Changes in the Brown Seaweed Ascophyllum Nodosum (L.) Le Jol. Plant Morphology and Biomass Produced by Cutter Rake Harvests in Southern New Brunswick, Canada

Shoots and clumps of shoots of the commercial brown seaweed Ascophyllum nodosum (“rockweed”) add to the benthic complexity of the intertidal environment, providing an important habitat for invertebrates and vertebrates. To protect the structure of this habitat, management plans for the rockweed harvest of southern New Brunswick include restrictions on gear type and exploitation rates limited to 17% of the harvestable biomass. However, owing to physical and environmental factors, the harvest is not homogeneous, creating patches of exploitation ranging from 15 to 50%.A direct relationship existed between clump vulnerability, weight and length in a controlled harvest at 50% exploitation within 8 m by 8 m plots. At this exploitation rate, the gear rarely impacted clumps below 50 g or 60 cm in length. Clumps larger than 300 g and 130 cm were reduced by up to 55% of their length and 78% of their biomass. The overall impacts of the harvest on intertidal habitat is however of short duration as biomass recovers after a year of the experimental harvest. The rapid recovery is mostly due to a stimulation of growth and branching of the suppressed shoots of the clumps. Some harvested plots showed biomass even higher than initial levels, suggesting an increase in productivity at least during the first year after the harvest.     

Biomass and height of Ascophyllum nodosum after two decades of continuous commercial harvesting in eastern Canada

With the increasing demand for seaweed resources worldwide, management must ensure that the harvest of wild seaweed stocks is sustainable. We evaluate the impact of over 25 years of commercial harvesting of Ascophyllum nodosum in eastern Canada by comparing the biomass and height of the seaweed in the late 1990s to the late 2010s over a broad spatial scale spanning the provinces of Nova Scotia and New Brunswick. There has been no significant decrease in the biomass of A. nodosum in either province, and biomass has increased in some regions of New Brunswick during that period. The average height of A. nodosum has decreased by 7.8 cm in Nova Scotia while it increased by 13.8 cm in New Brunswick. Biomass of A. nodosum in unharvested sectors was 7% higher than that in harvested sectors while height of A. nodosum in unharvested sectors in New Brunswick is similar to the values observed in harvested sectors. Over the same period, water temperature has increased in both provinces and, in recent years, has at times exceeded the optimal growing temperature for A. nodosum within bays in Nova Scotia. We conclude that the current management and harvest of A. nodosum in eastern Canada are sustainable and maintain the biomass and height of A. nodosum beds but that control sites are necessary to offer adequate comparisons as environmental conditions are changing.

     

Changes in composition of rockweed (Ascophyllum nodosum) beds due to possible recent increase in sea temperature in Eastern Canada

Ascophyllum nodosum (rockweed) is the main economic resource of the seaweed industry in the Atlantic Provinces of Canada. The annual harvest steadily increased since 1995, reaching a historic peak of 37,000 tonnes in 2007. Due to a high demand for fertilizers and animal feed supplements derived from rockweed, this trend seems likely to continue. The current management plan for the sustainable harvest of the A. nodosum resource is considered conservative. The resource has been managed with a precautionary approach since 1995 to protect the integrity of the habitat. Acadian Seaplants Limited (ASL) has been granted approximately 90% of the government-issued licenses to harvest A. nodosum resources in the Maritimes. The Canadian approach is based on an annual harvest from a given bed and not strip-and-return after several years. Since 1995, ASL has proactively undertaken extensive annual surveys and research on biomass productivity of this renewable resource to establish acceptable annual exploitation rates. Historically, the rockweed beds of southwestern Nova Scotia (NS) have been almost 99% pure A. nodosum, with a minor component of Fucus vesiculosus. However, since 2004, a steady increase in F. vesiculosus, with a peak of 4.6% of the total biomass in 2008, was recorded. This coincided with one of the mildest winters on record for the Maritimes. This increase in temperature seemed to be also responsible for an unusual recruitment of the blue mussel Mytilus edulis in rockweed beds in some areas of southern New Brunswick (NB) in 2006, causing the detachment of up to 30% of the seaweed biomass in some harvesting sectors. Another phenomenon observed in southwestern NS during 2003 and 2004 was extensive ice damage on rockweed beds produced by an early melting of the ice, with losses of up to 90% of the rockweed biomass in some areas.     

