Field Collection and Laboratory Maintenance of Canopy-Forming Giant Kelp to Facilitate Restoration.

Canopy-forming kelps are essential foundation species, supporting biodiversity and providing ecosystem services valued at more than USD$500 billion annually. The global decline of giant kelp forests due to climate-driven ecological stressors underscores the need for innovative restoration strategies. An emerging restoration technique known as 'green gravel' aims to seed young kelps over large areas without extensive underwater labor and represents a promising restoration tool due to cost-effectiveness and scalability. This video article illustrates a protocol and tools for culturing giant kelp, Macrocystis pyrifera. It also provides a resource for further studies to address the successes and limitations of this method in field settings. We outline field and laboratory-based methods for collecting reproductive tissue, sporulating, inoculating, rearing, maintaining, and monitoring substrates seeded with early life stages using the 'green gravel' technique. The protocol simplifies and centralizes current restoration practices in this field to support researchers, managers, and stakeholders in meeting kelp conservation objectives.

When species don't move together: Non-concurrent range shifts in Eastern Pacific kelp forest communities.

Species range shifts due to changing ocean conditions are occurring around the world. As species move, they build new interaction networks as they shift from or into new ecological communities. Typically, species ranges are modeled individually, but biotic interactions have been shown to be important to creating more realistic modeling outputs for species. To understand the importance of consumer interactions in Eastern Pacific kelp forest species distributions, we used a Maxent framework to model a key foundation species, giant kelp (Macrocystis pyrifera), and a dominant herbivore, purple sea urchins (Strongylocentrotus purpuratus). With neither species having previously been modeled in the Eastern Pacific, we found evidence for M. pyrifera expansion in the northern section of its range, with no projected contraction at the southern range edge. Despite its known co-occurrence with M. pyrifera, models of S. purpuratus showed a non-concurrent southern range contraction and a co-occurring northern range expansion. While the co-occurring shifts may lead to increased spatial competition for suitable substrate, this non-concurrent contraction could result in community wide impacts such as herbivore release, tropicalization, or ecosystem restructuring.

Shading by giant kelp canopy can restrict the invasiveness of Undaria pinnatifida (Laminariales, Phaeophyceae).

The spread of non-indigenous and invasive seaweeds has increased worldwide, and their potential effects on native seaweeds have raised concern. Undaria pinnatifida is considered among the most prolific non-indigenous species. This species has expanded rapidly in the Northeast Pacific, overlapping with native communities such as the iconic giant kelp forests (Macrocystis pyrifera). Canopy shading by giant kelp has been argued to be a limiting factor for the presence of U. pinnatifida in the understory, thus its invasiveness capacity. However, its physiological plasticity under light limitation remains unclear. In this work, we compared the physiology and growth of juvenile U. pinnatifida and M. pyrifera sporophytes transplanted to the understory of a giant kelp forest, to juveniles growing outside of the forest. Extreme low light availability compared to that outside (~0.2 and ~4.4 mol photon ⋅ m-2 ⋅ d-1 , respectively) likely caused a "metabolic energy crisis" in U. pinnatifida, thus restricting its photoacclimation plasticity and nitrogen acquisition, ultimately reducing its growth. Despite M. pyrifera juveniles showing photoacclimatory responses (e.g., increases in photosynthetic efficiency and lower compensation irradiance, Ec ), their physiological/vegetative status deteriorated similarly to U. pinnatifida, which explains the low recruitment inside the forest. Generally, our results revealed the ecophysiological basis behind the limited growth and survival of juvenile U. pinnatifida sporophytes in the understory.

Artificial intelligence convolutional neural networks map giant kelp forests from satellite imagery.

