Giant kelp vegetative propagation: Adventitious holdfast elements rejuvenate senescent individuals of the Macrocystis pyrifera “integrifolia” ecomorph

Recent findings on holdfast development in the giant kelp highlighted its key importance for Macrocystis vegetative propagation. We report here for the first time the development of adventitious holdfasts from Macrocystis stipes. Swellings emerge spontaneously from different areas of the stipes, especially in senescent or creeping individuals. After being manually fastened to solid substrata, these swellings elongated into haptera, which became strongly attached after 1 month. Within 4 months, new thalli increased in size and vitality, and developed reproductive fronds. Our results suggest the usage of these structures for auxiliary attachment techniques. These could act as a backup, when primary holdfasts are weak, and thus improve the survival rate of the giant kelp in natural beds.     

Ecological effects of harvesting Lessonia (Laminariales,Phaeophyta) in central Chile

Lessonia nigrescens and L. trabeculata are economically important canopy-forming kelps in Chile. Experimental harvesting of stipes above the first dichotomy reduces stipe movement and inter-stipe friction, allowing the development of a heavy epiphytic load and increased grazing. Complete stipe removal leads to holdfast death as neither species is able to simultaneously regenerate all stipes. The invertebrate fauna inside the holdfast does not respond to upper canopy changes, but mortality does occur in partial or complete plant removals. Kelp removal also affects inter-plant distances, results in increased access of grazers to the outside and inside of kelp holdfasts, reduces recruitment of other algal species, and modifies the morphology of L. trabeculata such that the plants become more susceptible to removal by water movement.     

Climate‐driven substitution of habitat‐forming species leads to reduced biodiversity within a temperate marine community

Aim

In marine ecosystems, habitat‐forming species (HFS) such as reef‐building corals and canopy‐forming macroalgae alter local environmental conditions and can promote biodiversity by providing biogenic living space for a vast array of associated organisms. We examined community‐level impacts of observed climate‐driven shifts in the relative abundances of two superficially similar HFS, the warm‐water kelp Laminaria ochroleuca and the cool‐water kelp Laminaria hyperborea.

Location

Western English Channel, north‐east Atlantic

Methods

We compared algal and invertebrate assemblages associated with kelp stipes and holdfasts, across multiple sites and sampling events. Significant differences were recorded in the structure of assemblages between the host kelp species at each site and event.

Results

Assemblages associated with stipes of the cool‐water HFS were, on average, >12 times more diverse and supported >3600 times more biomass compared with the warm‐water HFS. Holdfast assemblages also differed significantly between species, although to a lesser extent than those associated with stipes. Overall, assemblages associated with the warm‐water HFS were markedly impoverished and comprised far fewer rare or unique taxa.

Main conclusions

While previous research has shown how climate‐driven loss of HFS can cause biodiversity loss, our study demonstrates that climate‐driven substitutions of HFS can also lead to impoverished assemblages. The indirect effects of climate change remain poorly resolved, but shifts in the distributions and abundances of HFS may invoke widespread ecological change, especially in marine ecosystems where facilitative interactions are particularly strong.     

STUDIES ON THE GIANT KELP,MACROCYSTIS. II. REPRODUCTION

Neushul , M. (U. Washington, Seattle.) Studies on the giant kelp, Macrocystis. II. Reproduction. Amer. Jour. Bot. 50(4): 354–359. Illus. 1963.—The reproduction of Macrocystis pyrifera was studied in the sea and in the laboratory. The estimated minimum time needed for the completion of the sexual life history of Macrocystis pyrifera in La Jolla, California, is from 12 to 14 months. Young sporophytes begin fruiting when they have from 4 to 8 stipes and a somatic frond weight of from 8 to 10 kg. The behavior of abscised sporophylls in laboratory tanks, as well as their morphology and coloration, strongly suggests an interrelationship between the translocation of photosynthetic products and fructification.     

