Effects of harvesting onAscophyllum nodosum (L.) Le Jol. (Fucales,Phaeophyta): a demographic approach

Although populations ofAscophyllum nodosum are harvested commercially, little is known about the effects on demographic vital rates (growth, reproduction, survival). This study examines the effects of harvesting season and harvesting intensity on growth, reproduction and mortality of intact fronds in four size classes and in fronds truncated by the harvest. Knowledge of size-specific vital rates was used to evaluate the response of the population to harvesting.Harvesting season and harvesting intensity did not exert a significant effect on growth. Growth in plots not subject to harvesting was less than in harvested plots. No major differences in growth, reproduction and survival between intact and severed fronds emerged. The number of fronds attaining reproduction was enhanced by increased harvesting intensity and by cutting in summer. Harvesting did not seem to induce breakage, and breakage appeared higher in the uncut plots. Most harvesting treatments did not influence survivorship and survivorship was similar among all size classes. Growth rates were inversely related to sizes of fronds.Assessment of variation across size classes yielded more accurate estimates of growth rates than those of previously used methods. Accurate size class specific-growth rates will be a useful criterion when regulating intervals between harvests. Furthermore, assessment of size-specific vital rates allows identification of the frond size classes most relevant to the preservation of resources. Because of their fast growth rates and abundance, fronds in class 1, and, to a lesser extent, class 2, are responsible for most of the population regrowth after harvest. In contrast, classes 3 and 4 contribute little to recovery. This finding provides a strong basis for a harvesting strategy that targets the largest fronds.Author for correspondence     

POPULATION PARAMETERS AND PHOTOSYNTHETIC CAPABILITIES OF CORALLINE RHODOLITHS: BIOGEOGRAPHIC IMPLICATIONS

Rhodoliths are bed forming, unattached coralline algae that incorporate large quantities of carbonate into their thalli in oceans worldwide. Lithophyllum margaritae and Neogoniolithon tricotomun are common Rhodoliths from the Gulf of California, however, little is known about their biogeographic distribution, and their ecological and physiological characteristics. As a consequence, the objective of this study was to define the temporal population dynamics, growth rates and photosynthetic capabilities of L. margaritae and N. tricotomun from the Gulf of California. The Gulf of California is characterized by annual temperature fluctuations that can exceed 20° C. Rhodolith beds exceeding one km in length were observed from the intertidal zone to a depth of approximately 20 m at the southern Gulf. Percent cover estimates for spherical unattached individuals of L. margaritae and N. tricotomun ranged from 10 to 100%. Growth rates were determined by staining the rhodoliths with Alizarine red and evaluating the linear increment of calcium carbonate after an incubation period in the field. Annual growth rates averaged 1.8 mm year⁻¹ in L. margaritae and 3.4 mm year⁻¹ in N. tricotomun , however, growth rates varied seasonally. The photosynthetic response of both species as a function of temperature was evaluated in the laboratory from 10° to 30° C. Maximum net photosynthetic rates peaked at approximately 25 to 30° C in both species, suggesting that maximum growth rates occur during the warmest months of the year. These results lead to a better understanding of the biogeographic ranges of subtidal coralline algae.     

Scaling of Frond Form in Hawaiian Tree Fern Cibotium glaucum: Compliance with Global Trends and Application for Field Estimation

Large‐fronded tree ferns are critical components of many tropical forests. We investigated frond and whole‐plant allometries for Hawaiian keystone species Cibotium glaucum, for prediction and to compare with global scaling relationships. We found that C. glaucum fronds maintain geometric proportionality across a wide range of plant and frond sizes. These relationships result in strong allometries that permit rapid field estimation of frond size from simple linear dimensions. C. glaucum frond allometries complied with intra‐ and interspecific global trends for leaf area versus mass established for much smaller‐leafed species, indicating ‘diminishing returns’ in photosynthetic area per investment in mass for larger fronds. The intraspecific trend was related to declining water content in larger fronds, but not to a significantly larger investment in stipe or rachis relative to lamina. However, C. glaucum complied with the global interspecific trends for greater allocation to support structures in larger leaves. Allometries for frond number and size versus plant height showed that as plants increase in height, frond production and/or retention progressively declines, and the increases of leaf size tend to level off. These frond and whole plant‐level relationships indicate the potential for estimating frond area and mass at landscape scale to enrich studies of forest dynamics.     

