Effects of proximity to an offshore hard-bottom reef on infaunal abundances

Rock outcrops represent an extensive, commercially important habitat along the southeast coast of North America, supporting large abundances of algae, invertebrates and fish. These rock outcrops have often been viewed as largely self-supporting systems, but some studies have suggested that the fish and invertebrate predators that use the ledges for shelter are strongly dependent on food from adjacent, soft-bottom habitats. We examined benthic macrofaunal abundances along two 75-m transects away from a rock ledge near Wrightsville Beach. North Carolina (depth ca. 30 m) in July 1990 and April 1991. The July 1990 samples indicated significantly higher abundances of total infauna, and of polychaetes, bivalves, isopods, and scaphopods, at a distance of 75 m from the rock ledge. In April 1991, when macrofaunal abundances were generally higher than in the previous summer, distributions of major taxa with distance from the rock ledge were more variable. Caging studies on macrofaunal abundances 10 m from the ledge, and video observations of fish abundances at 10 and 75 m from the ledge, provide support for a trophic link between the rock ledge and the adjacent soft-bottom communities. Our studies suggest that there are potentially important indirect effects of predator-prey interactions among the rock ledge-associated predators and soft-bottom prey.     

Widespread mass mortalities of the green sea urchin in Nova Scotia,Canada

Mass mortalities of sea urchins exceeding 84 000 t live weight or 2 900 t organic weight occurred during the autumns of 1980 and 1981 along the southern coast of Nova Scotia, Canada. The kill was nearly complete along 160 km of shoreline (straight-line distance) and was inter-mittent along another 280 km. Sea urchins in a variety of rocky habitats were affected, including areas exposed to and sheltered from ocean swell, with both dense and no macrophyte cover, and from 0–13 m depth. The cause of the mortalities was presumed to be a biological agent and was transferable in the laboratory. Colonization of the habitat by subtidal macroalgae and a subsequent large increase in benthic primary production is expected to follow.     

Use of pelagic prey subsidies by demersal predators in rocky reefs: insight from movement patterns of lingcod

Allochthonous subsidies of energy and nutrients can affect community structure in patchy marine habitats, including rocky reefs, and their ecological consequences may depend on the mechanism of energy transfer. Lingcod (Ophiodon elongatus) are demersal predators that trophically link nearshore rocky reefs with offshore pelagic habitats through consumption of pelagic fishes. We quantified lingcod habitat use and movement patterns to make inferences about the temporal and spatial conditions under which lingcod may acquire pelagic prey. Lingcod maintained small home ranges (21,272 ± 13,630 m²) within a rocky reef in the San Juan Archipelago, Washington; eight of nine individuals used rocky habitat exclusively. Depths occupied by lingcod (0–50 m) coincided with pelagic fish distribution on the rocky reef; however, diel patterns in lingcod activity varied inversely with occurrence of pelagic fishes on the reef. Our findings suggest that the pelagic subsidy to lingcod is not strongly mediated through directed off-reef foraging by lingcod.     

The reproduction of some littoral molluscs from Macquarie Island (Sub-Antarctic)

The reproduction of 9 species of littoral molluscs from the sub-Antarctic Macquarie Island was examined. The mode of larval development is reported for all species, and the reproductive patterns for 7 are described from collections taken monthly over a period of 1 year. Two species release gametes for external fertilization, whereas the other 7 have non-pelagic development via brooding or the laying of egg cases; some of these findings were deducted from examination of gonads rather than by direct observation. Features of reproduction important in any correlation between a littoral animal's reproduction and its habitat are discussed, and relationships between pelagic and non-pelagic development of the Macquarie molluscs and (a) distribution in the local environment and (b) habitat are drawn. The widely recognised correlation between species having a non-pelagic development and small number and large size of eggs is considered as a method for predicting the type of larval development (in terms of pelagic versus non-pelagic) from an examination of the ovary. It is suggested that further data on modes of development could be obtained from specimens of marine invertebrates collected throughout southern latitudes and that such data would enhance zoogeographical interpretations.     

