Southern California Kelp Forests Research Articles

Incidental take of Giant Sea Bass in the gill net fishery

The Giant Sea Bass is an apex predator of southern California kelp forests that historically supported large commercial and recreational fisheries but whose population was drastically reduced due to overfishing. Although regulations have been in place to protect Giant Sea Bass for over 30 years, there is concern for continued allowance of their incidental take in commercial gill net fisheries. Through landing records, logbooks, and federal observer data, we examined the commercial take of Giant Sea Bass to determine the: magnitude, location, age, and disposition of catch; whether this take was incidental or targeted; and its impact on the current population. We found that Giant Sea Bass landings declined since gill netting was banned in state waters in 1994 and their overall landings decreased by 75% since becoming protected in 1981. Over time, a greater proportion of older fish was seen in the landings with an increase in mature individuals over 13 years of age since 2000. We found no evidence that commercial fishers are targeting Giant Sea Bass. On average, Giant Sea Bass were landed on 4.2% of gill net trips from 1994 to 2021. Additionally, over the past decade the majority of Giant Sea Bass landings came from only 15 permit holders, but this made up only ~2% of their total landings in pounds as these are the most active permittees, bringing in the highest landings of White Seabass or California Halibut. Assessing gill net observer data, we found the mortality of released Giant Sea Bass is high at 72%, although despite this level of mortality, multiple fishery independent sources indicate Giant Sea Bass populations are increasing, suggesting that current protection measures may be effective. Despite these findings, it is challenging to interpret the full impact of the commercial fishery on Giant Sea Bass as an accurate measure of total discards is unknown without consistent observer coverage.

Spatial ecology of the Giant Sea Bass, Stereolepis gigas, in a southern California kelp forest as determined by acoustic telemetry.

The fisheries history of the Giant Sea Bass, Stereolepis gigas (Telostei: Polyprionidae), is closely linked to its spatial ecology. Its overharvest is directly associated with formation of spatially distinct spawning aggregations during summer, while its subsequent recovery is hypothesized to be the result of spatially explicit gear restrictions. Understanding the spatial ecology of Giant Sea Bass is a key part of efforts to assess contemporary threats such as commercial harvest and incidental catch by recreational fisheries. In this study, we used acoustic telemetry to characterize Giant Sea Bass space use in the La Jolla kelp forest using an acoustic array that encompasses two marine protected areas (MPAs) and heavily trafficked recreational fishing grounds. Five of the seven fish we tagged remained in the La Jolla array for at least 6 months. Two fish were resident across multiple years, with one fish consistently detected for 4 years. Only one fish was detected in the broader network of regional acoustic receivers, moving north approximately 8 km to Del Mar. Most tagged fish had home ranges and core use areas indicating they spend considerable time outside MPAs, particularly in areas with high recreational fishing activity. During spawning season we detected fish less frequently in the La Jolla array and recorded higher movement rates. While the current MPA network in La Jolla by no means offers complete protection to this fish, it does appear to support long-term persistence of some individuals in a region of exceptionally high recreational fishing pressure.

The Blob marine heatwave transforms California kelp forest ecosystems

Ocean warming has both direct physiological and indirect ecological consequences for marine organisms. Sessile animals may be particularly vulnerable to anomalous warming given constraints in food acquisition and reproduction imposed by sessility. In temperate reef ecosystems, sessile suspension feeding invertebrates provide food for an array of mobile species and act as a critical trophic link between the plankton and the benthos. Using 14 years of seasonal benthic community data across five coastal reefs, we evaluated how communities of sessile invertebrates in southern California kelp forests responded to the “Blob”, a period of anomalously high temperatures and low phytoplankton production. We show that this event had prolonged consequences for kelp forest ecosystems. Changes to community structure, including species invasions, have persisted six years post-Blob, suggesting that a climate-driven shift in California kelp forests is underway.

