Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.     

Community metabolism of intertidal flats in the Ems-Dollard estuary

To obtain an insight into the flux of carbon through intertidal sediments of the Ems-Dollard estuary, the annual cycles of gross benthic primary production and community respiration were measured at six stations, together with a set of environmental parameters. In a stepwise multiple regression analysis it was shown that temperature alone and temperature plus viable bacteria explained 50 and 70% respectively of the observed variation in community respiration. Other variables, including the rate of primary production and amount of organic carbon in the sediment were less important. The rate of primary production could not be fitted adequately into a multiple regression equation. The annual values of community respiration (177–794 gO₂·m^-2·yr^-1) and primary production (82–628 gO₂·m^-2·yr^-1) were within the range of published values. except for one station in the vicinity of a wastewater outfall, which had an extreme production (average 984 gO₂·m^-2·yr^-1). At four stations, annual community respiration exceeded primary production by 40%. It is concluded that the main carbon flux within the sediment, from CO₂ to benthic primary producers, to benthic consumers and from there to CO₂ again,was completed within a month or so, leaving untouched the large bulk of organic matter within the sediment. Possible effects of wastewater discharges on community metabolism are discussed.Publication No. 43 of the project Biological Research in the Ems-Dollard Estuary     

Seasonal changes in the structure of an intertidal community

Quantitative (0.25 m²) samples of macrofaunal (>1.0 mm) invertebrates were taken in each season from one habitat of an intertidal sandbar in the North Inlet estuary near Georgetown, South Carolina, USA. During all seasons the community inhabiting the sample site was numerically dominated by two species of haustoriid amphipods (Acanthohaustorius millsi and Pseudohaustorius caroliniensis). Seasonal changes at the community level were clearly controlled by the population dynamics of the numerically dominant species, and qualitative information on life histories was important to the interpretation of analyses' results.This work was supported by the Environmental Technology Center of Martin Marietta Corporation and the Belle W. Baruch Institute for Marine Biology and Coastal Research. It is Contribution No. 138 of the Belle W. Baruch Institute for Marine Biology and Coastal research.     

Parasitism can influence the intertidal zonation of non-host organisms

We investigated habitat utilization in Blainvilles beaked whale, Mesoplodon densirostris, in the northern Bahamas and, as such, this is the first analysis of fine-scale habitat utilization of any member of the genus Mesoplodon. We divided the area into 500×500 m grid squares and each square was assigned a classification for presence or absence of Blainvilles beaked whales, along with details of water depth, seabed gradient and seabed aspect. All squares where Blainvilles beaked whales occurred had gradients from 68 to 296 m/km and depths from 136 to 1,319 m and most faced northeast compared with 0–526 m/km, 10–3,000 m and all aspects for the whole study area. Generalized additive models and classification trees indicated that, in order of importance, aspect, gradient and depth were related to occurrence of Blainvilles beaked whales within the study area. We hypothesize that the relationships between habitat utilization and these topographic variables relates to the effects of interactions between a deepwater current and the seabed topography on preferred prey. Locally, prey animals may be concentrated in areas with a northeast aspect, intermediate gradients and depths between 200 and 1,000 m where the Deep Western Boundary Current is forced towards the surface by the local topography. These are the areas where Blainvilles beaked whales preferentially occurred.Communicated by O. Kinne, Oldendorf/Luhe     

A diatom assemblage living below the surface of intertidal sand flats

The assemblages of species living on and in intertidal sand flats at Barnstable Harbor, Massachusetts (USA) are briefly described. An endopsammic assemblage of diatoms, mainly Amphora species, living below the surface is described for the first time in such a marine situation. Some aspects of its ecology and relationship to the other diatom assemblages living on the sediments are discussed.     

