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Plankton are widely considered to be at the mercy of ocean currents, even after decades of research revealing that plankton regulate dispersal by positioning themselves in surface and bottom currents flowing in different directions. The degree of effectiveness of these behaviors remains controversial, because tiny plankters cannot be tracked at sea. Here, we experimentally tested the effectiveness of 3 vertical positioning behaviors in nature by developing a biomimetic robot that emulates them. We conducted a challenging test by deploying them in complex circulation during strong upwelling winds and wind relaxation and reversal events. Behavior alone dramatically affected transport. Transport trajectories of robots with 3 different behaviors diverged markedly while those sharing the same behavior were very similar. Moreover, all 3 behaviors produced trajectories that matched previously modeled projections during both upwelling and relaxation conditions at the study site: shallow plankton disperse far, deep plankton move little, and plankton migrating from depth during the day to the surface at night travel an intermediate distance. The ability of weakly swimming plankton to control their fate and replenish populations in a dynamic ocean is of central importance to the ecology and evolution of marine life and to the management of resources in a changing climate.

Larval dispersal phases are a key determinant of population dynamics in recruitment-limited, coastal upwelling regions. Larvae were long considered to be highly susceptible to offshore transport, except in the lee of headlands where eddies form during upwelling conditions. We examined the spatial variation of benthic invertebrate larval assemblages in relation to the retentive upwelling shadow in northern Monterey Bay (California, USA) during strong upwelling (August 2013) and weak upwelling (October 2013). We characterized the spatial variation in physical characteristics of the water column, determined the cross-shore and depth distributions of invertebrate larvae in relation to the upwelling shadow, and examined how these physical and biological patterns change with upwelling strength. Larval abundances and environmental data (water temperature, salinity, chlorophyll a concentration) were collected simultaneously using a plankton pump and profiling CTD at 3 depths: above, within, and below the chlorophyll a maximum layer. Larvae were primarily detected near the bottom. Larvae of most taxa were positively associated with a subsurface chlorophyll a maximum layer in August, but not in October when this layer was near the surface. Adult habitat distribution was related to the spatial distribution of larvae. Larvae of nearshore taxa occurred in the inner bay, while larvae of predominately offshore taxa occurred in the outer bay. Taxa with similar adult habitat (nearshore versus offshore) co-occurred in water samples. In addition, larvae of offshore taxa were commonly associated with offshore water types. Thus, the distribution of larvae within northern Monterey Bay appears to be strongly influenced by adult habitat distribution, vertical positioning of larvae in the water column, and upwelling strength.

Morphological defences of plankton can include armour, spines and coloration. Spines defend from gape-limited fish predators, while pigmentation increases visibility to fishes but defends from ultraviolet radiation (UVR). Planktonic crab larvae (zoeae) exhibit inter- and intraspecific variability in the lengths of defensive spines, extent of pigmentation and body size. The determinants of this variability and the relationships among these traits are largely unknown. Larvae may employ generalized defences against the dual threats of UVR and predation or specialized defences against their primary threat, with an unknown role of allometric or phylogenetic constraints. Generalization would result in longer spines compensating for the increased predation risk imposed by darker pigments, while specialization would lead to more investment in either defence from predation (long spines) or UVR (dark pigments), at the expense of the other trait. We examined (a) the relationship between spine lengths and pigmentation, (b) the scaling of spine lengths with body size, and (c) phylogenetic constraint in spine lengths, pigmentation, and body size, among and within 21 species of laboratory-hatched and 23 species of field-collected crab larvae from Panama and California. We found a negative relationship between spine length and pigmentation among species from laboratory and field. Within species, we found a marginally significant negative relationship among field-collected larvae. Spine lengths showed positive allometric scaling with carapace length, while spine and carapace lengths, but not pigmentation, had significant phylogenetic signals. The negative relationship we observed between pigmentation and spine length supports our defence specialization hypothesis. Positive allometric scaling of spine lengths means larger larvae are better defended from predators, which may indicate that larvae face greater predation risk as they grow larger. Phylogenetic constraint may have arisen because related species encounter similar predation threats. Conversely, phylogenetic constraint in the evolution of spine lengths may induce convergent behaviours resulting in related species facing similar predation threats. Our results improve understanding of the evolution of the larval morphology of crabs, morphological defences in the plankton and evolutionary responses of morphology to multiple spatially segregated selective forces. A free Plain Language Summary can be found within the Supporting Information of this article.

Dispersal of benthic species in the sea is mediated primarily through small, vulnerable larvae that must survive minutes to months as members of the plankton community while being transported by strong, dynamic currents. As climate change alters ocean conditions, the dispersal of these larvae will be affected, with pervasive ecological and evolutionary consequences. We review the impacts of oceanic changes on larval transport, physiology, and behavior. We then discuss the implications for population connectivity and recruitment and evaluate life history strategies that will affect susceptibility to the effects of climate change on their dispersal patterns, with implications for understanding selective regimes in a future ocean. We find that physical oceanographic changes will impact dispersal by transporting larvae in different directions or inhibiting their movements while changing environmental factors, such as temperature, pH, salinity, oxygen, ultraviolet radiation, and turbidity, will affect the survival of larvae and alter their behavior. Reduced dispersal distance may make local adaptation more likely in well-connected populations with high genetic variation while reduced dispersal success will lower recruitment with implications for fishery stocks. Increased dispersal may spur adaptation by increasing genetic diversity among previously disconnected populations as well as increasing the likelihood of range expansions. We hypothesize that species with planktotrophic (feeding), calcifying, or weakly swimming larvae with specialized adult habitats will be most affected by climate change. We also propose that the adaptive value of retentive larval behaviors may decrease where transport trajectories follow changing climate envelopes and increase where transport trajectories drive larvae toward increasingly unsuitable conditions. Our holistic framework, combined with knowledge of regional ocean conditions and larval traits, can be used to produce powerful predictions of expected impacts on larval dispersal as well as the consequences for connectivity, range expansion, or recruitment. Based on our findings, we recommend that future studies take a holistic view of dispersal incorporating biological and oceanographic impacts of climate change rather than solely focusing on oceanography or physiology. Genetic and paleontological techniques can be used to examine evolutionary impacts of altered dispersal in a future ocean, while museum collections and expedition records can inform modern-day range shifts.

Various physical mechanisms are implicated in the transport of zooplankton to the outer edge of the surf zone, which is the final barrier before reaching the adult habitat of many meroplanktonic organisms. To explore these physical mechanisms, we measured the abundance of zooplankton in the surf zone hourly for 3 consecutive days during strong internal tidal forcing while concurrently measuring winds, currents, and seawater temperature. Strong temperature changes in the water column that were associated with internal tidal bores, as well as onshore coastal winds, coincided with peaks in abundance of barnacle cyprids, gastropods, and bryozoan larvae in the surf zone. This study supports the hypothesis that both internal tidal bores and onshore winds can accumulate zooplankton nearshore, and that these transport mechanisms may act in concert.

