Integration of Local and Regional Perspectives on the Species Richness of Coral Assemblages

SYNOPSIS. We have evaluated the relationship between regionalspecies richness and the number of species occurring withinlocal, quantitatively sampled assemblages of scleractinian corals.Our data have been extracted from the published literature describingrichness patterns from over 100 locations around the world.In general, we find a positive relationship between local andregional richness. Local richness is not independent of regionalrichness as posited by conventional theory and there is no hardupper limit indicating saturation. Instead, local coral assemblagesare regionally enriched. This result suggests that these assemblagesare open to regional sources of species. The degree of regionalenrichment is geographically variable. In the Indo-Pacific,assemblages in speciose regions appear to be less open and muchmore sensitive to local depth and habitat gradients than thosein more depauperate regions. Other large-scale geographicaland historical effects on local richness in the Indo-Pacificinclude the degree of isolation from high-diversity regionsand distance from the equator. In contrast, local richness inthe relatively homogeneous and depauperate western Atlanticis insensitive to the large-scale variables we examined. Asin most ecological communities, membership in local assemblagesof corals is not absolutely limited (by biotic interactionsor local environmental factors) nor is it totally open to regionalpools of species. Understanding the dynamics of coral communitieswill require integrating the local ecological perspective withlarge-scale phenomena (i.e., physical TECO processes [Myers,1994] and evolutionary history [Hugueny et al. 1997]). Suchan integration will necessarily encompass multiple spatial andtemporal scales.     

Coral Community Adaptability to Environmental Change at the Scales of Regions, Reefs and Reef Zones

SYNOPSIS. Projected global increases in temperature, sea level,storminess and atmospheric carbon dioxide (CO₂) are likely tocause changes in reef coral communities which the present humangeneration will view as deleterious. It is likely coral communitytrajectories will be influenced as much by the reduction inintervals between extreme events as the projected increasesin means of environmental parameters such as temperature, atmosphericCO₂ and sea-level. Depressed calcification rates in corals causedby reduced aragonite saturation state of water may increasevulnerability of corals to storms. Moreover, reduction in intervalsbetween storms and other extreme events causing mass mortalityin corals (coral predators, diseases, bleaching) are likelyto more frequently "set back" reef coral communities to earlysuccessional stages or alternate states characterized by non-calcifyingbenthos (plants, soft corals, sponges). The greater the areaand the longer the duration of dominance of putative "coral/corallinealgae" zones of coral reefs by non-calcifying stages, the lesswill be the reefs capacity to accrete limestone bulk lockedup in the big skeletal units of late successional stages (i.e.,very large old corals). Averaged over decades to centuries,the effects of such changes on the coral community's carryingcapacity for other biota such as fish are unpredictable. A "shiftingsteady-state mosaic" null model may provide a useful conceptualtool for defining a baseline and tracking changes from thatbaseline through time.     

Effects of Habitat Connectivity on the Abundance and Species Richness of Coral Reef Fishes: Comparison of an Experimental Habitat Established at a Rocky Reef Flat and at a Sandy Sea Bottom

Coral reef fish assemblages are widely recognized for the coexistence of numerous species, which are likely governed by both coral diversity and substratum complexity. However, since coral reefs provide diverse habitats due to their physical structure and different spatial arrangements of coral, findings obtained from an isolated habitat cannot necessarily be applied to fish assemblages in other habitats (e.g. continuous habitats). The aim of this study, therefore, was to determine by a field experiment whether habitat connectivity (spatial arrangement of coral colonies) affects abundance and species richness of fishes in an Okinawan coral reef. The experiment consisted of transplanted branching coral colonies at a 4m×8m quadrat at both a rocky reef flat and sandy sea bottom. Generally, the abundance of fishes was greater at the sandy sea bottom, especially for three species of pomacentrids, one species of labrids, one species of chaetodontids and two species of apogonids. Species–area curves showed that the species richness of fishes was significantly greater in the quadrat at the sandy sea bottom at 3, 6 and 9 months after the start of the experiment. The rate of increase in abundance of fishes per area was significantly greater in the quadrat at the sandy sea bottom over the study period. The results of rarefaction analyses showed that the rate of increase in species richness per abundance was significantly higher in the quadrat at the sandy sea bottom in the juvenile settlement period, indicating that the magnitude of dominance by particular species was greater at the sandy sea bottom habitat. Our findings suggest that habitat connectivity affects the abundance and species richness of coral reef fishes, i.e. the isolated habitat was significantly more attractive for fishes than was the continuous habitat. Our findings also suggest that the main ecological factors responsible for organization of fish assemblage at a continuous habitat and at an isolated habitat are different.     