Climate change is producing shifts in the distribution and abundance of marine species. Such is the case of kelp forests, important marine ecosystem-structuring species whose distributional range limits have been shifting worldwide. Synthesizing long-term time series of kelp forest observations is therefore vital for understanding the drivers shaping ecosystem dynamics and for predicting responses to ongoing and future climate changes. Traditional methods of mapping kelp from satellite imagery are time-consuming and expensive, as they require high amount of human effort for image processing and algorithm optimization. Here we propose the use of mask region-based convolutional neural networks (Mask R-CNN) to automatically assimilate data from open-source satellite imagery (Landsat Thematic Mapper) and detect kelp forest canopy cover. The analyses focused on the giant kelp Macrocystis pyrifera along the shorelines of southern California and Baja California in the northeastern Pacific. Model hyper-parameterization was tuned through cross-validation procedures testing the effect of data augmentation, and different learning rates and anchor sizes. The optimal model detected kelp forests with high performance and low levels of overprediction (Jaccard's index: 0.87 ± 0.07; Dice index: 0.93 ± 0.04; over prediction: 0.06) and allowed reconstructing a time series of 32 years in Baja California (Mexico), a region known for its high variability in kelp owing to El Niño events. The proposed framework based on Mask R-CNN now joins the list of cost-efficient tools for long-term marine ecological monitoring, facilitating well-informed biodiversity conservation, management and decision making.

Morphological, genotypic and metabolomic signatures confirm interfamilial hybridization between the ubiquitous kelps Macrocystis (Arthrothamnaceae) and Lessonia (Lessoniaceae).

Macrocystis pyrifera and Lessonia spicata are economically and ecologically relevant brown seaweeds that recently have been classified as members of two separated families within Laminariales (kelps). Here we describe for the first time the Macrocystis pyrifera x Lessonia spicata hybridization in the wild (Chiloe Island, Southeastern Pacific), where populations of the two parents exist sympatrically. Externally, this hybrid exhibited typical features of its parents M. pyrifera (cylindrical and flexible distal stipes, serrate frond margins and presence of sporophylls) and L. spicata (rigid and flat main stipe and first bifurcation), as well as intermediate features between them (thick unfused haptera in the holdfast). Histological sections revealed the prevalence of mucilage ducts within stipes and fronds (absent in Lessonia) and fully developed unilocular sporangia in the sporophylls. Molecular analyses confirmed the presence of the two parental genotypes for ITS1 nrDNA and the M. pyrifera genotype for two predominantly maternally inherited cytoplasmic markers (COI and rbcLS spacer) in the tissue of the hybrid. A metabolome-wide approach revealed that this hybrid is more chemically reminiscent to M. pyrifera. Nevertheless, several hits were identified as Lessonia exclusive or more remarkably, not present in any of the parent. Meiospores developed into apparently fertile gametophytes, which gave rise to F1 sporophytes that reached several millimeters before suddenly dying. In-vitro reciprocal crossing of Mar Brava gametophytes from both species revealed that although it is rare, interfamilial hybridization between the two species is possible but mostly overcome by pseudogamy of female gametophytes.

Short-term effects of hypoxia are more important than effects of ocean acidification on grazing interactions with juvenile giant kelp (Macrocystis pyrifera).

Species interactions are crucial for the persistence of ecosystems. Within vegetated habitats, early life stages of plants and algae must survive factors such as grazing to recover from disturbances. However, grazing impacts on early stages, especially under the context of a rapidly changing climate, are largely unknown. Here we examine interaction strengths between juvenile giant kelp (Macrocystis pyrifera) and four common grazers under hypoxia and ocean acidification using short-term laboratory experiments and field data of grazer abundances to estimate population-level grazing impacts. We found that grazing is a significant source of mortality for juvenile kelp and, using field abundances, estimate grazers can remove on average 15.4% and a maximum of 73.9% of juveniles per m2 per day. Short-term exposure to low oxygen, not acidification, weakened interaction strengths across the four species and decreased estimated population-level impacts of grazing threefold, from 15.4% to 4.0% of juvenile kelp removed, on average, per m2 per day. This study highlights potentially high juvenile kelp mortality from grazing. We also show that the effects of hypoxia are stronger than the effects of acidification in weakening these grazing interactions over short timescales, with possible future consequences for the persistence of giant kelp and energy flow through these highly productive food webs.

Kelp forests at the end of the earth: 45 years later.