Impact of sewage on the distribution,abundance and community structure of rocky intertidal macro-organisms

The biological effects of a low-volume domestic sewage discharge were studied near Wilson Cove, San Clemente Island, California (USA), from February to June, 1972. There were fewer species and less cover near the outfall (7 macro-invertebrates, 17.6%; 13 macrophytes, 91.7%) than in nearby “unpolluted” control areas (9 macroinvertebrates, 9.2%; 30 macrophytes, 103.4%). The outfall biota was less diverse than that of the controls, as shown by 5 different diversity indices. A great reduction in community stratification (spatial heterogeneity) and, hence, community complexity occurred near the outfall; this reduction in stratification was primarilly due to the absence of Egregia laevigata, Halidrys dioica, Sargassum agardhianum and Phyllospadix torreyi. These were replaced in the mid-intertidal near the outfall by a low turf of blue-green algae, Ulva californica, Gelidium pusillum and small Pterocladia capillacea, and in the lower intertidal by Serpulorbis squamigerus covered with Corallina officinalis Var. chilensis. A statistically-based determination of assemblages or groups of organisms (i.e., cluster analysis) revealed 3 discrete outfall and 3 discrete control area groups; 3 assemblages contained samples from both areas. The distributional patterns of these groups indicate that near the outfall the degree of dilution of discharged sewage is more important in regulating zonation than is tidal height. The enhancement of the suspension feeder Serpulorbis squamigerus and the omnivores Ligia occidentalis, Pachygrapsus crassipes and Anthopleura elegantissima in the outer fringe of the outfall plume hypothetically is due to their ability to utilize sewage as a food source. A critical effect of the outfall may be to decrease environmental stability thereby favoring rapid-colonizers and more sewage-tolerant organisms. The outfall macrophytes were characterized by relatively higher net primary productivities, smaller growth forms, simpler and shorter life histories, and most were components of early successional stages.     

High intertidal community organization on a rocky headland in Maine,USA

A mosaic patchwork of the barnacle Balanus balanoides L., the mussel Mytilus edulis L., and the alga Fucus vesiculosus L. was found in the transitional region between the mid and high intertidal zones on a rocky headland on Mount Desert Island, Maine, USA. The development of the mosaic was observed by following recruitment and survival of B. balanoides in denuded patches located at the same tidal level along a 60 m stretch of shore. Barnacle recruitment was least under canopies of F. vesiculosus and greatest in open areas kept moist at low tide by surf. Barnacle survival after settlement was least under the F. vesiculosus canopy due to the whiplash effect of the algal fronds in the surf and greatest in open areas free from competition from mussels. In open areas, early mortality was correlated with settlement density. In areas of dense settlement (60 spat cm^-2) up to 90% mortality resulted within 5 months from crowding associated with growth. In older individuals crowding produced hummocks of elongated, weakly attached barnacles which were more prone to removal by surf than uncrowded barnacles. Mussels exerted competitive dominance over barnacles for space and the presence of mussel beds prevented further barnacle recruitment. Mussels suffered extensive mortality during winter storms when surf removed dense mats of weakly attached mussels. The patchy distribution of mussels and barnacles results from irregular rock substrata producing numerous environmental patches with respect to wave exposure and drainage at low tide, and from densitydependent mortality of both mussels and barnacles which creates patches of new colonizable space within each environmental patch.     

Changes in barnacle population structure along an intertidal community gradient in the Florida Keys

Population distributions, densities and percentage deaths and individual size variations of the barnacles Chthamalus stellatus and Tetraclita squamosa, and the distribution of the vermetid snail Spirogluphus annulatus are measured to clarify determinants the Florida Keys (USA). The species composition of the pilings community and barnacle population densities correspond to physical gradients dependent on piling distance from shore. Interspecific competition for space is insignificant in determining C. stellatus presence or absence on pilings. Intraspecific crowding in highdensity zones of adult T. squamosa is evidenced by a significant decrease in basal diameter as density increases. The peak percentage of dead C. stellatus coincides with peak densities. The percentage of dead T. squamosa is at a low point at peak densities due to the ability of T. squamosa cyprids to settle and survive on adult shells in high-density regions. Feasible perturbation experiments for testing the importance of interspecific competition in determining the densities and vertical distributions of the species are discussed. The results of such experiments can be used to test the assumptions and predictions of the widely applied competition coefficient measure proposed by Levins (1968). Several theoretical deficiencies of Levins' measure are discussed.     