Environmental context dependency in species interactions

Ecological interactions are not uniform across time and can vary with environmental conditions. Yet, interactions among species are often measured with short-term controlled experiments whose outcomes can depend greatly on the particular environmental conditions under which they are performed. As an alternative, we use empirical dynamic modeling to estimate species interactions across a wide range of environmental conditions directly from existing long-term monitoring data. In our case study from a southern California kelp forest, we test whether interactions between multiple kelp and sea urchin species can be reliably reconstructed from time-series data and whether those interactions vary predictably in strength and direction across observed fluctuations in temperature, disturbance, and low-frequency oceanographic regimes. We show that environmental context greatly alters the strength and direction of species interactions. In particular, the state of the North Pacific Gyre Oscillation seems to drive the competitive balance between kelp species, asserting bottom-up control on kelp ecosystem dynamics. We show the importance of specifically studying variation in interaction strength, rather than mean interaction outcomes, when trying to understand the dynamics of complex ecosystems. The significant context dependency in species interactions found in this study argues for a greater utilization of long-term data and empirical dynamic modeling in studies of the dynamics of other ecosystems.

Detrital supply suppresses deforestation to maintain healthy kelp forest ecosystems.

Herbivores can reach extraordinary abundances in many ecosystems. When herbivore abundance is high, heavy grazing can severely defoliate primary producers and, in some cases, even drive ecosystem to undergo regime shifts from a high productivity state to a denuded, low productivity state. While the phenomenon of herbivore-driven regime shifts is well documented, we only partially understand the mechanisms underlying these events. Here, we combine herbivory experiments with 21 years of long-term monitoring data of kelp forest ecosystems to test the hypothesis that herbivores drive regime shifts when herbivory exceeds primary production. To test this hypothesis, we quantified how the foraging habits of an important group of marine herbivores-sea urchins-change with increases in sea urchin biomass and trigger regime shifts to a foundation species, giant kelp (Macrocystis pyrifera). Using experiments, we quantified how the grazing capacity of urchins increases as urchin biomass increases, then we combined these estimates of urchin grazing capacity with estimates of kelp production to predict when and where urchin grazing capacity exceeded kelp production. When grazing capacity exceeded kelp production, sea urchins caused a 50-fold reduction in giant kelp biomass. Our findings support the hypothesis that the balance between herbivory and production underlies herbivore-driven regime shifts in Southern California kelp forests and provides insight into when and where urchins are likely to force regime shifts in kelp forest ecosystems.

Experimental assessment of the impacts of ocean acidification and urchin grazing on benthic kelp forest assemblages

Ocean acidification (OA) is likely to differentially affect the biology and physiology of calcifying and non-calcifying taxa, thereby potentially altering key ecological interactions (e.g., facilitation, competition, predation) in ways that are difficult to predict from single-species experiments. We used a two-factor experimental design to investigate how multispecies benthic assemblages in southern California kelp forests respond to OA and grazing by the purple sea urchin, Strongylocentrotus purpuratus. Settlement tiles accrued natural mixed assemblages of algae and invertebrates in a kelp forest off San Diego, CA for one year before being exposed to OA and grazing in a laboratory experiment for two months. Space occupying organisms were identified and pooled into six functional groups: calcified invertebrates, non-calcified invertebrates, calcified algae, fleshy algae, sediment, and bare space for subsequent analyses of community structure. Interestingly, communities that developed on separate tile racks were unique, despite being deployed close in space, and further changes in community structure in response to OA and grazing depended on this initial community state. On Rack 1, we found significant effects of both pCO2 and grazing with elevated pCO2 increasing cover of fleshy algae, but sea urchin grazers decreasing cover of fleshy algae. On Rack 2, we found a ~ 35% higher percent cover of sediment on tiles reared in ambient pCO2 but observed ~27% higher cover of bare space in the high pCO2 conditions. On Rack 3, we found an average of 45% lower percent cover of calcified sessile invertebrates at ambient pCO2 than in high pCO2 treatments on Rack 3. Net community calcification was 137% lower in elevated pCO2 treatments. Kelp sporophyte densities on tiles without urchins were 74% higher than on tiles with urchins and kelp densities were highest in the elevated pCO2 treatment. Urchin growth and grazing rates were 49% and 126% higher under ambient than high pCO2 conditions. This study highlights consistent negative impacts of OA on community processes such as calcification and grazing rates, even though impacts on community structure were highly context-dependent.

Marine Protection Induces Morphological Variation in the California Moray, Gymnothorax mordax.

