SCALE‐DEPENDENT PATTERNS OF DISTURBANCE AND RECOVERY IN GIANT KELP FORESTS

We studied spatial variability in giant kelp (Macrocystis pyrifera) forests at 84 sites along the west coast of North America in order to assess the impacts of the 1997–98 El Niño. Our sites spanned the geographic range of giant kelp in the Northern Hemisphere and were surveyed just before, immediately following, several months after, more than one year after, and nearly two years after the El Niño. Interspersion of sample units allowed us to compare the effects of this disturbance among spatial scales ranging from a few meters to more than a thousand kilometers. Variance components analyses revealed that El Niño shifted the relative importance of factors that regulate giant kelp communities from factors acting at the scale of a few meters (local control) to factors operating at hundreds of kilometers (regional control). Moreover, El Niño resulted in a near‐to‐complete loss of giant kelp populations throughout nearly two‐thirds of the species' range. Evaluation of these effects along with oceanographic data (at the “appropriate” spatial scales), along with closer examination of giant kelp populations in the most severely impacted region (Baja) suggested that the among‐region differences in giant kelp survival was due, at least in part, to El Niño‐induced differences in ocean climate. Giant kelp recovery following El Niño was also scale‐dependent, but driven by factors different from those of the disturbance. Here, we present results for several species of macroalgae in an attempt to relate the importance of El Niño to that of other processes in creating scale‐dependent patterns of variability.     

Estimating scale-dependency in disturbance impacts: El Niños and giant kelp forests in the northeast Pacific

Recent discussions on scaling issues in ecology have emphasized that processes acting at a wide range of spatial and temporal scales influence ecosystems and thus there is no appropriate single scale at which ecological processes should be studied. This may be particularly true for environmental disturbances (e.g. El Niño) that occur over large geographic areas and encompass a wide range of scales relevant to ecosystem function. However, it may be possible to identify the scale(s) at which ecosystems are most strongly impacted by disturbances, and thus provide a measure by which their impacts can be most clearly described, by assessing scale-dependent changes in the patterns of variability in species abundance and distribution. This, in turn, may yield significant insight into the relative importance of the various forcing factors responsible for generating these impacts. The 1997–98 El Niño was one of the strongest El Niños ever recorded. I examined how this event impacted giant kelp populations in the northeast Pacific Ocean at 90 sites ranging from central Baja California, Mexico to central California, USA. These sites spanned the geographic range of giant kelp in the Northeast Pacific and were surveyed just before, immediately following, several months after, more than 1 year after, and nearly 2 years after the El Niño. I used a hierarchical sample design to compare these impacts at five spatial scales spanning six orders of magnitude, from a few meters to more than 1,000 km. Variance Components Analyses revealed that the El Niño shifted control over giant kelp abundance from factors acting at the scale of a few meters (local control) to factors operating over hundreds to thousands of kilometers (regional control). Moreover, El Niño resulted in the near-complete loss of all giant kelp throughout one-half of the species range in the northeast Pacific Ocean. Giant kelp recovery following El Niño was far more complex and variable at multiple spatial scales, presumably driven by numerous factors acting at those scales. Recovery returned local control of giant kelp populations within 6 months in southern California, and within 2 years in Baja California.     

Little evidence for climate effects on local-scale structure and dynamics of California kelp forest communities

Climatic variables such as temperature and precipitation play an important role in controlling local and regional scale differences in population dynamics and species distributions, and large-scale climatic events such as El Niño southern oscillation (ENSO) have been shown to affect population dynamics of key species in many ecosystems, particularly in kelp forests. Few studies have been able to evaluate the consequences of climate variables on the structure and dynamics of biological communities, in large part because the lack of data at appropriate spatial and temporal scales has made it difficult to adequately address local-scale responses of species and communities to such events over relevant time scales. Here, we combined an unprecedented dataset of kelp forest species' abundances from the Channel Islands, California with data for several local, regional, and global scale climatic variables to evaluate the temporal and spatial scale at which one can detect community-wide effects of climate variables, in particular ENSO events. We found large and significant local-scale differences in community structure, but these differences were not related to differences in climatic variables. Moreover, giant kelp abundance, which has been shown to be highly sensitive to water temperature and storm disturbance, was a poor predictor of community differences, and all communities tended to decline in abundance over the 20-year sampling period, suggesting a press perturbation to the system such as PDO cycles or sustained fishing pressure. Although ENSO events can have dramatic impacts on the abundance and distribution of giant kelp itself across the range of the species, such events appear to have little effect on local-scale kelp forest community structure or dynamics.     