Disentangling the links between habitat complexity and biodiversity in a kelp‐dominated subantarctic community

Habitat complexity is one of the most important factors modulating species diversity. This feature comprises several interrelated attributes, such as number, size, and spatial arrangement of complexity‐forming elements. However, the separate and joint effects of these attributes on diversity and community structure are still not well understood. Here, we assess the relationships between several structural‐complexity attributes of the subantarctic kelp Lessonia flavicans and species richness, total abundance, and structure of kelp‐associated macrobenthic communities. We predicted that longer thalli and larger holdfasts favor greater species richness and total abundance of invertebrate organisms. To test the prediction, an observational sampling program was established in two sites of the Strait of Magellan. Uni‐ and multivariate analyses revealed both positive and negative effects of kelp structural‐complexity attributes on diversity. Holdfast diameter and maximum frond length, followed by thallus wet weight, had the strongest positive fits to species richness and total abundance; the number of stipes, on the other hand, was negatively associated with both response variables. Longer fronds were associated with greater abundances of spirorbid polychaetes. Larger holdfasts supported larger abundances of Nereididae and Terebelidae polychaetes and the limpet Nacella mytilina. Contrarily, kelps with longer fronds and more stipes supported fewer amphipods. In this way, we demonstrate that different dimensions of habitat complexity can have contrasting effects on diversity and community structure, highlighting the fundamental role of multiple dimensions of kelp habitat complexity for local biodiversity.     

Temporal variation and feeding ecology of juvenile Sebastes in rocky intertidal tidepools of northern California, with emphasis on Sebastes melanops Girard

This study examined the seasonal use of rocky intertidal areas by young-of-the-year (YOY) Sebastes spp. at three locations in northern California, 2003 to 2005. Six species of Sebastes were noted during the study with black rockfish Sebastes melanops comprising >99% of the total number sampled. Sebastes melanops were noted in rocky intertidal areas from May to August with peak abundances occurring in May or June. Variation in recruitment and feeding was evident among years. Diet analysis, using the index of relative importance (% I _RI) and the modified index (% I _M), showed that YOY S. melanops and Sebastes mystinus consumed a variety of prey items with harpacticoid copepods and gammaridean amphipods being the dominant prey categories. The rocky intertidal appears to serve as an important habitat for YOY S. melanops in northern California.     

Evaluation of remote sensing techniques for monitoring giant kelp populations

Photographs and maps of the floating canopy of the giant kelp, Macrocystis pyrifera, provide an important data source to monitor nearshore water quality in southern California. Declines in water quality related to turbidity from coastal development, ocean discharges, and non-point source runoff have caused reductions in the areal extent of these kelp beds. Historically the kelp beds have been monitored by a variety of methods including small format infrared and color photography. New digital remote sensing instruments combined with geographical information system (GIS) databases offer an efficient method for collecting and analyzing data on changes in kelp bed size and location. SPOT satellite imagery has been found to provide adequate resolution for mapping the larger beds of giant kelp along the California coast. Beds smaller than 10 ha are not resolved well with SPOT imagery and need to be mapped with a resolution greater than the 20 m pixel size provided by the SPOT multispectral imagery. Imagery from a prototype of the Positive Systems ADAR system, an airplane mounted multispectral video sensor, provided a spatial resolution of 2.3 m in 4 spectral bands. ADAR imagery taken on 2 October 1991 of the San Onofre Kelp Bed in northern San Diego County showed 39% more kelp than small format color infrared photography made during the same time period.     

EFFECTS OF SHADING BY ADULTS ON THE GROWTH OF BLADE-STAGE MACROCYSTIS PYRZFERA (PHAEOPHYTA) DURING AND AFTER THE 1982–1984 EL NIÑO1