Coral Reefs on the Edge? Carbon Chemistry on Inshore Reefs of the Great Barrier Reef

While increasing atmospheric carbon dioxide (CO₂) concentration alters global water chemistry (Ocean Acidification; OA), the degree of changes vary on local and regional spatial scales. Inshore fringing coral reefs of the Great Barrier Reef (GBR) are subjected to a variety of local pressures, and some sites may already be marginal habitats for corals. The spatial and temporal variation in directly measured parameters: Total Alkalinity (TA) and dissolved inorganic carbon (DIC) concentration, and derived parameters: partial pressure of CO₂ (pCO₂); pH and aragonite saturation state (Ω_ar) were measured at 14 inshore reefs over a two year period in the GBR region. Total Alkalinity varied between 2069 and 2364 µmol kg⁻¹ and DIC concentrations ranged from 1846 to 2099 µmol kg⁻¹. This resulted in pCO₂ concentrations from 340 to 554 µatm, with higher values during the wet seasons and pCO₂ on inshore reefs distinctly above atmospheric values. However, due to temperature effects, Ω_ar was not further reduced in the wet season. Aragonite saturation on inshore reefs was consistently lower and pCO₂ higher than on GBR reefs further offshore. Thermodynamic effects contribute to this, and anthropogenic runoff may also contribute by altering productivity (P), respiration (R) and P/R ratios. Compared to surveys 18 and 30 years ago, pCO₂ on GBR mid- and outer-shelf reefs has risen at the same rate as atmospheric values (∼1.7 µatm yr⁻¹) over 30 years. By contrast, values on inshore reefs have increased at 2.5 to 3 times higher rates. Thus, pCO₂ levels on inshore reefs have disproportionately increased compared to atmospheric levels. Our study suggests that inshore GBR reefs are more vulnerable to OA and have less buffering capacity compared to offshore reefs. This may be caused by anthropogenically induced trophic changes in the water column and benthos of inshore reefs subjected to land runoff.     

Complementarity of Rotating Video and Underwater Visual Census for Assessing Species Richness,Frequency and Density of Reef Fish on Coral Reef Slopes

Estimating diversity and abundance of fish species is fundamental for understanding community structure and dynamics of coral reefs. When designing a sampling protocol, one crucial step is the choice of the most suitable sampling technique which is a compromise between the questions addressed, the available means and the precision required. The objective of this study is to compare the ability to sample reef fish communities at the same locations using two techniques based on the same stationary point count method: one using Underwater Visual Census (UVC) and the other rotating video (STAVIRO). UVC and STAVIRO observations were carried out on the exact same 26 points on the reef slope of an intermediate reef and the associated inner barrier reefs. STAVIRO systems were always deployed 30 min to 1 hour after UVC and set exactly at the same place. Our study shows that; (i) fish community observations by UVC and STAVIRO differed significantly; (ii) species richness and density of large species were not significantly different between techniques; (iii) species richness and density of small species were higher for UVC; (iv) density of fished species was higher for STAVIRO and (v) only UVC detected significant differences in fish assemblage structure across reef type at the spatial scale studied. We recommend that the two techniques should be used in a complementary way to survey a large area within a short period of time. UVC may census reef fish within complex habitats or in very shallow areas such as reef flat whereas STAVIRO would enable carrying out a large number of stations focused on large and diver-averse species, particularly in the areas not covered by UVC due to time and depth constraints. This methodology would considerably increase the spatial coverage and replication level of fish monitoring surveys.     

