Larval supply and juvenile recruitment of coral reef fishes to marine reserves and non-reserves of the upper Florida Keys,USA

For marine organisms, decoupling between the planktonic larval stage and the benthic-associated juvenile stage can lead to variable patterns of population replenishment, which have the potential to influence the effectiveness of marine reserves. We measured spatial and temporal variability in larval supply and recruitment of fishes to coral reefs of different protection levels and tested whether protection level influenced the relationship between supply and recruitment. We sampled pre-settlement larvae and newly settled recruits from four reefs (two reserves and two non-reserves) in the Florida Keys National Marine Sanctuary, USA. Replicate point measures of larval supply over 14 months and 17 monthly measurements of recruitment varied significantly among months and sites. Sites with the same protection level had significantly different patterns of larval supply as well as larval and recruit diversity, but recruitment magnitude differed only by protection level, where densities were greater at reserves. Differences in larval supply among sites included two particularly large peaks in larval abundance at one site, possibly associated with the observed passage of small-scale oceanographic features. To examine whether relationships between larval supply and recruitment varied by protection level, we selected one species that was present in both the light trap samples and the monthly recruitment surveys. Recruitment of the bicolor damselfish Stegastes partitus was significantly and positively related to larval supply at three of the four sites thus, protection level did not influence this linkage. Since local variability among sites can lead to spatial differences in population replenishment, characterization of larval supply and recruitment to potential marine reserve sites may help to identify optimal locations in a region and contribute to more effective reserve design.     

Associations between the abundance of piscivorous fishes and their prey on coral reefs: implications for prey-fish mortality

Few studies have examined predator-prey relationships in diverse communities such as those found on coral reefs. Here we examined patterns of spatial and temporal association between the local abundance of predator and prey fishes at Lizard Island on the Great Barrier Reef, Australia. We predicted that the nature of this association would have implications for patterns of prey-fish mortality. Strong positive relationships between prey and piscivore abundance were found throughout the study. Greater densities of predators and of prey were found on patch-reef habitats, compared with contiguous reef-slope habitats. Declines in prey-fish abundance on patch reefs were density-dependent and correlated with the densities of predators. The relative roles of recruitment and piscivore movement in determining patterns of predator and prey abundance were assessed from surveys of recruit densities and an intensive programme of tagging two species of rock-cod, Cephalopholis cyanostigma and C. boenak (Serranidae), over 2 years. Patterns of recruitment explained little of the variation in the abundance and distribution of piscivorous fish. If movement explains large-scale patterns of distribution, this was not evident from the tagging study. The two rock-cod species were highly sedentary, with individuals on patch reefs seldom moving among reefs. Individuals on reef slopes were also highly site-attached, although they moved greater distances than those on patch reefs. Although the mechanisms responsible remain to be determined, this study demonstrated strong associations between the abundance of piscivorous fish and their prey on coral reefs. This relationship appeared to be an important factor in producing density-dependent declines in the abundance of prey. Received: 30 April 2000 / Accepted: 22 September 2000     

Density dependence vs. independence, and irregular population dynamics of a swallow-wort fruit fly

Although most long-term studies of consumer-resource (e.g., predator-prey) interactions select species showing cyclic population dynamics, strong consumer-resource interactions can also produce irregular, noncyclic dynamics. Here, we present a case in which a seed predator, the tephritid fruit fly Euphranta connexa, shows fluctuations in density of more than two orders of magnitude over a 22-year period. To explain these fluctuations, we analyzed a stage-specific data set to quantify the density-dependent and density-independent components of larval survivorship and realized fecundity. Both larval survivorship and realized fecundity were strongly density dependent. Larval survivorship dropped from 0.62 at low larval density to 0.081 at high larval density, whereas fecundity dropped from 84.3 to 0.32 eggs per individual, more than a 100-fold decrease. We divided density-independent variation in E. connexa population dynamics into components for variability in (1) larval survivorship, (2) realized fecundity, and (3) annual fruit abundance. Of these components, 96% of the density-independent variance in per capita population growth rates was caused by fluctuations in fruit abundance. This highlights the importance of the strong consumer-resource interactions in driving fluctuations in E. connexa abundance. It also demonstrates that E. connexa dynamics are remarkably simple, and aside from the 4% of unexplained variance in per capita population growth rates, our understanding of E. connexa dynamics is remarkably complete.     

