Interactions among sea otters, sea stars, and suspension-feeding invertebrates in the western Aleutian archipelago

The structuring and organizing effects of apex predators on ecosystems are becoming increasingly well documented. The enhancement of kelp forests via sea otter predation on herbivorous sea urchins is among the earliest and best known examples. This study provides evidence for direct and indirect trophic interactions among sea otters, predatory sea stars, and filter-feeding mussels (Mytilus trossulus) and barnacles (Semibalanus cariosis). In western Massacre Bay at Attu Island (173°E, 53°N), subtidal transects showed sea star body size and biomass density declined markedly between 1983 and 1994 as sea otters reinhabited this area. Mussels and barnacles translocated from the rocky intertidal zone to shallow subtidal habitats to assess loss rates from sea star predation showed lower mortality rates after the arrival of sea otters. Prey mortality rates in subtidal caged controls were consistently low and similar to those of intertidal controls in both years. These findings elucidate a trophic pathway by which sea otters can influence ecosystems separate from the well-known sea otter/sea urchin/macroalgae cascade.     

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.     

Demography of the intertidal brown alga Pelvetia fastigiata in Southern California,USA

The demography of Pelvetia fastigiata was studied from 1973 through 1977 in the intertidal zone near La Jolla, California, USA. Populations were usually small and relatively stable as new recruits replaced older individuals that were lost; however, in 1977 high recruitment substantially increased total abundances. The survivorship of recruits was similar regardless of the season or year of recruitment. Mortality rates decreased with age, and the mean life expectancy continued to increase for at least 1 yr. Losses of juveniles were high; 50% lived 80 d and only 9% survived to the age of first reproduction (about 1.5 yr). Plants present at the start of the study had higher survivorship rates than new recruits, 19 and 7%, respectively, living more than 3 yr. Individuals within aggregations of large plants lived longer than most dispersed individuals, and aggregations remained intact throughout the study in spite of low recruitment. Established individuals had low mortality rates and populations probably relied on these persistent individuals for reproduction.     

Changes in sea urchin populations after the destruction of kelp beds

In St. Margaret's Bay, Nova Scotia, Canada, there are large areas in which sea urchins (Strongylocentrotus droebachiensis) have eliminated beds of kelp (Laminaria spp.). Sites were identified where destruction of kelp beds had taken place 1, 2, 3, 3.5 and 4 years ago. With increase of time since kelp disappearance, the sea urchins showed decreased growth rate, reduced gonad size, but an increase in numbers resulting from high recruitment rates in the first two years after kelp bed destruction. These sea urchin populations, by their browsing, effectively prevent the regeneration of kelp. There is, as yet, no evidence of the sea urchin populations being starved out to allow the kelp to return.     

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.     

A general bioeconomic simulation model for annual-crop marine fisheries

A generalized bioeconomic simulation model of annual-crop marine fisheries is described and its use in marine fisheries management is demonstrated. The biological submodel represents the recruitment of new organisms into the fishery, the movement of organisms from one fishing area to another and from one depth to another, the growth of organisms and the mortality of organisms resulting both from natural causes and from fishing. The economic submodel represents the fishing effort exerted on each resource species, the monetary costs of fishing, the value of the harvest and the rent (or excess profits) to the fishery.Basic dynamics of the model results from changes in the number of organisms in the fishery over time, which can be summarized as a set of difference equations of the general form ΔN/Δt = R + I ? E ? M ? F where ΔN/Δt is the net change in number of organisms in the fishery over time, R is recruitment, I is immigration, E is emigration, M is natural mortality and F is fishing mortality. R is a driving variable, whereas I, E, M and F are functions of the state of the system at any given point in time. The model can be run in a deterministic or stochastic mode. Values for parameters affecting rates of recruitment, movement, growth, natural mortality and fishing mortality can be selected from uniform, triangular or normal distributions.Use of the model within a fisheries-management framework is demonstrated by evaluating several management alternatives for the pink shrimp (Penaeus duorarum) fishery on the Tortugas grounds in the Gulf of Mexico. Steps involved in use of the model, including parameterization, validation, sensitivity analysis and stochastic simulations of management policies, are explained.     