Description of a Tetraclita stalactifera panamensis community on a rocky intertidal Pacific shore of Panama

The midlittoral zone at Paitilla Beach, Panama, Is occupied by a group of organisms closely associated with Tetraclita stalactifera panamensis. Upon death, this cirripede provides housing for a number of species bound by common substrata and interacting with each other due to their close physical proximity; the faunistic group associated with each barnacle qualifies then, as a small, well-defined community which represents a subsample of the more comprehensive midlittoral community dominated by T. stalactifera panamensis. In patches where it is abundant, T. stalactifera panamensis occupies 28% of the surface area available. It has a relative abundance of only 1% of the specimens found in the zone, but all the other 95 species (with an average of 26, 810 individuals/m²) identified from the area are associated either with the parietal canals, or the external or internal surfaces of T. stalactifera panamensis tests. Each one of these structures represents a habitat which provides surface for attachment, shelter, or both, to 37 species of polychaetes, 32 of mollusks, 20 of crustaceans, 3 of actiniarians, 2 of pycogonids, 2 of sipunculans and an unknown number of tanaid, nemertean and turbellarian species. The single most diverse space-niche is represented by the parietal canals (H=0.783), occupied mostly by polychaetes. dead barnacles harbor an average of 131 individuals per test, which compose a fauna of higher diversity (H=0.678) than that associated with live barnacles (H=0.163). The latter contain 53 specimens per test on the average. Although there is considerable overlapping of species associated with the different T. stalactifera panamensis space-niches, mollusks are clearly more abundant on internal surfaces, polychaetes in parietal, canals; Balanus spp. on external surfaces, and anemones at the base of the parapet on the external surfaces of the tests. Over 220 species of macro-invertebrates were identified from the rocky intertidal at Paitilla. Most of the species are concentrated in the lower (157 species, 1824 specimens/m²) and midlittoral (96 species, 26, 810 specimens/m²) intertidal zones. Splash zone is depauperate (6 species, 1578 specimens/m²). Of the entire fauna identified from the intertidal, 73 species (33%) occur in more than one zone and 147 (67%) are restricted to a particular belt. This last category of zone-restricted species breaks down into 42% for the lower intertidal, 17% for the midlittoral, 6% for the intermediate zone, and 2% for the splash zone.     

The primary productivity of marine macrophytes from a rocky intertidal community

This study represents the first report of primary production rates for Southern California intertidal producers. The production rates of 18 marine macrophytes near Wilson Cove, San Clemente Island, are close to those for other marine algal communities. No apparent relationship was revealed between the productivity of an alga and the division to which it belongs; however, productivity was associated with growth form. Encrusting prostrate forms were the lowest producers in terms of g C/m²/h and g C/g dry weight/h; sheet-like and finely-branched forms showed a greater productivity than coarsely-branched forms. Gelidium pusillum and Ulva californica had considerably greater production rates than the other algae measured.     

Spatial structure of an intertidal molluscan assemblage on a sheltered sandy beach

The intertidal molluscan fauna on a sandy-mud beach in Newport Bay, California, USA, is divisible into two vertically distinct species assemblages which correspond to the midlittoral zone and sublittoral fringe observed on rocky coastlines and exposed sandy beaches. The species assemblage comprising the midlittoral zone is unusual, however, in that the numerically dominant species are not confined to this zone but range downward through the sublittoral fringe. The two species assemblages are vertically separated at +0.5 to +1.0 ft (+0.15 to +0.30 m; relative to 0.0 tidal datum at mean lower low water), which is higher than previously observed for the separation between the midlittoral zone and sublittoral fringe on rocky shores and exposed sandy beaches. The species composition of some of the samples at +0.5 and +1.0 ft (+0.15 and +0.30 m) was intermediate between samples higher and lower on the beach, while several samples were devoid of molluscs altogether. Additionally, the surface sedimentary environment changes in terms of an increased percent silt-clay fraction and higher sorting coefficients below +0.5 ft (+0.15 m). Since infaunal zonation is correlated with tidal height at the substrate surface, environmental factors operative at the surface are probably most important in influencing the zonation on this beach.     