We investigated whether cross-shore distributions of coastal phytoplankton to the surf zone are controlled by hydrodynamics and their biological characteristics. Data from a rip-channeled beach indicate that concentrations of phytoplankton are higher in the surf zone than offshore. To examine how phytoplankton is transported toward the shore, we used a coupled biophysical model, comprised of a 3D physical model of coastal dynamics and an individual-based model (IBM) for tracking phytoplankton on the rip-channeled beach. Waves and wind in the biophysical model were parameterized by the conditions during the sampling period. Previous studies indicated that growth rates of phytoplankton can be enhanced by high turbulence, which might contribute to high phytoplankton concentration in the surf zone. Some numerical and laboratory works showed that turbulence can also increase the downward velocity of phytoplankton, which could be carried by onshore bottom currents and remain in the surf zone. Furthermore, we adapted the IBM with the theoretical model of diurnal vertical migration (DVM) for phytoplankton. The theoretical DVM works as follows: in the morning, phytoplankton cells adhere to air bubbles and stay at the surface and close to the shore in the daytime because onshore wind and surface current direction is usually onshore; in the late afternoon, the cells switch their attachment from air bubbles to sand grains and sink to the bottom where the water flow is normally onshore at night. Finally, depth-varying growth of phytoplankton was also incorporated into the DVM module. Simulations using neutral passive particles do not give the expected results of observed patterns. All tested mechanisms, i.e., wind- and wave-driven currents, rip-current circulation, turbulence-driven growth and sinking, DVM, and depth-varying growth, enhanced onshore phytoplankton migration and cell concentrations in the surf zone, indicating that both biological traits and physical factors can be essential to phytoplankton cross-shore transport and spatial variability. Our model is open to be modified and re-parameterized, followed by further analysis and validation, so that it can be more adequate for ecological assessment of coastal areas.

Planktonic larvae are thought to be very susceptible to offshore advection in upwelling regimes, increasing dispersal and decreasing recruitment. However, larvae of 42 species of nearshore benthic crustaceans primarily developed on the inner shelf at locations both in (98.5%) and away (99.8%) from a perennial upwelling center in the upwelling season of a recruitment-limited region characterized by persistent, strong, upwelling. During three cross-shelf cruises conducted at each location, larvae of 21 species remained on the inner shelf at both sites by occurring beneath seaward-flowing surface currents while larvae of other species migrated to midshelf (four species) or offshore (14 species) by initially developing near the surface. Postlarvae apparently returned to shore either deep in landward-flowing upwelled water or near the surface where behavior allows them to be transported shoreward by internal waves, diel wind cycles or wind relaxation events. Thus, recruitment limitation in upwelling regimes does not appear to be caused by larval mortality from offshore transport, requiring new research directions to advance our understanding of population dynamics, structure and connectivity.

The Intermittent Upwelling Hypothesis (IUH) posits that subsidies of larvae and phytoplankton to intertidal communities should vary unimodally along a gradient of upwelling from persistent upwelling to persistent downwelling with most subsidies occurring where upwelling is of intermediate strength and intermittent. Furthermore, the hypothesis states that larvae and phytoplankton are transported far offshore by strong, persistent upwelling and fail to subsidize nearshore communities, whereas weak upwelling or downwelling reduces nutrients for phytoplankton production limiting food for larvae and nearshore communities. We review studies conducted at sea and onshore and reanalyze published data to test the IUH and evaluate alternative hypotheses. To test the hypothesis, we examine five predictions that must hold if the IUH is true. (1) Larvae should inhabit the surface Ekman layer where they are transported offshore during upwelling. Larvae of many intertidal taxa occur deeper in the water column where currents flow shoreward during upwelling. (2) Larvae of nearshore species should occur farther offshore during upwelling than during relaxation or downwelling. Larvae of many nearshore species remain within several kilometers of shore during both conditions. (3) Larval settlement in intertidal communities should be lower during upwelling than relaxation or downwelling. Daily larval settlement has not observed to be higher during relaxation or downwelling events; settlement has most often been seen to vary with the fortnightly tidal cycle likely due to onshore larval transport by internal tides. (4) Larval settlement and recruitment in intertidal communities should be lower in areas of strong, persistent upwelling than where upwelling is weaker and less persistent. Recruitment of mussels and barnacles to artificial and natural substrates did not vary with the strength of upwelling, but did vary inversely with two measures of desiccation potential, and directly with indicators of surf zone hydrodynamics; larval recruitment was higher where surf zones were more dissipative with rip currents. (5) Phytoplankton subsidies to nearshore communities should be highest where upwelling is moderate and intermittent. Like larval subsidies, phytoplankton subsidies varied spatially with surf zone hydrodynamics rather than upwelling. This reconsideration of the evidence for the IUH finds the hypothesis unsupported.

The California Current System is characterized by strong coastal upwelling that drives high primary production with implications for life in the plankton. We investigated the role of environmental variability at local, regional, and basin-wide scales in determining decapod larval distributions across space and time over 8 yr (2004 to 2011). We focused on an area of the California Current off the coast of central California, USA, characterized by a strong upwelling jet, a retentive zone with high primary production, a buoyant bay outflow plume, and proximity to both open coast and bay habitats. While multiyear studies of holoplankton distributions have revealed strong effects of basin-scale environmental variability, we did not find that basin-scale environmental changes resulted in major shifts in meroplankton distributions. Instead, meroplankton distributions in the California Current remained consistent across years — the oceanographic environment affected meroplankton distributions along complex shoreline topography but not across the shelf. Chlorophyll fluorescence and variables associated with regional upwelling were most correlated with larval distributions, indicating an association between larval distributions and primary production. We also found that environmental variability did not explain much of the variability in the larval distributions (only 5 to 20%), indicating that larval behaviors and demographic variables mediated the role of physical forcing in determining larval distributions. Larval distributions provide us with clues to determine larval transport and survival, adding to our understanding of how marine populations are connected, identifying threats to their persistence, and informing effective marine conservation and resource management planning.

Larvae of intertidal invertebrates need to cross the surf zone to settle in their adult habitat. Onshore transport of invertebrate larvae and detritus at a steep beach was simulated with a biophysical larval tracking model. Hydrodynamic model calculations were performed for 24 h after a 24 h spin-up stage with bathymetry and averaged wave data obtained during the summer of 2011 at Carmel River State Beach, California, and with and without onshore wind. The physical model output was then transferred to a Lagrangian larval tracking model using several types of particles representing larvae. A southward alongshore current controlled particle distribution in the middle and north of the domain. At the southern shore, negatively buoyant particles were trapped by eddies generated between the alongshore current and shore, while positively buoyant particles were carried onshore by wind-driven surface currents. The concentration of modeled detritus in the surf zone was positively correlated with that of negatively buoyant larvae. Additionally, the concentrations of detritus and competent larvae within the surf zone were negatively correlated with wave height, consistent with the observations of the accompanying field study. Some eddies contributed to forming high particle concentration patches by trapping them in the surf zone. More small eddies were generated closer to the shore with smaller waves, leading to high larval and detrital concentration in the surf zone. As waves increased in size, fewer and larger eddies formed, predominantly outside the surf zone, and consequently fewer larvae and detritus particles entered or stayed in the surf zone.