RECOVERY AND GENETIC DIVERSITY OF THE INTERTIDAL KELP LESSONIA NIGRESCENS (PHAEOPHYCEAE) 20 YEARS AFTER EL NIÑO 1982/831

Massive mortality in kelp beds of the Pacific coasts of North and South America was caused by the rise in surface seawater temperature during the El Niño Southern Oscillation (ENSO) event of 1982/83, the strongest in the four and half previous centuries. In northern Chile a stretch of 600 km of coastline showed massive mortality of the intertidal kelp species Lessonia nigrescens Bory, of which only a few individuals managed to survive. Kelps and their associated biodiversity recovered but kelp beds re-colonization in general was variable in time and space seemingly very slow along northern Chilean coasts. Here we show, effectively, that northward re-colonization advanced less than 60 km in 20 years. Conversely, kelp beds of the Northern Hemisphere recovered 300 km in only six months after the same ENSO event. Genetic diversity in the two most affected populations of L. nigrescens shows half of the heterozygosity and polymorphism with respect to that observed in six non affected populations. In addition, geographically separated populations seem highly isolated as evidenced by high and significant fixation indices (all F_ST values over 0.4).     

IMPROVED BODY CONDITION OF WEANED HAWAIIAN MONK SEAL PUPS ASSOCIATED WITH EL Niño EVENTS: POTENTIAL BENEFITS TO AN ENDANGERED SPECIES

Efforts to enhance recovery of endangered Hawaiian monk seals ( Monachus schauinslandi ) require an understanding of factors influencing population dynamics. This study examines relationships between body condition and survival of monk seal pups at French Frigate Shoals and Laysan Island and El Niño events. Girth measurements and mass estimates were used as indicators of pup body condition, and survival was evaluated from weaning to age 2. Linear models and logistic regression analysis were used to evaluate El Niño effects. Temporal trends in mean girth, mass, and survival were identified at both study sites. After accounting for temporal trends, girths were 3.7 cm and 2.7 cm greater during El Niño years at French Frigate Shoals and Laysan Island, respectively. Average mass estimates were significantly greater during El Niño years at French Frigate Shoals (2.6 kg increase), but were not significantly different at Laysan Island (1.8 kg increase). Weaned pups born at French Frigate Shoals during El Niño years survived significantly better, but this effect was not detected at Laysan Island. El Niño events probably affect pup condition and survival, but these parameters need to be monitored during future El Niño events to clarify these relationships.     

MONITORING THE PERSISTENCE OF GIANT KELP AROUND SANTA CATALINA ISLAND USING A GEOGRAPHIC INFORMATION SYSTEM

Geographic information systems (GIS) facilitate monitoring and analysis of population distributions at spatial and temporal scales differing from those employed in conventional field monitoring. This study utilizes a GIS-based gap analysis of a network of marine reserves around Santa Catalina Island relative to the regional ecology, disturbance regime, and persistence of giant kelp ( Macrocystis pyrifera ), a keystone species in the nearshore, marine environment. Catalina's orientation and greatly-dissected coastline create diverse microhabitats with respect to storm exposure, temperature, light regime and topographic factors. GIS overlay methods applied to multi-temporal kelp distribution maps generated a model representing the spatial "persistence" of kelp. Correlations between the kelp's geographic distribution and persistence, the disturbance regime and physical variables conferring resistance to or recovery from it were drawn. This analysis identified regions of persistent kelp under disturbance regimes markedly different from those in the existing reserves, suggesting the designation of additional reserves in unprotected areas is ecologically warranted.     