Widespread density-dependent seedling mortality promotes species coexistence in a highly diverse Amazonian rain forest

Negative density-dependent mortality can promote species coexistence through a spacing mechanism that prevents species from becoming too locally abundant. Negative density-dependent seedling mortality can be caused by interactions among seedlings or between seedlings and neighboring adults if the density of neighbors affects the strength of competition or facilitates the attack of natural enemies. We investigated the effects of seedling and adult neighborhoods on the survival of newly recruited seedlings for multiple cohorts of known age from 163 species in Yasuni National Park, Ecuador, an ever-wet, hyper-diverse lowland Amazonian rain forest. At local scales, we found a strong negative impact on first-year survival of conspecific seedling densities and adult abundance in multiple neighborhood sizes and a beneficial effect of a local tree neighborhood that is distantly related to the focal seedling. Once seedlings have survived their first year, they also benefit from a more phylogenetically dispersed seedling neighborhood. Across species, we did not find evidence that rare species have an advantage relative to more common species, or a community compensatory trend. These results suggest that the local biotic neighborhood is a strong influence on early seedling survival for species that range widely in their abundance and life history. These patterns in seedling survival demonstrate the role of density-dependent seedling dynamics in promoting and maintaining diversity in understory seedling assemblages. The assemblage-wide impacts of species abundance distributions may multiply with repeated cycles of recruitment and density-dependent seedling mortality and impact forest diversity or the abundance of individual species over longer time scales.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Early life histories in the bryozoan<Emphasis Type="Italic"> Schizobrachiella sanguinea</Emphasis>: a case study

Mediterranean populations of Schizobrachiella sanguinea seem to span a wide range of contrasting benthic habitats. We have studied adult, larval, and recruit populations of S. sanguinea from sublittoral (approximately 10 m depth) communities at Blanes (Spain, NW Mediterranean). At the study site, the species colonises semi-obscure caves and, to a lesser extent, communities of sciaphilic algae. Our first goal was to show the periods of brooding, larval release, and recruitment. Plankton was collected about twice a week between July 1998 and June 2000 over an artificial reef. A total of 303 larvae were collected in 23 of the 102 days of sampling. At the study site the species shows a single, annual larval release period, which occurs from late March to June–July every year. Recruits were found from late May to late August 1999 and in late May 2000. A recruitment peak was observed in July 1999 and again in late May 2000. The presence of brooding adults corresponded to the periods of larval supply and recruitment. Our second goal was to investigate the causes of the variable distribution of the species along several communities in the north-west Mediterranean Sea. We describe adult distribution and the dynamics of larval supply, recruitment of early (15 days from settlement) and late (4 months from settlement) recruits. Patterns of larval presence and early recruit distribution along the communities on the reef did not reflect the adult distributions, whereas this was closely matched by the distribution of late recruits. Post-recruitment mortality strongly affected recruits of the PA (photophilic algae) and SA (sciaphilic algae) communities, whereas it was very low in the communities of SOC (semi-obscure caves). Mortality of recruits was density independent and colony survivorship in the SA communities, where recruitment was the highest, appeared strongly affected by competition with brown algae. Like the majority of cheilostomate bryozoans, S. sanguinea releases a coronate larva capable of active swimming and positive reaction to light. Such swimming abilities may allow larvae to actively select the substrate on which to settle and to avoid unsuitable substrates. It seems that larval behaviour does not explain the preference of S. sanguinea for sciaphilic habitats. High post-recruitment mortality due to various factors, especially competition with fast-growing algae, seems particularly relevant in determining the adult distributions observed.Communicated by S.A. Poulet, Roscoff     