The kelp forests of southern South America are some of the least disturbed on the planet. The remoteness of this region has, until recently, spared it from many of the direct anthropogenic stressors that have negatively affected these ecosystems elsewhere. Re-surveys of 11 locations at the easternmost extent of Tierra del Fuego originally conducted in 1973 showed no significant differences in the densities of adult and juvenile Macrocystis pyrifera kelp or kelp holdfast diameter between the two survey periods. Additionally, sea urchin assemblage structure at the same sites were not significantly different between the two time periods, with the dominant species Loxechinus albus accounting for 66.3% of total sea urchin abundance in 2018 and 61.1% in 1973. Time series of Landsat imagery of the region from 1998 to 2018 showed no long-term trends in kelp canopy over the past 20 years. However, ~ 4-year oscillations in canopy fraction were observed and were strongly and negatively correlated with the NOAA Multivariate ENSO index and sea surface temperature. More extensive surveying in 2018 showed significant differences in benthic community structure between exposed and sheltered locations. Fish species endemic to the Magellanic Province accounted for 73% of all nearshore species observed and from 98-100% of the numerical abundance enumerated at sites. Fish assemblage structure varied significantly among locations and wave exposures. The recent creation of the Yaganes Marine Park is an important step in protecting this unique and biologically rich region; however, the nearshore waters of the region are currently not included in this protection. There is a general lack of information on changes in kelp forests over long time periods, making a global assessment difficult. A complete picture of how these ecosystems are responding to human pressures must also include remote locations and locations with little to no impact.

Copper pollution exacerbates the effects of ocean acidification and warming on kelp microscopic early life stages.

Ocean warming (OW), ocean acidification (OA) and their interaction with local drivers, e.g., copper pollution, may negatively affect macroalgae and their microscopic life stages. We evaluated meiospore development of the kelps Macrocystis pyrifera and Undaria pinnatifida exposed to a factorial combination of current and 2100-predicted temperature (12 and 16 °C, respectively), pH (8.16 and 7.65, respectively), and two copper levels (no-added-copper and species-specific germination Cu-EC50). Meiospore germination for both species declined by 5-18% under OA and ambient temperature/OA conditions, irrespective of copper exposure. Germling growth rate declined by >40%·day-1, and gametophyte development was inhibited under Cu-EC50 exposure, compared to the no-added-copper treatment, irrespective of pH and temperature. Following the removal of copper and 9-day recovery under respective pH and temperature treatments, germling growth rates increased by 8-18%·day-1. The exception was U. pinnatifida under OW/OA, where growth rate remained at 10%·day-1 before and after copper exposure. Copper-binding ligand concentrations were higher in copper-exposed cultures of both species, suggesting that ligands may act as a defence mechanism of kelp early life stages against copper toxicity. Our study demonstrated that copper pollution is more important than global climate drivers in controlling meiospore development in kelps as it disrupts the completion of their life cycle.

Giant kelp, Macrocystis pyrifera, increases faunal diversity through physical engineering.

Foundation species define the ecosystems they live in, but ecologists have often characterized dominant plants as foundational without supporting evidence. Giant kelp has long been considered a marine foundation species due to its complex structure and high productivity; however, there is little quantitative evidence to evaluate this. Here, we apply structural equation modelling to a 15-year time series of reef community data to evaluate how giant kelp affects the reef community. Although species richness was positively associated with giant kelp biomass, most direct paths did not involve giant kelp. Instead, the foundational qualities of giant kelp were driven mostly by indirect effects attributed to its dominant physical structure and associated engineering influence on the ecosystem, rather than by its use as food by invertebrates and fishes. Giant kelp structure has indirect effects because it shades out understorey algae that compete with sessile invertebrates. When released from competition, sessile species in turn increase the diversity of mobile predators. Sea urchin grazing effects could have been misinterpreted as kelp effects, because sea urchins can overgraze giant kelp, understorey algae and sessile invertebrates alike. Our results confirm the high diversity and biomass associated with kelp forests, but highlight how species interactions and habitat attributes can be misconstrued as direct consequences of a foundation species like giant kelp.