The effect of shading by an adult canopy on blade-stage Macrocystis pyrifera (L.) C. A. Agardh was estimated by comparing the average growth rate of individuals under a canopy to that of individuals in a canopy gap. This comparison was made in 1983 during a strong El Niño and again in 1986 after the El Niño. Estimated nutrient concentrations in 1983 were two orders of magnitude below those in 1986, whereas ambient light levels were over 3 times higher. The kelp canopy caused similar proportional light reductions (20–30%) during both years. Blades grew 18% slower under the canopy than in the clearing in 1983 and about 77% slower under the canopy in 1986. Blade-stage individuals grew at the same rates in clearings in 1983 and 1986. Regardless of shading, the average growth rate of blade-stage kelp under the ambient, low-nutrient conditions of 1983 was higher than that later observed for multifronded juveniles during the same El Niño. The growth of blade-stage kelp was more like that of larger juveniles growing under high-nutrient conditions. The difference may be due to greater concentrations of nutrients very near the sea floor where single blades are growing compared to concentrations higher in the water column where larger kelp have most of their tissues.     

Diurnal, horizontal and vertical dispersal of kelp-associated fauna

The kelp Laminaria hyperborea is host for a rich fauna of mobile invertebrates. Dispersal patterns of these animals may be crucial for their availability to visual predators like fish, which are known to search for food in these productive habitats. Diurnal, horizontal and vertical dispersal within and out of the kelp forest were studied by analysing colonization of artificial substrata placed among kelps. The species composition of the fauna was different on three parts of the kelp: lamina, stipes (with epiphytes) and holdfast. The artificial substrata were colonized mainly by species associated with kelp. More species and individuals colonized the artificial substrata at night than during the day. The most abundant faunal groups on the kelps and the artificial substrata were amphipods and gastropods. Both groups dispersed at a significantly higher rate at night than during the day. Rapid horizontal dispersal out of the kelp forest was found. The artificial substrata were also colonized quickly by kelp fauna at all vertical levels inside and above the kelp forest. However, species associated with the kelp holdfast tended to disperse close to the bottom, while stipe-associated fauna moved through all parts of the kelp forest and even above the canopy layer. A high dispersal rate appears common amongst the mobile species living on kelp and seems to be advantageous, despite the risks involved in emigrating from the habitat and being exposed to predators. Higher activity at night may reduce availability to predators.     

Patterns of abundance and assemblage structure of epifauna inhabiting two morphologically different kelp holdfasts

The mobile fauna associated with two sympatric kelp species with different holdfast morphology (Saccorhiza polyschides and Laminaria hyperborea) was compared to test for differences in the assemblage structure of holdfast-associated mobile epifauna. A total of 24,140 epifaunal individuals were counted from 30 holdfasts of each kelp species. Overall epifaunal abundances exceeded faunal abundances previously reported from holdfasts of other kelps. Three taxonomic groups, Amphipoda, Mollusca, and Polychaeta, accounted for ca. 85% of all individuals. Total abundances increased with the amount of habitat available, quantified either as the volume or the area provided by the holdfasts. The multivariate structure of the epifaunal assemblage did not differ between holdfasts of the two kelp species. However, epifaunal assemblages responded differentially to the habitat attributes provided by each type of kelp holdfast: multivariate variation in the assemblage structure of epifauna was mostly explained by holdfast area and volume for L. hyperborea, and by the surface-to-volume ratio for S. polyschides holdfasts. Therefore, the physical attributes of biogenic habitats, here kelp holdfasts that better predict patterns in the assemblage structure of associated fauna can differ according to their different physical morphology, even though the overall assemblage structure of associated fauna was similar.     

Seasonal variation in epifaunal communities associated with giant kelp (Macrocystis pyrifera) at an upwelling‐dominated site