Population dynamics of the sea urchin Strongylocentrotus purpuratus in a Central California kelp forest: recruitment,mortality, growth,and diet

The population dynamics of Strongylocentrotus purpuratus inhabiting dense mats of geniculate coralline algae in a shallow central California Macrocystis pyrifera forest was examined. Sea otters had occupied the area for over two decades. Sea urchin density and size distribution were sampled during 1984–1986 to determine recruitment and mortality patterns. Growth rates were obtained from tetracycline-labeled individuals and changes in sizefrequency modes. Periodic collections were made to examine gut contents and the relationship of diet to food availability. The interaction of recruitment and mortality resulted in a dynamic population structure. Mean densities ranged from 6.5 to 12.7 urchins 0.25 m^-2. The population consisted primarily of urchins up to 40 mm in test diameter in a size distribution which changed from unimodal to bimodal and back over the 2 yr study. Mortality was temporally variable and related to test diameter. Growth rates were somewhat lower than most previous reports for the species and suggested that most of the population was made up of 1 and 2 yr-old individuals. Diet consisted largely of geniculate coralline algae, with fleshy brown algae becoming important when available as drift. This study showed high densities of small urchins can exist in a California kelp forest inhabited by sea otters, but regular recruitment may be necessary to maintain such populations.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Temporal variation in community interfaces: kelp-bed boundary dynamics adjacent to persistent urchin barrens

We examined, over 2 years, factors affecting the temporal stability of the lower limit of kelp beds (Alaria esculenta) at five subtidal sites in the Mingan Islands, northern Gulf of St. Lawrence. The position of the lower limit of the beds varied markedly among sites and over time and was largely controlled by the green sea urchin, Strongylocentrotus droebachiensis, which formed dense (up to 500 individuals m^–2) feeding fronts at the lower edge of the beds. These aggregations advanced over the kelp most rapidly during the summer (at rates as high as 2.5 m month^–1), and there appeared to be a threshold urchin biomass of ~5 kg m^–2 below which the fronts could not substantially reduce the limit of the beds. The fronts consisted mainly of large individuals, whereas smaller urchins predominated in the barrens zone below the kelp beds. At one site, we recorded large seasonal shifts in overall urchin densities, with large increases and decreases during the summer and winter, respectively. An urchin exclusion experiment indicated that algal recruitment in the barrens was two orders of magnitude greater in the absence than in the presence of urchins. The kelp Agarum cribrosum greatly restricted urchin movements, and the greater temporal stability of the kelp bed at one site appeared related to the gradual replacement of Alaria esculenta in the lower kelp bed by a large stand of Agarum cribrosum. We propose that perturbations by abiotic factors (e.g., ice scouring and water motion) trigger important but localized changes in urchin densities that, in turn, largely determine the limits of kelp bed distribution in this region of the Atlantic where urchin barrens are a persistent community state.Communicated by R.J. Thompson, St. Johns     

A natural history study of Encope grandis and Mellita grantii,two sand dollars in the northern gulf of California,Mexico

Populations of the sand dollars Encope grandis and Mellita grantii were studied at Playa Hermosa in the northern Gulf of California, Mexico. Intertidal distribution and abundance were estimated for the two species from 1969 to 1972. They occurred in the lower intertidal (-0.4 to -1.3 m) and on into the subtidal. Density of E. grandis varied from a high of 380/m² in the spring of 1970 to a low of 0.1/m² in the fall of 1971. Density of M. grantii ranged from a high of 56/m² in 1970 to a low of 0.8/m² in 1971. Growth curves were constructed based on shifts in the modes of the size distributions, and rates of mortality were estimated from the growth parameters and size distributions. It is estimated that E. grandis would attain 95% of its maximum size (74 mm) in about 6 years with an annual mortality rate of 18%. M. grantii would reach 95% of its maximum size (38 mm) in about 5 years with an annual mortality rate of 58%. The populations of these sand dollars appear to be limited by the physical environment.     