Reconciling the influence of global climate phenomena on macrofaunal temporal dynamics at a variety of spatial scales

Determining the relative importance of environmental forces on population dynamics is a fundamental question for ecologists. Growing concern over the ecological effects of climate change emphasizes the importance of defining whether broad-scale environmental forces uniformly act upon local populations (hierarchy theory) or cross-scale interactions influence local responses (multiscale theory). This study analyses 13 years of data on species abundances at six sites within a large harbour to determine the effect of the El Niño Southern Oscillation (ENSO). Environmental variables both directly and indirectly related to ENSO were observed to be important predictors of the temporal dynamics of abundance in many species, but the observed effects were not consistent across sites or species. While nearly all species were affected by large temporal and spatial scale variability, smaller temporal scale, location-specific environmental variables (such as wind-generated wave exposure and turbidity) were also generally important, increasing the variability explained by our models by up to 25%. As with many other broad-scale variables, generality of response to ENSO is affected by interactions across time and space with smaller scale heterogeneity. This study therefore suggests that the degree of interaction between broad-scale climatic factors, such as ENSO, with smaller scale variability, will determine the consistency of responses over large spatial scales, and control our ability to predict effects of climate change on coastal and estuarine communities.     

Dynamics of the shallow-water fish assemblage of the Patos Lagoon estuary (Brazil) during cold and warm ENSO episodes

In southern Brazil, cold ( La Niña ) and warm ( El Niño ) episodes of the El Niño Southern Oscillation (ENSO) phenomenon cause drought and high rainfall, respectively. The low precipitation and freshwater outflow associated with La Niña during 1995–1996 were associated with an increase in the abundance of marine species in the Patos Lagoon estuary. During the 1997–1998 El Niño , high precipitation and river discharge were associated with low abundance of marine species in the estuary. ANOVA results showed a higher abundance during La Niña than El Niño for estuarine resident (RES) and estuarine dependent (DEP) fishes. During La Niña catch per unit of effort (CPUE) of RES increased from the marine to estuarine area, but during El Niño CPUE increased at the marine area and diminished during summer and autumn in some estuarine sites. DEP fishes had an opposite abundance pattern. During La Niña , these fishes were abundant at the coastal marine area and along some estuarine sites, but during El Niño , CPUE remained almost the same at the marine area but dropped along some estuarine sites. These different abundance patterns for dominant fish groups yielded a positive interaction between stations and climatic events. With higher river discharge and the consequent decline of dominant euryhaline fishes, such as Mugil platanus and Atherinella brasiliensis , freshwater species increased in abundance and richness in the shallow waters of the stuary. The ENSO phenomenon influences precipitation and estuarine salinity in southern Brazil and thereby seems to have a strong influence on recruitment, immigration, and emigration dynamics of fish species living within and adjacent to estuarine habitats.     

Effects of large-scale climatic fluctuations on survival and production of young in a Neotropical migrant songbird, the yellow warbler Dendroica petechia

Migrant songbirds are vulnerable to changes in climatic conditions on both the breeding and wintering grounds. For North American Neotropical migrants, the El Niño/Southern Oscillation (ENSO), via its effects on global precipitation and temperature, modulates the productivity of their temperate and tropical terrestrial ecosystems used during the course of their annual cycle. We evaluated how a densely nesting population of yellow warblers Dendroica petechia in a riparian forest in southern Manitoba, Canada, responded to the El Niño/Southern Oscillation (ENSO) between 1992 and 2001. Standardized mist netting was used to estimate apparent annual survival of adults and production of young. Both adult survival and the production of young were positively correlated with the Southern Oscillation Index (SOI). More specifically, values of both these demographic parameters were lowest during El Niño years and highest during La Niña years. These findings demonstrate the influence of climate on populations of Neotropical migrants in North America. The more frequent El Niños predicted to result from future global climate change could negatively affect populations of yellow warblers and other Neotropical migrants breeding in this region.     