Zoogeographical aspects of rocky-intertidal molluscan fauna of the Pacific coasts of Japan

Intertidal molluscan faunas were studied on the exposed rocky shores of the Pacific coasts of Japan (excluding Hokkaido) to determine distribution patterns and latitudinal gradients of taxa. Collections were made at spring low tide in 1980. Five types of distribution patterns were found: tropical, tropical-temperate, warm-temperate, temperate, and boreal. Many examples of parallel species displacement, the term which Thorson (1957) proposed and generalized, were found. Latitudinal gradients of molluscan taxon were observed. Frequency of the Archaeogastropoda in gastropods increases northward, while frequency of the Mesogastropoda and the Neogastropoda decreases. Frequency of predatory gastropods decreases northward, and that of herbivorous ones increases. Commonness indices show that the molluscan fauna of the study sites situated within the wide area between the Kanto and Kyushu are closely related with each other, and they can be regarded as one province. The Tohoku district is less related with this region, and the Okinawa district is the least related.     

Intertidal assemblages on artificial and natural habitats in marinas on the north-west coast of Italy

Following the progressive expansion of human populations, the number of artificial habitats is increasing in shallow waters of urbanised coastal areas. The comparison of assemblages between natural and artificial habitats is necessary to determine whether there is real loss and fragmentation of natural habitats. This study investigates the changes in intertidal assemblages caused by marinas for tourists, along exposed rocky shores on the north-west coast of Italy. Marinas, being made of transplanted boulders and by internal wave-sheltered seawalls, introduce different types of artificial habitats in a relatively small area. Intertidal assemblages on breakwaters, seawalls and adjacent rocky shores were compared at three locations, thousands of metres apart. To assess the generality of patterns through time, natural and artificial habitats were sampled at three different times, over a period of about 2 years. Data were analysed by means of multivariate and univariate analyses to test the hypotheses that assemblages and abundances of single taxa differed among habitats, consistently among locations and times of sampling. Furthermore, the variability of assemblages at the scales of tens of centimetres and metres was compared among habitats. Assemblages on seawalls were largely distinct from those on rocky shores or breakwaters. Seawalls, which supported a smaller number of species than breakwaters and rocky shores, were dominated by encrusting algae and lacked common species such as Rissoella verrucosa and Patella rustica. The abundance of main-space occupiers did not differ between breakwaters and rocky shores, but there were differences in variability of assemblages at both the spatial scales investigated. This study provides evidence for differences between intertidal assemblages supported by artificial habitats at marinas and those on adjacent rocky shores. Differences in habitat-structure (and/or wave-exposure in the case of seawalls) could explain the occurrence of distinct intertidal assemblages. Despite the nature and magnitude of these differences, varied according to the type of artificial habitat considered, neither breakwaters nor seawalls could be considered surrogates of rocky shores.Communicated by R. Cattaneo-Vietti, Genova     

Comparative species composition and relative abundance of decapod crustaceans in marine habitats of Panamá

The community structure of the decapod crustacean fauna of 7 tropical, shallowwater, marine habitats (sandy beaches, mangrove swamps and rocky intertidal habitats on both the Pacific and Caribbean coasts of Panama, and Pocillopora damicornis coral habitat of the Bay of Panama) were examined and analyzed for species composition and relative abundances. Collections from the 7 habitats yielded 4361 individuals, representing 236 species. The number of species per habitat was (Pacific, Caribbean): sandy beach (16, 7); mangrove (20, 17); P. damicornis (53); rocky intertidal (78, 67). There were more species represented by more individuals in the Pacific habitats. An index of faunal similarity was calculated for each pair (Pacific-Caribbean) of habitats. This index is the number of ecologically similar congeneric species which occurred in both habitats expressed as a percentage of the total number of species present in the pair of habitats. For the sandy beach communities there were three Pacific species which were similar to three Caribbean species, a similarity of 6/23 or 26%. The index of similarity for the mangrove communities is 54% and for the rocky intertidal communities it is 37%. The P. damicornis community has affinities with the Pacific rocky intertidal community (18%), with that of the Caribbean rocky intertidal (16%) and with that of Indo-West Pacific pocilloporid corals (20%). A few specialized species dominated each of the communities. The habitats and the number of species accounting for over half of the individuals present are (Pacific, Caribbean): sandy beach (1, 1); mangrove (4, 4); P. damicornis (5); and rocky intertidal (3, 6). Most of the species in each community were represented by one or a few individuals.     