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

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

Foliose algal assemblages and deforested barren areas: phlorotannin content,sea urchin grazing and holdfast community structure in the Aleutian dragon kelp,Eualaria fistulosa

In the Aleutian Archipelago, two distinct organizational states of kelp forest communities exist, foliose algal assemblages and deforested barren areas. The canopy-forming kelp Eualaria fistulosa can be found in both states, although it is much less abundant in the deforested state. In contrast, sea urchins also occur in both states, but they are considerably more abundant in the deforested state. This study determined whether the protective phlorotannin content in E. fistulosa sporophylls that originated in foliose algal assemblages would differ from those that originated in deforested areas. We also examined sea urchin grazing rates on these sporophylls to determine whether there were any differences in grazing rates related to the origin of the sporophylls. Next, we determined whether taxon richness and abundance of E. fistulosa holdfast communities would differ depending on the origin of the holdfast. Our results showed that sporophylls that were collected in barren areas generally had higher phlorotannin content than sporophylls found in foliose algal assemblages, although phlorotannin content varied across the study area. Grazing rates on sporophyll tissue and holdfast community structure did not differ between foliose algal assemblages and barren areas. These results show that phlorotannin content is greatest in areas under high grazing pressure and that phlorotannins may possibly protect E. fistulosa and the holdfast communities from being grazed in barren areas.     

The importance of sandflat morphology to recruitment of the intertidal snail Nassarius pauperatus during ten consecutive years at three sites in South Australia

This project was desinged to confirm the temporal consistency of inter-site differences in recruitment of an intertidal gastropod, and to test the hypothesis that these differences are accounted for by differences in shore topography. Annual recruitment of the intertidal sanil Nassarius pauperatus (Lamarck) was estimated at two sandflats in Gulf St Vincent, South Australia, over a period of 12 yr (1979 to 1990). At one sandflat the gradient was slight, the zone containing N. pauperatus was wide, and there were extensive pools of water at low tide, whilst at the other sandflat the gradient was steeper, the zone containing N. pauperatus was narrower, and there were fewer or no pools. Although recruitment at both sites varied temporally, the density of recruits was always higher at the first site. These data, together with data covering 10 yr (1981–1990) from a sandflat where the gradient of the substratum was decreasing due to accretion of sediment, suggest that shore topography is an important determinant of relative recruitment for N. pauperatus in South Australia.     

Genetic differentiation in the intertidal zone in populations of the alga Enteromorpha linza (Ulvales: Chlorophyta)

Genetic differentiation within the intertidal zone was examined in six populations of the asexually reproducing alga Enteromorpha linza growing in the Long Island Sound, USA. Four of the five populations sampled in 1981 showed significant differentiation between high and low intertidal positions with respect to the GOT-2 locus. The pattern of differentiation was consistent for samples collected at several times during the year with some seasonal modifications. Four additional polymorphic loci, resolved in 1982, identified a total of 13 five-locus genotypes or clones. Four of the six populations sampled in 1982 showed significant differences in clone frequency between high and low intertidal positions. Laboratory experiments revealed differences in response to temperature among the clones. At 24°C a high intertidal associated clone showed an increase in growth, while low intertidal associated clones showed decreased growth compared to growth at 15°C. These results suggest that the microgeographic differentiation observed for E. linza in the intertidal zone may in part be due to the differential adaptation of clones to different intertidal environments. Additional demographic information is needed for individual clones in order to determine the role longevity, reproductive output, recruitment and interclonal competition play in maintaining the observed differentiation.     