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.

Photosynthesis and nitrogen uptake of the giant kelp Macrocystis pyrifera (Ochrophyta) grown close to salmon farms.

Finfish aquaculture is an activity that has experienced an explosive global development, but presents several environmental risks, such as high nitrogen outputs with potential eutrophication consequences. Therefore, the integration of seaweed aquaculture with the aim of decreasing nitrogen emissions associated with intensive salmon farming has been proposed as a bioremediation solution. Ecophysiological knowledge about seaweeds cultured close to farming cages is, however, still rudimentary. We experimentally studied the growth and physiological responses of Macrocystis pyrifera (Linnaeus) C. Agardh in a suspended culture system near a commercial salmon farm at three culture depths in order to understand its productivity performance. The results showed maximum growth responses at intermediate depths (3 m) as opposed to near the surface (1 m) or at a deeper culture level (6 m). At 6 m depth, light limitations were detected, whereas the sporophytes growing at 1 m depth responded to high irradiances, especially in late spring and summer, where they were more intensely exposed to decay of photosynthesis than individuals from other depths. Accordingly, photosynthetic pigment concentrations (chlorophyll a and c, and fucoxonthin) were higher during low-light seasons (winter and early spring) but decreased during the summer. On the other hand, although both nitrogen uptake and Nitrate Reductase (NR) activity varied seasonally, increasing significantly in spring and summer, these variables were not affected by culture depth. Therefore, the optimal culture depth of M. pyrifera near salmon farms appears to be a physiological integration between nitrogen supply and demand, which is modulated by plant acclimation to the seasonal change in light and temperature. The results allow to discuss about the environmental constrains of M. pyrifera in an ecophysiological context to improve the understanding of its aquaculture, and to contribute relevant information on the use of this species in bioremediation.

Cooperation induces other cooperation: Fruiting bodies promote the evolution of macrocysts in Dictyostelium discoideum.

Biological studies of the evolution of cooperation are challenging because this process is vulnerable to cheating. Many mechanisms, including kin discrimination, spatial structure, or by-products of self-interested behaviors, can explain this evolution. Here we propose that the evolution of cooperation can be induced by other cooperation. To test this idea, we used a model organism Dictyostelium discoideum because it exhibits two cooperative dormant phases, the fruiting body and the macrocyst. In both phases, the same chemoattractant, cyclic AMP (cAMP), is used to collect cells. This common feature led us to hypothesize that the evolution of macrocyst formation would be induced by coexistence with fruiting bodies. Before forming a mathematical model, we confirmed that macrocysts coexisted with fruiting bodies, at least under laboratory conditions. Next, we analyzed our evolutionary game theory-based model to investigate whether coexistence with fruiting bodies would stabilize macrocyst formation. The model suggests that macrocyst formation represents an evolutionarily stable strategy and a global invader strategy under this coexistence, but is unstable if the model ignores the fruiting body formation. This result indicates that the evolution of macrocyst formation and maintenance is attributable to coexistence with fruiting bodies. Therefore, macrocyst evolution can be considered as an example of evolution of cooperation induced by other cooperation.

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.

Short- and long-term acclimation patterns of the giant kelp Macrocystis pyrifera (Laminariales, Phaeophyceae) along a depth gradient.

The giant kelp, Macrocystis pyrifera, is exposed to highly variable irradiance and temperature regimes across its geographic and vertical depth gradients. The objective of this study was to extend our understanding of algal acclimation strategies on different temporal scales to those varying abiotic conditions at various water depths. Different acclimation strategies to various water depths (0.2 and 4 m) between different sampling times (Jan/Feb and Aug/Sept 2012; long-term acclimation) and more rapid adjustments to different depths (0.2, 2 and 4 m; short-term acclimation) during 14 d of transplantation were found. Adjustments of variable Chl a fluorescence, pigment composition (Chl c, fucoxanthin), and the de-epoxidation state of the xanthophyll cycle pigments were responsible for the development of different physiological states with respect to various solar radiation and temperature climates. Interestingly, the results indicated that phlorotannins are important during long-term acclimation while antioxidants have a crucial role during short-term acclimation. Furthermore, the results suggested that modifications in total lipids and fatty acid compositions apparently also might play a role in depth acclimation. In Aug/Sept (austral winter), M. pyrifera responded to the transplantation from 4 m to 0.2 m depth with a rise in the degree of saturation and a switch from shorter- to longer-chain fatty acids. These changes seem to be essential for the readjustment of thylakoid membranes and might, thus, facilitate efficient photosynthesis under changing irradiances and temperatures. Further experiments are needed to disentangle the relative contribution of solar radiation, temperature and also other abiotic parameters in the observed physiological changes.

