Physiological ecology of photosynthesis in Prasiola stipitata (Trebouxiophyceae) from the Bay of Fundy,Canada

The physiological ecology of Prasiola stipitata was examined in situ from two supralittoral sites in the Bay of Fundy (Nova Scotian, Canada) during November 2011, when the population was undergoing major expansion. Photosynthetic parameters (effective quantum yield, Φ_PSII, maximum quantum yield, F_v/F_m, and relative electron transport rate, rETR) were evaluated using chlorophyll fluorescence of PSII. A largely shaded and continuously moist population showed no change in Φ_PSII from one hour after sunrise to sunset in which natural irradiance varied between 3 and 300 μmol photons m^?2 s^?1. High irradiance (up to 1800 μmol photons m^?2 s^?1) had no apparent negative impacts on either quantum yield or rETR, but high desiccation in the field reduced quantum yield to almost zero. When thalli were brought into the laboratory, no change in F_v/F_m was observed up to 60% dehydration; however, there was a steep decline in F_v/F_m between 60% and 85% dehydration. Thalli showed complete recovery of F_v/F_m within one hour of reimmersion in seawater after 2 days of desiccation. After 15 days of desiccation full recovery required 24 h and after 30 days of desiccation thalli showed only partial recovery. These observations confirm the adaptation to photosynthesis in high irradiances and the rapid recovery following extreme desiccation observed in other Prasiola species.     

Species richness and diversity in different functional groups across environmental stress gradients: a model for marine rocky shores

We present a model predicting how the species richness and diversity within benthic functional groups should vary across the full environmental stress gradient across which a regional biota from marine rocky shores can occur. Built upon previous models, our model makes predictions for sessile species (macroalgae and filter feeders), herbivores, and carnivores. We tested some of its predictions by surveying vertical (intertidal elevation) and horizontal (wave exposure and ice scour) stress gradients in northern Nova Scotia, Canada. Because of harsh winter conditions, these coasts only depict approximately intermediate‐to‐high yearly levels of stress that the cold‐temperate, rocky intertidal biota from the northwestern Atlantic can experience. The observed trends matched predictions for sessile species in 75% of the studied gradients, and showed a moderate agreement for herbivores and carnivores only when they were combined as mobile consumers. Agreement meant that both richness and diversity increased from the most stressful to the most benign habitats that can be found in northern Nova Scotia. Also as predicted, sessile species generally showed a faster rate of increase in richness than mobile consumers. Our model also predicted a higher overall richness for sessile species than for mobile consumers, which was true by a factor of 3. Therefore, our model may constitute a useful tool to understanding community composition as a function of abiotic stress, which may in turn facilitate studies on community functioning. Model predictions for lower stress ranges could be tested on more southern shores where the same regional biota occurs.     

Species richness and diversity across rocky intertidal elevation gradients in Helgoland: testing predictions from an environmental stress model

Environmental stress affects species richness and diversity in communities, but the precise form of the relationship is unclear. We tested an environmental stress model (ESM) that predicts a unimodal pattern for total richness and diversity in local communities across the full stress gradient where a regional biota can occur. In 2008, we measured richness and diversity (considering all macrobenthic species) across the entire intertidal range on three rocky shores on Helgoland Island, Germany. Intertidal elevation is known to be positively related to abiotic stress. Since Helgoland is between the northern and southern biogeographic boundaries for the cold-temperate NE Atlantic intertidal biota, it exhibits low stress levels for this biota at low elevations and high stress at high elevations because of long (>6 h) emersion times. Thus, we predicted a unimodal trend for richness and diversity across elevation. On all three shores, richness increased from high to middle elevations, but remained similar between middle and low elevations. Diversity followed the same trend on one shore and different trends (although also non-unimodal) on the other two. Evenness explained the trend differences between richness and diversity. Overall, our study yielded little support for the ESM. Reasons for richness and diversity not decreasing at low elevations may be related to influences of mostly subtidal species, Helgoland’s intertidal range, or sampling resolution. Our study also suggests that the ESM must be developed further to differentiate between richness and diversity. We offer recommendations to improve future ESM research using intertidal systems.     