Management and production of the brown algae Ascophyllum nodosum in the Canadian maritimes

Commercial exploitation of Ascophyllum nodosum (rockweed) along the coast of Nova Scotia began in the late 1950s when it was used as a raw material for manufacturing alginate and “kelp” meal. Today, this resource is used as a biostimulant extract for crops and animal feed supplements and is the main economic resource of the seaweed industry in the Maritime Provinces and Canada. The management of rockweed, by dividing the resource into many sectors, permitted the assessment of yield per unit area of bed. In Nova Scotia, mechanical harvesting of A. nodosum operated on a pulse of 2- to 3-year schedule providing yields of 21.9 to 47.7?wet?t ha^?1 averaging 35.3?±?7.6?wet?t ha^?1. Acadian Seaplants Limited (ASL) has become the dominant player in the region since 1995 with more than 75% of the total biomass under its leases and more than 90% of the total landing of rockweed in the last 13?years. Sustained harvests at ASL leases and under an annual harvest schedule using hand cutting methods have averaged 17.4?±?2.6?t ha^?1. Exploitation rates above 35% of the harvestable crop lead to a pulse harvest strategy and the need to move infrastructure year after year. In New Brunswick, a new approach to management began in 1995 with a regulated 17% exploitation rate. Thus, the yield in that province is 14.3?wet?t ha^?1. The current summer standing crop for this region has been calculated at 352,723?wet?t, covering an area of 4,960?ha, with an average biomass of 71.3?t ha^?1. The harvest in the region reached peak landings in 2010 with just over 40,100?t. The consistent yield per hectare of A. nodosum beds is proof of good management practices and an ecologically sustainable harvest in the Canadian Maritimes.     

Changes in the population structure of Ascophyllum nodosum (L.) Le Jolis due to mechanical harvesting

The use of a Norwegian suction cutter to harvest populations of the brown alga Ascophyllum nodosum (L.) Le Jolis in southwestern Nova Scotia started in 1985. The impact of this type of mechanical harvest on the algal population structure was evaluated. Changes in the length and density of individual plants (clumps) within 0.25 m^–2 quadrats, as well as the length of individual shoots within clumps were monitored before and after an experimental harvest. The mechanical harvest cut mainly the longer plants, thus changed the initial bimodal size structure of the population to unimodal. There was a 20 to 36% plant mortality, reducing the plant density from 92.6 to 73.6 individuals m^–2. Pre-harvest size distribution of the tagged shoots was skewed to the right and became more normal after the harvest. Tagged shoots in the harvested quadrats suffered a 42% mortality as compared to 11% of those in the control quadrats. An understanding of the impact of the mechanical harvesting on the harvested population is essential in the design of a management strategy. Sources of variation in the impact of mechanical harvest include the tide level at time of harvest, length of time the machine operated in one site, skill of the machine operator, and sharpness of the machine cutting blades.     

Comparison of the structure of populations of Ascophyllum nodosum (Fucales,Phaeophyta) at sites with different harvesting histories

Changes in the structure of the Ascophyllum nodosum population at Pubnico, southwest Nova Scotia, Canada, at an experimental site subjected to mechanical harvest and at two control sites never subjected to mechanical harvest were monitored from 1991 to 1994. A bimodal population structure measured in terms of plant length was characteristic of all these sites before the experiment. The population structure of the experimental site became unimodal immediately after experimental harvest by machine and remained unimodal for the subsequent two years. However, a bimodal population structure began to appear in the third year. A bimodal population structure remained evident at the control site with bedrock as the substratum but was less evident at the other control site where the substratum is made up of boulders and cobbles. Movement of loose rocks with rockweeds still attached may have contributed to the less distinct modal structure of this control site. Other sites with different harvesting histories monitored in the summer of 1992 showed some interesting patterns. A unimodal population structure was evident in Argyle Sound and Pubnico Point South and at Charlesville, which had been harvested one and two years before, respectively. A bimodal population structure was more evident at Frenchman's Point, which had been harvested three years prior. The rate of change from a unimodal to a bimodal population structure may depend on the intensity of harvest. Extensive canopy removal in intensively harvested areas may be conducive to an influx of recruits and to regeneration from the holdfast. Hence, plant length modal structure may be a useful measure of the relative state of recovery of a harvested population.     