Heat waves,baby booms,and the destruction of kelp beds by sea urchins

Large populations of sea urchins, Strongylocentrotus droebachiensis (Müller), destroyed kelp beds along the Atlantic coast of Nova Scotia in the 1960's and 1970's. The origin of these large sea urchin populations is not understood. We have investigated the potential influence of variable growth and development of the planktonic larvae of sea urchins (in response to temperature and food abundance) on recruitment of benthic juveniles. The adult sea urchins were collected at Sandy Cove, Digby County, Nova Scotia, Canada, in December 1986. Temperature strongly affected larval size and the growth of the echinus rudiment within the range 3° to 9°C, and larvae grew most rapidly at 14°C. Food abundance had a smaller effect on larval growth, and these effects were apparent only at high temperature. Larvae fed the same concentration of two different algal food species grew and developed similarly. Correspondence between spring temperature variation and qualitative variation in sea urchin recruitment, as well as strong temperature effects on larval growth in culture, and the occurrence of a large, positive temperature anomaly in June 1960, all suggest that temperature effects on larval growth and development may have led to intense sea urchin recruitment in 1960 and the appearance of large adult populations 4 to 6 yr later. This result invites further research.     

Habitat modification affects recruitment of abalone in central New Zealand

The habitat experienced during early life-history stages can determine the number and quality of individuals that recruit to adult populations. In a field experiment, biogenic habitat complexity was manipulated (presence or absence of foliose macroalgae) at two depths (2–3 m and 5–6 m) and the habitat-dependent effects on recruitment of the black foot abalone (Haliotis iris) were examined at three field sites along the south coast of Wellington, New Zealand (41°20′S, 174°47′E), between July and November 2005. Recruit density (<5 weeks post-settlement) was measured on cobbles covered with crustose coralline algae. Habitats of low complexity (barrens treatments) had consistently greater densities of recruits than habitats of high complexity (algae treatments). However, recruits in algae habitats were larger, and for deep habitats, there was greater survival in algae habitats compared with barrens habitats. While depth had no significant effect on early recruit (<2 weeks post-settlement) density, late recruit (<5 weeks post-settlement) density was greater in shallow habitats, and so it seems recruit survival was greater in shallow habitats. In this experiment, algal habitat complexity had strong effects on early recruit abundance, but habitat-dependent variations in recruit growth and survival may modify initial patterns of abundance and determine recruitment to adult abalone populations.     

Spatiotemporal variability in recruitment around Iriomote Island, Ryukyu Archipelago, Japan: implications for dispersal of spawning corals

Spatiotemporal recruitment patterns of scleractinian corals were investigated around Iriomote Island, Ryukyu Archipelago, Japan, in relation to adult coral cover in 2005 and 2006. Although almost all corals were broadcasting spawners, the relationship between recruitment and adult coral cover differed among coral families (Acroporidae, Poritidae, and Pocilloporidae), likely due to differences in embryonic development time. For spawning pocilloporid corals, whose larvae develop relatively more rapidly, recruitment was higher at sites where adult coral cover was higher. In contrast, recruitment was not related to adult coral cover in acroporid and poritid corals, whose embryonic development times were relatively slow. Moreover, recruitment of acroporid corals varied between years, and recruitment was greater at leeward compared to windward reefs for a few days after spawning. These results suggest that embryonic development time and wind-driven surface currents affect larval dispersal and subsequent recruitment patterns at a local scale. Based on embryonic development time, some spawning corals are more likely to have higher rates of self-seeding than others. Our results predict that among spawning corals, local populations of acroporid and poritid corals, whose larvae potentially disperse over long distances and recruit in neighboring reefs, are more resilient to local disturbances than those of pocilloporid corals, whose recruitment relies upon local stock.     

Some biological interactions affecting intertidal populations of the kelp Egregia laevigata

Near Santa Barbara, California (USA), the large laminarian kelp Egregia laevigata (Setchell) occurred from the lower intertidal zone to subtidal depths. In the intertidal zone there was a large recruitment of E. laevigata in the spring. The kelp were largely excluded from a zone in the lowest intertidal zone where the surf grass Phyllospadix sp, grew, but were abundant on all other rock surfaces. Experiments demonstrated that, while neither grazers nor sessile organisms significantly reduced recruitment, E. laevigata of the previous year-class did so. There were also interactions among E. laevigata of the same year-class, expressed as a density-dependent mortality of very small algae, and as faster growth rates and greater number of branches of kelp at low densities. These processes tended to make both numbers and biomass of E. laevigata uniform. The mortality rates of large E. laevigata were so high that, in some locations, no kelp survived for more than 8 months. E. laevigata was, therefore, essentially an annual and opportunistic species in the intertidal zone.     