Larvae in the coastal ocean are transported toward shore by a variety of mechanisms. Crossing the surf zone is the last step in a shoreward migration and surf zones may act as semipermeable barriers altering delivery of larvae to the shore. We related variation in the structure of intertidal barnacle populations to surf zone width (surf zone hydrodynamics proxy), wave height, alongshore wind stress (upwelling proxy), solar radiation, and latitude at 40 rocky intertidal sites from San Diego, California to the Olympic Peninsula, Washington. We measured daily settlement and weekly recruitment of barnacles at selected sites and related these measures to surf zone width. Chthamalus density varied inversely with that of Balanus, and the density of Balanus and new recruits was negatively related to solar radiation. Across the region, long-term mean wave height and an indicator of upwelling intensity and frequency did not explain variation in Balanus or new recruit densities. Balanus and new recruit densities, daily settlement, and weekly recruitment were up to three orders of magnitude higher at sites with wide (>50 m), more dissipative surf zones with bathymetric rip currents than at sites with narrow (

Nearly all of our understanding of rocky inter-tidal ecology comes from studies conducted at low tide. To study inter-tidal organisms at high tide, we anchored waterproof digital GoPro® video cameras in wave-exposed tidepools and recorded the daytime movements of the black turban snail, Tegula funebralis, over the tidal cycle between May and August 2012 near Bodega Bay, California. Overall, snails moved more quickly and presumably foraged more during low tides and on days with warmer air and perhaps water temperatures. This is similar to other ectotherms that exhibit increased metabolic rates, movement and foraging in warmer conditions. Snails also moved less during flood and high tides, may have moved downward in tidepools at flood tides, and showed evidence of reduced activity on days with larger waves. This inactivity and refuge seeking may have been a strategy to avoid dislodgment by waves. Analysis of snail trajectories showed foraging bouts indicated by alternating zig-zagging and straight movement. There was no effect of temperature, wave height, or tidal phase on distribution of snail turning angles, suggesting that they may have foraged consistently but moved faster during warm conditions and low tides, thereby grazing a larger area. This is one of few direct recordings of inter-tidal organisms on wave-exposed rocky shores during high tide. The methods used here are easily transferable to other studies, which are needed to increase our understanding of behaviors that structure rocky shore communities during high tide.

Ecological connectivity can influence the distributions of diversity and productivity among ecosystems, but relationships among multiple marine ecosystems remain relatively uncharacterized. Sandy beaches are recipient ecosystems that support coastal food webs through deposits of drift macrophytes (wrack), and serve as test cases for exploring within-seascape connectivity. We present results from the first comprehensive survey of geographic and temporal patterns of wrack cover and composition on beaches along the North Central Coast of California and test the role of local donor ecosystems and physical factors in predicting wrack distribution. We surveyed wrack at 17 beaches in August 2010, and monthly at a subset of 10 beaches for 13 months. We estimated explanatory variables of (1) local donor ecosystem cover (kelp forests, rocky intertidal, and bays and estuaries), (2) biomass transport, and (3) beach morphology. Regression analyses were used to evaluate relationships among the cover of six key wrack categories and the explanatory variables above, for two time periods. We found persistent geographic variation in wrack composition and detected significant relationships between wrack cover and cover of local donor ecosystems for five of the six wrack categories (Nereocystis, Zostera, Postelsia, mixed red algae, and mixed brown algae). Transport mechanisms (wind exposure, swell exposure) or attributes of the recipient ecosystem (beach width, beach slope) explained additional spatial variation for three of the six wrack categories (Zostera, Phyllospadix, and mixed red algae). Our results support the concept of considering ecological connectivity (particularly the role of donor ecosystems upon which recipient ecosystems rely) in the design and management of protected areas.

Trait-mediated indirect interactions (TMIIs) are an important component of food web structure and dynamics. We determined whether TMIIs occur in rocky tidepool communities on the west coast of the USA. In the laboratory, both adults and juveniles of the keystone predator Pisaster ochraceus and adults of a smaller predatory seastar Leptasterias spp. caused the abundant herbivorous snail Tegula funebralis to stop foraging and flee the water, inducing a positive TMII on micro-and macroalgae. Snails preferred 3 common species of macroalgae (Ulva lactuca, Cladophora columbiana and Porphyra spp.) over 4 others, indicating that seastars might provide the strongest benefits to these species in tidepools. In the laboratory, snails responded rapidly to both species of predatory seastars and many more snails responded than could be eaten; thus, there is a potential for TMIIs to occur in natural populations. Snails responded to waterborne cues from P. ochraceus by reducing grazing and leaving still water, and reducing grazing in laminar flow (0.5 l min-1), resulting in TMII effects at least as far as 75 cm away. Adult P. ochraceus and Leptasterias spp. introduced to tidepools during low tide induced many snails to flee the tidepools. Considerable individual variation occurred in the responses of snails. Medium and large snails mediated TMIIs and hungry snails were marginally less responsive to seastars potentially altering TMII strength in nature. Thus, we demonstrated that TMIIs could occur in natural tidepools and showed how predator and algal identity, predator and prey size, water flow and prey hunger level may influence these TMIIs.

Characterizing the behavior of larvae prior to settlement is integral to understanding population dynamics because coastal oceanography may facilitate or limit settlement. Otolith microchemistry can be used to determine patterns of fish movement, although there is a limited understanding of how this tool can be applied in coastal marine systems. Our goal in this study was to evaluate the application of otolith microchemistry to characterize water mass associations of settlement-stage marine fish in a coastal upwelling region using a 3-step approach. First, we characterized seawater chemistry of coastal water mass types across multiple years, finding differences in the chemical signatures of strong upwelling, weak upwelling, and relaxation events. Second, we experimentally determined the effect of temperature on the partitioning of trace elements in otoliths for 2 rockfishes (Sebastes spp.) to find that the effect of temperature on otolith partition coefficients was element-and species-specific. Finally, we compared coeval changes in seawater and otolith chemistry of settlement-stage rockfishes that were exposed to naturally variable conditions over an upwelling-relaxation cycle. We subsequently evaluate whether laser ablation inductively coupled plasma mass spectrometry effectively measures otolith chemistry over ecologically relevant time scales. We discovered that elemental concentrations in otoliths respond rapidly to changes in seawater chemistry and reflect equivalent proportional changes. This study provides evidence that elemental signatures are valuable tools for reconstructing larval histories of marine fish in coastal upwelling regions.

Fine-scale spatial and temporal variation in the genetic composition of benthic recruits, known as chaotic genetic patchiness, is often observed in marine and estuarine species with planktonic larvae. Several explanations have been proposed for chaotic genetic patchiness, including sweepstakes reproductive success, variability in larval source, and natural selection. In a survey of the green shore crab Hemigrapsus oregonensis in Bodega Bay, California, USA, allele frequencies at a mitochondrial single nucleotide polymorphism were found to differ significantly among samples of first-stage zoeae and between zoeae and adults. Sweepstakes reproductive success is unlikely to be responsible because the fecundity of this species is too low and there was no reduction in genetic diversity among zoeae. In principle, influxes of larvae from genetically distinct populations over 500 km to the north could have caused these differences; however, coalescent estimates indicated that gene flow from these distant populations has been very low and it is unlikely that first-stage zoeae would have been transported such great distances. The possibility remains that natural selection, directly or indirectly, is responsible for the observed patchiness in mitochondrial allele frequencies.

Several studies on trait-mediated indirect interactions (TMIIs) have shown that predators can initiate trophic cascades by altering prey behaviour. Although it is well recognized that individual prey state alters antipredator and foraging behaviour, few studies explore whether this state-dependent prey behaviour can alter the strength of the ensuing tritrophic cascade. Here, we link state-dependent individual behaviour to community processes by experimentally testing whether hunger level and body size of prey altered antipredator behaviour and thus changed the strength of trophic cascades between predators and primary producers. In rocky intertidal tide pools on the California Coast, waterborne cues from the predatory seastar Leptasterias spp. (Stimpson) can cause the herbivorous snail Tegula (Chlorostoma) funebralis (A. Adams) to reduce grazing and flee tide pools, resulting in positive indirect effects on tide pool microalgae. However, we show that the strength of this behaviourally-mediated cascade may be contingent on prey hunger level and body size. During short field experiments at low tide, medium-sized snails that were either newly collected from the field or fed for 1�week in the laboratory mediated strong TMIIs because they grazed less when seastars were present. In contrast, no TMIIs occurred when medium-sized snails had been starved for 1�week because they continued grazing regardless of seastar presence. Newly collected small snails fled from seastars but did not mediate cascades because they ate little algae. Despite reaching an apparent size refuge from predation, many newly collected large snails fled from seastars, but those individuals that remained tended to graze the algae more quickly, resulting in unexpected negative indirect effects of seastars on algae cover. The implication of this pattern for the natural system is unclear. Because average hunger level and size of snails vary over time and space in nature, a mosaic of TMII strength may exist. Overall, the strength of tritrophic TMIIs in tide pools depended on individual prey state, supporting model predictions and adding to sparse empirical evidence. This outcome suggests that patterns occurring system-wide over the long term may be influenced by the state-dependent decisions made by the individuals present.