Kelp forests are highly productive and species‐rich benthic ecosystems in temperate regions that provide biogenic habitat for numerous associated species. Diverse epifaunal communities inhabit kelp sporophytes and are subject to variations in the physical environment and to changes experienced by the kelp habitat itself. We assessed seasonal variations in epifaunal invertebrate communities inhabiting giant kelps, Macrocystis pyrifera, and their effects on this seaweed. Six seasonal samplings were conducted over a year at an upwelling‐dominated site in northern‐central Chile where physical conditions are known to fluctuate temporally. More than 30 taxa were identified, among which peracarid crustaceans stood out in both diversity and abundance. Species richness and abundance differed among sporophyte sections (holdfast and fronds) and throughout the year. The frond community was dominated by two grazers (the amphipod Peramphithoe femorata and the isopod Amphoroidea typa), while suspension feeders, grazers, and omnivores (the amphipod Aora typica, the isopod Limnoria quadripunctata, and polychaetes) dominated the holdfasts. Abundances of the dominant species fluctuated throughout the year but patterns of variation differed among species. The most abundant grazer (P. femorata) had highest densities in summer, while the less abundant grazer (A. typa) reached its peak densities in winter. Interestingly, the area of kelp damaged by grazers was highest in autumn and early winter, suggesting that grazing impacts accumulate during periods of low kelp growth, which can thus be considered as ‘vestiges of herbivory past.’ Among the factors determining the observed seasonal patterns, strong variability of environmental conditions, reproductive cycles of associated fauna, and predation by fishes vary in importance. Our results suggest that during spring and early summer, bottom‐up processes shape the community structure of organisms inhabiting large perennial seaweeds, whereas during late summer and autumn, top‐down processes are more important.     

Inhibition of kelp recruitment by turfing algae and consequences for an Australian kelp community

In the sublittoral kelp forests of New South Wales, dictyotalean turfing algae dominate the substratum in areas where kelp canopies have been removed by storms. The effects of the presence of these algae on the recruitment of kelp plants and the subsequent structure of the benthic community were determined. Areas of turfing algae in clearings were experimentally cleared of turf at a time of dense kelp settlement, and areas of the kelp canopy were also removed. These treatments were compared with unmanipulated areas of turf in clearings and areas under the kelp canopy. The number of kelp recruits in each replicate plot was recorded throughout time as were the abundances of most of the macroscopic and microscopic species living on the substratum. Kelp was found to recruit very quickly to areas of substratum which had the overlying kelp canopy removed, but few recruits appeared under the natural canopy, or in areas with turfing algae, or in clearings where turfing algae had been removed. The results showed that whilst the presence of a natural kelp canopy and/or turfing algae inhibited the recruitment of kelps, removal of turfing species from clearings did not facilitate kelp recruitment. This indicated that the presence of turf did not solely inhibit kelp recruitment, but some lingering influence of turf seems to affect the substratum in clearings such that kelp cannot recruit to such areas. Turfing species returned and dominated the substratum in clearings where turf had been removed. After an initial decline, encrusting species increased in abundances in clearings where kelp recruited as these recruits developed a canopy. Turf and encrusting species remained great in cover in the respective control treatments. The results are discussed in terms of alternate stable states within this community. Such concepts are concluded to be dependent upon the spatial and temporal scales of one's investigation.     

Changes in the fish fauna associated with a sub-Antarctic <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis> kelp forest in response to canopy removal

The fish fauna associated with a Macrocystis pyrifera forest and the effects of the canopy removal on this fauna were studied in the Beagle Channel, Tierra del Fuego. Seasonal changes and differences in the fish communities that inhabit the canopy and the holdfast were also studied. Two patches of kelp forest separated by 200 m were selected. In one, the canopy was removed periodically while the other was used as a control. Samples were collected seasonally, from autumn 1999 to 2001. Fish fauna in the water column was sampled using trammel nets and holdfast fish fauna was sampled by removing the complete holdfast. Different assemblages of fish species were captured in the water column (surface and bottom) and in the holdfast. In the former there were mainly pelagic and benthopelagic species and the latter predominantly demersal species. The principal effect of the canopy removal was a drop in the abundance (and total weight) of Paranotothenia magellanica (doradito), the species with the strongest relationship with the canopy, principally at the surface of the treated patch probably due to a reduction in the availability of refuges. A high degree of seasonality was observed for fish species’ total weight, abundance and diversity, with higher values in summer and autumn. Taking into account our data and available data on kelp growth in the same locality, we suggest of cutting the kelp forest (preferably those farther away from the coast) once a year in winter or early spring, in order to minimize impact on the fish community.     