Patterns of Coral Recruitment and Post-settlement Mortality on Bermuda's Reefs: Comparisons to Caribbean and Pacific Reefs

The recruitment of juvenile corals and post-settlement mortalityare important processes for coral population dynamics and reefcommunity ecology. I monitored juvenile coral recruitment andsurvival on a severely disturbed reef in Bermuda from 1981 to1989 and on adjacent healthy reefs from 1986 to 1990. Poritesastreoides was the dominant recruiting species at all sites,due to the release of brooded planulae that may settle rapidly.The dominant corals on Bermuda's reef, Diploria spp., were poorrecruiters, perhaps due to the broadcast mode of reproductionof these species. However, Diploria spp. have lower juvenilemortality rates compared to P. astreoides, which may explaintheir abundance on Bermuda's reefs. Brooding corals, primarily agariciids, were the dominant recruitson Atlantic reefs compared to high recruitment rates by spawningacroporids in the Pacific, which may be the result of differentenvironmental conditions and/or evolutionary trends in the twooceans. The latter group also suffered high post-settlementmortality compared to brooding coralsin both the Atlantic andthe Pacific. Massive corals in both oceans had generally lowrecruitment rates, related to their spawning mode of reproduction,and low rates of post-settlement mortality. The dominant roleof long-lived massive corals on the Atlantic and Pacific reefscan be understood in terms of their life-history strategy incomparison to the relatively short-lived Pacific acroporidsand Atlantic agariciids that rely on different strategies tomaintain their populations.     

Spatial and Temporal Patterns in Assemblages of Temperate Reef Fish

SYNOPSIS. For reef fish in temperate marine regions, such componentsof local assemblage diversity (i.e., within a reef) as speciesrichness, total fish density, and rank order of abundance canremain relatively constantthrough time. Long-term data (17 years)for assemblages on 2 reefs in Southern California revealed that,despite high turnover in rare species, overall species richnesswas affected only moderately by major oceanographicdisturbances.This resilience of the assemblage is in marked contrast to hightemporal variation in densities exhibited by many local populationsof individual species, and it suggests that measurements ofdiversity to indicate status of an assemblage should be usedwith caution. Here we consider various processes and factors,together with the spatial and temporal scales over which theyoperate, that can influence local diversity (and its estimation)of reef fishes. Mechanisms that can "buffer" local diversityof reef fishes include dispersal of young that inter-connectssubpopulations, high "inertia" in relative abundance and populationstructures (especially for long-lived species), and broad ecologicalrequirements of many species. These considerations suggest thatthe effect of disturbances on local diversity of reef fisheswill depend in part on the magnitude, duration, frequency andspatial scale of the perturbation. While long-term data arefew, available information suggests that, due to life historycharacteristics of the fish and the spatial and temporal scalesat which disturbances are likely to occur, assemblages of temperatemarine reef fish might be relatively resilient to environmentalperturbations     

Guide to authors preparing papers for the 7th International Coral Reef Symposium and for publication in Coral Reefs

This is a statement of the problem, along with a summary of the method adopted for solving the problem, the major results and conclusions, and an explanation regarding the importance of the research. The Abstract should not include phrases such as the results will be discussed....The organising committee for the 7th International Coral Reef Symposium, to be held in Guam from 22–26 June 1992, have adopted the Coral Reefs format for all papers that are to be published in the conference proceedings. Following are some suggestions for writing and structuring papers. These tips may prove helpful to the less-experienced author.     

物种丰富度垂直分布格局及影响机制

物种丰富度分布格局是一定地域内物种丰富度沿三维空间的立体分布,包括物种丰富度在经度、纬度和垂直梯度(海拔高度和海水深度)三个维度上的空间分异。近年来物种多样性的垂直分布格局与机制研究得到了生物地理学家和生态学家的重视。物种丰富度的垂直分布格局存在多种类型,但随海拔增加而物种数减少的单调递减模型和中海拔物种丰富度最高的单峰模型较为常见。目前在机制研究中验证较多的是气候稳定性、生物因子(种间相互作用)、能量、生境异质性、干扰、进化时间、物种分化速率、面积、中域效应(mid-domain effect)、生态位保守性(niche conservatism)等假说和机制。物种丰富度的分布格局是多方面因素综合作用的结果;由于地理、地形、气候、地质演化历史、物种库和进化历史、物种分化速率、干扰等差异,在不同地区存在着特别的物种丰富度空间分布格局和机制;处于同一地区的不同类群的物种也因进化扩散历史和生态适应能力不同而呈现多样化的分布格局。因此,对不同地区和类群的物种丰富度格局和机制进行研究应具体分析后才能得到可信结论。     