Impact of Culm Harvest on Seed Production in a Monocarpic Bamboo

Successful regeneration of bamboos from seed is a key issue in the ecology of many tropical regions and the livelihoods of their inhabitants. The gregarious monocarpy of many bamboos may be driven by a need to satiate seed predators by seeding in abundance at infrequent intervals. In long‐lived clonal monocarps, seed production is expected to be positively related to the success of the clone in generating more and larger ramets during its lifetime. Ramification may be constrained by harvesting of culms, but it is unclear whether the reduction in productivity is proportional to the loss of reproductive biomass. We counted the seed produced by 661 culms (ramet stems) sampled from 90 clumps of the gregariously monocarpic bamboo Schizostachyum dullooa that is intensively harvested by villagers in northeastern India. The smallest clumps had fewer culms and few or no culms more than one year old. Seed production was indeed positively related to culm size and the number of culms in a clump. First‐year culms were markedly more productive than older culms after controlling for culm diameter and clump size. There was a negative effect of clump size on productivity per culm which may occur because clumps that had been harvested heavily were able to exploit resources retained in rhizomes from harvested culms. Nevertheless, small clumps produced much less seed than larger clumps, generating a risk of unknown magnitude that heavily harvested stands of monocarpic bamboos may be unable to satiate seed predators during their single opportunity for reproduction.     

Harpacticoid copepod communities of floating seaweed: controlling factors and implications for dispersal

The structure of harpacticoid copepod communities of floating detached intertidal algae was studied by surveys and experiments. Seaweed clumps were sampled from Faxaflói, western Iceland, at various distances from the shore (0.2–14 km). Altogether 20 species of harpacticoids were found in these clumps, most of them typical phytal species. Multidimensional scaling ordination (MDS) separated the samples into three distinct clusters in which the communities were significantly different from each other. The distance from shore did not seem to affect the community structure. However, diversity and abundance of harpacticoids increased with size and complexity of the clumps. Experimental Ascophyllum nodosum fronds placed 300–600 m offshore were rapidly colonised by harpacticoids. Density and species diversity were, after 20 days afloat, higher than in the free-floating clumps. Species of the genus Harpacticus were successful colonisers, in particular Harpacticus chelifer and Harpacticus uniremis. In a survey of macrofauna (0.5-mm sieve) on floating algae clumps from various locations around Iceland, H. chelifer was present in all clumps studied, up to 117 km offshore. Its sex ratio was skewed towards females, often with a high ratio of ovigerous females. There was, however, no evidence that this species is using the algal clumps as a platform for reproduction.     

Analysis of the multi-stem clump structure ofLitsea japonica Juss. growing in a coastal dwarf forest

The multi-stem clump structure of a coastal dwarf forest dominated byLitsea japonica Juss. was investigated in order to clarify the sprouting characteristics and self-maintenance of clumps by stem alternation. The size and age distribution of multi-stem clumps were analyzed using cumulative relative frequency curves.L. japonica had a large number of stems and an even height distribution or young age-biased distribution of stems within a clump. These results indicated the sequential flushing of sprouts at high frequency. Height distribution within a clump ofL. japonica was relatively even compared to other species. This clump structure suggested the stable self-maintenance of individuals in all ranges of size and age without disturbances. It originated specific sprouting characteristics as a response to the severe stress of salty wind.Ardisia sieboldii Miq. had few stems within a clump. Although the stem height distribution of large individuals tended to be even, most clumps had a large size-biased distribution of stem height which indicated simultaneous sprouting. From this structure, sprouts of this species were thought to be of less significance in the stable self-maintenance of individuals thanL. Japonica.     

