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.     

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.     

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.     

THE IMPACT OF FROND CROWDING ON FROND BLEACHING IN THE CLONAL INTERTIDAL ALGA MAZZAELLA CORNUCOPIAE (RHODOPHYTA, GIGARTINACEAE) FROM BRITISH COLUMBIA, CANADA

The importance that frond crowding represents for the survival of fronds of the clonal intertidal alga Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (Rhodophyta, Gigartinaceae) was investigated in Barkley Sound, British Columbia, Canada. Frond density is high for this species, up to 20 fronds·cm^?2 in the most crowded stands. Frond crowding imposes a cost in the form of reduced net photosynthetic rates when fronds are fully hydrated as a result of reduced irradiance compared with experimental (not found naturally) low-density stands. However, the interaction between desiccation and irradiance alters this relationship between net photosynthetic rates and frond density. During a typical daytime low tide in spring, irradiance is 10–30 μmol·m^?2·s^?1 below the canopy of fronds, and frond desiccation (relative to total water content) can reach 43% at the end of the low tide. In contrast to natural stands, fronds from experimentally thinned stands are subjected to irradiances up to 2000 μmol·m^?2·s^?1 because of the spatial separation among fronds and can desiccate up to 81% at the end of the same low tide. Laboratory experiments showed that negative net photosynthetic rates occur between 40% and 80% desiccation at an irradiance of 515 μmol·m^?2·s^?1, and the literature suggests that strong bleaching could occur as a result. At 20 μmol·m^?2·s^?1 of irradiance and desiccation levels up to 40%, simulating understory conditions of natural stands, net photosynthetic rates are never negative. Experimental thinning of stands of M. cornucopiae done during spring effectively resulted in a stronger extent of frond bleaching compared with natural stands. Therefore, the cost of reduced net photosynthetic rates at high frond densities when fronds are fully hydrated is counterbalanced by the protective effects of frond crowding against extensive bleaching, essential for survival at the intertidal zone. Future research will have to demonstrate the possible relationship between the frequency and duration of negative net photosynthetic rates and the extent of frond bleaching.     

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.     

ONTOGENETIC CHANGES IN BUOYANCY,BREAKING STRENGTH,EXTENSIBILITY, AND REPRODUCTIVE INVESTMENT IN A DRIFTING MACROALGA TURBINARIA ORNATA (PHAEOPHYTA)1

Turbinaria ornata (Turner) J. Agardh is a tropical alga that disperses by detached, reproductively mature floating fronds. Material properties (breaking stress, breaking extension), buoyancy, and the proportion of reproductive tissue per frond were measured for juvenile, adult, and old fronds of T. ornata. Correlations between these factors indicate that as fronds age and become more reproductively mature, the tissue in their stipes (where they break) becomes weaker, more brittle, and the overall buoyancy of the frond increases. Measurement of drag force experienced by fronds from each ontogenetic stage allowed calculation of the environmental stress factor (ESF), which indicates the likelihood of detachment of a frond in the flow environment of its habitat. The ESF for fronds of each ontogenetic stage predicted that reproductively mature fronds (adult and old) break more readily than immature (juvenile) fronds. Increased proportions of reproductively mature fronds in floating rafts following storms compared with the proportion of mature fronds attached to the substratum support the ESF predictions. This combination of ontogenetic changes in material properties, buoyancy, and reproductive maturity in combination with the life history of T. ornata may contribute to the dispersal of this alga throughout French Polynesia.     

Reproduction ofChondrus crispus Stackhouse (Rhodophyta,Gigartinales) in sublittoral Prince Edward Island,Canada

The influence of size (as biomass), morphology and depth on reproductive patterns was examined in populations ofChondrus crispus in the sublittoral zone of Prince Edward Island, Canada. Fronds of ≥ 10 mg wet wt were considered, and a minimum biomass could not be related to reproduction. However, as frond biomass increased, the frequency of reproductive structures also increased. Reproductive maturity was related to the number of dichotomies. Fronds with less than two dichotomies were only occasionally reproductive, while those with two or more dichotomies could bear sori. Depth in the Prince Edward Island environment had no apparent influence on reproductive maturity, and there was no apparent relationship between depth and the vertical distribution of cystocarpic or tetrasporic fronds.     