Human,Oceanographic and Habitat Drivers of Central and Western Pacific Coral Reef Fish Assemblages

Coral reefs around US- and US-affiliated Pacific islands and atolls span wide oceanographic gradients and levels of human impact. Here we examine the relative influence of these factors on coral reef fish biomass, using data from a consistent large-scale ecosystem monitoring program conducted by scientific divers over the course of >2,000 hours of underwater observation at 1,934 sites, across ~40 islands and atolls. Consistent with previous smaller-scale studies, our results show sharp declines in reef fish biomass at relatively low human population density, followed by more gradual declines as human population density increased further. Adjusting for other factors, the highest levels of oceanic productivity among our study locations were associated with more than double the biomass of reef fishes (including ~4 times the biomass of planktivores and piscivores) compared to islands with lowest oceanic productivity. Our results emphasize that coral reef areas do not all have equal ability to sustain large reef fish stocks, and that what is natural varies significantly amongst locations. Comparisons of biomass estimates derived from visual surveys with predicted biomass in the absence of humans indicated that total reef fish biomass was depleted by 61% to 69% at populated islands in the Mariana Archipelago; by 20% to 78% in the Main Hawaiian islands; and by 21% to 56% in American Samoa.     

Reef Fishes in Biodiversity Hotspots Are at Greatest Risk from Loss of Coral Species

Coral reef ecosystems are under a variety of threats from global change and anthropogenic disturbances that are reducing the number and type of coral species on reefs. Coral reefs support upwards of one third of all marine species of fish, so the loss of coral habitat may have substantial consequences to local fish diversity. We posit that the effects of habitat degradation will be most severe in coral regions with highest biodiversity of fishes due to greater specialization by fishes for particular coral habitats. Our novel approach to this important but untested hypothesis was to conduct the same field experiment at three geographic locations across the Indo-Pacific biodiversity gradient (Papua New Guinea; Great Barrier Reef, Australia; French Polynesia). Specifically, we experimentally explored whether the response of local fish communities to identical changes in diversity of habitat-providing corals was independent of the size of the regional species pool of fishes. We found that the proportional reduction (sensitivity) in fish biodiversity to loss of coral diversity was greater for regions with larger background species pools, reflecting variation in the degree of habitat specialization of fishes across the Indo-Pacific diversity gradient. This result implies that habitat-associated fish in diversity hotspots are at greater risk of local extinction to a given loss of habitat diversity compared to regions with lower species richness. This mechanism, related to the positive relationship between habitat specialization and regional biodiversity, and the elevated extinction risk this poses for biodiversity hotspots, may apply to species in other types of ecosystems.     

Predicting Coral Species Richness: The Effect of Input Variables,Diversity and Scale

Coral reefs are facing a biodiversity crisis due to increasing human impacts, consequently, one third of reef-building corals have an elevated risk of extinction. Logistic challenges prevent broad-scale species-level monitoring of hard corals; hence it has become critical that effective proxy indicators of species richness are established. This study tests how accurately three potential proxy indicators (generic richness on belt transects, generic richness on point-intercept transects and percent live hard coral cover on point-intercept transects) predict coral species richness at three different locations and two analytical scales. Generic richness (measured on a belt transect) was found to be the most effective predictor variable, with significant positive linear relationships across locations and scales. Percent live hard coral cover consistently performed poorly as an indicator of coral species richness. This study advances the practical framework for optimizing coral reef monitoring programs and empirically demonstrates that generic richness offers an effective way to predict coral species richness with a moderate level of precision. While the accuracy of species richness estimates will decrease in communities dominated by species-rich genera (e.g. Acropora), generic richness provides a useful measure of phylogenetic diversity and incorporating this metric into monitoring programs will increase the likelihood that changes in coral species diversity can be detected.     