Gametophyte-sporophyte coalescence in populations of the intertidal carrageenophyte <Emphasis Type="Italic">Mazzaella laminarioides</Emphasis> (Rhodophyta)

Mazzaella laminarioides has consistently been reported as a typical coalescent/clump species with a triphasic life history of the Polysiphonia-type in which the haploid gametophyte is the predominant phase with respect to the diploid sporophyte. Preliminary observations of intertidal populations revealed that, in some instances, cystocarpic and tetrasporic fronds emerged from the same clump (G-T clumps), implying a coalescent process of haploid and diploid thalli by fusion of their corresponding adjacent basal holdfasts. Population surveys at three sites in Coliumo Bay, central Chile, were carried out to characterize frond demography as well as to asses the frequency of gametophyte-tetrasporophyte (G-T) coalescence. Visual and resorcinol methods were employed to determine the phases of the fronds collected over central transects of 15 randomly sampled clumps. Coalescence of G-T clumps was infrequent, with gametophytes dominating over tetrasporophyte thalli.     

Extracts of the Brown Seaweed <Emphasis Type="Italic">Ascophyllum nodosum</Emphasis> Induce Gibberellic Acid (GA<Subscript>3</Subscript>)-independent Amylase Activity in Barley

Extracts of the brown seaweed Ascophyllum nodosum have been used as a biostimulant to promote growth and productivity in a number of agricultural production systems. Although the extracts have been shown to improve seedling emergence and vigor in a variety of plants, including barley, the mechanism(s) of this growth-promoting effect is(are) largely unknown. In our study, A. nodosum extract induced amylase activity in barley seed-halves; a significant difference in amylase activity was observed in seeds without an embryo. The addition of activated charcoal to the treatment media negated the bioactivity of the extracts suggesting the organic nature of bioactive compounds in A. nodosum extracts. The extracts induced amylase activity in a gibberellic acid (GA)-deficient barley mutant (grd2). LC-MS-MS analysis failed to detect the presence of GA₃ in the extracts. ABA supplementation of the medium caused a significant reduction of amylase activity in GA-treated seeds compared with those treated with the A. nodosum extract. Taken together, our results suggest that the organic components of A. nodosum extract induce amylase activity independent of GA₃ and might act in concert with GA-dependent amylase production leading to enhanced germination and seedling vigor in barley. Being derived from a renewable resource, the bioactive compounds from A. nodosum could be used to improve crop productivity in sustainable agricultural systems.     

Sexual dimorphism in clonal growth forms and ramet distribution patterns in <Emphasis Type="Italic">Rumex acetosella</Emphasis> (Polygonaceae)

We investigated clonal traits in the dioecious herb Rumex acetosella to characterize sexual dimorphism in clonal forms and to correlate below-ground clonal patterns and above-ground ramet distributions. We recorded creeping root length, branching patterns, ramet and clump (caespitose ramets from the same position on the root) sprouting patterns, and biomass allocations in three females and males. We also estimated the patch size of flowering ramets within a quadrat. No sexual dimorphism was detected in the frequencies of branches and flowering ramets per root length. Male plants allocated proportionally more biomass to below-ground organs. Total root length did not differ between the sexes. Females sprouted more clumps with fewer flowering ramets per root length than males, which sprouted fewer clumps with more flowering ramets, which meant that clump sprouting patterns were phalanx-like in females and guerrilla-like in males. Flowering ramets were aggregately distributed in both females and males and patch sizes were similar between sexes, indicating that the spreader propagations were not found in the guerrilla-like males. We assumed that sexual dimorphism occurred in response to physiological integration for higher reproductive effort in females.     

Effect of epiphytism on reproductive and vegetative lateral formation in the brown, intertidal seaweed Ascophyllum nodosum (Phaeophyceae)

In the brown alga Ascophyllum nodosum (L) Le Jolis, a common species on sheltered Northern temperate rocky shores, gametes are produced in receptacles that emerge from small depressions (lateral pits) along the branched frond. These lateral pits are also the preferred settling site for the obligate epiphyte Polysiphonia lanosa (L) Tandy. Therefore, epiphytism can be expected to interfere with host reproductive output. The present study investigated the potential impact of the epiphyte on A. nodosum in two series of laboratory experiments that measured: (i) the direct shading of the host plant underneath an epiphyte canopy; and (ii) the development of receptacles in clean and epiphytised A. nodosum segments (excised from individual fronds) over a 6 month period. These experiments showed that light reaching emerged fronds underneath a dense epiphyte cover was reduced by 40%, and this was independent of the degree of desiccation the epiphyte experienced. Concurrently, in the growth study with epiphytised A. nodosum segments (segments with one clean and one epiphytised lateral pit) total receptacle biomass per epiphytised fragment was significantly reduced compared with clean segments (0.52 g and 1.25 g per gram of frond segment, respectively), although this effect was only significant in A. nodosum from sheltered shores. However, expressed as biomass per lateral pit, receptacle biomass in the remaining clean lateral pits in epiphytised segments was significantly increased in segments from both shores, demonstrating that A. nodosum can at least partially compensate for the loss of production resulting from epiphytism.     