QUANTIFYING GROWTH AND CALCIUM CARBONATE DEPOSITION OF CALLIARTHRON CHEILOSPORIOIDES (CORALLINALES,RHODOPHYTA) IN THE FIELD USING A PERSISTENT VITAL STAIN1

Growth and calcium carbonate deposition rates of the coralline alga Calliarthron cheilosporioides Manza were quantified by monitoring fronds in the intertidal zone that had been chemically labeled with the nontoxic fluorescent brightener Calcofluor white. This vital stain effectively labeled apical meristems of coralline thalli in the field: fronds exposed for only 5 min had detectable chemical marks at least 1.5 years later. By distinguishing portions of thalli that developed before and after exposure, this methodology permitted accurate measurement of growth and calcium carbonate deposition at each meristem. In Calliarthron, meristematic activity declined with increasing frond size. However, because growing fronds dichotomize, the total number of meristems and the deposition rate of new calcified tissue both increased with frond size. Growth rates reported here suggest that large fronds may not be as old as previously estimated. The Calcofluor white method may improve demographic studies of corallines by resolving growth and age of fronds in the field and may facilitate studies of climate change on calcium carbonate deposition in these ecologically important, calcifying algae.     

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.     

Influence of tissue source and growth rates on dry weight and carrageenan composition of Chondrus crispus (Gigartinales,Rhodophyta)

Carrageenan analyses were conducted on vegetative female clones of Chondrus crispus that were cultured to provide tissues with differing growth rates. Tissue dry weights increased from apex to base of fronds. Total carrageenan contents were lower in apical 1 to 2 cm segments than elsewhere in the frond, except when the alga was grown at high photon irradiances. Clone 373A contained more carrageenan than clone G8. The proportion of 0.3 M KCl-soluble polymers in the total native carrageenans varied from 44 to 92%, being highest in older tissues of fronds cultured at high photon irradiances. The apical 1 cm segments contained less KCl-soluble carrageenans than other tissues from the corresponding fronds. The KCl-soluble carrageenans, when alkali-modified and refractionated, afforded the expected kappa-iota carrageenan in > 79% yields. The remainder consisted of a polymer containing 23.1% SO₃Na and 8.4% 3,6-anhydrogalactose. Lambda carrageenan was not detected. Variations in carrageenan distribution between the apical region and other parts of the frond may reflect the increasing influence of medullary tissue developed as the immature cells differentiate.     

砷胁迫下不同砷富集能力植物内源生长素与抗氧化酶的关系

采用室内土培法,研究了砷胁迫(0,50,100和200 mg/kg)对凤尾蕨属中砷超富集植物大叶井口边草(Pteris cretica var.nervosa)和非砷超富集植物剑叶凤尾蕨(Pteris ensiformis)生物量、株高、叶片内源3-吲哚乙酸(IAA)含量、IAA氧化酶(IAAO)活性、砷含量、抗氧化酶(超氧化物歧化酶SOD、过氧化物酶POD、过氧化氢酶CAT)活性以及细胞膜脂过氧化产物丙二醛(MDA)含量的影响,并用逐步回归法分析了IAA含量与砷含量、抗氧化酶活性、MDA含量之间的关系。此外,研究了100 mg/kg砷处理下IAA、IAAO、3种抗氧化酶和MDA的时间动态。结果表明,与对照(不加砷)相比,加砷处理下大叶井口边草的株高和生物量未出现显著变化,但剑叶凤尾蕨在中、高浓度砷胁迫下则显著降低;中、高浓度砷胁迫均使2种植物叶片含砷量、IAA含量显著增加和IAAO活性显著下降,但这种改变在大叶井口边草中更为显著;加砷处理使大叶井口边草叶片3种抗氧化酶活性维持或增加,剑叶凤尾蕨中SOD和CAT活性虽能维持,但POD活性则显著下降,说明砷胁迫下大叶井口边草具有更强的抗氧化能力。逐步回归分析结果显示,2种植物叶片IAA含量均与砷含量成显著正相关,但在大叶井口边草中,IAA含量还与CAT活性成显著负相关。时间动态研究表明,第13天,大叶井口边草具有最高的IAA含量、最低的IAAO活性以及最低的CAT活性,而剑叶凤尾蕨中的变化规律则不明显。因此,叶片保持较高的IAA含量、较低的IAAO和CAT活性有助于大叶井口边草超量富集砷。     