A discrete-time logistic model of frond dynamics for Mazzaella parksii (Rhodophyta, Gigartinales)

Mathematical modelling is useful in population ecology and resource management. Logistic models have traditionally been applied to unitary organisms, but it is unclear whether they could be used at the frond (ramet) level for clonal seaweeds. This study shows that frond dynamics for the clonal seaweed Mazzaella parksii (=M. cornucopiae) can be described by a discrete-time logistic model. The model is realistic in that it includes density-dependence, which was previously demonstrated experimentally for this species, and only necessitates data on frond density measured at discrete time intervals. This may constitute a useful tool for the management of clonal seaweeds of economic importance that occur in dense stands. This revised version was published online in August 2006 with corrections to the Cover Date.     

RAMET DYNAMICS FOR THE CLONAL SEAWEED PTEROCLADIELLA CAPILLACEA (RHODOPHYTA): A COMPARISON WITH CHONDRUS CRISPUS AND WITH MAZZAELLA CORNUCOPIAE (GIGARTINALES)

Little is known about the dynamics and the ecological interactions among ramets (fronds) from populations of clonal red seaweeds. Small ramets are very difficult to tag, so their growth cannot be monitored directly. The temporal variation of the relationship between stand biomass and ramet density offers information on ramet performance. We calculated this relationship for an intertidal population of Pterocladiella capillacea (Gmelin) Santelices et Hommersand (Gelidiales) from Baja California, Mexico. Biomass and density were positively correlated on an annual basis, indicating that biomass accumulated without involving self-thinning among ramets. This contrasts with nonclonal seaweeds, for which self-thinning among individuals occurs during growth, but agrees with other clonal red seaweeds, such as Chondrus crispus Stackhouse and Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (both Gigartinales). The growth pattern for these members of the Gelidiales and of the Gigartinales holds despite differences in holdfast morphology and ramet branching degree and despite differences in the capacity of coalescence during early stages, known only for the Gigartinales. The positive slope for the dynamic biomass–density relationship, on a bilogarithmic scale, was statistically steeper for M. cornucopiae than for P. capillacea and for C. crispus. This suggests that the addition of new ramets during the growth season may be relatively more beneficial for biomass accumulation rates for M. cornucopiae. This would be expected for high-intertidal species subjected to strong abiotic stress, for which ramet crowding constitutes a key protection. Pterocladiella capillacea occurs at the mid-intertidal zone and C. crispus at the subtidal zone, so ramets would be relatively less important in that respect.     

Review of studies on biomass-density relationships (including self-thinning lines) in seaweeds: Main contributions and persisting misconceptions

Ecological models relating biomass and density are relatively simple to calculate and offer information on, for example, the interactions among organisms and size constraints. Biomass‐density relationships have mostly been studied for terrestrial plants, but recently they have also been increasingly investigated for seaweeds. Unfortunately, a number of misconceptions have limited the overall contribution of algal studies to biomass‐density theory in general. Aiming to improve this situation, the present paper first summarizes the current knowledge on biomass‐density theory, particularly focusing on the main concepts that, with varying degrees of validity, exist in the published literature: the self‐thinning rule (in its boundary and dynamic interpretations), the interspecific biomass‐density relationship, and the ultimate biomass‐density line. Afterwards, the present paper provides a critical review of past biomass‐density studies on seaweeds. The main contributions of studies on clonal and unitary species are discussed, while the misconceptions that persist to these days are identified in order to help future studies to be based on solid grounds.     