Coalescence in wild organisms of the intertidal population of Lessonia berteroana in northern Chile: management and sustainability effects

Lessonia berteroana is the subject of 85 % of the total harvest of Chilean brown seaweeds, representing close to 10 % of total kelp biomass worldwide harvested for alginic acid extraction. Frequent incidence of coalescent holdfasts was detected in natural populations, and this process gives rise to fused sporophytes. This study presents the coalescence events in natural populations in northern Chile. During 2011, in natural intertidal populations, 435 target sporophytes of different sizes were observed weekly and 63.90 % (278) showed physical signs of coalescent disks. We were able to distinguish five fusion modes depending on the number and size of each of the participant sporophytes and their spatial distribution. There was a progressive decrease of density and an increase in the number of stipes of each sporophyte over time. Two processes were recognized: active fusion of juvenile sporophytes and passive fusion of adult and senescent sporophytes. However, most of the coalescence processes were detected in juvenile sporophytes with a holdfast diameter of 0.5 to 2 cm. The minimum distance between pairs of coalescing sporophytes was 0.5 cm, and the maximum distance was 13 cm. For good harvesting practices, it is recommended that plants with over 20 cm holdfast diameter are harvested from natural populations because over this size, all of the plants have reproduced. Coalescence events would produce plants with legal size requirement; however, these plant units would not be reproductive, affecting the sustainability of this important coastal resource.     

An Analysis of Clumping in the Soil Amoeba Mayorella palestinensis*

SYNOPSIS. Mayorella palestinensis, a small soil amoeba, clumped during active growth in axenic, fluid culture media. Clumping was not associated with starvation (as in the case of cellular slime molds) or with encystation Hartmannella rhysodes was used as a control amoeba which did not clump to the same extent, but otherwise resembled M. palestinensis. Mixtures of both amoebae formed mixed clumps, without subsequent segregation of amoebae in the clumps. Clumping by M. palestinensis was temperature dependent, localized to the surface of the amoeba and required surface conformation. The requirement for a living system could be separated from the surface specificity of M. palestinensis and satisfied with living amoebae that did not clump by themselves, i.e., H. rhysodes, but did clump with dead M. palestinensis. It is proposed that amoeba-to-amoeba adhesion is similar to foreign substrate adhesion, and that M. palestinensis adheres to itself because its surface is more suitable for adhesion than other surfaces present in the culture. A competition for adhesion exists between surfaces so that clumps can be dissociated by providing a foreign surface that is more suitable for adhesion than the surface of the amoeba.     

SPATIAL DISTRIBUTION OF GENETIC INDIVIDUALS IN THICKETS OF ALNUS INCANA SSP. RUGOSA,A CLONAL SHRUB

Thickets of speckled alder (Alnus incana ssp. rugosa (Du Roi) Clausen) consist of numerous discrete clumps of stems. Presumably all stems in a single clump are part of a single genetic individual, but a genet could comprise more than one clump. Starch-gel electrophoresis was used to identify genetic individuals in four alder populations in central New York. A single genetic marker, a tetramer with three alleles, could discriminate five genotypes. Nearest neighbor analysis revealed that genotypes were distributed randomly. That is, the pattern of genotypes was statistically indistinguishable from a model where each clump is considered a unique individual and where clump genotypes are randomly distributed. Calculation of Morisita's index of dispersion confirmed that clumps of a single genotype were not aggregated. Although alder is capable of forming root suckers and offsets, lateral expansion of genets is apparently ineffective. Apparently, spatial distribution of genetic individuals within alder thickets is not influenced by clonal growth or by other factors acting to cause patterns in the genetic structure of plant populations.     