Convergent evolution of vascular optimization in kelp (Laminariales).

Terrestrial plants and mammals, although separated by a great evolutionary distance, have each arrived at a highly conserved body plan in which universal allometric scaling relationships govern the anatomy of vascular networks and key functional metabolic traits. The universality of allometric scaling suggests that these phyla have each evolved an 'optimal' transport strategy that has been overwhelmingly adopted by extant species. To truly evaluate the dominance and universality of vascular optimization, however, it is critical to examine other, lesser-known, vascularized phyla. The brown algae (Phaeophyceae) are one such group--as distantly related to plants as mammals, they have convergently evolved a plant-like body plan and a specialized phloem-like transport network. To evaluate possible scaling and optimization in the kelp vascular system, we developed a model of optimized transport anatomy and tested it with measurements of the giant kelp, Macrocystis pyrifera, which is among the largest and most successful of macroalgae. We also evaluated three classical allometric relationships pertaining to plant vascular tissues with a diverse sampling of kelp species. Macrocystis pyrifera displays strong scaling relationships between all tested vascular parameters and agrees with our model; other species within the Laminariales display weak or inconsistent vascular allometries. The lack of universal scaling in the kelps and the presence of optimized transport anatomy in M. pyrifera raises important questions about the evolution of optimization and the possible competitive advantage conferred by optimized vascular systems to multicellular phyla.

Connectivity structures local population dynamics: a long-term empirical test in a large metapopulation system.

Ecological theory predicts that demographic connectivity structures the dynamics of local populations within metapopulation systems, but empirical support has been constrained by major limitations in data and methodology. We tested this prediction for giant kelp Macrocystis pyrifera, a key habitat-forming species in temperate coastal ecosystems worldwide, in southern California, USA. We combined a long-term (22 years), large-scale (~500 km coastline), high-resolution census of abundance with novel patch delineation methods and an innovative connectivity measure incorporating oceanographic transport and source fecundity. Connectivity strongly predicted local dynamics (well-connected patches had lower probabilities of extinction and higher probabilities of colonization, leading to greater likelihoods of occupancy) but this relationship was mediated by patch size. Moreover, the relationship between connectivity and local population dynamics varied over time, possibly due to temporal variation in oceanographic transport processes. Surprisingly, connectivity had a smaller influence on colonization relative to extinction, possibly because local ecological factors differ greatly between extinct and extant patches. Our results provide the first comprehensive evidence that southern California giant kelp populations function as a metapopulation system, challenging the view that populations of this important foundation species are governed exclusively by self-replenishment.

Patch definition in metapopulation analysis: a graph theory approach to solve the mega-patch problem.