Positive indirect effects of reef fishes on kelp performance: the importance of mesograzers

It has been suggested that microcarnivorous reef fishes may play an important role in giant kelp forest communities by preventing infestations of mesograzers that could severely impact or potentially destroy recovering kelp forests after extreme disturbance events. However, these trophic linkages, specifically the direct and indirect effects of fishes on the biomass of mesograzers, grazing intensity, and the performance of giant kelp, have not been sufficiently quantified and evaluated as to their importance and in the absence of such disturbance events. We examined experimentally the effects of mesograzers on the growth and performance of giant kelp in the presence and absence of their fish predators near Santa Catalina Island, California (U.S.A.). Mesograzer biomass and grazing intensity were significantly higher when fishes were excluded from giant kelp, which in turn, lowered kelp performance. This pattern was consistent both on experimental plots of kelp as habitat isolates, and on a continuous reef. Moreover, the abundance of mesograzers was inversely related to the abundance of kelp perch among several kelp-forested reefs, suggesting that these effects can occur at larger spatial scales. Because of differences in the diet and behavior of two microcarnivorous fishes, the kelp perch and se?orita, we conducted an experiment manipulating each species and its density independently to determine their separate effects on mesograzers and kelp performance. Concurrently we examined the growth and mortality of juvenile kelp. Grazing intensity decreased, estimates of kelp performance increased, and the growth of juvenile kelp increased with increasing densities of fish but with no detectable effects between fishes. Our results demonstrate that these microcarnivorous fishes have positive indirect effects on kelp performance by reducing mesograzer biomass and grazing intensity, and the early life stages of other fishes also may be important. More specifically, these fishes have a positive effect on the density of fronds of giant kelp that can result in greater recruitment success and the abundance of kelp-associated invertebrates and fishes. Indeed, this study suggests that mesograzers have the potential to be one of the most important herbivores in kelp forest ecosystems.     

Density-dependent settlement and mortality structure the earliest life phases of a coral population

The local densities of heterospecifics and conspecifics are known to have profound effects on the dynamics of many benthic species, including rates of settlement and early post-settlement survivorship. We described the early life history of the Caribbean coral, Siderastrea radians by tracking the population dynamics from recently settled planulae to juveniles. Through three years of observation, settlement correlated with the abundance of other benthic organisms, principally turf algae (negatively) and crustose coralline algae (positively). In addition, adult density showed independent effects on coral settlement and early post-settlement survivorship. Settlement rates increased across low levels of adult cover and saturated at a maximum around 10% cover. Early post-settlement survivorship decreased with adult cover, revealing structuring density dependence in coral settlers. The earliest life stages of corals are defined by low survivorship, with survivorship increasing appreciably with colony size. However, recent settlers (one-polyp individuals, < 1-year-old) are more likely to grow into two-polyp juveniles than older single polyps (> 1-year-old) that were delayed in their development. The early benthic phase of corals is defined by a severe demographic bottleneck for S. radians, with appreciable density-dependent and density-independent effects on survivorship. For effective management and restoration of globally imperiled coral reefs, we must focus more attention on this little studied, but dynamic, early life history period of corals.     