CHANGES IN THE RELATIVE ABUNDANCE AND DISTRIBUTION OF GRAY WHALES (ESCHRICHTIUS ROBUSTUS) IN MAGDALENA BAY, MEXICO DURING AN EL NIÑO EVENT

The oceanographic changes during the El Niño and La Niña of 1998–1999 have provided an opportunity to document the effects of these events on the migratory patterns of gray whales ( Eschrichtius robustus ). Magdalena Bay is located at the southernmost point of the gray whale's annual breeding migration along the Baja California Peninsula in Mexico. Standardized surveys were conducted for three consecutive seasons (1997–1999) in Magdalena Bay, enabling an assessment of the changes in the relative abundance of gray whales with altered temperature. The average temperature of the bay in 1998 was 4.2°C higher than in 1997 and 5.8°C higher than in 1999. The number of whale sightings during that time period was inversely related to temperature, with nearly 15 times greater relative abundance observed in 1999 than in 1998. It is hypochesized that the decreasing temperatures associated with the La Niña event of 1999 may have induced more whales to travel farther south along their migration route. Alternative hypotheses are also expiored. Comparisons of the three years are discussed with consideration of the importance of environmental conditions versus area fidelity as determining factors in the location of gray whale calving and breeding activities.     

Notes on the effects of E1 Niño on the seabirds of the Mollendo district, southwest Peru, in 1983

In the austral summer of 1982–83 the cool Peru Coastal Current Zone of the Pacific Coast of South America was affected by the strongest and most prolonged incursion of the warm El Niño Current in at least 58 years. An account is given of the visual effects of the disturbance on the seabirds of the Mollendo district on the coast of southwest Peru. With one exception, local numbers of the endemic cool water species were drastically reduced, with little evidence of recovery up to April 1984. During the period when El Niño was at its height, several warm water species of seabirds, previously either virtually or completely unknown at Mollendo, were recorded.     

THE SECRET LIFE OF KELPS: PLANKTONIC PROCESSES AND POPULATION DYNAMICS

Understanding spatio‐temporal variability in recruitment is vital to studies of kelp population dynamics. Research on settlement and post‐settlement processes has suggested that arrival of kelp zoospores to suitable substrate is important in limiting kelp recruitment, yet the role of planktonic processes in kelp population dynamics has not been studied due to difficulties in sampling and identifying zoospores. I developed a method to estimate kelp zoospore abundance from in situ plankton samples and used it to study various processes regulating the availability of giant kelp (Macrocystis pyrifera) zoospores for settlement. My studies focused on (1) identifying temporal scales over which zoospore abundance is most variable, (2) describing physical and biological processes that regulate this variability, and (3) determining the relationship between zoospore abundance and settlement. I found that short‐term variability in zoospore abundance (<24 hrs) was not due to changes in supply but rather dispersion, caused by oscillating hydrodynamic forces (e.g. waves). Long‐term variability in zoospore abundance, however, was best explained by the size and density of reproductive adult plants, with zoospore abundance being most variable at the scale of days to months. Changes in adult reproductive condition caused rapid changes in zoospore abundance suggesting that the supply of kelp zoospores is sensitive to environmental regulation of adult physiology. Thus, unlike with marine animals, these results indicate that variability in kelp propagule supply, over scales most likely to affect subsequent settlement and recruitment, is more tightly coupled to demographic and reproductive mechanisms than to physical transport processes.     