Variations in larval settlement in coastal upwelling regions such as the California Current System (CCS) have been attributed to variations in physical forcing at various time and space scales, but existing findings are often conflicting and fail to explicitly consider larval transport and swimming behavior. Using virtual larvae in a realistic simulation of the CCS, temporal relationships between wind stress, temperature and nearshore settlement in central California are explored for several vertical swimming behaviors, given a pelagic larval duration (PLD) of 20 to 22 d. A robust negative correlation between upwelling-favorable, PLD-averaged wind stress and settlement was found at timescales of days to years for larvae exposed to the surface boundary layer (SBL), while settlement for larvae that remain below the layer throughout development is increased, and their overall settlement is ∼2 to 20 times higher. A 20 d running mean of the wind stress can account for 52 to 86% of logit-transformed settlement variance over the 6 yr study period. Wind stress and settlement are coherent at all intra-annual periods greater than the PLD. Monthly climatological cycles of PLD-averaged wind stress and transformed settlement are nearly identical for behaviors exposed to the SBL and constitute ∼80% of the monthly variance. Monthly anomalies of wind stress and settlement are also significantly correlated. Nearshore surface temperature is also well correlated with settlement, but significantly less so than wind stress on both seasonal and intra-seasonal timescales. Other PLDs ranging from 10 to 40 d displayed similar results. Physical forcing in this model is found to dominate intrinsic eddy variability in driving settlement.

Larvae of benthic marine organisms are released amid high densities of suspension feeding and predatory adults and are highly subject to being consumed, even by conspecifics or their own parent. During laboratory feeding trials conducted in June 2006, female shore crabs (Hemigrapsus oregonensis) from Stege Marsh in San Francisco Bay (37°54.530′ N, 122°19.734′ W) that released their larvae during the previous 24 h ate fewer conspecific larvae than females that had not recently released larvae, though the behavior was not repeated during similar trials in 2007. Additionally, the number of larvae eaten increased with increasing starvation time, and hungrier females showed a trend toward eating more larvae from a different species (Carcinus maenas) than larvae of conspecifics. Thus, suppression of suspension feeding may reduce conspecific predation of newly released larvae, but this response partially depends on hunger level. This is the first time crabs have been shown to suppress feeding to reduce cannibalism of larvae, and this behavior could affect reproductive success and population dynamics.

Larvae of many intertidal species develop offshore and must cross the surf zone to complete their onshore migration to adult habitats. Depending on hydrodynamics, the surf zone may limit this migration, especially on reflective rocky shores. As a logistically tractable analog of a rocky shore environment, we carried out a comprehensive biological and physical study of the hydrodynamics of a steep reflective sandy beach. Holoplankton and precompetent larval invertebrates were much less abundant within the surf zone than offshore, and their concentrations inside and outside the surf zone were not significantly correlated, suggesting that they were not entering the surf zone. Persistent offshore flow throughout the water column at the outer edge of the surf zone may prevent these organisms from entering the surf zone. In contrast, the concentrations of detritus and a competent larval invertebrate (i.e. cyprids), while also not significantly correlated with concentrations offshore, were frequently more concentrated in the surf zone than offshore. Within the surf zone, the concentration of detritus was significantly correlated with concentrations of competent larval invertebrates (barnacles, gastropods, polychaetes, and bopyrid amphipod) and organisms that may be associated with detritus (amphipods and harpacticoid copepods). These concentrations were significantly negatively correlated with average daily wave height. We hypothesize that detritus and larvae enter the surf zone near the bottom during calm wave conditions by a process of near-bottom streaming. Near-bottom streaming is associated with all surf zones and may be a general mechanism for onshore transport of larvae close to the coast.

Fundamental knowledge of mortality during the planktonic phase of the typical marine life cycle is essential to understanding population dynamics and managing marine resources. However, estimating larval mortality is extremely challenging, because the fate of microscopic larvae cannot be tracked as they develop for weeks in ocean currents. We used a two-pronged approach to provide reliable estimates of larval mortality: (1) frequent, long-term sampling where the combination of larval behaviors and recirculation greatly reduces larval transport to and from the study area, and (2) an improved method of calculating larval mortality that consists of a vertical life table with a negative binomial distribution to account for the notorious patchiness of plankton. Larval mortality rates of our study species (barnacles and crabs) were ≤0.14 larvae/d, which produce survivorships over an order of magnitude higher than commonly determined for marine larvae. These estimates are reliable because they were similar for species with similar dispersal patterns. They are conservative because they were conducted in a highly advective upwelling system, and they may be even lower in other systems using our approach. Until other systems can be tested, our improved estimates should be used to inform future models of population dynamics and the evolution of life histories in the sea.

Larvae of benthic marine organisms are released amid high densities of suspension feeding and predatory adults and are highly subject to being consumed, even by conspecifics or their own parent. During laboratory feeding trials conducted in June 2006, female shore crabs (Hemigrapsus oregonensis) from Stege Marsh in San Francisco Bay (37°54.530' N, 122°19.734' W) that released their larvae during the previous 24 h ate fewer conspecific larvae than females that had not recently released larvae, though the behavior was not repeated during similar trials in 2007. Additionally, the number of larvae eaten increased with increasing starvation time, and hungrier females showed a trend toward eating more larvae from a different species (Carcinus maenas) than larvae of conspecifics. Thus, suppression of suspension feeding may reduce conspecific predation of newly released larvae, but this response partially depends on hunger level. This is the first time crabs have been shown to suppress feeding to reduce cannibalism of larvae, and this behavior could affect reproductive success and population dynamics. © 2014 Blackwell Verlag GmbH.

Larval supply determines year-class strength of fisheries and complex ecological interactions among adults of benthic marine species. In upwelling regions, a latitudinal cline in the intensity and persistence of upwelling is thought to affect larval advection and recruitment, thereby regulating the intensity of interactions in adult populations and communities. We tested this hypothesis by determining the monthly cross-shelf abundances of nearshore benthic crustacean larvae throughout development during the peak upwelling season for 7 years in a region of intermittent upwelling and high recruitment (45°N). We tested whether larvae were found farther offshore during upwelling conditions, and we compared their interspecific cross-shelf distributions to previous results from a region of strong, persistent upwelling (38°N). We also compared larval abundances across the 2 upwelling regions for 1 year. Most species were retained nearshore, regardless of intra- and inter-annual variations in the intensity of upwelling. In both upwelling regions, larvae of each species consistently occurred at different distances from the shore. Further, there were no differences in nearshore larval abundance across upwelling regions for all but 1 larval stage of 1 species. Thus, latitudinal variations in the intensity and persistence of upwelling do not appear to affect larval survival, providing further evidence that nearshore processes may be a primary determinant of larval delivery to the rocky intertidal across these regions. © Inter-Research 2014.