The manner in which patches are delineated in spatially realistic metapopulation models will influence the size, connectivity, and extinction and recolonization dynamics of those patches. Most commonly used patch-definition methods focus on identifying discrete, contiguous patches of habitat from a single temporal observation of species occurrence or from a model of habitat suitability. However, these approaches are not suitable for many metapopulation systems where entire patches may not be fully colonized at a given time. For these metapopulation systems, a single large patch of habitat may actually support multiple, interacting subpopulations. The interactions among these subpopulations will be ignored if the patch is treated as a single unit, a situation we term the "mega-patch problem." Mega-patches are characterized by variable intra-patch synchrony, artificially low inter-patch connectivity, and low extinction rates. One way to detect this problem is by using time series data to calculate demographic synchrony within mega-patches. We present a framework for identifying subpopulations in mega-patches using a combination of spatial autocorrelation and graph theory analyses. We apply our approach to southern California giant kelp (Macrocystis pyrifera) forests using a new, long-term (27 years), satellite-based data set of giant kelp canopy biomass. We define metapopulation patches using our method as well as several other commonly used patch delineation methodologies and examine the colonization and extinction dynamics of the metapopulation under each approach. We find that the relationships between patch characteristics such as area and connectivity and the demographic processes of colonizations and extinctions vary among the different patch-definition methods. Our spatial-analysis/graph-theoretic framework produces results that match theoretical expectations better than the other methods. This approach can be used to identify subpopulations in metapopulations where the distributions of organisms do not always reflect the distribution of suitable habitat.

Genetic and experimental evidence for a mixed-age, mixed-origin bank of kelp microscopic stages in southern California.

Laboratory studies have demonstrated that the microscopic stages of kelps can rapidly resume development from a delayed state. Like terrestrial seeds or aquatic resting eggs, banks of delayed kelp stages may supplement population recovery after periods of stress, playing an important role for kelp populations that experience adult sporophyte absences due to seasonal or interannual disturbances. We found that removing the microscopic stages from natural rock substratum could prevent the appearance of juvenile kelp sporophytes for three months and the establishment of a diverse kelp assemblage for over four months within a southern California kelp forest. Juveniles were observed within one month in plots where microscopic stages were left intact, which may confer an advantage for the resulting sporophytes as they attain larger sizes before later recruiting neighbors. Microsatellite diversity was high (expected heterozygosity HE approximately 0.9) for juveniles and adults within our sites. Using a microsatellite-based parentage analysis for the dominant kelp, Macrocystis pyrifera, we estimated that a portion of the new M. pyrifera sporophyte recruits had originated from their parents at least seven months after their parents had disappeared. Similar delay durations have been demonstrated in recent laboratory studies. Additionally, our results suggest that zoospore dispersal distances > 50 m may be supported by including additional microsatellite loci in the analysis. We propose a mixed-age and, potentially, a mixed-origin bank of M. pyrifera gametophytes promotes maximal genetic diversity in recovering populations and reduces population genetic subdivision and self-fertilization rates for intact populations by promoting the survival of zoospores dispersed > 10 m and during inhospitable environmental conditions.

The importance of progressive senescence in the biomass dynam of giant kelp (Macrocystis pyrifera).

Temporal variation in primary producer biomass has profound effects on the structure and function of the surrounding ecological community. The giant kelp (Macrocystis pyrifera) exhibits strong intra-annual variation in biomass density, which is better explained by the demographic rates of fronds than by those of whole plants. To better understand the processes controlling the dynamics of giant kelp fronds we collected monthly time-series data of frond initiation and survival. These data were used to determine how frond loss and frond initiation rates were predicted by factors thought to affect the growth and survival of Macrocystis, including external environmental factors (i.e., wave height, day length, temperature, nutrient concentration, and neighborhood density) and intrinsic biological characteristics (i.e., frond age, plant size, and nutritional status). Our results revealed that frond dynamics were better explained by intrinsic biological processes rather than external environmental factors. A metric of frond age structure that incorporated progressive senescence was the best predictor of frond loss rate, accounting for 58% of the explained variation in frond loss. A similar analysis revealed that frond age structure was also the single best predictor of frond initiation rate, accounting for 46% of the explained variation. To further examine the importance of senescence in biomass dynamics, we used frond age-dependent mortality and frond initiation rates to predict biomass in subsequent months and found that the model explained 73% of the observed variation in biomass at our sites. Vegetation dynamics of many species including giant kelp are often considered largely in the context of external controls on resource availability and physical disturbance. Our results indicate that investigations of the processes controlling vegetation dynamics may benefit greatly from the inclusion of intrinsic biological factors such as age-dependent mortality and growth, which can outweigh the effects of external forcing in accounting for fluctuations in vegetation biomass.