Spatial variation and effects of habitat on temperate reef fish assemblages in northeastern New Zealand

Reef-associated fishes can respond to changes in habitat structure and the nature of their response can change with different spatial scales of observation. A structured hierarchical mensurative sampling design was used to sample temperate reef fish assemblages in northeastern New Zealand at several spatial scales over 2 years. The three spatial scales examined were tens of meters (transects), hundreds to thousands of meters (sites) and hundreds of kilometers (locations). We tested the hypothesis that fish assemblages differed between kelp forest habitat (relatively dense stands of the kelp, Ecklonia radiata (C. Agardh) J. Agardh, median depth=13.5 m) and barrens habitat (rocky reef dominated by turfing and encrusting red algae and the grazing urchin, Evechinus chloroticus (Valenciennes), median depth=6.7 m). Recently developed multivariate techniques were used to test for and quantify multivariate variation at different spatial scales. There were significant effects of habitat on the spatial distribution of fish assemblages, characterised by greater abundances or frequencies of Parika scaber, Chromis dispilus, Trachurus novaezelandiae, Nemadactylus douglasii, Bodianus unimaculatus, Odax pullus and Pseudolabrus miles in kelp forest habitat, and greater abundances or frequencies of Notolabrus celidotus, Notolabrus fucicola, Girella tricuspidata, Coris sandageri, Chironemus marmoratus, Parma alboscapularis, Scorpis violaceus and Kyphosus sydneyanus in barrens habitat. Some of the more common species, including Upeneichthys lineatus, Scorpis lineolatus and Cheilodactylus spectabilis showed no strong consistent effects of these two differing habitats on their distributions. There was, however, a significant Habitat×Locations interaction: effects of habitat did not occur at all locations. Variability was highest at the scale of individual transects and variability from site to site and from location to location was comparable. Spatial variation was large compared to inter-annual variation, which was minimal, and spatial patterns were consistent in the 2 years examined. Further experiments, including manipulations, are required to understand what mechanisms and processes might be driving these patterns. This study, coupled with results from previous studies, suggests that there may be a dynamic inter-play between effects of habitat on fish and effects of fish on biogenic habitat, such as kelp forests.     

Consistency and variation in kelp holdfast assemblages: Spatial patterns of biodiversity for the major phyla at different taxonomic resolutions

The focus of this study was to measure natural spatial variability in the biodiversity of fauna inhabiting kelp holdfasts in northeastern New Zealand at several spatial scales: from meters up to hundreds of kilometers. We wished to test the hypothesis that multivariate variation and biodiversity would vary significantly at different spatial scales in different ways for the major phyla in the holdfast community (Arthropoda, Annelida, Mollusca and Bryozoa). Biodiversity was considered in terms of richness, total abundance, structural composition (as measured by the Bray-Curtis dissimilarity measure) and taxonomic breadth for each major phylum and for the assemblage as a whole. We also examined the effect of taxonomic resolution on multivariate patterns. Species richness and total abundance increased with increases in holdfast volume. Multivariate variation was greatest at the smallest spatial scale for all phyla, but different phyla showed different patterns of multivariate variation at different spatial scales. Variations among locations at the largest spatial scale were primarily due to differences in the composition and richness of bryozoans and molluscs. Location effects became less and less distinct with decreases in taxonomic resolution. There were very few significant differences in richness or abundance for holdfasts of a given volume, taxonomic breadth did not vary significantly across locations, nor did the proportional abundances of phyla. These consistencies across large spatial scales in the absence of environmental impacts and results from other studies suggest that holdfast communities in New Zealand systems would provide a useful model assemblage against which future impacts may be detected as changes in proportions of component phyla. In addition, high variability detected at small and large scales at the species level, especially for bryozoans and molluscs, suggest that these communities may also provide unique opportunities for studying and understanding sources and functions of marine biodiversity.     

Population dynamics of a polychaete during three El Niño events: disentangling biotic and abiotic factors

The macrobenthic community in shallow soft-bottom areas in the Bay of Ancón, Peru, is characterised by low biodiversity due to low oxygen concentrations. During El Niño events, higher temperature and higher concentrations of dissolved oxygen induce a temporary increase in biodiversity. However, the structure and dynamics of the emerging communities vary strongly among events. The reasons for this variation are poorly understood, in particular the relative influence of abiotic vs biotic factors. To disentangle the roles of abiotic and biotic factors, population models based on detailed monitoring data of three El Niño events were developed focused on the population dynamics of one species in the community, the polychaete Sigambra bassi , which showed different responses in different El Niño events. Calculated and observed population dynamics are compared using root mean square deviation (RMSD). The results show that S. bassi abundance can be determined by abiotic environmental conditions. Besides, three biotic factors improved model performance in different El Niño events: negative density dependence in larval settlement, lower carrying capacity in the presence of other species and positive density dependence in adult survival. The results demonstrate how population models can be used to decode information hidden in long-term monitoring data of abiotic and biotic community parameters about factors driving population dynamics.     