Implications of movement behavior on mussel dislodgement: exogenous selection in a<Emphasis Type="Italic"> Mytilus</Emphasis> spp. hybrid zone

Rocky intertidal habitats often exhibit high levels of environmental heterogeneity, and the ability of organisms to move between microhabitats is likely to have a profound influence on their rates of mortality and overall fitness. Mussels within the Mytilus edulis complex are morphologically very similar, yet at sites where these species hybridize in southwest England, populations repeatedly show evidence of selection against individuals with alleles specific to M. edulis Linnaeus, in favor of those with alleles specific to M. galloprovincialis Lamarck. Differential movement rates of these two species were examined within simulated mussel beds (gravel substrate) in the winter (February) and summer (July) of 2001. M. edulis-like mussels moved more frequently and more quickly to the exterior of gravel beds than did M. galloprovincialis-like mussels. Coupled with measurements of attachment strength in the field conducted in July 2001, we used a wave force model to examine the probability of dislodgement for each species under a range of water velocities. Results suggest that by preferentially moving to the exterior of beds, M. edulis experiences higher dislodgement rates due to exposure to large hydrodynamic forces than do M. galloprovincialis. As a consequence of lower attachment strengths, M. edulis is also predicted to have higher mortality rates than M. galloprovincialis in interior portions of the bed. Thus, differential movement behavior may contribute to the differential genotype-specific mortality rates observed in the Mytilus spp. hybrid zone in southwest England, and is an example of behavior potentially modifying rates of exogenous selection in an intertidal hybrid zone.Communicated by J.P. Grassle, New Brunswick     

Population dynamics of the temperate Australian soft coral Capnella gaboensis

Populations of over 100 colonies of Capnella gaboensis Verseveldt, 1977 at each of four study sites in Sydney Harbour were monitored for recruitment, survival, and mortality from October 1982 to November 1984. Larval settlement and survival were observed in the laboratory. Successful in situ larval settlement of C. gaboensis was estimated to be in the order of 0.26%. The predominant mode of recruitment was from larvae. Asexual recruitment resulted only from the fission of large colonies, and not by fragmentation or by stolon development. Mortality of newly settled C. gaboensis in situ during their first year was estimated to be around 99.75%. For colonies more than 1 yr old, mortality was highest amongst juveniles. After colonies reach about 3 yr of age, they have a high expectation of further life, until they reach about 15 yr of age; life expectancy declines each year thereafter. These estimates of life expectancy are based upon the conservative assumption that the largest colonies are about 20 yr old. The likely causes of mortality for colonies in the field are: predation by urchins, ovulids and nudibranchs; abrasion by kelp; and overgrowth by algae and sponges. The causes of mortality for larvae and recruits in the field, however, can be predicted only on the basis of laboratory studies and observations of field conditions.     

The invasive gastropod Crepidula fornicata: reproduction and recruitment in the intertidal at its northernmost range in Wales, UK, and implications for its secondary spread

The establishment and spread of a non-native species in an introduced range depends to a large extent on the performance of the species under the prevailing environmental conditions. The spawning, larval and spatfall periods of the invasive gastropod Crepidula fornicata were monitored in the intertidal zone at its northernmost range in Wales, UK, between February 2010 and January 2011. The duration of the reproductive season was similar to that recorded from more southerly European populations. Spawning and larval release occurred throughout most of the year even at low seawater temperatures of <7 °C, but benthic recruitment was observed over a much shorter period at seawater temperatures >16 °C. Recruitment was low and likely controlled by post-settlement mortality. These observations suggest that C. fornicata’s northwards spread in Welsh waters will not be limited by seawater temperature negatively affecting reproduction, but by processes acting after larval release. These data show the importance of incorporating settlement and post-settlement processes into studies on recruitment success when aiming to predict the potential spread of a potentially harmful invader such as C. fornicata.     

Settlement and recruitment of sea urchins (Strongylocentrotus spp.) in a sea-urchin barren ground and a kelp bed: are populations regulated by settlement or post-settlement processes?