DEMOGRAPHY OF FRONDS OF SARGASSUM LAPAZEANUM FROM SOUTHERN BAJA CALIFORNIA, MEXICO, DURING EARLY STAGES

In terms of biomass, Sargassum lapazeanum (Phaeophyceae, Fucales) is one of the most important seaweeds in La Paz Bay, on the southeastern coast of the Baja California Peninsula, Mexico. This species can be found year-round in shallow subtidal sites. Standing biomass is lowest between fall and winter and highest between spring and summer. We are currently studying the annual demography of fronds as a necessary step to understanding the mechanisms of population regulation. Work is being done on a bed of about 90 m in length and three m in width. At this stage of abstract submission, we can report results for winter only. Random samples were collected in February and in March 2000 to estimate stand biomass, frond density, and size (frond length) structure. In addition, we labeled fronds with numbered plastic tags to estimate their rates of growth and of mortality during this period. Rates of recruitment were estimated from a combined analysis of the above. In February, fronds were all shorter than 6.5 cm. Between February and March, mean total frond density increased from 122 to 776 fronds m⁻² (n = 30 quadrats). The mean recruitment rate was 667 fronds m⁻², the mean growth rate was 0.5 cm day⁻¹ (n = 60 fronds), and the mean mortality rate was 43 fronds m⁻². From last year's preliminary observations, we expect to observe peaks of biomass and of reproduction in late spring, followed by negative growth rates and high mortality rates during summer.     

Regeneration and reproduction of Mazzaella cornucopiae (Rhodophyta, Gigartinaceae) after frond harvesting

The effects that different intensities of frond harvesting have on frond regeneration and subsequent production of reproductive structures were investigated for the red intertidal alga Mazzaella cornucopiae (Postels & Ruprecht) Hommersand from British Columbia, Canada. Harvesting was done by pruning fronds in the late spring (when stand biomass is highest) of 1993 at two intensities: total and partial collection of fronds, in this second case leaving all frond biomass less than 1 cm high in place. Holdfasts were not damaged. Total percent cover of thalli, frond density, mean frond length, and stand biomass for these experimental quadrats were statistically similar to values for control quadrats in the spring of 1994. These results suggest that one total harvest of fronds per year, done in late spring without damaging holdfasts, may give the highest sustainable yield of biomass. The effects of harvesting intensity on reproduction were variable and difficult to explain. Neither the appearance nor the abundance of cystocarpic fronds were affected by frond pruning, compared with control areas, but pruning did affect the appearance and the abundance of tetrasporic fronds. Partial pruning resulted in a longer presence of tetrasporic fronds, whereas total pruning was associated with their complete absence. Results are compared with those for the few other species of the Gigartinaceae for which experimental harvesting has been done. This revised version was published online in August 2006 with corrections to the Cover Date.     

Allometric estimation of total leaf area in the neotropical palm Euterpe oleracea at La Selva, Costa Rica

We estimated the magnitude of the total leaf area of the neotropical palm Euterpe oleracea and examined its allometry relative to the variation in stem height and diameter at La Selva Biological Station in Costa Rica. The allometric relationships between frond leaf area and frond length (from tip to base), and between frond leaf area and number of leaflets, were determined by natural logarithmic regressions to estimate the total area of each frond. Palm total leaf area was then estimated by adding the area of the composing fronds. We fit 14 separate regression models that related one or more of the morphological variables (number of fronds, diameter at breast height, stem height) to the total leaf area. Our results show that palm total leaf area is directly proportional to the number of fronds and palm size as reflected in stem height and diameter. Eight out of the 14 models had r ² values of >0.90 and incorporated a diverse combination of predictor variables. Simple linear regression models were more congruent with the observed values of total leaf area, whereas natural logarithmic models overestimated the value of total leaf area for large palms. Both approaches show a high degree of association among morphological characters in E. oleracea supporting the hypothesis that palms behave like unitary organisms, and are morphologically constrained by the lack of secondary meristems. To afford attaining canopy heights, woody palms need to show a high degree of phenotypic integration, shaping their growth and allometric relationships to match spatial and temporal changes in resources.     