DISTRIBUTION OF ALGAL EPIPHYTES ACROSS ENVIRONMENTAL GRADIENTS AT DIFFERENT SCALES: INTERTIDAL ELEVATION,HOST CANOPIES,AND HOST FRONDS1

Understanding epiphyte distribution in coastal communities is important because these organisms affect many others directly or indirectly. Yet, their distribution has been considerably less studied than that of their hosts and other primary‐space holders. Identifying major sources of variation in epiphyte abundance is thus still a need. Environmental gradients help predict species distribution and are pervasive on marine shores. In this study, we test the notion that environmental gradients across intertidal elevation, throughout host canopies, and along host fronds explain a large variation in the abundance of sympatric epiphytes. Our model system was the assemblage of Ascophyllum nodosum (L.) Le Jol. and its epiphytes Vertebrata lanosa (L.) T. A. Chr. [= Polysiphonia lanosa (L.) Tandy], Elachista fucicola (Velley) Aresch., and Pylaiella littoralis (L.) Kjellm. On the coast of Nova Scotia (Canada), we found evidence of a spatial segregation among these species at almost all scales. While the red epiphyte V. lanosa was more common at high‐ and midintertidal elevations (peaking at midelevations) and on middle segments of host fronds, the brown epiphytes E. fucicola and P. littoralis were more common at low elevations and restricted to distal segments of host fronds. Canopy habitat affected abundance only for V. lanosa, which was more common within the host canopy than on its periphery at midelevations. Since the studied gradients are related to predictable changes in abiotic factors, the identification of likely causes behind the observed patterns was facilitated. Our study ends by proposing abiotic and biotic factors that deserve priority in the experimental testing of the forces structuring this assemblage.     

ROLE OF SURFACE WOUNDS AND BROWN ALGAL EPIPHYTES IN THE COLONIZATION OF ASCOPHYLLUM NODOSUM (PHAEOPHYCEAE) FRONDS BY VERTEBRATA LANOSA (RHODOPHYTA)1

Ascophyllum nodosum (L.) Le Jol. forms extensive beds in wave‐sheltered, rocky intertidal habitats on the northwestern Atlantic coast. This fucoid seaweed is host to an obligate red algal epiphyte, Vertebrata lanosa (L.) T. A. Chr. [=Polysiphonia lanosa (L.) Tandy], and two facultative brown algal epiphytes, Elachista fucicola (Velley) Aresch. and Pylaiella littoralis (L.) Kjellm. Although V. lanosa can occur throughout most of the length of host fronds, it largely predominates in midfrond segments. The two brown algal epiphytes are restricted to distal segments. Through field experiments conducted in Nova Scotia, Canada, we tested the hypothesis that surface wounds are required for the colonization of distal segments of host fronds by V. lanosa. Distal tissues normally have a smooth surface because of their young age (A. nodosum fronds grow apically). By creating small wounds that mimicked grazing wounds distributed elsewhere on host fronds, we demonstrated that V. lanosa can colonize distal frond segments during the growth and reproductive season (summer and autumn). Approximately half of the artificial wounds were colonized by V. lanosa during this time. The experimental exclusion of both brown algal epiphytes from distal frond segments did not affect colonization by V. lanosa. Thus, we conclude that the absence of surface irregularities on distal segments of host fronds, specifically small wounds, is the main factor explaining the absence of V. lanosa there. We propose that further experimental work clarifying epiphyte distribution in host beds will enhance our ability to understand the functional role of epiphytes in intertidal ecosystems.     

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.     

The interspecific biomass–density relationship for terrestrial plants: where do clonal red seaweeds stand and why?

For crowded stands of terrestrial plants, ranging from mosses to trees, plant (or ramet, for clonal plants) density is negatively related to stand biomass. Stand biomass and ramet density were determined for Mazzaella cornucopiae and for Pterocladiella capillacea , two morphologically distinct intertidal clonal red seaweeds, to compare them with terrestrial plants. For these seaweeds, ramet densities were similar to the highest values reported for terrestrial plants (mosses, specifically). Stand biomass was higher than average values expected from the terrestrial interspecific biomass–density relationship, but lower than the limits expected from the terrestrial ultimate biomass–density line. These seaweeds show unexpectedly low ramet slenderness and high biomass packing per unit of volume, compared with the trend observed for terrestrial plants. Possible explanations for these differences are related to the particular physiology and habitat of intertidal clonal seaweeds.