Identifying biophysical mechanisms of larval transport is essential to understanding the delivery of larvae to adult habitats. In addition, harmful algal blooms (HABs) can be transported onshore from populations that form offshore. In summer 2011, we measured sea surface and bottom temperatures and daily phytoplankton abundance and intertidal cyprid (barnacle post larvae) settlement at Carmel River State Beach, California, USA. Using time-series analysis, we compared the abundance of Pseudo-nitzschia spp. and daily cyprid settlement to physical forcing mechanisms (e.g. internal tides and upwelling-relaxation events) that could generate onshore delivery. Minimum bottom water temperature was significantly cross-correlated with the spring-neap tidal cycle; minimum temperatures occurred between neap and spring tides, and maximum temperatures were recorded around neap tides. When the temperature data were transformed to remove the relationship between tides and temperature, we found significantly higher maximum sea surface temperatures during upwelling-relaxation events. We observed 4 pulses in Pseudo-nitzschia spp. abundance. Pseudo-nitzschia spp. chains were longest at the start of pulses and then decreased, suggesting that they had been transported to shore from a more productive site offshore, likely the upwelling front. Pulses occurred during periods of maximum sea surface temperature associated with upwelling-relaxation events. In contrast, cyprid settlement was significantly cross-correlated with the spring-neap tidal cycle, with settlement peaks occurring during fortnightly periods of cold bottom temperatures; onshore transport of cyprids appears to have been due to the internal tides. © Inter-Research 2014.

ABSTRACT: Explorations of the dynamics of nearshore regions of the coastal zone are missing from many efforts to understand larval transport and delivery to suitable habitats. Larval distributions in the coastal ocean are variable and depend on physical processes and larval behaviors, leading to biophysical interactions that may increase larval retention nearshore and bolster their return to natal sites. While recent evidence suggests that many larvae are retained within a few kilometers from shore, few studies incorporate measurements sufficiently close to shore to plausibly assess supply to the shoreline benthos. We measured cross-shore distributions of larvae of benthic crustaceans between 250 and 1100 m from shore (i.e. just beyond the surf zone) within the coastal boundary layer (CBL)-a region of reduced alongshore flow-and simultaneously quantified a suite of physical factors that may influence larval distributions. We found high larval abundance within the CBL, with a peak at 850 m from shore, and a decrease in abundance along the shoreward edge of the sampled transect. We also found distinctly different larval assemblages at outer stations within the CBL, as compared to inner stations that are more influenced by shoreline dynamics. These patterns persisted across sample dates, suggesting that the spatial structure of nearshore larval assemblages is at least somewhat robust to temporal changes in physical conditions. Thus, while larval abundance appears to be high within the CBL, larvae appear to be sparse within the narrow band of water adjacent to the surf zone. Low larval supply adjacent to suitable habitats has important implications for the coupling of supply and recruitment, and resulting dynamics of shoreline populations.© Inter-Research 2013. www.int-res.com.

Timing of larval release by many shore crabs is cued by environmental cycles to occur during nocturnal spring ebb tides, when larvae are transported away from high densities of planktivorous fishes in the dark. However, a recent laboratory study indicated that larval release may be weakly synchronized relative to this safe period in upwelling regions, potentially increasing fish predation. We determined the timing of larval release and predation in marshes in an upwelling region by sampling plankton and fishes during flood and ebb tides on either side of high slack tide. Larval release was weakly synchronized, peaking during spring and intermediate ebb tides in twilight and darkness. Almost all larvae (99.8%) were eaten at twilight during peak release, when they likely were more visible than at night. However, larvae comprised only 4.1% of the diets of the 3 fish species that ate them. These fish species were often absent when conditions would make larvae most vulnerable to predation, and they preferred other prey to welldefended larvae. Larvae released outside the safe period were eaten more than those that were released during the safe period, providing selection for the timing of larval release. However, despite the large numbers of larvae present outside of the safe period, predation by fishes was much lower than expected. Thus, the selective effects of fish predation may be relaxed, raising the possibility that the strength of fish predation as a selective force varies among coasts and other selective forces that may affect the timing of larval release. © Inter-Research 2013.

We explore the phytoplankton community structure and the relationship between phytoplankton diversity and productivity produced by a self-emergent ecosystem model that represents a large number of phytoplankton type and is coupled to a circulation model of the California Current System. Biomass of each modeled phytoplankton type, when averaged over the uppermost model level and for 5-years, spans 7 orders of magnitude; 13 phytoplankton types contribute to the top 99.9% of community biomass, defining modeled species richness. Instantaneously, modeled species richness ranges between 1 and 17 while the Shannon index reaches values of 2.3. Diversity versus primary productivity shows large scatter with low species richness at both high and low productivity levels and a wide range of values including the maximum at intermediate productivities. Highest productivity and low diversity is found in the nearshore upwelling region dominated by fast growing diatoms; lowest productivity and low diversity occurs in deep, light-limited regions; and intermediate productivity and high diversity characterize offshore, oligotrophic surface waters. Locally averaged diversity and productivity covary in time with the sign of correlation dependent on geographic region as representing portions of the diversity-productivity scatter. © 2012 Elsevier B.V.

The maintenance of marine populations depends on the completion of larval migrations between adult and larval habitats, but the ability of microscopic larvae to regulate their movements in such a dynamic environment has been debated. Using a comparative hypothesis testing approach and intensive hourly sampling throughout the water column, we determined the ability of larvae of 6 species with different swimming abilities (2 gastropods, 2 crabs, 2 fishes) to overcome strong tidal mixing during spring tides and regulate their vertical, and hence, horizontal position, in opposing depth-stratified currents in the upper Hudson River estuary, USA. The vertical distributions of nonmotile eggs and swimming larvae generally differed, suggesting that larvae regulated depth. Eggs were passively mixed by tidal currents, but larvae typically aggregated in stratified portions of the water column, which fostered retention in the upper estuary. However, the capacity to regulate depth depended on swimming ability and the degree of mixing. In a predominantly mixed tidal environment upstream, mixing overcame most larvae when current velocities were maximal during mid-ebb or mid-flood tides; tidal vertical migrations were not evident for any species, and diel vertical migrations were apparent for only 1 species. In a partially stratified water column downstream, diel vertical migrations were apparent for larvae of 3 of 4 invertebrate species, and tidal vertical migrations were apparent for the 2 fish species. The presence of all larval stages showed that all species regulated depth sufficiently well to remain in the upper estuary. Regardless of swimming ability, larvae were retained in the estuary by occurring near the level of no net motion, even without completing tidal vertical migrations. Continuous profiling of larvae and hydrodynamics is necessary to reveal the ability of larvae to regulate depth in tidal mixing and recruit to adult populations. © Inter-Research 2013.

Densities of abalone in southern California have been dramatically reduced by overfishing and disease, leading to the collapse of some populations, and low fertilization rates may be hindering population recovery. This is a pattern typical of abalone species globally. However, movement may produce clustered distributions that promote fertilization success in broadcast spawners, such as abalone, even at low regional densities. We translocated wild, adult pink abalone Haliotis corrugata to an existing pink abalone patch to create a high-density aggregation, and then used acoustic telemetry to characterize abalone movement and monitor aggregation maintenance for a period of 14 mo in the Point Loma kelp forest near San Diego, California. Abalone showed a minimal flight response to handling that did not differ between the resident or translocated groups. Most individuals exhibited small home ranges (median area 183 m2) and homing behavior consisting of regular back-and-forth movement to a single point. Nomadic movement was also observed in several individuals. Though site fidelity may help maintain aggregations, abalone density decreased, and nearest neighbor distances increased at our site to nearinitial levels after 18 mo via a combination of mortality, large movements of a few individuals, and small, incremental movements of most individuals. No coordinated movements that would suggest spawning behavior were observed. Translocation of wild abalone to produce aggregations may not result in high rates of fertilization success that promote population recovery. However, the homing behavior observed may provide more opportunities for mating than expectations based on static measures of density and aggregation state. © Inter-Research 2013.