Survival of juvenile Ecklonia radiata sporophytes after canopy loss

The understorey beneath a canopy of the kelp Ecklonia radiata often contains juvenile sporophytes of the same species. When canopy disturbance occurs, these juvenile sporophytes are exposed to new environmental conditions. If these juvenile sporophytes survive these new conditions, then they become a ready source of kelps to rapidly form a new canopy. This study investigated the potential of pre-existing juvenile sporophytes of E. radiata to survive post-disturbance conditions and contribute to the rapid formation of a new canopy. The potential of canopy recovery by recruitment of kelp from zoospores was also investigated. These processes were studied at different times during the summer and on reefs ranging in topographic complexity from simple, flat reefs to highly complex, rugose reefs. By tagging juvenile sporophytes after the adult kelp canopy was removed and monitoring them through time, it was demonstrated that most juveniles (> 50%) survived the change in conditions after canopy loss, with some juveniles going on to become members of a new canopy. Approximately 6-47% of tagged sporophytes died within 3-4 days after canopy removal possibly due to excessive photoinhibition and photostress as demonstrated by changes in photosynthetic performance (decreased alpha values) of juveniles. The potential contribution of juvenile sporophytes to the rapid formation of a new canopy appears to be dependent on the timing of canopy removal with late summer-autumn canopy loss favouring faster recovery. Topographically complex reefs had less short-term (7 days) survival of juvenile sporophytes than topographically simple reefs; however this difference was not carried through to the long-term (6 months) abundance of adult kelp in experimental clearings, which was greater on topographically complex reefs. Clearly, juvenile sporophytes in arrested development under existing canopies of the small kelp E. radiata are important for the rapid recovery of the kelp canopy once adults are lost through physical disturbance.     

Size-dependent resource limitation and foraging-predation risk trade-offs: growth and habitat use in young arctic char

Variation in growth and habitat use is closely connected to individual responses to habitat specific resource levels and predation risk. In three mountain lakes which differed in the density of young-of-the-year (YOY) arctic char ( Salvelinus alpinus ), we studied the growth, diets and habitat use of YOY char in relation to resource levels. With two model approaches, we further examined the extent to which YOY and 1-yr old char were resource limited and, if so, whether resource limitation was associated with habitat use and small char densities. Benthic prey dominated the diet of YOY char and YOY char habitat use was restricted to the near-shore habitat in all lakes. In all lakes were chironomid densities higher in the near-shore habitat than in the offshore benthic habitat whereas zooplankton densities were higher in the pelagic than in the near-shore habitat. Growth of YOY char in all lakes was close to predicted maximum growth despite large variation in YOY densities between lakes. Model results suggested that density dependent resource limitation in YOY char is unlikely to occur despite restricted near-shore habitat use. In contrast, strong density dependent resource limitation was predicted in 1-yr old char with a restricted habitat use to the near-shore habitat. Correspondingly, field data suggested that the habitat use of 1-yr old char was density dependent as the use of the offshore habitat increased earlier in time and to a larger extent at high densities. As small individuals are vulnerable to predation but constrained by their low food processing capacity relative to their encounter capacity, we suggest that resource limitation in small individuals should be less pronounced and habitat use should mainly depend on predation risk. A trade-off in habitat use between foraging gain and predation risk is therefore expected to be more likely for individuals, large enough to be resource limited but still small enough to be vulnerable to predation.     

Factors affecting the production of sporophylls in the giant kelp Macrocystis pyrifera (L.) C.Ag.

The effects of variation in vegetative biomass, stand density, and stand location on sporophyll production in the giant kelp Macrocystis pyrifera (L.) C.Ag. were examined in a multifactorial experiment off the coast of Santa Barbara, California. Results indicate that vegetative biomass greatly influences zoospore production in this kelp, since the removal of 75% of vegetative fronds led to a drastic decrease in sporophyll production. Sporophyll biomass was shown to be closely correlated to zoospore production (r = 0.995). Plants growing offshore produced significantly less sporophyll biomass per plant than those inshore. Site differences in sporophyll production may have been caused by differential shading resulting from the persistence of the surface canopy offshore and the senescence of it inshore. As a possible result, increased plant density significantly decreased sporophyll biomass offshore and had no effect inshore. The timing of events such as storms which remove vegetative biomass could affect the local population dynamics of this kelp by decreasing zoospore production.     