The effectiveness of marine protected areas (MPAs) on the general health and conservation of species, habitats, and community interactions is of great interest to researchers, managers, and recreationalists. However, the ecological and behavioral diversity of vertebrate predators of southern California kelp forests limits our ability to make general conclusions about MPA effectiveness across a variety of species. Identifying and studying species with extreme feeding habits or prey-capture strategies may offer greater insight into predator-prey relationships and reveal the trophic importance of an animal in the larger community. Moray eels (family Muraenidae) have been shown to have morphological and behavioral adaptations that allow them to consume large prey whole, identifying them as important predators. From 2015 to 2018, we studied the health and feeding behavior of a long-lived, elusive, and benthic kelp forest predator, the California moray eel (Gymnothorax mordax). We trapped eels inside and outside of Blue Cavern Onshore State Marine Conservation Area, an MPA on the northwest side of Santa Catalina Island, CA which prohibits the take of any species. Over 4 years, we captured 1736 eels. Overall, we found that morays were longer, older, heavier, had higher body condition, and were found in greater abundance within the MPA. Although fish comprised the majority of their summer diet, morays outside of the MPA were consuming a more diverse set of fish, while kelp bass comprised more than half of the diet for morays inhabiting the MPA. Additionally, we found that morays within the MPA had larger relative vertical gape distances (VGDs) and narrower heads. Our recapture data support the high site fidelity of morays, indicating that their diet and morphology are influenced by their local community. While the majority of morays are thriving in the MPA, as suggested by their robust sizes and longevity, high abundance appears to result in higher frequencies of cannibalism, the presence of an undescribed disease, and lower growth rates. Our results suggest that the MPA affects the life history of morays and may select for an alternative feeding strategy in which eels develop larger VGDs, smaller adductor muscles, and a specialized diet which is presumably influenced by the local environment. In addition, observations of cannibalistic behavior and species-specific disease provide us with important insight into natural factors that may still regulate populations removed from anthropogenic disturbances such as fishing.

Climate and fishing drive regime shifts in consumer-mediated nutrient cycling in kelp forests.

Globally, anthropogenic pressures are reducing the abundances of marine species and altering ecosystems through modification of trophic interactions. Yet, consumer declines also disrupt important bottom-up processes, like nutrient recycling, which are critical for ecosystem functioning. Consumer-mediated nutrient dynamics (CND) is now considered a major biogeochemical component of most ecosystems, but lacking long-term studies, it is difficult to predict how CND will respond to accelerating disturbances in the wake of global change. To aid such predictions, we coupled empirical ammonium excretion rates with an 18-year time series of the standing biomass of common benthic macroinvertebrates in southern California kelp forests. This time series of excretion rates encompassed an extended period of extreme ocean warming, disease outbreaks, and the abolishment of fishing at two of our study sites, allowing us to assess kelp forest CND across a wide range of environmental conditions. At their peak, reef invertebrates supplied an average of 18.3±3.0µmolNH4 + m-2 hr-1 to kelp forests when sea stars were regionally abundant, but dropped to 3.5±1.0µmolNH4 + m-2 hr-1 following their mass mortality due to disease during a prolonged period of extreme warming. However, a coincident increase in the abundance of the California spiny lobster, Palinurus interupptus (Randall, 1840), likely in response to both reduced fishing and a warmer ocean, compensated for much of the recycled ammonium lost to sea star mortality. Both lobsters and sea stars are widely recognized as key predators that can profoundly influence community structure in benthic marine systems. Our study is the first to demonstrate their importance in nutrient cycling, thus expanding their roles in the ecosystem. Climate change is increasing the frequency and severity of warming events, and rising human populations are intensifying fishing pressure in coastal ecosystems worldwide. Our study documents how these projected global changes can drive regime shifts in CND and fundamentally alter a critical ecosystem function.

Species insurance trumps spatial insurance in stabilizing biomass of a marine macroalgal metacommunity.

Because natural ecosystems are complex, it is difficult to predict how their variability scales across space and levels of organization. The species-insurance hypothesis predicts that asynchronous dynamics among species should reduce variability when biomass is aggregated either from local species populations to local multispecies communities, or from metapopulations to metacommunities. Similarly, the spatial-insurance hypothesis predicts that asynchronous spatial dynamics among either local populations or local communities should stabilize metapopulation biomass and metacommunity biomass, respectively. In combination, both species and spatial insurance reduce variation in metacommunity biomass over time, yet these insurances are rarely considered together in natural systems. We partitioned the extent that species insurance and spatial insurance reduced the annual variation in macroalgal biomass in a southern California kelp forest. We quantified variability and synchrony at two levels of organization (population and community) and two spatial scales (local plots and region) and quantified the strength of species and spatial insurance by comparing observed variability and synchrony in aggregate biomass to null models of independent species or spatial dynamics based on cyclic-shift permutation. Spatial insurance was weak, presumably because large-scale oceanographic processes in the study region led to high spatial synchrony at both population- and community-level biomass. Species insurance was stronger due to asynchronous dynamics among the metapopulations of a few common species. In particular, a regional decline in the dominant understory kelp species Pterygophora californica was compensated for by the rise of three subdominant species. These compensatory dynamics were associated with positive values of the Pacific Decadal Oscillation, indicating that differential species tolerances to warmer temperature and nutrient-poor conditions may underlie species insurance in this system. Our results illustrate how species insurance can stabilize aggregate community properties in natural ecosystems where environmental conditions vary over broad spatial scales.

