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.     

Cell wall chemistry and tissue structure underlie shifts in material properties of a perennial kelp

Laminaria is an abundant kelp genus in temperate nearshore ecosystems that grows with a circannual ‘stop-start’ pattern. Species of Laminaria play important ecological roles in kelp forests worldwide and are harvested commercially as a source of food and valuable extracts. In order to evaluate seasonal differences in tissue properties and composition, we compared the material properties, histology and cell-wall composition of overwintering blades with newly synthesized, actively growing blades from Laminaria setchellii. We found that overwintering blades were fortified with a thicker cortex and increased cell wall investment, leading to increased material strength. Overwintering tissues were composed of higher proportions of cellulose and fucose-containing polysaccharides (i.e. FCSPs, fucoidans) than newly formed blades and were found to possess thicker cell walls, likely to withstand the waves of winter storms. Chemical cell wall profiling revealed that significant proportions of fucose were associated with cellulose, especially in overwintering tissues, confirming the association between cellulose and some fucose-containing polysaccharides. Changes in material properties during the resting phase may allow these kelps to retain their non-growing blades through several months of winter storms. The results of this study demonstrate how one species might regulate its material properties seasonally, and at the same time shed light on the mechanisms that might control the material properties of kelps in general.     

GIANT KELP (MACROCYSTIS PYRIFERA, PHAEOPHYCEAE) RECRUITMENT NEAR ITS SOUTHERN LIMIT IN BAJA CALIFORNIA AFTER MASS DISAPPEARANCE DURING ENSO 1997–1998

During the ENSO event of 1997–1998, density and population structure were evaluated in a Macrocystis pyrifera forest located in Bahía Tortugas, Baja California, Mexico, near the southern limit of the species' distribution in the Northern Hemisphere. Observations in Bahía Tortugas were made quarterly from January 1997 to September 1998 using SCUBA diving surveys. No macroscopic plants were found in the Bahía Tortugas area from October 1997 to April 1998, a local absence of at least 7 months. Aerial surveys further suggest regional disappearance along most of the Baja California coast during the event. Unexpectedly, plants were found in Bahía Tortugas again in July 1998, in spite of the widespread disappearance of the species less than a year earlier. Long-distance spore dispersal was an unlikely cause of the recruitment because: 1) the nearest spore source was more than 100 km away; 2) recruitment appeared to be simultaneous at many sites and occurred rapidly after the cessation of the ENSO event; and 3) the recruits occurred in the same areas as before disappearance. We suggest that a microscopic stage that was not visible during dive surveys survived the stressful conditions of ENSO and caused the recruitment event, supporting the hypothesis that a bank of microscopic forms can survive conditions stressful to macroscopic algae.     

CARBON ALLOCATION IN MACROCYSTIS PYRIFERA (PHAEOPHYTA): INTRINSIC VARIABILITY IN PHOTOSYNTHESIS AND RESPIRATION1

Measurements of net photosynthesis (PS, O₂ evolution), dark respiration (R, O₂ consumption), and light and dark carbon fixation (¹⁴C) were conducted on whole blades, isolated blade discs, sporophylls, apical scimitars and representative portions of stipe and holdfast of the giant kelp Macrocystis pyrifera L.C. Ag. On a dry weight basis, highest net PS rates were observed in apical scimitar segments and whole blades (3.81 and 3.07 mgC · g dry wt^?1· h^?1, respectively), followed by sporophylls (1.42 mgC·g dry wt^?1· h^?1) and stipe segments (0.15 mgC·g dry wt^?1· h^?1). No PS capacity was observed in holdfast material. Respiration rates showed similar ranking ranging from 1.22 mgC·g dry wt^?1·h^?1 for apical scimitar to 0.18–0.22 mgC·g dry wt^?1· h^?1 for holdfast material. Considerable within blade variability in both PS and R was also found. Steepest PS and R gradients on both an areal and weight basis were found within immature blades followed by senescent and mature blade material. Highest net PS rates were associated with the blade tips ranging from 3.08 (mature blades) to 10.3 mgC·dry wt^?1·h^?1 (immature blades). Highest rates of R generally occurred towards the basal portions of blades and ranged from 1.03–1.80 mgC·g dry wt^?1·h^?1 for immature blades. The variability within and between blades was high, with coefficients of variation approaching 50%. The observed patterns can be related to the decreasing proportionment of photosynthetic tissue and increasing proportionment of structural tissue as occurs from the blade tip to the blade base. Rates of light carbon fixation (LCF) revealed longitudinal profiles similar to oxygen measurements for the different blade types, with the absolute rates being slightly lower. Patterns of dark carbon fixation (DCF) were less easily interpreted. Highest rates of DCF (0.04–0.06 mgC·g dry wt^?1·h^?1) occurred at the basal portions of immature and senescent blades. Longitudinal profiles of total chlorophyll (a + c) on both an areal and weight basis were very similar to the profiles of PS. Normalized to chlorophyll a, PS displayed an unusual longitudinal profile in immature tissue; however, such profiles for mature and senescent tissues were similar to those for PS on an areal basis. It was demonstrated that it is difficult, if not impossible, to select single tissue discs that are representative of whole blades. The metabolic longitudinal profiles reveal a characteristic developmental pattern; the previous working definitions of immature, mature, and senescent blades, based on morphology and frond position thus have a physiological basis.     