Plant and frond dynamics of the giant kelp,Macrocystis pyrifera,forming a fringing zone in the Falkland Islands

Fluctuations in plant and frond characteristics are described for Macrocystis pyrifera (L.) C. Agardh (Laminariales, Phaeophyta) forming a fringing zone in the Falkland Islands. Giant kelp plants were sampled along a transect in the austral autumn (May 1986) and late spring (December 1986) which, according to previous frond weight analysis, were the times when extremes in population parameters were expected. Plant density and holdfast wet weights were similar for both seasons, but plants had more fronds and the fronds weighed more in spring than in autumn. Consequently, in autumn the frond biomass (1·1 wet kg m^?2) and productivity (34·1 wet g m^?2 d^?1) were lower than in spring, when a biomass of 5·0 wet kg m^?2 and a productivity of 72·4 wet g m^?2 d^?1 were recorded. Production of new fronds and loss of old fronds were determined at monthly intervals between April 1986 and March 1987. New frond production rates followed fluctuations in the quantity of light and varied between 0·08 and 0·48 fronds per plant per day. Frond loss rates did not show a seasonal pattern and fluctuated between 0·05 and 0·42 fronds per plant per day. It is suggested that the Falkland Islands Macrocystis population is more stable than most other giant kelp beds at high latitudes, because of the absence of winter storms.     

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.     

Comparative spatial distribution, size, biomass and growth rate of two varieties of bracken fern (Pteridium aquilinum L. Kuhn) in a neotropical montane habitat

A comparative field study of caudatum and arachnoideum, the two Pteridium aquilinum varieties of the caudatum subspecies known to grow in the neotropics, was performed in a montane savanna habitat of the Venezuelan Andes that was affected by wildfire. Frond size, ramet density and spatial distribution, blade and rhizome biomass, and frond elongation and expansion rates were measured in separate, isolated stands each containing only one bracken variety and covering approximately the same area (540 m²). In addition to clearly discernible morphological differences, caudatum and arachnoideum were found to possess distinct features: caudatum tends to develop open stands of relatively shorter blades of 76.6±0.89 cm (±SE) of rachis length and lower ramet density (1.6 fronds m^-2, max.=7 fronds m^-2) whereas arachnoideum grows into longer, more expanded fronds 124±1.6 cm tall and significantly higher ramet density (5.1 fronds/m², max.=14.6 fronds m^-2). The sum of aerial and underground biomass was found to be notably larger for arachnoideum (8522±614 Kg/ha) than for caudatum (1929±131 Kg ha^-1) in stands growing under the same habitat conditions. Therefore the spatial distribution of arachnoideum appeared considerably more compact than that of caudatum. Blade growth rates and development time were also very different. Newly emerged caudatum croziers developed into mature blades within 42 to 48 days following an inverse exponential curve whereas arachnoideum blades required 70 to 75 days to reach maturity following a linear development. All the above dissimilarities are interpreted as the hitherto unreported indication of diverging growth strategies of two cohabitant bracken varieties following fire.     

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.     

The effects of control on the biomass, carbohydrate content and bud reserves of bracken (Pteridium aquilinum (L.) Kuhn), and an evaluation of a bracken growth model

This paper examines the initial effects of bracken control on frond numbers and biomass, and the biomass, carbohydrate reserves and bud densities of bracken stands cut once per year, twice per year, subject to a single application of asulam or left untreated. The seasonal dynamics of these parameters are displayed; carbohydrate and biomass are both removed from the rhizome system to produce frond tissue, and are replenished at the end of the growing season. Asulam application reduced densities of both active and dormant buds, and both frond biomass and density. It did not significantly reduce rhizome biomass or carbohydrate reserves in the two years after treatment. Cutting, either once or twice per year reduced both rhizome biomass and rhizome carbohydrate reserves, as well as bud densities, though the latter were reduced in proportion to biomass. Cutting twice a year reduced the production of fronds, both in numbers and biomass. The collected data were used to evaluate a model of bracken growth, and subsequently to improve estimates of some of the model parameters. The model simulations of control treatments were compared to field data. The effects of cutting once per year and spraying with asulam were predicted accurately, but the bracken stand was more resilient to cutting twice per year than would be expected from model predictions. The combination of cutting and spraying is discussed as a potential tool in land management and the deficiencies of the model are discussed in relation to the need for future research into the biology of bracken.     