Experiments Reveal Limited Top‐down Control of Key Herbivores in Southern California Kelp Forests

Experiments Reveal Limited Top‐down Control of Key Herbivores in Southern California Kelp Forests

Experiments reveal limited top-down control of key herbivores in southern California kelp forests.

Predator responses to gradients in prey density have important implications for population regulation and are a potential structuring force for subtidal marine communities, particularly on rocky reefs where herbivorous sea urchins can drive community state shifts. On rocky reefs in southern California where predatory sea otters have been extirpated, top-down control of sea urchins by alternative predators has been hypothesized but rarely tested experimentally. In laboratory feeding assays, predatory spiny lobsters (Panulirus interruptus) demonstrated a saturating functional response to urchin prey, whereby urchin proportional mortality was inversely density-dependent. In field experiments on rocky reefs near San Diego, California, predators (primarily the labrid fish California sheephead, Semicossyphus pulcher) inflicted highly variable mortality on purple urchin (Strongylocentrotus purpuratus) prey across all density levels. However, at low to moderate densities commonly observed within kelp forests, purple urchin mortality increased to a peak at a density of ~11 urchins/m2 . Above that level, at densities typical of urchin barrens, purple urchin mortality was density-independent. When larger red urchins (Mesocentrotus franciscanus) were offered to predators simultaneously with purple urchins, mortality was density-independent. Underwater videography revealed a positive relationship between purple urchin density and both the number and richness of fish predators, but these correlations were not observed when red urchins were present. Our results demonstrate highly variable mortality rates across prey densities in this system and suggest that top-down control of urchins can occur only under limited circumstances. Our findings provide insight into the dynamics of alternate community states observed on rocky reefs.

Temporal and spatial patterns in behavioral responses of marine predators to a sudden influx of abalone prey (Haliotis rufescens)

Predator–prey interactions exist on a variety of spatial and temporal scales; one of the earliest measurable responses to changes in these interactions is behavior. We examined the behavior of southern California kelp forest predators in response to a concentrated increase in the abundance of abalone during a restoration stocking experiment. We tested three hypotheses: (1) kelp forest predator density will increase following abalone stocking, (2) variations in predator characteristics will create an unequal impact on abalone, and (3) predation intensity will be greatest early in the experiment. Octopus discovered and exploited the influx of prey within the first week following stocking; their densities surged and then returned to pre-stocking levels after 2 months. This was not observed with any other predator. Damage from crustacean, fish, and octopus predation was observed on the recovered abalone shells, but were not correlated with predator densities. A larger percentage of recovered small shells had evidence of crustacean and fish predation, indicating there may be size-specific impacts of predators on abalone. Our results demonstrate that restoration stocking experiments can quantitatively test the predatory community’s response to reintroductions, as well as predation risk of newly stocked prey species when exposed to a diverse suite of predators.

Soundscape fishing: Spatial variability in a low-frequency fish chorus in the southern California kelp forest

The kelp forests off the coast of southern California support a diverse assemblage of fishes, many of which are known to produce sound. Here, the spatial variability of a low-frequency (325–545 Hz) fish chorus recorded at three sites near the kelp forests off La Jolla, California, is described. This chorus dominated the dusk soundscape at all sites in May/June 2015, 2016, and 2017. During these times, spectral levels around 400 Hz increased by approximately 30 dB over a period of 3 h between 19:00 and 22:00 local time. The location of the fish chorus was estimated during each year using beamforming and time difference of arrival (TDOA) techniques on signals recorded by either a two-element 30-m aperture linear seafloor array or an array with four-elements, 20-m aperture in a tetrahedral-shaped configuration. This location was relatively constant during the chorusing each night. Environmental factors such as temperature, macroalgae assemblage and bottom cover, and geological features were investigated as possible drivers of the spatial distribution of the chorus. [Research supported by California Sea Grant (R/HCME-28) and a Natural Sciences and Engineering Research Council of Canada (NSERC) Postgraduate Scholarship-Doctoral (PGS D-3).]