Phylogeography of the diamond turbot (Hypsopsetta guttulata) across the Baja California Peninsula

We compared morphology and sequenced nuclear and mitochondrial genes from 11 populations of a previously genetically unstudied “Baja California disjunct” species, the diamond turbot (Hypsopsetta guttulata). This species exhibits very limited adult movement and restriction to soft-bottom habitats but has a moderately long pelagic larval duration. Therefore, if pelagic larval duration is correlated with gene flow between Gulf of California and Pacific populations, we expect a reduced level of genetic and morphological differentiation. However, if adult habitat and ecology have more effect on gene flow, we expect the populations in the two bodies of water to be more highly differentiated. We used logistic regression to compare morphological features and phylogenetic and population genetic analyses to compare nucleotide sequence data. Gulf of California H. guttulata are different from Pacific populations in morphology and both mitochondrial and nuclear gene sequences. MtDNA shows reciprocal monophyly, and nuclear sequences from the Gulf of California formed a monophyletic group. Population genetic analyses also suggest further population subdivision within the Pacific and within the Gulf of California. We argue that adult ecology has a significant effect on migration rates among populations in the Pacific Ocean and the Gulf of California.     

Spatial variability in habitat associations of pre- and post-settlement stages of coral reef fishes at Ishigaki Island,Japan

Successful settlement of pelagic fish larvae into benthic juvenile habitats may be enhanced by a shortened settlement period, since it limits larval exposure to predation in the new habitat. Because the spatial distribution of marine fish larvae immediately prior to settlement versus during settlement was unknown, field experiments were conducted at Ishigaki Island (Japan) using light trap sampling and underwater visual belt transect surveys to investigate the spatial distribution patterns of selected pre- and post-settlement fishes (Acanthuridae, Pomacentridae, Chaetodonidae and Lethrinidae) among four habitats (seagrass bed, coral rubble, branching coral and tabular coral). The results highlighted two patterns: patterns 1, pre- and post-settlement individuals showing a ubiquitous distribution among the four habitats (Acanthuridae) and pattern 2, pre-settlement individuals distributed in all habitats, but post-settlement individuals restricted to coral (most species of Pomacentridae and Chaetodontidae) or seagrass habitats (Lethrinidae). The first pattern minimizes the transition time between the larval pelagic stage and acquisition of a benthic reef habitat, the latter leading immediately to a juvenile lifestyle. In contrast, the second pattern is characterized by high settlement habitat selectivity by larvae and/or differential mortality immediately after settlement.     

Multiple trans-Arctic passages in the red alga Phycodrys rubens: evidence from nuclear rDNA ITS sequences

In order to investigate how episodes of geological and climatic change have influenced the distribution and evolutionary diversification of Arctic to cold temperate-North Atlantic seaweed species, intraspecific genetic variation was analyzed among isolates of the sublittoral, benthic red alga Phycodrys rubens (collected between June 1992 and January 1994). Rooted phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer (ITS) sequences and the plastid encoded Rubisco spacer sequences suggest that P. rubens invaded the North Atlantic from the Pacific shortly after the opening of the Bering Strait (3 to 3.5 million years ago), colonizing both the western and eastern Atlantic coasts. Based on these data we further hypothesize that P. rubens survived along the European coasts during the more recent Pleistocene glaciations, while becoming locally extinct along the North American Atlantic coasts. Following retraction of the last ice sheet, the western Atlantic coast was colonized a second time from the Pacific. The presence of two distinct genetic types (based on ITS and Rubisco sequences) along the European coasts is postulated to be a result of isolation and subsequent differentiation. This is likely because ice-free areas are known to have existed in northern Scotland and Norway during the last glaciation. The presence of an East Atlantic genetic type along the West Atlantic coast is believed to be a recent introduction (caused by human activity) of P. rubens to Newfoundland.     