Distance decay of community dynamics in rocky intertidal sessile assemblages evaluated by transition matrix models

It is well known that the similarity in species composition between two communities decays with the geographic distance that separates them. It is thus likely that the similarity in the dynamics of two communities also decays with distance, because the distance–decay relationship is fundamental in nature. However, the distance–decay relationships of community dynamics have not yet been revealed. We used transition matrix models to evaluate distance–decay relationships of seasonal community dynamics (from spring to summer) in rocky intertidal sessile assemblages along the Pacific coast of Japan between 31°N and 43°N. We evaluated the distance–decay relationships of whole-community dynamics and of three dynamics-related components—recruitment, disturbance, and species interaction (competition and facilitation)—for communities separated by distances ranging from several meters to thousands of kilometers. The similarity of the recruitment dynamics among communities declined rapidly with distance within the fine spatial scale, but only moderately within larger scales. The similarity of the disturbance dynamics was independent of distance, and the similarity of species interaction declined slightly with increasing distance. The similarity of whole-community dynamics declined rapidly with distance at a fine spatial scale and moderately at larger scales. The fact that the distance–decay relationship of whole-community dynamics was similar to that of recruitment may suggest that recruitment processes are the most important determinant of spatial variability of community dynamics at our study sites during the study period.     

Inter-hemispheric synchrony of forest fires and the El Niño-Southern Oscillation

Fire histories were compared between the south-western United States and northern Patagonia, Argentina using both documentary records (1914–87 and 1938–96, respectively) and tree-ring reconstructions over the past several centuries. The two regions share similar fire–climate relationships and similar relationships of climatic anomalies to the El Niño–Southern Oscillation (ENSO). In both regions, El Niño events coincide with above-average cool season precipitation and increased moisture availability to plants during the growing season. Conversely, La Niña events correspond with drought conditions. Monthly patterns of ENSO indicators (southern oscillation indices and tropical Pacific sea surface temperatures) preceding years of exceptionally widespread fires are highly similar in both regions during the 20th century. Major fire years tend to follow the switching from El Niño to La Niña conditions. El Niño conditions enhance the production of fine fuels, which when desiccated by La Niña conditions create conditions for widespread wildfires. Decadal-scale patterns of fire occurrence since the mid-17th century are highly similar in both regions. A period of decreased fire occurrence in both regions from c. 1780–1830 coincides with decreased amplitude and/or frequency of ENSO events. The interhemispheric synchrony of fire regimes in these two distant regions is tentatively interpreted to be a response to decadal-scale changes in ENSO activity. The ENSO–fire relationships of the south-western USA and northern Patagonia document the importance of high-frequency climatic variation to fire hazard. Thus, in addition to long-term trends in mean climatic conditions, multi-decadal scale changes in year-to-year variability need to be considered in assessments of the potential influence of climatic change on fire regimes.     