SELF‐THINNING AND SIZE INEQUALITY DYNAMICS IN A CLONAL SEAWEED (SARGASSUM LAPAZEANUM,PHAEOPHYCEAE)1

Fronds of clonal seaweeds with extensive holdfasts relative to frond size are known not to self‐thin during growth, even in crowded stands. We tested whether frond self‐thinning would occur for such a seaweed since these traits are more similar to those of unitary seaweeds, which do self‐thin in crowded conditions. We used Sargassum lapazeanum Setch. et N. L. Gardner (Fucales, Phaeophyceae) from the Pacific coast of Mexico, for which we first confirmed its clonal nature by performing a regeneration experiment in culture tanks. During the growth season (winter to late spring), S. lapazeanum stand biomass increased, while frond density and size inequality (Gini coefficient for frond biomass) decreased. These results indicate that self‐thinning occurred at the frond level. We propose a conceptual model for frond dynamics for clonal seaweeds in general. In stands of clonal species with small fronds and relatively extensive holdfasts (particularly when holdfasts are perennial), frond dynamics would be determined mostly by intraclonal regulation, which seems to prevent excessive crowding from occurring. Such species display a positive biomass–density relationship during the growth season. On the contrary, in stands of clonal species with large fronds relative to holdfast size, frond dynamics would be determined mostly by interactions among genets. For such species, self‐thinning may be detected at the frond level in crowded stands, resulting in a negative biomass–density relationship during growth.     

STRUCTURE AND DYNAMICS OF A POPULATION OF PALMARIA PALMATA (RHODOPHYTA) IN NORTHERN SPAIN1

Palmaria palmata (Linnaeus) O. Kuntze (Rhodophyta, Palmariaceae) is a seaweed commercially harvested for human consumption. Its population density, size structure, and frond dynamics were investigated from May 1999 to May 2001 at one intertidal locality in the northern coast of Spain, which is within the southern distributional boundary of the species in the eastern Atlantic coasts. The effect of size, age, and the life‐history phase (haploid vs. diploid) on frond growth and mortality were also evaluated. The study was carried out by mapping and monitoring fronds in the field. New fronds (macroscopic recruits or sprouts) appeared in spring, but subsequent mortality of these young fronds and detachment of the host plant led to lower density values in January. Palmaria palmata exhibited a distinctive seasonal growth cycle, with positive net growth from March to August and breakage from August to March. Interannual differences were also detected, with higher net growth in 2000 than in 1999. Net growth was apparently independent of age, reproductive status (fertile vs. reproductive), and life‐history phase (haploid vs. diploid) but was dependent on size, as longer fronds showed minor growth or greater breakage than small ones. Mortality, on the other hand, was more dependent on age than on size in the period analyzed (March–May 2000). Results of the study indicate that both size and age should be included as state variables and temporal changes in transition probabilities considered in the development of demographic models of the species.     

A pilot-scale study of the vegetative propagation and suspended cultivation of the carrageenophyte alga Gigartina skottsbergii in southern Chile

Different propagation techniques for cultivation of vegetative Gigartina skottsbergii fronds were tested using a system of suspended ropes, to which inoculants were attached. Our results showed that triangular fragments obtained from the circular G. skottsbergii thalli produced harvestable frond of 800 cm² after 8 months. In contrast, inoculants of intact juvenile fronds of comparable size needed at least 10 months to reach the same size. A control experiment with spores developing on an artificial substrate showed that 20 months were needed to reach a surface of 500 cm², confirming the superiority of our fragment culture system. A pilot study demonstrated that with a density of six fronds m⁻¹ of farming line, the proposed system can be economically interesting for local fishermen.     

ABSCISIC ACID EFFECTS ON FRONDS AND ROOTS OF LEMNA MINOR L.

Plants of Lemna minor L. were grown in axenic culture in order to investigate the direct effect of abscisic acid (ABA) on individual fronds and roots. Both frond and root growth rates were inhibited by 60%, and total growth of both organs was reduced by 30% with ABA concentrations of 2 mg/l. Abscisic acid inhibited frond reproduction, and treated fronds tended to remain attached to parents. Reduction of root elongation and frond expansion occurred within 1–4 hr. Prolonged exposure to ABA inhibited both cell enlargement and cell division in the roots. The results are discussed in the light of current views on the control of growth with endogenous levels of promoter and inhibitor hormones.     

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.     