Genetic population structure of the seagrass Thalassodendron ciliatum in sandy and rocky habitats in southern Mozambique

The genetic diversity of the dioecious seagrass Thalassodendron ciliatum in six populations in southern Mozambique was studied by means of random amplified polymorphic DNA (RAPD). Samples were taken from sandy and rocky habitats over a span of 880 km. Analysis of molecular variance (AMOVA) showed that most of the genetic variability (71.6% of total genetic variation) was observed within populations. There were also significant genetic differences among populations within each habitat (sandy and rocky). We did not find any significant overall genetic difference between sandy versus rocky populations, indicating that the morphological differences between plants from these two habitats are not maintained by reproductive isolation. There was no significant correlation between geographical and genetic distance, which is discussed in the light of current patterns along the coast. All sampled populations consisted of several genetically distinct individuals, indicating that sexual reproduction is widespread.     

Sympatric species of threespine stickleback differ in their performance in a spatial learning task

Increasing evidence suggests that cognitive function is under selection in diverse taxa and that this results in different cognitive abilities in taxa experiencing different selective environments. For example, species inhabiting spatially complex environments might be expected to have good spatial learning ability. We investigated whether local habitat conditions influence learning by comparing the performance of two recently diverged species of threespine stickleback (Gasterosteus aculeatus complex) in a spatial learning task. The two species reside sympatrically in several lakes. Benthics occupy the spatially structured vegetated littoral zone, whereas limnetics occupy the spatially homogenous open-water pelagic zone. We trained fish to locate a hidden reward in a T-maze and asked whether they did so by learning a body-centred pattern of movement (turn left or right) or by using plant landmarks. Both benthics and limnetics used turn and landmark cues, but benthics learnt the task almost twice as quickly as limnetics. This difference is consistent with the hypothesis that benthic and limnetic sticklebacks are equipped with spatial learning abilities well suited to the spatial complexity of their littoral and pelagic habitats. Our findings add to the understanding of the evolution of learning.     

Spread ofCyclope neritea (Mollusca: Gastropoda) along the north-eastern Atlantic coasts in relation to oyster culture and to climatic fluctuations

The geographic distribution ofCyclope neritea (Linné) (Mollusca: Gastropoda) along the Atlantic coasts of Portugal, Spain and France is reported. Analysis of information published over the last 90 yr indicates that, since 1983,C. neritea has extended its range northwards from Arcachon Bay along the coast of the Bay of Biscay. The spread of this small gastropod is directly related to the commercial transport of the Pacific cup oyster (Crassostrea gigas Thunberg) to the Atlantic from Mediterranean oyster-culture sites, and to its encountering climatic conditions suitable for its settlement and reproduction in its new habitat. Its transport by human intervention explains the rapid spread ofCyclope neritea during 1983 and 1984, despite its lack of a planktonic stage or of effective natural migration by adults. Climatic conditions in the southern parts of the Bay of Biscay are similar to those of the French Mediterranean lagoons, explaining the easy acclimatization of this eurythermic gastropod. Also, coastal air and sea-surface temperature records over the last two decades (1971–1988) have revealed a slight warming of the southern parts of the Bay of Biscay. This has resulted in a more strongly marked latitudinal thermal gradient along the Bay of Biscay and has furthered survival and reproduction ofCyclope neritea emigrants. A detailed analysis of commercial supplies of cup oysters to the Bay of Biscay from the Mediterranean coast indicates that theC. neritea emigrants undoubtedly originated from the salt-water lagoons of Salse-Leucate and Thau (French Mediterranean coast).     