Linking light attenuation and suspended sediment loading to benthic productivity within an Arctic kelp‐bed community1

Annual growth and productivity of kelp in the Stefansson Sound Boulder Patch, located along the Arctic coast of Alaska, is regulated almost entirely by PAR received during the summer open‐water period. Increased water turbidity during summer, often in response to storm activity, has been linked to low levels of ambient PAR and measurable decreases in kelp elongation. However, the relationship between PAR and water transparency has not been quantified, which compromises efforts to assess the effects of changing climate and weather conditions on kelp production. During the 2001–2002 summer periods, the inherent optical properties (IOPs) of Stefansson Sound waters were measured in conjunction with total suspended sediments (TSS) concentrations, which differed significantly between the 2 years, for input into a radiative transfer equation (RTE). In both years, the highest TSS levels (24.2 and 18.5 mg · L^?1 in 2001 and 2002, respectively) occurred in nearshore areas and were coincident with increased beam attenuations (13.8 and 8.3 m^?1). Lower TSS concentrations and attenuations were measured offshore. Data input to the RTE provided a TSS‐concentration‐specific attenuation coefficient that was used in a productivity model to estimate annual kelp productivities throughout the Boulder Patch based on modeled irradiance and averaged site‐specific TSS concentrations. Production estimates varied across the Boulder Patch but were lower in 2001 (0.12–0.34 g C · g dwt^?1 · year^?1, where dwt stands for dry weight) compared to 2002 (0.24–0.80 g C · g dwt^?1 · year^?1). Production in both years was greater in offshore locations with lower TSS loads. Results suggest that PAR availability during the summer is heavily influenced by TSS concentrations, and that changes in storm intensity and frequency, associated with current warming trends, may have significant effects on the primary production of these unique benthic algal communities.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.     

REPRODUCTIVE LONGEVITY OF DRIFTING KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE) IN MONTEREY BAY,USA1

Drifting Macrocystis pyrifera (L.) C. Agardh sporophytes have long been viewed as the primary long‐distance dispersal vector; yet, few data exist that support the ability of reproductive viable sporophytes to actually travel the presumed hundreds to thousands of kilometers. This study addressed the reproductive longevity of experimental and naturally occurring M. pyrifera drifters. Temporal variability in sporophyte size and reproduction was estimated for experimental drifting sporophytes that were tethered to surface buoys and compared with attached plants (controls). Reproductive viability was also studied for beach‐cast drifters (BCD), and naturally drifting sporophytes observed during field surveys in Monterey Bay. Detached drifting sporophytes were tracked with radio transmitters to follow drifter trajectories and to measure drifting speed. Experimental drifters (ED) experienced a 74% reduction in frond length after 35 days, a 76% reduction in average frond number after 70 days, and a reduction in average sorus area by 83% after 28 days. Although zoospore production was reduced following detachment, sporophytes remained fertile with high zoospore germination success as long as sori were present (125 days). Zoospore production and germination success for natural and BCD was similar to ED. The average displacement of radio‐tagged drifters was 7.12 km·day^?1, suggesting that a sporophyte adrift for 125 days disperses viable propagules (zoospores) over 890 km (±363). Dispersal of propagules is important for population restoration, distribution, and genetic diversity. Such dispersal distances are long enough to connect potentially all Northern Hemisphere Macrocystis populations across a generational timescale and may facilitate inter‐hemispheric gene flow.     