Genetic diversity has emerged as an important source of variation in the ecological properties of populations, but there are few studies of genetic diversity effects on colonisation processes. This relative scarcity of studies is surprising given the influence of colonisation on species coexistence, invasion, and population persistence. Here, we manipulated relatedness in experimental populations of colonising larvae in four sessile marine invertebrates. We then examined the influence of coloniser relatedness on the number, spatial arrangement and phenotype of colonisers following permanent settlement. Overall, relatedness influenced colonisation in all four species, but the effects of relatedness on colonisation differed among species. The variable responses of species to manipulations of relatedness likely reflect differences in intensity of inter- and intra-specific competition among adults, as well as the differential consequences of larval behaviours for each species. Relatedness appears to play an underappreciated role in the colonisation process, and we recommend that future studies of genetic diversity effects consider not only adult stages - the focus of most work to date - but also the importance of genetic diversity in early life history stages. © 2012 The Authors. Oikos © 2012 Nordic Society Oikos.

Trace elemental signatures incorporated into calcified structures in fish and mollusk larvae have been used to reveal population connectivity and larval dispersal trajectories. To determine whether trace element analysis could be applied to larvae that do not permanently retain calcified structures, we raised larvae of the porcelain crab Petrolisthes cinctipes from multiple locations along the northern California, USA, coast in a common water source for up to 8 wk. We analyzed 20 elements in extractions of soft tissues from embryos and the 2 larval and 1 postlarval stages. Elemental signatures of individuals in each developmental stage were compared to an atlas of signatures from each collection site using discriminant analysis to determine whether larvae could be accurately assigned to their site of origin in 2 ways. First, postlarvae were classified using a natal site atlas constructed using embryonic signatures from each site, and this classification had poor success (average 39.7% correct). Second, larvae of each stage were classified using a natal site atlas constructed using signatures of larvae from that same stage. This yielded considerably better classification success (81.7% correct overall). Thus, the same trace element signatures were not consistently maintained from embryos to postlarvae, but differences in signatures among natal sites were maintained during the larval period. Trace element signatures in soft tissues could be useful in tracking dispersal between stages and determining how many sites, rather than which sites, contributed to a cohort of larvae or settlers. © Inter-Research 2013.

Dispersal is fundamental to understanding the ecology and evolution of species and effectively managing resources, but information on dispersal is rare for the vast majority of marine life that develops as miniscule larvae in the plankton. Until recent evidence to the contrary, it was widely suspected that larvae developing in productive upwelling regimes along eastern ocean margins are susceptible to cross-shelf transport by strong, dynamic currents and often are unable to replenish populations. We now show that interspecific differences in depth preference likely play a role in regulating differences in offshore migration from adult populations. Two open-coast species (Petrolisthes cinctipes and Pachygrapsus crassipes) did not undertake endogenously timed tidal vertical migrations but an estuarine species (Hemigrapsus oregonensis) did, which would facilitate seaward dispersal. Surprisingly, none of the 3 species undertook diel vertical migrations in the laboratory, even though all do in the field. Diel vertical migrations in marine and freshwater species elsewhere in the world are cued by light and reduce fish predation, but they may be cued by turbulence in upwelling regimes, thereby reducing transport from adult populations. Thus, larvae of species that hatch in different locations and develop at different distances from the shore exhibit diverse larval swimming behaviors that regulate transport in dynamic upwelling regimes in previously undescribed ways, which has implications for population connectivity, local adaptation and resource management. Copyright © 2013 Inter-Research.

Natural trace element signatures have increasingly been used to track the dispersal of marine larvae, and these studies require an atlas of potential source populations with distinctive elemental signatures. To determine whether natal site atlases could be used repeatedly and to identify site characteristics that yield the best results, we built atlases in 5 consecutive years using embryos of the porcelain crab Petrolisthes cinctipes that were collected from 15 sites spanning 190 km of the open coast of northern California, USA. We analyzed the elemental composition of the embryos using a discriminant function optimization procedure to determine the suite of elements that resulted in the best reclassification success for individual sites and groups of sites each year. No single element or group of elements succeeded in discriminating the origins of embryos every year, and the reclassification success of the atlas varied at all spatial scales among years. Average reclassification success at the site level ranged annually from 39.5 to 54.3% correct, and combining sites into 2 or 3 areas improved the overall reclassification success to 72.5 to 97.7% correct. Sites with (1) distinctive elemental compositions of rocks, (2) unusual habitats, (3) consistent freshwater input, (4) consistent anthropogenic inputs, or (5) complex local oceano graphy had the highest reclassification success (up to 86.7% correct), but interannual variation in runoff reduced the temporal stability of the atlas. To improve population connectivity estimates, future trace element studies should consider these 5 key characteristics when selecting sites and anticipate temporal variation in natal site signatures. © Inter-Research 2013.

Many organisms live in spatially delimited habitat patches, where local extinctions are compensated by dispersal and recolonization of empty habitat patches. To predict the dynamics of such metapopulations in an era of large environmental changes, it is essential to understand the key abiotic factors affecting local occurrence and temporal variation in patch occupancy. Here, we investigated the metapopulation dynamics of the marine copepod Tigriopus californicus, which is restricted to high intertidal and supralittoral rock pools. We monitored populations of T. californicus in northern California for almost 2 yr and observed pronounced seasonal changes in patch occupancy with

Intertidal crabs in diverse habitats worldwide release larvae synchronously during nocturnal spring high tides. This expedites seaward transport of the larvae to beyond high density areas of predatory fishes under the cover of darkness. We found that 4 species of intertidal crabs along the west coast of the USA shared this reproductive timing pattern. As in other mixed semidiurnal tidal regimes, biweekly patterns of larval release were more closely synchronized with the tidal amplitude cycle than the lunar cycle, and some crabs released larvae in daylight. However, unlike other places in the world, larval release was weakly synchronized to environmental cycles regardless of interspecific differences in vertical distributions on the shore. We provide evidence that weak synchrony in the timing of larval release in upwelling regimes can result from exposure to environmental variation over long incubation periods of externally brooded embryos. According to the prevailing paradigm, weaker synchrony in the timing of larval release will increase predation by planktivorous fishes in upwelling regimes. Weak synchrony in the timing of larval release should increase larval mortality in a wide array of animals that brood embryos in the intertidal zone, regardless of the selective force operating, and it could contribute to recruitment limitation in upwelling regimes. © Inter-Research 2011.