Trophic importance of subtidal metazoan meiofauna: evidence from in situ exclusion experiments on soft and rocky substrates

In coastal marine ecosystems, predation might affect spatial distribution and population dynamics of benthic assemblages. Here, by means of experimental exclusion of potential predators, we compared the effects of epibenthic predation on metazoan meiofaunal assemblages on soft and rocky substrates. Different patterns of abundance were observed in uncaged versus caged plots, across habitats. In caged soft substrates, the abundance of Nematodes, Copepods and Polychaetes increased by 56, 45, 57%, respectively, in the first 3 months. An increase in the number of meiofaunal taxa was also observed. The exclusion of predators from rocky substrates showed less clear patterns. It did not affect the number of taxa while a decrease in meiofaunal abundance was observed. Our results suggest that the exclusion of epibenthic predators had clear effect on total metazoan meiofaunal abundance and on the number of taxa, only in soft bottoms. The different impact of predation across habitats can be potentially explained by differences in terms of spatial variability and substrate complexity. We estimated that, coarsely, more than 75% of total metazoan meiofaunal production can be channeled to higher trophic levels through predation on soft-bottoms. Among meiofaunal taxa, Polychaetes and Nematodes provided the major contribution to benthic energy transfers. These results suggest the trophic relevance of metazoan meiofauna in coastal food webs and claim for the refinement of further experiments for the quantification of its role in different ecological systems.     

Global biodiversity patterns of benthic marine algae

Species richness patterns are remarkably similar across many marine taxa, yet explanations of how such patterns are generated and maintained are conflicting. I use published occurrence data to identify previously masked latitudinal and longitudinal diversity gradients for all genera of benthic marine macroalgae and for species in the Order Bryopsidales. I also quantify the size, location, and overlap of macroalgal geographic ranges to determine how the observed richness patterns are generated. Algal genera exhibit an inverse latitudinal gradient, with biodiversity hotspots in temperate regions, while bryopsidalean species reach peak diversity in the tropics. The geographic distribution of range locations results in distinct clusters of range mid-points. In particular, widespread taxa are centered within tight latitudinal and longitudinal bands in the middle of the Indo-Pacific and Atlantic Oceans while small-ranged taxa are clustered in peripheral locations, suggesting that variation in speciation and extinction are important drivers of algal diversity patterns. Hypotheses about factors that regulate diversity contain underlying assumptions about the size and location of geographic ranges, in addition to predictions as to why species numbers will differ among regions. Yet these assumptions are rarely considered in assessing the validity of the prevailing hypotheses. I assess a suite of hypotheses, suggested to explain patterns of marine diversity, by comparing algal-richness patterns in combination with the size and location of algal geographic ranges, to the richness and range locations predicted by these hypotheses. In particular, the results implicate habitat areas and ocean currents as the most plausible drivers of observed diversity patterns.     

Marine habitat mapping in support of Marine Protected Area management in a subarctic fjord: Gilbert Bay, Labrador, Canada

This paper presents an approach that allows production of benthic substrate and habitat maps in fjord environments. This approach is used to support the management of the Gilbert Bay Marine Protected Area (MPA) in southeastern Labrador, Atlantic Canada. Multibeam sonar-derived bathymetry, seabed slope, and acoustic reflectance (backscatter) were combined using supervised classification methods and GIS with ground-truthed benthic sampling in order to derive maps of the substrates and main benthic habitats. Six acoustically distinct substrate types were identified in the fjord, and three additional substrate types without a unique acoustic signature were recognized. Ordination by multidimensional scaling and analysis of similarity generalized these to four acoustically distinct habitat types. Greatest within-habitat (alpha) diversity was found in the coralline-algae encrusted gravel habitat. Greatest between-habitat (beta) diversity was found in the management Zones 1 and 2, which have the highest level of protection. The study confirmed that the zoning plan for the MPA, which was designed to protect spawning and juvenile fish habitat for a local genetically distinct population of Atlantic cod, afforded highest levels of protection to areas with highest habitat diversity.     