Macrozoobenthic community structure in the Ría de Foz,an intertidal estuary (Galicia,Northwest Spain)

The community structure of the macrobenthic (>1 mm) fauna of an intertidal soft-bottom estuary, the Ría de Foz (Galicia, Northwest Spain) was investigated on 99 occasions during June, September and December 1984 and March 1985. The infaunal data were subjected to classification and ordination techniques. The analyses indicated that sediment characteristics (grain size, percentage silt-clay and organic-matter content) and tidal height are the most important factors governing the distribution and abundance of the intertidal communities. Three faunal assemblages were identified and examined in terms of species dominance, constancy and fidelity. In the clean sands of the outer estuary two main assemblages were distinguished; one, corresponding to the boreal LusitanianTellina community, inhabited the middle and lower tidal levels; the other, typical of a biocoenosis of sands and slightly muddy mediolittoral sands, inhabited the upper tidal level. The third benthic assemblage occupied the muddy sands and mud of the inner estuary and displayed some features in common with theCardium edule-Scrobicularia community described by Thorson.     

Desiccation as a factor in the intertidal zonation of barnacles

Four species of balanomorph barnacles, Balanus crenatus Brugière, B. balanoides (L.), Elminius modestus Darwin and Chthamalus stellatus (Poli), were studied to assess the susceptibility of intertidal barnacle species to desiccation. Known sized samples of barnacles were exposed to controlled desiccating conditions and subsequent survival and water loss were determined. It is clear that the ability to live high on the shore is dependent on a reduction of the overall permeability to water loss. Because of greater surface area to volume ratios, small stages are particularly prone to desiccation. In normal intertidal emersion periods, small stages of B. crenatus particularly, and also of B. balanoides and E. modestus which are similar in their desiccation resistance, would be susceptible to desiccation at normal temperatures and low humidities. Large barnacles would be more prone to death from high temperatures when the tide is out. The spat of C. stellatus, although surviving much longer than spat of larger dimensions of the other species, must also be prone to prolonged emersion conditions at high shore levels.     

Respiration as a function of oxygen concentration in intertidal barnacles

Respiration as a function of oxygen concentration was studied in two species of intertidal barnacles: Balanus amphitrite amphitrite ¹ (Darwin) and B. tintinnabulum tintinnabulum (L.). A critical oxygen tension was observed in both species below which the respiratory regulation broke down. In B. amphitrite amphitrite the critical oxygen tension was 2.5 ml O₂/l, and in B. tintinnabulum tintinnabulum 3.5 ml O₂/l. Species differences and habitat relations were observed in both species in their respiratory adaptation to oxygen concentration. B. amphitrite amphitrite which inhabits oxygen-deficient areas was able to regulate to much lower concentrations than B. tintinnabulum tintinnabulum which inhabits oxygen-rich open intertidal regions.     

Phytoplankton community structure and primary production in small intertidal estuarine-bay ecosystem (eastern English Channel,France)

From May 2002 to October 2003, a fortnightly sampling programme was conducted in a restricted macrotidal ecosystem in the English Channel, the Baie des Veys (France). Three sets of data were obtained: (1) physico-chemical parameters, (2) phytoplankton community structure illustrated by species composition, biovolume and diversity, and (3) primary production and photosynthetic parameters via P versus E curves. The aim of this study was to investigate the temporal variations of primary production and photosynthetic parameters in this bay and to highlight the potential links with phytoplankton community structure. The highest level of daily depth-integrated primary production Pz (0.02–1.43 g C m⁻² d⁻¹) and the highest maximum photosynthetic rate P ^B _max (0.39–8.48 mg C mg chl a ⁻¹ h⁻¹) and maximum light utilization coefficient α^B [0.002–0.119 mg C mg chl a ⁻¹ h⁻¹ (μmol photons m⁻² s⁻¹)] were measured from July to September. Species succession was determined based on biomass data obtained from cell density and biovolume measurements. The bay was dominated by 11 diatoms throughout the year. However, a Phaeocystis globosa bloom (up to 25 mg chl a m⁻³, 2.5 × 10⁶ cells l⁻¹) was observed each year during the spring diatom bloom, but timing and intensity varied interannually. Annual variation of primary production was due to nutrient limitation, light climate and water temperature. The seasonal pattern of microalgal succession, with regular changes in composition, biovolume and diversity, influenced the physico-chemical and biological characteristics of the environment (especially nutrient stocks in the bay) and thus primary production. Consequently, investigation of phytoplankton community structure is important for developing the understanding of ecosystem functioning, as it plays a major role in the dynamics of primary production.     