Sea Urchin Behavior in a Southern California Kelp Forest: Food, Fear, Behavioral Niches, and Scaling Up Individual Behavior

ABSTRACT Red and purple sea urchins (Mesocentrotus francisanus and Strongylocentrotus purpuratus) cohabit the west coast of North America and exhibit behavioral switching between sheltering, when food is abundant, and emergence and overgrazing, when food is scarce. To better understand individual urchin foraging behavior, we conducted a series of time-lapse behavioral studies within and at the edge of a resilient sea urchin barren. Photographs were taken at 15-min intervals for weeklong periods to observe behavior (1) in different microtopographic settings, (2) in response to food additions, and (3) along a spatial gradient from the leading edge of a sea urchin grazing front to ∼100 m behind it. Movement was limited for both species when crowded or in complex microtopography. Consistent differences in sheltering behaviors and diel movement patterns were observed between species in the presence and absence of food indicating behavioral niche differentiation. Red sea urchins responded to food falls at dista...

Protection of large predators in a marine reserve alters size-dependent prey mortality.

Where predator-prey interactions are size-dependent, reductions in predator size owing to fishing has the potential to disrupt the ecological role of top predators in marine ecosystems. In southern California kelp forests, we investigated the size-dependence of the interaction between herbivorous sea urchins and one of their predators, California sheephead (Semicossyphus pulcher). Empirical tests examined how differences in predator size structure between reserve and fished areas affected size-specific urchin mortality. Sites inside marine reserves had greater sheephead size and biomass, while empirical feeding trials indicated that larger sheephead were required to successfully consume urchins of increasing test diameter. Evaluations of the selectivity of sheephead for two urchin species indicated that shorter-spined purple urchins were attacked more frequently and successfully than longer-spined red urchins of the same size class, particularly at the largest test diameters. As a result of these size-specific interactions and the higher biomass of large sheephead inside reserves, urchin mortality rates were three times higher inside the reserve for both species. In addition, urchin mortality rates decreased with urchin size, and very few large urchins were successfully consumed in fished areas. The truncation of sheephead size structure that commonly occurs owing to fishing will probably result in reductions in urchin mortality, which may reduce the resilience of kelp beds to urchin barren formation. By contrast, the recovery of predator size structure in marine reserves may restore this resilience, but may be delayed until fish grow to sizes capable of consuming larger urchins.

Effects of predators on sea urchin density and habitat use in a southern California kelp forest

Grazing sea urchins can reduce kelp abundance and therefore strongly affect kelp forest community structure. Despite the ecological importance of sea urchins, direct field studies on the role that urchin predators play in shaping urchin populations are rare for southern California. We conducted surveys and manipulative experiments within kelp forests near San Diego, CA, (32–51′28″N, 117–16′00″W) from 2006 to 2009 to determine whether predators such as sheephead (Semicossyphus pulcher) and spiny lobsters (Panulirus interruptus) may be linked to purple urchin (Strongylocentrotus purpuratus) and red urchin (Strongylocentrotus franciscanus) distribution and habitat use, as well as purple urchin density-dependent mortality. Purple urchins were less dense and more cryptic inside a local marine protected area (MPA) that contained high predator abundance than in nearby heavily fished areas, whereas red urchins rarely were found outside the MPA. Urchin proportional mortality was inversely density dependent during the day when sheephead were active, despite fish aggregations in plots of high urchin density, but was density independent during the night when lobsters were active. Urchin mortality was reduced under understory algal cover during the day, but not during the night. Examining whether urchin mortality from predation is density dependent and how habitat complexity influences this relationship is imperative because behavioral changes and increases in urchin populations can have vast ecological and economic consequences in kelp forest communities.

Exploitation and recovery of a sea urchin predator has implications for the resilience of southern California kelp forests.