I sampled recruitment of very small sea urchins (Strongylocentrotus spp.) by using the anesthetic magnesium chloride to remove individuals from substrata collected in sea-urchin barren grounds (barrens) and kelp beds at Naples Reef near Santa Barbara, California, USA. Preliminary sampling found low numbers of newly settled individuals(<0.6 mm test diam) from April–July in 1984 and 1985, and in April, 1986. In early May, 1986, I found many newly settled seaurchins (0.3 to 0.6 mm, 5 to 17 d old), and I compared the densities of the cohort on several types of natural substrata in barrens and kelp-bed habitats. Newly settled individuals of both purple sea urchins (S. purpuratus) and red sea urchins (S. franciscanus) were present in similar, high densities (1 000 S. purpuratus m^-2) on foliose red algal turf, a dominant substratum ofthe kelp bed, and on crustose coralline algae, the dominant substratum of an adjacent barrens. Larvae of S. purpuratus reared and tested in the laboratory showed high rates of settlement on both red algal turf and on crustose coralline algae, but significantly lower rates on rock. Larvae also settled in response to a partiallypurified extract of coralline algae. The reduced settlement on natural rock surfaces relative to either algal treatment and the significant settlement in response to the extract of coralline algae indicate that larvae discriminate between natural substrata and probably respond to a settlement cue other than, or in addition to, a simple microbial (bacterial) film. The similar densities of young recruits of S. purpuratus on dominant substrata of barrens and kelp bed show that, at least in this case, differential settlement cannot explain the high densities of sea urchins in the barrens habitat. Movement between barrens and kelp bed is unlikely given the small sizes of the newly recruited sea urchins relative to the large distances often involved. Reduced post-settlement mortality of newly settled individuals in the barrens remains the most likely mechanism leading to the higher densities of sea urchins in barrens relative to kelp-bed habitats.     

Demographic variation in the kelp<Emphasis Type="Italic"> Laminaria hyperborea</Emphasis> along a latitudinal gradient

We compared populations of Laminaria hyperborea from different regions along the Norwegian coast (58–71°N). The aim of the study was to examine possible regional differences in population demography and to relate these to latitudinal gradients in ambient temperature and light. A number of population parameters were examined in understory and canopy kelp individuals. We found latitudinal differences in stipe growth rate and growth pattern, mortality, longevity, recruitment and density. Stipe growth rate was higher for young individuals in mid-Norway than in south and north Norway. Mean stipe length was related to temperature and light, and correlated significantly with indices combining temperature during the growth season and daylength in summer. Mortality decreased and longevity increased with increasing latitude, probably because of a temperature decrease with increasing latitude and a general reduction in metabolic rate. Greater longevity may also cause the observed decrease in recruitment rate with increasing latitude, since there are fewer opportunities for understory individuals to replace dead canopy individuals. The development of particularly large kelp in mid-Norway appears to be explained by high growth rates and not a particularly long life span, as earlier assumed. This has consequences for decisions of the optimal harvesting regime for the species in the region, with regard to yield and maintenance of associated biodiversity.Communicated by M. Kühl, Helsingør     

Life-history features of the marine intertidal limpet Notoacmea scutum (Gastropoda) in central California

Growth of individually marked specimens of Notoacmea scutum (Rathke) was followed for 15 months (April, 1978 to June, 1979) at a field site in central California, USA. Growth rates were highly seasonal, being highest from late spring through early summer and lowest during winter. Growth rate also varied significantly with intertidal height: limpets lower in the intertidal zone grew faster. Both relationships may have been related to the relative abundance of food or to the amount of time available for foraging. Recently settled limpets, which were present throughout the year, grew to approximately 17 mm in length by the end of their first year; by then, they had also reached the size at which mature gonads were consistently found (16 mm). Limpets were almost 26 mm long by the end of the second year. Substantial mortality occurred throughout the year, but it was particularly severe during winter, when a large portion of the reproductive population was lost. This pronounced winter mortality was probably not caused by invertebrate predators such as seastars. Most limpets at the study site were less than 2 yr old.