Diel and tidal cycles regulate larval dynamics in salt marshes and mangrove forests

Variation in the release and recruitment of larvae of estuarine invertebrates affects the distribution and abundance of adults, as well as trophic interactions in both the plankton and the benthos. Larval release and supply are often timed to environmental cycles such as the diel and tidal cycles. Here, we determined using plankton tows whether the abundance of larvae spanning salt marsh and mangrove habitats across the intertidal landscape varied with diel and tidal cycles. Using three different sampling designs across two sites and within each of two estuaries over a 12-month period, we covered a range of spatial and temporal scales. This allowed us to test the general prediction that densities of meroplankton in the water column would be greater during nocturnal ebb tides than during other phases of the diel or tidal cycle. As predicted, nocturnal ebb tides yielded the highest densities of meroplanktonic larvae and were dominated by first-stage crab zoeae and this finding was most pronounced in the salt marsh. Throughout the course of the year, greater numbers of meroplankters consistently occurred during the ebb tide compared with the flood tide. The densities of other taxa (e.g. gastropods and polychaetes) showed no clear trends with diel or tidal cycles. This study highlights the effects of these pervasive physical cycles on the timing of larval release and supply in the salt marsh–mangrove complex, and emphasises their contribution to the trophic interactions and the dynamics of benthic populations within estuaries.     

Temporal patterns of settlement, recruitment and post-settlement losses in a rocky reef fish assemblage in the South-Western Mediterranean Sea

Post-settlement events can significantly alter the density distribution of settlers and subsequently the adult population structure. The temporal and inter-annual variability of settlement and the effects of mortality on recruitment were investigated across 2 years in the vicinities of Cabo de Palos–Islas Hormigas Marine Reserve by visual census and light trap sampling. Settlement was seasonal with greater species richness and abundance in summer. Although temporal synchronization was observed between larval supply and settlement, densities of settlers could not be predicted from post-larval abundances. Timing of settlement was consistent between years but with high inter-annual variation in abundance. High mortality (~80 %) and general decoupling between post-larval and settlement stages suggest that early mortality is driving such patterns. Nevertheless, indications of habitat-mediated mortality were found for benthic species such as Symphodus and Diplodus, highlighting the importance of habitat in shaping population demography.     

Competitive organization of the soft-bottom macrobenthic communities of southern California lagoons

The sandy-bottom macrobenthic community of Mugu Lagoon, a relatively pristine southern California (USA) marine lagoon, demonstrated (1) nearly constant community composition over 37 months of observation, (2) relatively little temporal variability in the population densities of the most abundant species over 37 months, and (3) a pattern of depth stratification in which very little vertical overlap existed among the six most abundant species. The only two species whose vertical distributions overlapped broadly showed horizontal spatial segregation, each abundant in different areas within the sand habitat. These community characteristics imply the importance of biological factors in structuring the sand benthos. The relatively large volume required for living space by these macrofauna suggests that competition for space may be the biological factor most important in determining the observed temporal and spatial abundance patterns. The muddy-sand community and the mud community of Mugu Lagoon also revealed similar patterns of stratification: new abundant species replaced species at the same sedimentary level while not greatly affecting species populations at other non-overlapping levels. In the sand community of Tijuana Slough, two of the abundant species of Mugu Lagoon's sand community were nearly absent as an apparent result of human over-exploitation. Probably in response, densities of species living at the sedimentary levels normally occupied by the missing species were much higher than would be predicted if competition for space were unimportant. In field experiments, removal of the deposit feeder Callianassa californiensis resulted in high recruitment of Sanguinolaria nuttallii, whereas control areas showed no S. nuttallii recruitment. Experiments also suggest that negative intraspecific interactions between Cryptomya californica individuals may explain the observed rapid emigration from areas of artificially high density. Perhaps the relatively great environmental predictability of southern California lagoons has permitted competitive interactions to play a singnificant role in determining the temporal and spatial abundance patterns of the soft-bottom macrobenthos.     