Larvae are considered to be highly susceptible to offshore transport, thereby limiting recruitment in productive upwelling regions, except in the recirculation features that form in the lee of headlands. We show that 19 of 20 taxa of nearshore crustaceans were at least as common on the open coast as they were in the lee of a headland in a recruitment-limited region characterized by strong upwelling. Nine of these taxa occurred in high concentrations throughout development, indicating that larvae completed development nearshore in a coastal boundary layer of reduced Ekman transport. Larvae of 9 more taxa were prevalent as early stage larvae, and later stages were uncommon, indicating that larvae migrated offshore. The relative abundance of larval stages largely matched the vertical distributions of larvae in this study and a previous cross-shelf survey at this site, indicating that larval behavior plays a substantial role in determining whether larvae complete development nearshore or migrate offshore later in development. Larvae of taxa that occurred nearshore in high concentrations throughout development resided below a shallow Ekman layer of seaward-flowing surface waters, whereas most larvae of the other taxa occurred closer to the surface. Whether larvae undertook ontogenetic and diel vertical migrations also appeared to determine whether they were retained nearshore. Thus larval retention may be more widespread in upwelling regions than is widely believed, and the cause of recruitment limitation along upwelling coasts needs to be determined to further advance our understanding of the connectivity, dynamics and structure of coastal populations. © Inter-Research 2010.

Larvae are thought to be highly vulnerable to offshore transport of upwelling regions, limiting recruitment to infrequent wind relaxation and downwelling events. However, larvae could also be transported onshore by upwelled bottom waters, onshore wind-forcing or internal tides throughout the water column. We determined the relative importance of these hypothetical mechanisms for the timing of recruitment of 8 invertebrate taxa during the peak upwelling season in a region of strong, persistent upwelling. Recruitment was determined for 5 yr at an open embayment near the surface and bottom of the water column to examine the interaction of behavioral and physical processes regulating larval recruitment. Postlarvae consistently recruited near the surface or near the bottom depending on the species. Onshore delivery of larvae during wind relaxations and reversals did not best explain recruitment patterns in our area for most taxa. Only mussels consistently recruited more during these events, and they recruited in bottom rather than surface waters. Six crab taxa recruited primarily during upwelling. Recruitment of 7 taxa was intermittently correlated with the maximum tidal range, suggesting that internal waves also may deliver larvae onshore. Thus larvae may recruit by multiple processes in upwelling regions rather than being limited to infrequent relaxation events, leaving the mechanisms responsible for observed spatial patterns in larval recruitment and adult densities unexplained. Comprehensive studies of the behavioral and physical processes underlying larval recruitment and post-settlement mortality are needed to explain observed temporal and spatial variation in population dynamics and community structure in upwelling regions. © Inter-Research 2009.

Larvae are thought to be highly vulnerable to offshore transport in productive upwelling regions, which increases cross-shelf and alongshore dispersal, limits recruitment, and reduces the strength of community interactions. We investigated whether the last planktonic stages of shallowwater crabs (1) occur far offshore during the peak upwelling season in a region of strong upwelling that is recruitment limited, (2) occur farther offshore during upwelling than relaxation conditions, (3) occur farthest from shore at a major headland where currents are deflected offshore, and (4) recruit less during years of stronger upwelling. Crab postlarvae were collected from between 1 and 70 km from shore at 3 locations across an upwelling cell during upwelling and relaxation conditions in northern California, USA, for 3 yr, and recruitment was measured at 2 sites during 2 of these years. Postlarvae of all species collected were most abundant on the inner continental shelf (84%) and were scarce in the open ocean. Postlarvae of 3 taxa only occurred on the inner shelf, whereas postlarvae of the other 3 taxa collected occurred in low abundances on the outer shelf even during prolonged upwelling. They were abundant close to shore at the Point Reyes headland where flow is deflected offshore. Postlarvae did not occur farther from shore during a year of very strong upwelling, and 4 of 7 taxa recruited more then. Thus, crab postlarvae do not appear to be advected far offshore or recruit less during strong upwelling conditions, and the cause and extent of recruitment limitation warrants further investigation. © Inter-Research 2009.

Understanding interactions between invasive species and recipient communities is essential to determining whether invasive species will become established and spread. In this study, we explored the role of competition and the specific mechanisms of interaction in limiting the spread of the Mediterranean bay mussel Mytilus galloprovincialis within a Pacific Northwest invasion front. We examined the role of direct (interference) and indirect (exploitation) mechanisms of competition among M. galloprovincialis and 2 native mussels (M. trossulus and M. californianus). As the fastestgrowing organisms are often competitively dominant in space-limited systems, such as rocky intertidal communities, we used changes in relative performance (growth and survival) in monocultures and polycultures to assess interactions among mussels. Performance of M. galloprovincialis was always greater than that of the 2 native species of mussels in both field and laboratory manipulations of species composition and density, indicating that interspecific competition did not strongly limit the growth or survival of the invader. Moreover, the presence of M. galloprovincialis consistently led to both reduced growth and survival of M. trossulus. Laboratory studies of mussel feeding and behavior revealed M. galloprovincialis to be a robust interference competitor. The invader restricted movement, smothered and interfered with filter feeding of the 2 native species of mussels. Rather than limiting invasion, interference competition gave M. galloprovincialis a competitive advantage over the native mussels. Our results suggest M. galloprovincialis may have contributed to the displacement of M. trossulus along much of its historic southern range. ©Inter-Research 2009.

Trophic relationships among native and exotic species produce novel direct and indirect interactions that can have wide-ranging community level effects and perhaps confer invasion resistance. We investigated whether native predators have the potential to directly limit the spread of the exotic mussel Mytilus galloprovincialis or mediate interactions among native and invasive mussels at a rocky intertidal invasion front in northern California. Lower survival of M. galloprovincialis in transplanted cultures exposed to predators indicated that the invader was more vulnerable to predators than the numerically dominant native M. californianus. Survival and per capita mortality rate in monocultures and polycultures did not vary for either M. galloprovincialis or M. californianus, suggesting that predator-mediated apparent competition and associational defense did not occur. Complementary laboratory feeding trials determined which among 3 intertidal predators preferred the exotic to 2 native species of mussel. The whelk Nucella ostrina was most selective, consuming the thinner shelled mussels (M. galloprovincialis and the native M. trossulus) rather than the thickershelled native species M. californianus. The crab Cancer antennaiius and the sea star Pisaster ochraceus showed no preferences among mussel species. N. ostrina were commonly observed among field-transplanted mussels; thus whelk prédation may be especially important in limiting the establishment of the invasive mussel. However, 15% of M. galloprovincialis remained intact in the field after 1 yr, suggesting that prédation alone may not inhibit establishment of the invader. A tenuous balance between larval settlement and early post-settlement prédation likely characterizes the invasion front. © Inter-Research 2009.

Home range has been estimated for a limited number of marine fishes; however, the use of space and timing of activities within the home range has rarely been studied. In addition, understanding movement patterns of exploited fish species has been identified as a crucial science gap, impeding informed marine reserve design. We used a radio-acoustic positioning telemetry (VRAP) system to monitor detailed movements of 10 blue rockfish Sebastes mystinus around shallow rock pinnacles and stands of bull kelp Nereocystis leutkeana in central California in September 2002. The mean home range was 8783 m2 ± 1137 SE; however, activity was highly concentrated in 1 to 3 core areas within each home range. Mean core areas measured 1350 m2 ± 286 SE, but accounted for ∼83 % of activity. All core areas were centered over rock pinnacles where rockfish were highly aggregated. Individuals exhibited high site fidelity and made only brief radial excursions away from these centers or moved directly from one pinnacle to the next along defined corridors. Patterns of diel activity and nocturnal sheltering corresponded closely with nautical twilight. Cores overlapped, but estimated locations of nocturnal shelters differed significantly among individuals. Movement patterns were correlated with wind velocity, upwelling index, water temperature and habitat structure. © Inter-Research 2006.