Growth studies of macroalgae in a microcomputer-assisted spray cultivation system

Apical growth rates of Furcellaria lumbricalis (Huds.) Lamour. and Ahnfeltia plicata (Huds.) Fries (Gigartinales: Rhodophyceae), and Ascophyllum nodosum (L.) Le Jol. (Fucales: Phaeophyceae), were studied in spray culture, applying independent microcomputer systems for control, data-logging, and apical growth measurements. The system functioned with a minimum of maintenance. Sea-water temperature and light seemed to be more important for growth than nutrient supply under the prevailing conditions. Basal shoots (young, undivided parts of the plants close to the holdfast) of Furcellaria lumbricalis had growth rates three times higher than the apices (apical meristem on the distal end of mature plants) from the older parts of the thallus. For Ascophyllum nodosum apical growth rates of basal shoots, divided and undivided apices were similar. Whereas apical growth rate of A. nodosum was normal, the biomass yield was low. A. nodosum cultivated simultaneously in tanks and in continuous spray had similar growth rates. The spray cultivation technique facilitated control of the experimental conditions, and handling of the plants during the experimental work while excluding epiphytes.     

Sex ratio from germination through maturity and its reproductive consequences in the liverwort Sphaerocarpos texanus

Summary The dioecious winter ephemeral liverwort, Sphaerocarpos texanus disperses spore tetrads consisting of two males and two females. I examined the subsequent sex ratio of S. texanus at different stages in its life cycle to detect possible mechanisms affecting deviations from a 1:1 sex ratio and the effect of sex ratio on reproductive success. As S. texanus occurs in pure male, pure female and mixed sex clumps, I examined the proportions and sizes of these, the reproductive success of pure female and mixed sex clumps in the field and the sex ratio of germinating plants in a growth chamber. In both the field and the growth chamber the most abundant clump type was pure female followed by mixed sex and pure male clumps. These abundance patterns suggest that males have a lower survival rate than females before emergence and this lower survival rate continues through the gametophytic stage. This disadvantage may be due to the higher susceptibility of males to environmental conditions, to their competitive inferiority to females, and/or to differential resource allocation to the sexes within the spore tetrad. The female biased sex ratio at germination is consistent with predictions from sex ratio theory. Further my field data indicate that males may gain a survival benefit from growing in a mixed sex clump and both males and females benefit reproductively when they occur in mixed sex clumps.     

An assessment of the effect of harvest time and harvest method on biomass loss for Miscanthus × giganteus

This study examines the amount of biomass loss occurring in Miscanthus × giganteus crop at harvest. The study assesses loss incurred as a direct result of the harvest systems employed to collect the material along with examining how the time of harvest effects the amount of loss occurring over the spring harvest window. Pre harvest losses of 4.8–5.1% were measured prior to harvest. There was no significant difference between pre harvest loss and post harvest loss when a self‐propelled forage harvester fitted with a maize harvesting header was used to harvest the crop. The use of a conditioner mower and baler significantly increased crop losses to 9.4–14.1%. This demonstrates that correct selection of the harvest system can significantly increase biomass recovery. Additional losses were measured at headlands when the mower/baler system was used, but headland losses will not occur when self‐propelled forage harvesters are utilized. Losses were significantly greater in the area beside the swath after the baler pass when compared to prior to baling. This study has shown that correct selection of harvest systems can significantly increase biomass recovery, with no significant difference in pre harvest loss or harvest loss occurring as a result of cutting the M. × giganteus crop at different dates during the harvest window (March 1st, March 25th, April 21st).     

EFFECTS OF AVIAN SEED DISPERSAL ON THE GENETIC STRUCTURE OF WHITEBARK PINE POPULATIONS

We used allozyme analysis to examine family structure, the spatial patterning of related individuals, in two populations of whitebark pine (Pinus albicaulis), a subalpine conifer that commonly displays a multistem form. The individual stems within clumps are genetically distinct individuals, having arisen from separate seeds. Individuals within a clump are genetically more similar than individuals in different clumps, but individuals in neighboring clumps do not appear to be more similar than individuals in distant clumps. This family structure appears to be a direct result of the seed-caching behavior of Clark's nutcrackers (Nucifraga columbiana), the primary dispersal agent for whitebark pine seeds.