Predation,recruitment and the dynamics of communities of coral-reef fishes

I investigated the ability of predators to influence the patterns of species richness and abundance of non-piscivorous fishes on small, artificial reefs replenished by natural recruitment. Periodic removal of predators effectively reduced the species richness and abundance of predators on removal reefs. The difference between the number of predators on control and removal reefs was greatest immediately following the removal of predators and attenuated between removals. During periods of recruitment, species richness and total abundance of recently-recruited, non-piscivorous fishes were generally greater on predator-removal reefs than on control reefs. Species richness and total abundance of resident non-piscivorous fishes were not affected by the removal of predators in the first year of the experiment. Both abundance and species richness of residents, however, were greater on the removal reefs during the second year of the experiment. The difference in the responses of the two age classes to the removal of predators suggests that predators may affect community patterns of older age classes through time-lagged effects on the survivorship of younger age classes. At the end of the experiment, species richness was positively related to abundance for recruits and residents. The effects of removing piscivorous fishes on the abundance of non-piscivorous fishes were similar for species considered separately. A greater number of species of recruit and resident fishes were more abundant on reefs from which predators had been removed. These data suggest that predators can play an important role in structuring communities of fishes on coral reefs.     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Oviposition,larval abundance,in situ larval growth and recruitment of the herbivorous gastropodLacuna vincta in kelp canopies in Barkley Sound,Vancouver Island (British Columbia)

We investigated recruitment of the herbivorous gastropodLacuna vincta (Montagu, 1803) in the canopies ofMacrocystis integrifolia andNereocystis luetkeana beds in Barkley Sound, Vancouver Island (British Colombia), from 1987 to 1989. Four factors influencing intensity and patterns of recruitment were studied: (1) seasonality of oviposition, (2) larval abundance, (3) growth of larvae in the field and (4) larval settlement. Egg masses were abundant on low intertidal algae but were scarce in kelp canopies. Although egg masses could be found almost year-round, a distinct and intense period of oviposition occurred during winter and spring. Intracapsular development lasted 2.5 to 3.5 wk before planktotrophic veligers emerged. The duration of the planktonic period, 7 to 9 wk, was determined through an in situ study of cohorts ofLacuna spp. larvae present in the plankton between January and June 1988. The general timing of the onset of the spring peak recruitment period was predicted from these cohorts. Primary periods of recruitment ofL. vincta in the canopy occurred in April–May (average density up to 383.9 juveniles m^–2 blades), with a second period of lower intensity in the late summer—fall period. We observed similar trends between abundance of advanced larvae (> 500µm) in the plankton and recruitment rates in kelp canopies. Although adults were occasionally observed in the canopy, newly metamorphosed juveniles consistently dominated the habitat. The persistance of small juveniles (0.7 to 1.5 mm), rapid declines in density shortly after recruitment, and SCUBA observations of drifting individuals suggest that juveniles migrate to the under-canopy or low intertidal area after a brief period of growth on kelp blades.     

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Genetic heterogeneity among adult and recruit red sea urchins, Strongylocentrotus franciscanus

Allozyme electrophoresis was used to characterize genetic variation within and among natural populations of the red sea urchin Strongylocentrotus franciscanus. In 1995 to 1996, adult urchins were sampled from twelve geographically separated populations, seven from northern California and five from southern California (including Santa Rosa Island). Significant population heterogeneity in allelic frequencies was observed at five of six polymorphic loci. No geographic pattern of differentiation was evident; neighboring populations were often more genetically differentiated than distant populations. Northern and southern populations were not consistently distinguishable at any of the six loci. In order to assess within-population genetic variation and patterns of recruitment, large samples were collected from several northern California populations in 1996 and 1997, and were divided into three size classes, roughly representing large adults (>60 mm), medium-sized individuals (31 to 60 mm, “subadults”) and individuals <2 yr of age (≤30 mm test diam, referred to as “recruits”). Comparisons of allelic counts revealed significant spatial and temporal differentiation among size-stratified population samples. Recruit samples differed significantly from adult samples collected at the same locale, and showed extensive between-year variation. Genetic differentiation among recruit samples was much higher in 1997 than in 1996. Between-year differences within populations were always greater for recruits than for adults. Potential explanations for the differentiation of recruit samples include pre- and post-settlement natural selection and high interfamily variance in reproductive success or “sweepstakes” recruitment. Unless recruit differentiation can be attributed to an improbable combination of strong and spatially diverse selection, such differentiation across northern California populations indicates that the larval pool is not well mixed geographically (even on spatial scales <20 km), despite long planktonic larval duration. Received: 6 July 1999 / Accepted: 25 January 2000