Non-timber forest product harvesting in alien-dominated forests: effects of frond-harvest and rainfall on the demography of two native Hawaiian ferns

Non-timber forest products (NTFP) represent culturally and economically important resources for millions of people worldwide. Although many NTFP are harvested from disturbed habitats and therefore subject to multiple pressures, few quantitative studies have addressed this issue. Similarly few NTFP studies have assessed seasonal variation in demographic rates even though this can confound harvest effects. In Hawaiȁ8i, the wild-gathered ferns, Microlepia strigosa and Sphenomeris chinensis, represent highly important cultural resources but declining populations have led to conservation concerns. Both ferns are harvested from disturbed, alien-dominated forests and contemporary Hawaiian gathering practices often consist of harvest and concurrent weeding of alien invasive species. We assessed the effects of concurrent frond-harvest and alien species weeding on frond structure, density, and rates of production by comparing experimentally harvested vs. control plots, and documented relationships between frond demographic patterns and precipitation. Gathering practices had no impact on frond density of either species or on most other demographic parameters over the short term. Exceptions included a significant decrease in the density of the longest S. chinensis fronds and a significant decrease in M. strigosa frond production when fronds were gathered without alien weeding. However, seasonal and annual changes in frond density and production occurred across all plots of both species and were significantly correlated with precipitation. The relatively low harvest effects for both species are likely due to several factors including short frond longevity and the strict criteria used by gatherers to select harvestable fronds. The potential for sustainable harvest in the context of alien-dominated forests is discussed.     

Frond elongation rates of shallow waterMacrocystis pyrifera (L.) Ag. In northern Baja California,Mexico

Macrocystis pyrifera (L.) Ag.frond elongation rates were measured during autumn-winter, spring and summer in a shallow water (7.5 m depth) kelp bed in Bahía Papalote, northern Baja California. Frond elongation was maximum during spring and minimum during winter. Frond growth was significantly correlated with solar radiation, and was highest in the smallest size fronds (0–2 m). Average frond elongation rate varied between 0.3–11% d^–1 during the study period. The relationship between average frond length and elongation rate followed an exponential curve with a negative slope during autumn-winter and summer, but was best described by a straight line during spring. Standard growth rates were obtained by a graphic method. Standard growth rates had intermediate values between those reported for southern California and southern Baja California.M. pyrifera growth cycle shows a different trend from what has been previously reported.     

Succession and growth rates of encrusting crustose coralline algae (Rhodophyta,Cryptonemiales) in the upper fore-reef environment off Ishigaki Island,Ryukyu Islands

Observations were made on the succession and growth rates of crustose coralline algae growing in situ on artificial substrata in a shallow fore-reef environment on Ishigaki Island, Ryukyu Islands. Succession in well-illuminated environments manifests itself as a gradual replacement of species having very thin thalli by those having larger and thicker thalli. The species Porolithon onkodes, Paragoniolithon conicum and Lithophyllum insipidum achieved dominance by competitive interactions of overgrowing margins. The thicker species recruit quickly (within the first few months), but because of their slow growth rate do not displace the pioneer species that have very thin thalli until after the latter begin to die. Regardless of seasonal temperature fluctuations, which exceed 10 °C, the coralline algal succession is the same for each season. The maximum lateral growth rates of the major species range between 2.9 and 3.9 mm/month. Vertical growth rates of Porolithon onkodes, the thickest species, are the most rapid (more than 2 mm/year at maximum) relative to those of other species. Accretion rates of entire coralline algal cover on ungrazed substrata range from 1.0 to 1.2 mm/year (not allowing any lag time for recruitment), whereas those of grazed substrata are lower. These results are consistent with species which are ecological equivalents and live in similar environments on Caribbean reefs.     

Aragonite infill in overgrown conceptacles of coralline Lithothamnion spp. (Hapalidiaceae,Hapalidiales, Rhodophyta): new insights in biomineralization and phylomineralogy

New empirical and quantitative data in the study of calcium carbonate biomineralization and an expanded coralline psbA framework for phylomineralogy are provided for crustose coralline red algae. Scanning electron microscopy (SEM) and energy dispersive spectrometry (SEM‐EDS) pinpointed the exact location of calcium carbonate crystals within overgrown reproductive conceptacles in rhodolith‐forming Lithothamnion species from the Gulf of Mexico and Pacific Panama. SEM‐EDS and X‐ray diffraction (XRD) analysis confirmed the elemental composition of these calcium carbonate crystals to be aragonite. After spore release, reproductive conceptacles apparently became overgrown by new vegetative growth, a strategy that may aid in sealing the empty conceptacle chamber, hence influencing the chemistry of the microenvironment and in turn promoting aragonite crystal growth. The possible relevance of various types of calcium carbonate polymorphs present in the complex internal structure and skeleton of crustose corallines is discussed. This is the first study to link SEM, SEM‐EDS, XRD, Microtomography and X‐ray microscopy data of aragonite infill in coralline algae with phylomineralogy. The study contributes to the growing body of literature characterizing and speculating about how the relative abundances of carbonate biominerals in corallines may vary in response to changes in atmospheric pCO₂, ocean acidification, and global warming.     