Urban marine ecology in southern California: the ability of riprap structures to serve as rocky intertidal habitat

Increasing human populations along marine coastlines has lead to increasing urbanization of the marine environment. Despite decades of investigations on terrestrial ecosystems, the effect of urbanization on marine life is not well understood. Riprap is the rocky rubble used to build jetties, breakwaters, and armored shorelines. Roughly 30% of the southern California shoreline supports some form of riprap, while 29% of the shoreline is natural rocky substrate. Astonishingly few studies have investigated this anthropogenic rocky habitat even though it rivals a natural habitat in area on a regional scale along a coastline that has been extensively studied. In this study, I compared the diversity and community structure of exposed rocky intertidal communities on four riprap and four natural sites in southern California. I ask the following questions: (1) does diversity or community composition differ between intertidal communities on riprap and natural rocky habitats in southern California, (2) if so, which organisms contribute to those differences, (3) which physical factors are contributing to these differences, and (4) do riprap habitats support higher abundances of invasive species than natural habitats? On average, riprap and natural rocky habitats in wave exposed environments in southern California did not differ from each other in diversity or community composition when considering the entire assemblage. However, when only mobile species were considered, they occurred in greater diversity on natural shores. These differences appear to be driven by wave exposure. The presence of invasive species was negligible in both natural and riprap habitats.     

Biogeography of Recently Extinct Marine Species: Implications for Conservation

At least fifteen marine and coastal animal species have become extinct since the end of the Pleistocene. Analyses of the number of marine biogeographical provinces occupied by these species show that, contrary to the prevailing view that extinction is usually associated with a small range, at least five species (33%) had large ranges encompassing parts of two or more provinces. At least eight species occurred in areas that served as geographical refuges for taxa during the Pliocene and early Pleistocene. These refuges, in which primary planktonic productivity is generally very high, include the northwestern Atlantic and North Pacific. Extinctions in the northwestern Atlantic (four species) may be partly responsible for the subsequent success of human-introduced species in subtidal and open-coast intertidal habitats. Regions of high productivity deserve priority among marine areas to be protected from overexploitation and habitat destruction.     

Demography and phenology of the intertidal green algaCodium setchellii: The enigma of local scarcity on sand-influenced rocky shores

Demographic and phenological attributes that characterize rare and scarce species have not been extensively investigated for marine benthic organisms. From 1984 to 1987, 1 investigated the population ecology of the low intertidal, psammophytic green algaCodium setchellii Gardner that forms low-density populations on sand-influenced rocky benches on N.E. Pacific shores. To address why the alga is not more common in the habitat in which it is most abundant, I quantified sand fluctuations and population dynamics ofC. setchellii at several sites along the central coast of Oregon, USA. Sand movement was highly unpredictable both spatially and temporally on a local scale. AlthoughC. setchellii became fertile in late autumn to winter, the alga and rocky surfaces were often deeply buried by sand in these seasons. Algal recruitment was also spatially and temporally variable, and mortality of small thalli was high. Thallus growth was generally slow, and the alga had the capacity to persist at a given size for long periods. On sand-influenced rocky shores, mortality of largethalli was low, with some thalli persisting for several years. In laboratory experiments,C. setchellii lost substantial biomass during 4 to 5 wk sand burials, whereas two sympatric species of psammophytic algae lost little to no biomass. The within-site scarcity ofC. setchellii may be a consequence of (1) largely unpredictable environmental conditions that reduce algal recruitment and survival, and (2) apparent lack of specialized anatomical or reproductive adaptations to persist on sand-scoured rocky shores. The alga's persistence may be due to its slow growth and long lifespan.     

Intensive fishing of marine consumers causes a dramatic shift in the benthic habitat on temperate rocky reefs

Intensive fishing can cause dramatic, long-lasting shifts in benthic habitat. This study used three approaches to test whether overharvesting of blacklip abalone (Haliotis rubra) can cause a shift in benthic habitat to a configuration that is unsuitable for abalone, on the east coast of Tasmania, Australia. After 18 months of removing abalone from rocks, encrusting red algae (ERA) became overgrown by filamentous and foliose algae, sessile invertebrates and accumulated sediment. The differences in the community composition between locations, sites nested within locations and rocks were minor. Throughout the study, abalone were largely associated with areas of rock covered in ERA but avoided other habitats. A transplant experiment demonstrated that abalone preferred areas of rock covered in ERA but move away from overgrown rocks. These results suggest overharvesting of abalone results in a shift to benthic habitat poorly preferred by abalone. This could form a positive feedback loop that limits recovery of abalone populations and ERA.