Size-structured predator-prey interactions can be altered by the history of exploitation, if that exploitation is itself size-selective. For example, selective harvesting of larger sized predators can release prey populations in cases where only large individuals are capable of consuming a particular prey species. In this study, we examined how the history of exploitation and recovery (inside marine reserves and due to fisheries management) of California sheephead (Semicossyphus pulcher) has affected size-structured interactions with sea urchin prey in southern California. We show that fishing changes size structure by reducing sizes and alters life histories of sheephead, while management measures that lessen or remove fishing impacts (e.g. marine reserves, effort restrictions) reverse these effects and result in increases in density, size and biomass. We show that predation on sea urchins is size-dependent, such that the diet of larger sheephead is composed of more and larger sized urchins than the diet of smaller fish. These results have implications for kelp forest resilience, because urchins can overgraze kelp in the absence of top-down control. From surveys in a network of marine reserves, we report negative relationships between the abundance of sheephead and urchins and the abundance of urchins and fleshy macroalgae (including giant kelp), indicating the potential for cascading indirect positive effects of top predators on the abundance of primary producers. Management measures such as increased minimum size limits and marine reserves may serve to restore historical trophic roles of key predators and thereby enhance the resilience of marine ecosystems.

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.

Synchrony in dynamics of giant kelp forests is driven by both local recruitment and regional environmental controls.

Populations of many species display spatially synchronous fluctuations in abundance. Synchrony is most commonly attributed to three processes: factors that influence recruitment (e.g., dispersal, early survival), large-scale environmental variability, and spatially autocorrelated trophic interactions. However it is often difficult to link population synchrony to a specific dominant process, particularly when multiple synchronizing forces are operating. We utilized a new satellite-based data set of giant kelp (Macrocystis pyrifera) canopy biomass to examine population synchrony in southern California kelp forests on spatial scales ranging from 50 m to 300 km and temporal scales ranging from 1 to 11 years. We examined the relationship between synchrony and distance for adult kelp populations, kelp recruits, sea urchin abundance (a major grazer of kelp), and environmental variables known to influence kelp population dynamics. Population synchrony in giant kelp decreased with distance between populations: an initial rapid exponential decrease between 50 m and 1.3 km was followed by a second, large-scale decrease between distances of 1.3 km and 172 km. The 50-m to 1.3-km spatial scale corresponded to the scales of synchrony in the abundance of sea urchins and young kelp recruits, suggesting that local drivers of predation and recruitment influence small-scale synchrony in kelp populations. The spatial correlation patterns of environmental variables, particularly wave height, were similar to the synchrony-distance relationship of kelp populations from 1.3 km to 172 km, suggesting that regional environmental variability, i.e., the Moran effect, was the dominant process affecting synchrony at larger spatial scales. This two-step pattern in the relationship between kelp biomass synchrony and distance was apparent in each of the 11 years of our study. Our results highlight the potential for synthesizing approaches from both landscape and population ecology in order to identify the multiple processes that generate synchrony in population dynamics.

Trophic cascades induced by lobster fishing are not ubiquitous in southern California kelp forests.

Fishing can trigger trophic cascades that alter community structure and dynamics and thus modify ecosystem attributes. We combined ecological data of sea urchin and macroalgal abundance with fishery data of spiny lobster (Panulirus interruptus) landings to evaluate whether: (1) patterns in the abundance and biomass among lobster (predator), sea urchins (grazer), and macroalgae (primary producer) in giant kelp forest communities indicated the presence of top-down control on urchins and macroalgae, and (2) lobster fishing triggers a trophic cascade leading to increased sea urchin densities and decreased macroalgal biomass. Eight years of data from eight rocky subtidal reefs known to support giant kelp forests near Santa Barbara, CA, USA, were analyzed in three-tiered least-squares regression models to evaluate the relationships between: (1) lobster abundance and sea urchin density, and (2) sea urchin density and macroalgal biomass. The models included reef physical structure and water depth. Results revealed a trend towards decreasing urchin density with increasing lobster abundance but little evidence that urchins control the biomass of macroalgae. Urchin density was highly correlated with habitat structure, although not water depth. To evaluate whether fishing triggered a trophic cascade we pooled data across all treatments to examine the extent to which sea urchin density and macroalgal biomass were related to the intensity of lobster fishing (as indicated by the density of traps pulled). We found that, with one exception, sea urchins remained more abundant at heavily fished sites, supporting the idea that fishing for lobsters releases top-down control on urchin grazers. Macroalgal biomass, however, was positively correlated with lobster fishing intensity, which contradicts the trophic cascade model. Collectively, our results suggest that factors other than urchin grazing play a major role in controlling macroalgal biomass in southern California kelp forests, and that lobster fishing does not always catalyze a top-down trophic cascade.