Plant response to overcrowding – Lemna minor example

Plants adopt various strategies in response to increasing density. We tested that response in two populations of Lemna minor L. – a free floating aquatic plant that frequently experiences intraspecific competition for space. Surface area of fronds and colonies, colony size (the number of fronds per colony), the rate of reproduction (based on the number of produced fronds) and growth rate (enlargement of surface area of all colonies) were the analysed factors presumably affected by density. The study was performed in natural stands and in experimental conditions with the use of two contrasting plant densities. Plants growing in natural conditions produced fronds of smaller and less variable surface area as a response to overcrowding but the number of fronds per colony was unrelated to plant density. Stable experimental conditions facilitated formation of fronds and colonies larger than in the field but frond detachment decreasing colony size was more intensive at high than at low density. This strategy allowed plants to more efficiently occupy limited available space. No self-thinning was observed during experimental cultures. Due to increasing frond area in cultures, growth rate was always higher than the rate of plant reproduction. Both were strongly negatively affected by high density. Performed calculations indicate that density-dependent growth inhibition starts when L. minor colonies cover the available water surface with a mono-layer mat. Some types of responses were found to significantly differ between analysed populations, which was also shown by genetic differences tested with he ISSR-PCR technique. Possible causal relationship between plant strategies and their genomic structure needs, however, further studies.     

Variation on a theme of herbivory: Corallina–hermit crab relationship on a temperate‐subtropical rocky shore

Elucidation of species interactions involving multiple processes is an important task in ecology, as different processes may operate in non‐apparent, opposing directions. While herbivory is a major form of interspecific interaction affecting plants, calcareous algae are generally thought to be protected from herbivory due to CaCO₃ deposition. However, field observations and experiments in temperate‐subtropical intertidal rockpools showed that hermit crabs (Paguroidea, Decapoda) did not ingest Corallina but cut short its fronds. We tested the implications of this frond‐cutting behaviour with field experiments where hermit crab density and Corallina frond length were artificially manipulated. Hermit crab guts were fuller and contained more microscopic animals when Corallina fronds were shorter, suggesting that frond‐cutting affects accessibility to food resources that exist in the basal part of algal mat. Corallina showed compensatory growth when fronds were artificially reduced in length/density, simulating the cutting by hermit crabs. Frond density increased, probably due to increased exposure to light of the basal part of Corallina coupled with a reduction in apical dominance. The present study revealed a direct, non‐trophic relationship between hermit crabs and Corallina in which the former affected the latter for gaining access to food resources. Overall, the combination of positive and negative effects of hermit crab led to a positive, facilitative effect on Corallina. In terms of the effect on algae or plants, this relationship is functionally analogous to herbivory, without involving direct consumption. We discuss the problem of extending the concept of ecological engineering to herbivory and herbivory‐like phenomena and suggest the term ‘pseudo‐herbivory’ to describe relations in which animals have non‐trophic but herbivory‐like effects on plants/algae, as seen in the hermit crab–Corallina relationship.     

TRANSLOCATION OF 14C IN MACROCYSTIS INTEGRIFOLIA (PHAEOPHYCEAE)1,2

Translocation patterns in the giant kelp, Macrocystis integrifolia Bory, were investigated in situ using ¹⁴C tracer; sources and sinks were identified. Export was first detected after 4 h of labeling; experiments were routinely 24 h continuous ¹⁴C application. Mature blades exported ¹⁴C to young blades on the same frond and on younger fronds, as well as to sporophylls and frond initials at the bases of the fronds. Blades <0.3 m from the apex imported and did not export; this distance did not change seasonally. In spring export from blades 0.3–1.25 m from the apex was exclusively upwards; older blades also exported downwards. In fall downward export began 0.5 m from the apex, and blades >2 m from the apex exported exclusively downwards. Carbon imported by frond initials, young fronds, and sporophylls in fall may partly be stored for growth in early spring. No translocation was seen in very young plants until one blade (secondary frond initial) bad been freed from the apical blade; this blade exported to the apical blade for a time, but imported when it began to develop into a frond. The second and third formed blades on the primary fronds (sporophylls also exported when <0.3 m from the apex, and later stopped. Frond initials and sporophylls on later-formed fronds did not export at all. The translocation pattern in M. integrifolia differs from that previously reported in M. pyrifera in seasonal change and in distances from the apex at which the changes take place.