AN EVALUATION OF METHODS USED TO ASSESS INTERGENERIC HYBRIDIZATION IN KELP USING PACIFIC LAMINARIALES (PHAEOPHYCEAE)1

Kelp intergeneric laminarialean hybridizations and hybridization protocol were assessed using seven northeast Pacific kelp species: Alaria marginata Postels and Ruprecht, Costaria costata (C. A. Agardh) Saunders, Eisenia arborea Areschoug, Laminaria saccharina (L) Lamouroux, Lessoniopsis littoralis (Tilden) Reinke, Macrocystis integrifolia Bory, and Nereocystis leutkeana (Mertens) Postels and Ruprecht. Survival and development of sporophyte morphologies derived from selfings, separate males and females, and reciprocal crosses were evaluated over 30 weeks of cultivation. All cultures were initiated from cloned gametophytes. Two closely related species, Laminaria angustata Kjellman and L. japonica Areschoug, demonstrated the efficacy of long‐term (up to 30 years) cloned gametophytes in hybridization studies. Sporophyte morphologies appeared in 34%–69% of control and hybridization trials, and 6%–16% of all trials produced sporophytes in control and hybridization conditions that persisted through 30 weeks of cultivation. Sporophytes in control and hybridization conditions could appear normal or abnormal. Usually, the morphology of sporophytes in hybridizations and female controls resembled the female parent, whereas the sporophytes in male controls often had an abbreviated morphology, lacking definitive generic features. Species‐specific rDNA internal transcribed spacer molecular primers were used to determine the parentage of five putative hybrids. Only the L. japonica♀/L. angustata♂ hybrid bore both parental genomes. That negative controls could produce persistent and normal‐appearing sporophytes negates their value and emphasizes the importance of molecular confirmation in hybridization studies. These findings were applied to critique the only known wild intergeneric hybrid, Pelagophycus/Macrocystis.     

High acclimation potential in floating Macrocystis pyrifera to abiotic conditions even under grazing pressure – a field study

The persistence of floating seaweeds, which depends on abiotic conditions but also herbivory, had previously been mostly tested in outdoor mesocosm experiments. In order to investigate if the obtained mesocosm results of high seaweed persistence under natural environmental conditions and under grazing pressure can be extrapolated to field situations, we conducted in situ experiments. During two summers (2007 and 2008), Macrocystis pyrifera was tethered (for 14 d) to lines in the presence and absence of the amphipod Peramphithoe femorata at three sites (Iquique, Coquimbo, Calfuco). We hypothesized that grazing damage and seaweed persistence vary among sites due to different abiotic factors. By incubating the sporophytes in mesh bags, we were either able to isolate (grazing) or exclude (control) amphipods. To test for a mesh bag artifact, a set of sporophytes was incubated without mesh bags (natural). Mesh bags used to exclude herbivores influenced sporophyte growth and physiological performance. The chlorophyll a (Chl a) content depended largely on grazers and grazed sporophytes grew less than natural and control sporophytes within the two summers. A decrease in Chl a content was found for the sites with the highest prevailing irradiances and temperatures, suggesting an efficient acclimation to these sea surface conditions. Our field‐based results of sporophyte acclimation ability even under grazing pressure widely align with previous mesocosm results. We conclude that M. pyrifera and other temperate floating seaweeds can function as long‐distance dispersal vectors even with hitchhiking mesoherbivores.     

The importance of replicating genomic analyses to verify phylogenetic signal for recently evolved lineages

Genomewide SNP data generated by nontargeted methods such as RAD and GBS are increasingly being used in phylogenetic and phylogeographic analyses. When these methods are used in the absence of a reference genome, however, little is known about the locations and evolution of the SNPs. In using such data to address phylogenetic questions, researchers risk drawing false conclusions, particularly if a representative number of SNPs is not obtained. Here, we empirically test the robustness of phylogenetic inference based on SNP data for closely related lineages. We conducted a genomewide analysis of 75 712 SNPs, generated via GBS, of southern bull‐kelp (Durvillaea). Durvillaea chathamensis co‐occurs with D. antarctica on Chatham Island, but the two species have previously been found to be so genetically similar that the status of the former has been questioned. Our results show that D. chathamensis, which differs from D. antarctica ecologically as well as morphologically, is indeed a reproductively isolated species. Furthermore, our replicated analyses show that D. chathamensis cannot be reliably distinguished phylogenetically from closely related D. antarctica using subsets (ranging in size from 400 to 10 000 sites) of the 40 912 parsimony‐informative SNPs in our data set and that bootstrap values alone can give misleading impressions of the strength of phylogenetic inferences. These results highlight the importance of independently replicating SNP analyses to verify that phylogenetic inferences based on nontargeted SNP data are robust. Our study also demonstrates that modern genomic approaches can be used to identify cases of recent or incipient speciation that traditional approaches (e.g. Sanger sequencing of a few loci) may be unable to detect or resolve.