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.     

The fine-scale genetic structure of the two-spotted spider mite in a commercial greenhouse

The fine-scale genetic structure of Tetranychus urticae Koch was studied to estimate local gene flow within a rose tree habitat in a commercial greenhouse using seven microsatellite markers. Two beds of rose trees with different population densities were selected and 18 consecutive quadrats of 1.2 m length were sequentially established in each bed. Heterozygote deficiency was positive within quadrats, which was most likely a result of the Wahlund effect because the mites usually form small breeding colonies. Low population density and frequent inbreeding could also accelerate genetic differentiation among the breeding colonies. A short-range (2.4–3.6 m) positive autocorrelation and clear genetic cline among quadrat populations was detected within a bed. This suggests that gene flow was limited to a short range even if population density was substantially increased. Therefore, large-scale dispersal such as aerial dispersal contributed very little to gene flow in the greenhouse.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 1. The fronds

The encroachment of bracken (Pteridium aquilinum (L.) Kuhn) into areas previously dominated by heather represents a threat to the ecology, agricultural economy and landscape value of many UK upland areas, including the moorland of the North York Moors National Park. The morphology of bracken, within a mature stand and at several bracken-heather interfaces, has been studied at a number of sites within the National Park. Differences have been found in the frond growth of bracken in a mature stand, at stationary stand margins, and at advancing stand margins where bracken is encroaching into heather. Frequency of fronds present on bracken rhizome growing at a stationary stand margin close to the interface with heather (1–2 m behind the boundary) are approximately the same as those found within a mature stand. At advancing margins (again 1–2 m behind the boundary), maximum frond densities were often found to exceed those present in either a mature stand or at a stationary margin. Frond numbers decline rapidly at the stand margins as distance from the stand increases. This is especially true where the front is stationary and bracken is not encroaching into heather at a significant rate. Maximum frond heights in a mature stand consistently exceed those at stand margins (even 1–2 m into the stand) and are greater at stationary margins than at advancing margins. Outlying fronds at the edges of bracken stands are generally present in greater numbers, and further into the area dominated by heather, where the margin is advancing. Heights of outlying fronds fall as distance from the bracken stand increases, as does stipe length. Fronds at the edges of bracken stands emerge each spring before those further into the stand and are therefore particularly vulnerable to frost damage. Outlying fronds are not, however, the first to emerge. Early emerging fronds reach their maximum height and eventually become senescent before later emerging fronds. Whilst most fronds emerge before the end of June a few fronds continue to emerge throughout the summer. Frond densities close to the edges of bracken stands (1–2 m into the stand) are comparable to those in a mature stand. At advancing stand margins frond densities generally exceed those in a mature stand, suggesting that a large number of potential entry points for foliage-applied herbicides are available for bracken control at the stand margins. The ratio of potential uptake points to biomass of rhizome is also greatest at the edges of the stand, and the canopy 1–2 m into the stand is usually almost completely closed. It is possible therefore, the efficacy of herbicides could be improved by the use of small scale applications, using tractors or hand-held sprayers, close to the margins of bracken stands.     

Crowding in clonal seaweeds: Does self-thinning occur in Mastocarpus papillatus shortly before stand biomass peaks?

Fronds from crowded stands of clonal seaweeds, particularly those in which holdfasts are mostly perennial and are the major source of new fronds every year, are thought not to undergo self-thinning during the growth season, unlike those from crowded stands of unitary seaweeds. For clonal seaweeds, it is not known, however, what happens at the very end of the growth season, when crowding is highest for the year. By sampling twice more frequently than previously done for similar species, the possible occurrence of frond self-thinning was tested for Mastocarpus papillatus (Rhodophyta, Gigartinales, Petrocelidaceae) from western Canada during the growth season (spring) of 2003. Initially, stand biomass increased together with frond density, as found previously for similar clonal seaweeds. Shortly before stand biomass peaked for the year (June), frond density remained statistically unchanged. Thus, the increased sampling precision of this study confirms that fronds of these clonal seaweeds do not undergo self-thinning, not even shortly before crowding is highest. Frond size inequality for M. papillatus remained statistically similar during the growth season, which is also consistent with a model of no self-thinning. There are similarities in biomass–density dynamics and in size inequality dynamics between clonal seaweeds and clonal vascular plants.