LET THERE BE LIGHT (BUT NOT TOO MUCH): MOLECULAR EVOLUTION OF LIGHT-HARVESTING COMPLEXES

Understanding spatio-temporal variability in recruitment is vital to studies of kelp population dynamics. Research on settlement and post-settlement processes has suggested that arrival of kelp zoospores to suitable substrate is important in limiting kelp recruitment, yet the role of planktonic processes in kelp population dynamics has not been studied due to difficulties in sampling and identifying zoospores. I developed a method to estimate kelp zoospore abundance from in situ plankton samples and used it to study various processes regulating the availability of giant kelp ( Macrocystis pyrifera ) zoospores for settlement. My studies focused on (1) identifying temporal scales over which zoospore abundance is most variable, (2) describing physical and biological processes that regulate this variability, and (3) determining the relationship between zoospore abundance and settlement. I found that short-term variability in zoospore abundance (<24 hrs) was not due to changes in supply but rather dispersion, caused by oscillating hydrodynamic forces (e.g. waves). Long-term variability in zoospore abundance, however, was best explained by the size and density of reproductive adult plants, with zoospore abundance being most variable at the scale of days to months. Changes in adult reproductive condition caused rapid changes in zoospore abundance suggesting that the supply of kelp zoospores is sensitive to environmental regulation of adult physiology. Thus, unlike with marine animals, these results indicate that variability in kelp propagule supply, over scales most likely to affect subsequent settlement and recruitment, is more tightly coupled to demographic and reproductive mechanisms than to physical transport processes.     

GRAZING ON GIANT KELP MICROSCOPIC PHASES AND THE RECRUITMENT SUCCESS OF ANNUAL POPULATIONS OF MACROCYSTIS PYRIFERA (LAMINARIALES,PHAEOPHYTA) IN SOUTHERN CHILE1

The giant kelp Macrocystis pyrifera (L.) C. Agardh is widely distributed in the Northern Hemisphere and Southern Hemisphere, yet it exhibits distinct population dynamics at local to regional spatial scales. Giant kelp populations are typically perennial with the potential for year‐round reproduction and recruitment. In southern Chile, however, annual giant kelp populations exist and often persist entirely on secondary substrata (e.g., shells of the slipper limpet Crepipatella fecunda [Gastropoda, Calyptraeidae]) that can cover up to 90% of the rocky bottom. In these populations, the macroscopic sporophyte phase disappears annually during winter and early spring, leaving a 3–4 month period in which a persistent microscopic phase remains to support the subsequent year’s recruitment. We tested the effects of a suite of grazers on the recruitment success of this critical microscopic phase at two sites in southern Chile. Field experiments indicated that the snail Tegula atra negatively impacted M. pyrifera sporophyte recruitment, but that recruitment was highest in the presence of sessile female limpets, C. fecunda. Conversely, small male C. fecunda (biofilm grazers) did not regulate kelp recruitment. Laboratory observations showed that C. fecunda males only grazed on microscopic kelp gametophytes and small (<250 μm) sporophytes, rejecting larger sporophytes, whereas T. atra grazed on all the kelp stages. Recruitment to the C. fecunda treatments far exceeded that to bare rock in the absence of grazers but was not due to the physical presence of C. fecunda shells. We concluded that the key to M. pyrifera recruitment success in southern Chile is its capacity to colonize secondary substrates provided by the slipper limpet C. fecunda.     

Scales of spatial patterns of distribution of intertidal invertebrates

Few comparative studies of spatial patterns at different scales have examined several species in the same habitat or the same species over a range of habitats. Therefore, variability in patterns among species or among habitats has seldom been documented. This study quantifies spatial patterns of a suite of intertidal snails and a species of barnacle using a range of statistical techniques. Variability in densities was quantified from the scale of adjacent quadrats (over a distance of centimeters) to tens of kilometers. Significant differences in abundances occurred primarily at two spatial scales. Small-scale differences were found at the scales of centimeters or 1–2 m and, for many species on many shores, these accounted for most of the variability in abundances from place to place. These are likely to be determined by behavioural responses to small-scale patches of microhabitat. Large-scale differences in abundance were also found in most species at the scale of hundreds of meters alongshore. These are likely to be due to variation in recruitment (and/or mortality) because of limited dispersal by adults of these species. There was little or no additional variation among shores, separated by tens of kilometers, than was shown among patches of shore separated by hundreds of meters. Identification of the scale(s) at which significant differences in abundance are found focus attention on the processes (and the scales at which these processes operate) that influence patterns of distribution and abundance. Some of the advantages and disadvantages of various procedures are discussed.