In July 2002, we determined the life-history phase of 864 frond samples from four populations from Barkley Sound, using the resorcinol-acetal test. For these populations, gametophytes were significantly more numerous than tetrasporophytes, and the proportion of gametophytes was significantly higher at higher elevations. The proportion of gametophytes varied directly (although less markedly) with the degree of wave exposure, but significance could not be assessed for this factor. All of these patterns were spatially consistent, as they held for two different rocky points with similar physical characteristics. One of these points had also been surveyed in 1994-1995, which allows us to conclude that the population-wise gametophyte predominance and the positive relationship between elevation and the proportion of gametophytes are stable features of this species in this area. However, the average proportion of gametophytes was similar between the studied levels of wave exposure in 1994–1995; such a difference revealed as non-significant when data were combined with those for 2002. This suggests that wave exposure is not important in determining the proportion of life-history phases for this species. Comparisons with other species are done in search of general patterns for the Gigartinaceae.     

Seasonal population dynamics of Neoechinorhynchus qinghaiensis in the carp, Gymnocypris przewalskii przewalskii, from Qinghai Lake, China

Studies on the seasonal population dynamics of Neoechinorhynchus qinghaiensis (Acanthocephala: Neoechinorhynchidae) in its fish host Gymnocypris przewalskii przewalskii in the Qinghai Lake, China, were carried out with samples taken in May 1991, August 1992, November 1992 and February 1993. Prevalences were higher than 44% in all seasons. The mean intensity of infection was above 124 worms per fish. The maximum intensity of worms recovered from a single fish was 1402 in the autumn of 1992. Differences in the mean abundance, mean intensity and prevalence are not statistically significant relative to season and this is likely to be related to the stable temperatures recorded at the bottom of Qinghai Lake. Over-dispersed distributions of N. qinghaiensis in the host population, due to heterogeneity and feeding habits, were observed in all seasons. The size composition of both sexes of N. qinghaiensis showed males to be less than 3.5 mm and females between 0.5 and 4.25 mm, with the main recruitment phase in the worm populations occurring in the autumn, extending through winter and spring with the lowest recruitment occurring in the summer. The maturation and copulation of worms were mainly focused in the summer season. The sex ratio of female to male was both high in winter (1.51:1) and spring (1.48:1). The higher proportion of females and the change in the worm sex ratio in winter can be attributed to the reduced longevity of male worms. As immature male worms exhibit a higher proportion of the worm population than females in all seasons, further studies are needed to determine if such a situation compensates for the shorter life span of males.     

Demography and population dynamics of the mouse opossum (Thylamys elegans) in semi-arid Chile: seasonality, feedback structure and climate

Here we present, to the authors' knowledge for the very first time for a small marsupial, a thorough analysis of the demography and population dynamics of the mouse opossum (Thylamys elegans) in western South America. We test the relative importance of feedback structure and climatic factors (rainfall and the Southern Oscillation Index) in explaining the temporal variation in the demography of the mouse opossum. The demographic information was incorporated into a stage-structured population dynamics model and the model's predictions were compared with observed patterns. The mouse opossum's capture rates showed seasonal (within-year) and between-year variability, with individuals having higher capture rates during late summer and autumn and lower capture rates during winter and spring. There was also a strong between-year effect on capture probabilities. The reproductive (the fraction of reproductively active individuals) and recruitment rates showed a clear seasonal and a between-year pattern of variation with the peak of reproductive activity occuring during winter and early spring. In addition, the fraction of reproductive individuals was positively related to annual rainfall, while population density and annual rainfall positively influenced the recruitment rate. The survival rates were negatively related to annual rainfall. The average finite population growth rate during the study period was estimated to be 1.011 +/- 0.0019 from capture-recapture estimates. While the annual growth rate estimated from the seasonal linear matrix models was 1.026, the subadult and adult survival and maturation rates represent between 54% (winter) and 81% (summer) of the impact on the annual growth rate.     