Delivery mechanisms for crab postlarvae along upwelling coasts have been inferred from weekly sampling, but more frequent sampling is needed to better relate highly variable oceanographic conditions to postlarval supply. Settlement of 8 crab taxa was measured in Bodega Bay, California every 2 d from 12 May through 3 July, 2002. Abundance of postlarval and juvenile settlers was cross-correlated with physical variables. Four upwelling and 4 relaxation events were recorded as fluctuations in wind stress and sea temperature during this period. Transitions to and from upwelling conditions in Bodega Bay were brief and temperature changes occurred rapidly. The strongest correlations between crab abundance and physical variables indicative of upwelling and relaxation conditions were observed for Cancer magister, which settled primarily during relaxation-favorable conditions, and for Cancer antennarius/productus, which settled primarily during upwelling-favorable conditions, suggesting interspecific differences in delivery of postlarvae to adult habitat. Weak correlations with upwelling-favorable conditions were observed for Pugettia producta/richii and Pagurus spp. Settlement of Hemigrapsus nudus, Petrolisthes cinctipes, and P. eriomerus exhibited significant correlations with changes in tidal height. These results highlight the existence of multiple taxon-specific delivery mechanisms of closely related taxa in one small geographic region. © Inter-Research 2006.

Oceanic currents and larval accumulation potentially have large impacts on the choice of locations for marine reserves. Larval settlement of benthic invertebrates was greater in the lee than on the windward side of a small headland during the height of upwelling in central California during 2001 and 2002. Strong upwelling during the study was indicated by mean seasonal Bakun indices of 149 to 176 m3 s-1 per 100 m of coastline in 2001-2002. Weekly sampling of near-surface and near-bottom settlement in the lee of Bodega Head from August 2000 to September 2001 revealed that most larvae of 7 crab taxa settled during spring and summer, which coincides with the upwelling season. Comparison of sites in the protected (lee) and exposed (windward) sides of the headland (2 sites each) during the peak settlement season in 2001 showed that most larvae settled in the lee of the headland, including 91% of crabs, 89% of barnacles, and 80% of mussels in weekly samples. During 2002, weekly sampling at 1 protected and 1 exposed site also demonstrated that most settlement occurred in the lee of the headland, including 74% of crabs, 82% of barnacles, and 65% of mussels. Crabs settled mostly at the surface, whereas barnacles and mussels primarily settled near the bottom, indicating that postlarvae in both surface and bottom waters accumulate in the lee of the headland. Larval accumulation zones should be included in networks of marine reserves to supply adult populations with propagules in recruitment-limited upwelling regions. © Inter-Research 2006.

The utility of the lined shore crab Pachygrapsus crassipes as an indicator of small-scale variation in the condition of salt marshes was determined by conducting detailed mark-recapture studies of the spatial and temporal patterns of movement in 2 marshes in northern and southern California, and by conducting a stable isotope study along a nutrient gradient at one of the marshes. Crabs (1158) were captured from creek bank and vegetated marsh habitats, tagged, and released. Most crabs were recaptured in the same trap where they were originally tagged, and on average, they traveled less than 5 m from the site of first capture. The mean distance traveled did not increase significantly with the time between captures. A high percentage of crabs (62.1%) tagged near the creek bank remained there; fewer tagged crabs moved between the creek bank and the marsh plain (18.2%) or remained on the marsh plain (12.6%). Crabs moved least along creek banks and farthest when crossing creeks, although few of them did the latter (6.2%). During spring tides, crabs moved from tidal creeks onto the marsh plain. The nitrogen isotope data mirrored the gradient in nutrient input, supporting the conclusion that P. crassipes remained and fed within a localized area. Thus, the lined shore crab can serve as an indicator of small-scale differences in contaminant exposure. Furthermore, semiterrestrial species that are readily sampled and move little are abundant around the world, making them ideal indicators of habitat condition. © Inter-Research 2006.

We estimated juvenile abundance and predation potential for young-of-the-year (YOY) blue crabs Callinectes sapidus in upper- and lower-salinity marsh and submerged aquatic vegetation (SAV) 'nursery habitats' of Mobile Bay, Alabama, USA, and surrounding waters, and combined this information with previously published post-larval abundance data from these habitats to evaluate the relative roles of pre- and post-settlement events in determining the abundance of YOY crabs. We found little evidence for a significant relationship between megalopal supply and juvenile abundance, except shortly after a few very large, episodic pulses of postlarvae. However, even after large settlement events, in less than 14 d densities of YOY crabs had declined in a density-dependent manner to prior 'background' levels. These declines in density were probably due to high predation rates that produced as much as 95% mortality on a daily basis. As many previous studies had found, greater predation risk was recorded for crabs on unvegetated substrate than for those in SAV or marsh grass. Overall, losses to predators were high at our study sites compared to those reported in similar studies along the Atlantic coast of the United States. In both years of the study (1990 and 1991), the largest numbers of juvenile crabs were found in poly- and mesohaline SAV and salt marsh habitats. In Year 1, no significant correlations were observed between either marsh stem density or SAV vegetation biomass and YOY crab abundance, although at one location in the second year there was a significant positive correlation between the number of juveniles and mid-bay stem density. No significant differences in crab size (carapace width, CW) were found among locations in either year. In comparison with other, better-studied mid-Atlantic coast estuaries in the US (e.g., the Chesapeake and Delaware bays), very large numbers of megalopae (frequently 1 to 2 orders of magnitude greater) invade the Mobile Bay system, as well as other Gulf Coast estuaries. However, YOY abundance in marsh and SAV 'nursery habitats' was of the same order of magnitude as that reported for mid-Atlantic estuaries. We conclude that although postlarval supply was very large, post-settlement loss to predators was the dominant factor influencing blue crab population dynamics in our north central Gulf of Mexico study area.

We estimated juvenile abundance and predation potential for young-of-the-year (YOY) blue crabs Callinectes sapidus in upper- and lower-salinity marsh and submerged aquatic vegetation (SAV) 'nursery habitats' of Mobile Bay, Alabama, USA, and surrounding waters, and combined this information with previously published post-larval abundance data from these habitats to evaluate the relative roles of pre- and post-settlement events in determining the abundance of YOY crabs. We found little evidence for a significant relationship between megalopal supply and juvenile abundance, except shortly after a few very large, episodic pulses of postlarvae. However, even after large settlement events, in less than 14 d densities of YOY crabs had declined in a density-dependent manner to prior 'background' levels. These declines in density were probably due to high predation rates that produced as much as 95% mortality on a daily basis. As many previous studies had found, greater predation risk was recorded for crabs on unvegetated substrate than for those in SAV or marsh grass. Overall, losses to predators were high at our study sites compared to those reported in similar studies along the Atlantic coast of the United States. In both years of the study (1990 and 1991), the largest numbers of juvenile crabs were found in poly- and mesohaline SAV and salt marsh habitats. In Year 1, no significant correlations were observed between either marsh stem density or SAV vegetation biomass and YOY crab abundance, although at one location in the second year there was a significant positive correlation between the number of juveniles and mid-bay stem density. No significant differences in crab size (carapace width, CW) were found among locations in either year. In comparison with other, better-studied mid-Atlantic coast estuaries in the US (e.g., the Chesapeake and Delaware bays), very large numbers of megalopae (frequently 1 to 2 orders of magnitude greater) invade the Mobile Bay system, as well as other Gulf Coast estuaries. However, YOY abundance in marsh and SAV 'nursery habitats' was of the same order of magnitude as that reported for mid-Atlantic estuaries. We conclude that although postlarval supply was very large, post-settlement loss to predators was the dominant factor influencing blue crab population dynamics in our north central Gulf of Mexico study area.