Short- and long-term effects of eutrophication on the secondary production of an intertidal macrobenthic community

Secondary production of a macrobenthic community at an intertidal mudflat was estimated for 33 successive months. Sampling was carried out along a eutrophication gradient, including non-eutrophied Zostera meadows, an intermediate muddy area, and a strongly eutrophied sand-muddy flat, where macroalgal blooms of Enteromorpha spp. usually occur. The Zostera meadows were always the most productive habitat (145–225 g ash-free dry weight m^–2 year^–1). In the short term, the macroalgal bloom benefited the total estuarine production by enhancing the annual production in the eutrophied area. Nevertheless, our results show that this increase was short lived and in no way sufficient to match the production in the Zostera meadows. In the long term, the present study provides evidence that the disappearance of macrophyte beds, as a result of ongoing eutrophication, constitutes a major threat to the sustainability of the estuarine ecosystem.Communicated by S.A. Poulet, Roscoff     

Effects of physical disturbance on an estuarine intertidal community: field and mesocosm results compared

The response of an estuarine macrobenthic community to different intensities of physical disturbance was examined using comparable field and mesocosm experiments. In the field study, areas of sediment (2500 cm²) were disturbed by surface-raking at a range of disturbance intensities: no disturbance, once every 2 mo, once every month, once every 2 wk, once or three times every week. These disturbance regimes were also imposed on buckets of sediment (855 cm²) collected from the field study site and maintained within a mesocosm system. In the univariate data analysis, neither field nor mesocosm communities displayed any trends in relation to the frequency of disturbance. Diversity indices were generally reduced in the mesocosm communities compared with the field communities. Multivariate analysis indicated significant differences between the field communities subjected to the most intense disturbance and those communities disturbed every ±2 wk or less. These differences were attributed to reductions in the five numerically dominant species in the high- disturbance treatments. More statistically significant differences were observed between the different disturbance intensities in the mesocosm experiment than in the field study. The implications of differences between field and mesocosm experiments results are discussed. Received: 14 June 1999 / Accepted: 3 December 1999     

Moisture as a determinant of habitat quality for a nonbreeding neotropical migratory songbird

Identifying the determinants of habitat quality for a species is essential for understanding how populations are limited and regulated. Spatiotemporal variation in moisture and its influence on food availability may drive patterns of habitat occupancy and demographic outcomes. Nonbreeding migratory birds in the neotropics occupy a range of habitat types that vary with respect to moisture. Using carbon isotopes and a satellite-derived measure of habitat moisture, we identified a moisture gradient across home ranges of radio-tracked Northern Waterthrush (Seiurus noveboracensis). We used this gradient to classify habitat types and to examine whether habitat moisture correlates with overwinter mass change and spring departure schedules of Northern Waterthrush over the late-winter dry season in the tropics. The two independent indicators of moisture revealed similar gradients that were directly proportional to body mass change as the dry season progressed. Birds occupying drier habitats declined in body mass over the study period, while those occupying wetter habitats increased in body mass. Regardless of habitat, birds lost an average of 7.6% of their mass at night, and mass recovery during the day trended lower in dry compared with wet habitats. This suggests that daily incremental shortfalls in mass recovery can lead to considerable season-long declines in body mass. These patterns resulted in consequences for the premigratory period, with birds occupying drier habitats having a delayed rate of fat deposition compared with those in wet habitats. Taken together with the finding that males, which are significantly larger than females, are also in better condition than females regardless of habitat suggests that high-quality habitats may be limited and that there may be competition for them. The habitat-linked variation in performance we observed suggests that habitat limitation could impact individual and population-level processes both during and in subsequent periods of the annual cycle. The linkage between moisture and habitat quality for a migratory bird indicates that the availability of high-quality habitats is dynamic due to variation in precipitation among seasons and years. Understanding this link is critical for ascertaining the impact of future climate change, particularly in the Caribbean basin, where a much drier future is predicted.