PHOTOSYNTHETIC RESPONSES TO LIGHT AND TEMPERATURE OF THE HETEROMORPHIC MARINE ALGA MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Physiological differentiation of the heteromorphic life-history phases of the red alga Mastocarpus papillatus (C. Agardh) Kützing was assessed. Photosynthetic responses to light and temperature of the erect, foliose gametophyte were compared to those of the crustose tetrasporophyte. Plants of both phases were collected from four locations on the Pacific coast of Baja California, Mexico, and California, USA, between 32–4l° N latitude. Within each location, the chlorophyll-specific, light-saturated photosynthetic rates of gametophytes were generally five times greater than those of tetrasporophytes. Initial slopes of photosynthesis: irradiance curves were greater for gametophytes than for tetrasporophytes. The crust and the blade from each location were similar with respect to dark respiration rates. For tetrasporophytes from all locations, the photosynthetic temperature optima were between 12–15° C. The photosynthetic temperature optima for gametophytes ranged from 15–17° C for plants from Trinidad Head (41° N) to 22–25° C for plants from Punta Descanso (32° N). Both gametophytes and tetrasporophytes from the northernmost location had significantly higher photosynthetic rates than the same phase from the other three locations. The photosynthetic responses to light support models for the life history in which life history phases have different ecological roles. The gametophyte, thought to be specialized for rapid growth and competition, may allocate more resources to photosynthetic machinery, hence the higher maximum photosynthetic rates. The tetrasporophyte, thought to be specialized for resistance to herbivores, may allocate more resources to structural or chemical defenses in preference to photosynthetic machinery. Consequently, the tetrasporophyte has lower photosynthetic capacity.     

SEASONAL LIGHT AND TEMPERATURE INTERACTION EFFECTS ON DEVELOPMENT OF LAMINARIA SACCHARINA (PHAEOPHYTA) GAMETOPHYTES AND JUVENILE SPOROPHYTES1

In previous studies, Laminaria saccharina L. (Lamour.) sporophytes were found to exhibit two major peaks of sporogenesis and an annual life cycle in Long Island Sound, New York. Young sporophytes were observed shortly after the sporogenesis peaks in early autumn and spring, but most of the mature sporophytes decayed during summer. A new study was conducted to determine if the spring sporogenesis activity contributed to the recruitment observed in autumn through oversummering of gametophytic and juvenile sporophytic stages, as previously suggested. Reproduction and growth in gametophytes and growth in juvenile sporophytes were studied under crossed gradients of light and temperature. Periodic outplantings of substrata seeded with gametophytic and sporophytic stages to the field were conducted to assess actual survival. The optimum temperature and light conditions for gametophyte development, growth and reproduction varied with the time of year meiospores were obtained. Most of this variation was attributable to temperature. A seasonal adaptation to temperature in most developmental stages was observed. Higher temperatures resulted in greater numbers of male gametophytes. Gametophytic stages could develop at all times, suggesting that oversummering in this stage was possible. Juvenile sporophytes had a narrower optimum temperature range and again photon fluence rate contributed little to observed variances. Out planting of sporophytic stages at various times during the year indicated only sporophytes prepared from autumn and winter could survive summer conditions. The thalli of these plants grew rapidly in spring and eroded back to the meristematic region in summer. Most of these plants then quickly became reproductive, resulting in another autumn sporogenesis peak. Gametophytic and sporophytic outplantings prepared from spring meiospores did not survive the summer. Thus, the recruitment observed in autumn can only be the result of the previous autumn's sporogenesis activity. The sporogenous activities of spring and early summer appear to be unimportant, despite the fact that all reproductive indices are superior at those times.     

Chemical races in the red alga Laurencia nipponica (Rhodomelaceae, Ceramiales)

Genetic variation in the synthesis of halogenated secondary metabolites in the Japanese marine red alga Laurencia nipponica Yamada (Rhodomelaceae, Ceramiales) has been investigated in laboratory crossing experiments and chemical analyses, F₁ tetrasporophytes and F₁ gametophytes resulting from crosses within chemical races produced major metabolites characteristic of these races. F₁ tetrasporophytes derived from reciprocal interracial crosses produced: (i) both parental types of secondary metabolites; (ii) either of the parental types; or (iii) a further major compound in addition to both parental types or in addition to either of the parental types. The latter cases suggest that hybrid-specific products were formed by the combined enzymatic complements of the parents, as F₁ gametophytes derived from these interracial F₁ tetrasporophytes yielded one or other of their parental products in an approximate 1:1 ratio. The population structure was analyzed at localities in Hokkaido, where two of the chemical races occur sympatrically. At Usujiri (Minami-kayabe), where the prepacifenol race and the laureatin race were sym-patric, hybrid gametophytes (recombination type) were found in high frequency in addition to hybrid tetra sporophytes, which strongly suggests that a new, pre-pacifenol/laureatm race is beginning to be produced by natural hybridization and recombination. By contrast, at Oshoro Bay, where the laurencin race and the epi-lauraliene race grew together, the interracial hybrids were rare: only a few tetrasporophytes (probably F₁ generation) were found, suggesting that racial integrity may be retained by habitat segregation and/or the absence of recombination-type gametophytes.     