The influence of herbivores on primary producers can vary spatially and interact with disturbance

Identifying the major drivers of ecosystem change remains a central area of ecological research. Although top–down drivers of change have received particular focus, we still have little understanding of how consistently these factors control an ecosystem's shift in both directions, between different ecosystem states. Using a crossed experiment in a shallow embayment in southeastern Australia, we investigated the roles of disturbance (kelp removal) and sea urchin herbivory (via increased density) to determine their contributions to shifts away from a kelp‐dominated community. In a second experiment, done in urchin barren areas at two sites, we tested whether reductions in ambient sea urchin densities allowed an algal shift in the reverse direction. In both experiments, we observed that high densities of sea urchins could negatively influence kelp and macroalgal abundance. However, in the kelp bed, a moderate or severe disturbance resulted in a comparable algal response, irrespective of urchin density. The influence of sea urchins also varied dramatically between the two urchin barren sites. Here, reducing urchin densities resulted in algal recovery at one site, but at the other site, substantial colonisation of barren areas by canopy‐forming brown algae and Ulvales occurred across all (low, medium, and high) urchin density treatments. Our findings illustrate multiple pathways of urchin barren creation and algal recovery, and reveal that shifts both to and from an urchin barren state can occur irrespective of herbivore pressure. These alternate pathways can operate over short spatial distances or with different regimes of disturbance.     

Spatial and temporal variability of the benthic assemblages associated to the holdfasts of the kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis> in the Straits of Magellan,Chile

Macrocystis pyrifera (L.) C. Agardh is a characteristic macroalga in the Magellan region covering almost 30% of the shallow coastal waters. The focus of this study was to analyse the spatial and seasonal patterns in macrofauna communities associated to the holdfasts of Macrocystis pyrifera at two study sites in the Straits of Magellan, South Chile. In total, 114 species from 10 major taxa were isolated from the holdfasts. MDS clearly separated the holdfast fauna collected in different seasons, with autumn and winter collections being richer in terms of species richness and abundance as compared to the spring and summer situation. MDS also clearly separated the holdfast associated faunas of the two study sites, Bahía Laredo and Fuerte Bulnes. The community structure and species composition of the associated macro-invertebrates and vertebrates appeared rather heterogeneous, probably due to the extremely heterogeneous environmental conditions along the entire coastline of the Subantarctic Magellan region.     

Utilization of holdfast fragments for vegetative propagation of Macrocystis integrifolia in Atacama, Northern Chile

We attempted to propagate Macrocystis integrifolia (giant kelp) by fragmentation and regeneration of holdfasts, which consist of creeping stolons with lateral haptera. Stolons from a natural kelp population in Bahia Chasco (Atacama, North Chile) were cut into fragments, each containing one or more lateral stipe and frond initials. Fragments were attached to longlines with elastic bands. We used two additional types of inoculants as controls: (1) natural recruits from the local parent population and (2) laboratory-cultured young sporophytes. Length, reproductive phenology, and mortality were determined monthly. Our results confirmed the feasibility of M. integrifolia propagation by reattachment of stolon fragments, and we obtained up to eight new individuals from one parent holdfast. Individuals from holdfast regeneration formed sporangial sori 3 months earlier than control specimens from recruits and laboratory culture, while all three types gave similar values for mortality. Holdfast morphology of regenerates differed strongly depending on their origin: stolon fragments continued growth and developed new haptera except in the cut surfaces, natural recruits formed typical M. integrifolia holdfast morphs, and sporophytes originating from laboratory culture produced minor stolons with many haptera. Implications of our results on ecology, repopulation, and aquaculture are discussed.