Variation in gametophyte dominance in populations of Chondrus verrucosus (Gigartinaceae,Rhodophyta)

We describe the abundance, including spatial and temporal variability, of phases of the isomorphic Chondrus verrucosus Mikami from Japan. Chondrus verrucosus occurred in a dense (～90% cover) and temporally stable bed on a small, isolated rocky outcrop (Oyakoiwa) in Shizuoka Prefecture. Small vegetative fronds were always much more abundant than large vegetative and fertile fronds over the spring to late summer periods in 1999 and 2000. Over the same period, fertile carposporophytic fronds were generally more abundant than fertile tetrasporophytic fronds, and fertile male fronds appeared infrequently at low densities. Using the resorcinol‐acetal test, we determined the proportion of gametophytes and tetrasporophytes in three populations of C. verrucosus: Oyakoiwa and Noroshi (Shizuoka) in the summers of 1999 and 2000 and Kamehana Point (Miyagi) in autumn 2000. All populations had a significantly higher proportion of gametophytes than tetrasporophytes in both years, although gametophytic proportions were lower at Noroshi (～70%) than at Oyakoiwa (～80%) and Kamehana Point (～97%). However, examination of all isolated individuals sampled on Noroshi showed equal proportions of each phase in 1999, but gametophyte dominance (74%) in 2000. Differences in dispersal and spore production between phases are discussed as mechanisms potentially contributing to variation in gametophyte dominance.     

Cultivation of the red alga Chondracanthus chamissoi: sexual reproduction and seedling production in culture under controlled conditions

The red alga Chondracanthus chamissoi (Gigartinales) is endemic to the southern-central region of South America. In the Pacific Ocean, it is distributed from north-central Peru to Chiloe Island. This species is of economic importance because it is edible and used for carrageenan production. The tetrasporophyte phase was grown in the laboratory, obtaining male and female gametophytes that were incubated under different photoperiod, pH, salinity and temperature conditions. These gametophytes developed and generated reproductive structures that led to in vitro maturation. Subsequently, fertilisation occurred and formation of cystocarps was observed. Finally, carpospores were released and the formation of sporophytes completed the life history of this species under laboratory conditions. Reproductive phase growth rates were recorded for each of the different culture conditions used. Sporophytes reached the highest daily growth rate (22%), while gametophyte’s daily growth rate was slower (9%). This research confirms, in vitro, the assumption that C. chamissoi has a sexual triphasic life history Polysiphonia type with isomorphic gametophytes and tetrasporophytes. The development of the complete life history took 20 months in the laboratory.     

Seasonal and spatial patterns of population density in the marine macroalga Mazzaella splendens (Gigartinales, Rhodophyta)

Insight into demographic processes that operate at larger spatial scales can be achieved through studying local populations when a particular species of interest is examined over time, by many investigators, in a variety of locations. On the west coast of North America, Mazzaella splendens (Setchell et Gardner) Fredericq is such a species of interest. A synthesis of local demographic studies of M. splendens from the late 1960s to the present reveals a pattern that is potentially common to the larger natural populations. This is the pattern: population density is high in summer and low in winter for both alternate free‐living life history phases of M. splendens. The magnitude of this seasonal change decreases in increasingly wave‐exposed habitats. In wave‐sheltered habitats there is a seasonal alternation from summer haploid to winter diploid dominance. This alternation gradually changes to constant diploid dominance as wave exposure in the habitat increases. Changes in population density are primarily a function of appearances and disappearances of perennating basal crusts (genets), as modules are produced or lost, rather than differential module production by genets of one phase over those of the other. To test the generality of this pattern, we examined seasonal changes in density, in local populations of M. splendens, in both a wave‐sheltered and a wave‐exposed habitat at Second Beach, Barkley Sound. Greater seasonal fluctuation in population density at wave‐sheltered, compared to wave‐exposed habitats is supported as a pattern potentially common to the natural populations of M. splendens. A change from summer haploid dominance in wave‐sheltered areas to summer diploid dominance in wave‐exposed areas is similarly supported. All changes in population density were the result of appearances and disappearances of genets rather than differential module production by haploid versus diploid basal crusts, also consistent with previous observations. A seasonal alternation in phase dominance, however, was absent from the wave‐sheltered site at Second Beach, Barkley Sound for 3 consecutive years. Seasonal alternation in phase dominance of M. splendens appears dependent on local conditions and is not common to all natural populations.