A View From Both Ends: Shifts in Herbivore Assemblages Impact Top-Down and Bottom-Up Processes on Coral Reefs

Ecosystems - A fundamental goal in ecology is to understand the role of consumers in top-down (TD) and bottom-up (BU) processes that affect the functioning of ecosystems. Consumers ingest organic...     

Ecohydrodynamics of Cold-Water Coral Reefs: A Case Study of the Mingulay Reef Complex (Western Scotland)

Ecohydrodynamics investigates the hydrodynamic constraints on ecosystems across different temporal and spatial scales. Ecohydrodynamics play a pivotal role in the structure and functioning of marine ecosystems, however the lack of integrated complex flow models for deep-water ecosystems beyond the coastal zone prevents further synthesis in these settings. We present a hydrodynamic model for one of Earth''s most biologically diverse deep-water ecosystems, cold-water coral reefs. The Mingulay Reef Complex (western Scotland) is an inshore seascape of cold-water coral reefs formed by the scleractinian coral Lophelia pertusa. We applied single-image edge detection and composite front maps using satellite remote sensing, to detect oceanographic fronts and peaks of chlorophyll a values that likely affect food supply to corals and other suspension-feeding fauna. We also present a high resolution 3D ocean model to incorporate salient aspects of the regional and local oceanography. Model validation using in situ current speed, direction and sea elevation data confirmed the model''s realistic representation of spatial and temporal aspects of circulation at the reef complex including a tidally driven current regime, eddies, and downwelling phenomena. This novel combination of 3D hydrodynamic modelling and remote sensing in deep-water ecosystems improves our understanding of the temporal and spatial scales of ecological processes occurring in marine systems. The modelled information has been integrated into a 3D GIS, providing a user interface for visualization and interrogation of results that allows wider ecological application of the model and that can provide valuable input for marine biodiversity and conservation applications.     

Patterns in Fluorescence over a Caribbean Reef Slope: the Coral Genus Madracis

Patterns of fluorescence and colony tissue, colour were studied (field observations and epifluorescence microscopy) in six species of the coral genus Madracis over depth from 10 to 60 m at a reef slope in Curaçao. Two functions showed up: (1) Decrease in number of colourmorphs (n = 25) with depth suggests a photo-protective function where short wavelengths (e.g. UV) are transformed to long wavelengths, (2) Green fluorescence, observed in four species over their entire depth range, transforms radiation to wavelengths useful for photosynthesis. The observed patterns in fluorescence between species did not correspond to the current taxonomic classification. Our results do not support the usefulness of fluorescence as a taxonomic tool in corals.     

Patterns of Species Richness and Composition in Re-Created Grassland

The success of many prairie restorations is not well documented. A restoration begun in 1975 at the Fermi National Accelerator Laboratory near Chicago, Illinois allows assessment of restoration efforts as well as changes through time. Data are presented on species richness and composition for 13 restorations planted in successive years between 1975 and 1990 and two remnant prairies. Presence of species was recorded using a stratified random design. Species richness at several scales and non‐metric multidimensional scaling ordination were used to assess trends in the vegetation. Species richness declined through time at all scales examined and was always less in the restored prairies than that found in the remnant prairies. Species composition changed with time but not in the direction of the composition found in the remnants. Our understanding of the maintenance of species richness is not sufficient to allow the re‐creation of patterns of species found in remnant grassland communities.     

Species Richness and Evidence of Random Patterns in Assemblages of South American Titanosauria during the Late Cretaceous (Campanian–Maastrichtian)

The Titanosauria were much diversified during the Late Cretaceous, but paleobiological information concerning these sauropods continues to be scarce and no studies have been conducted utilizing modern methods of community analysis to infer possible structural patterns of extinct assemblages. The present study sought to estimate species richness and to investigate the existence of structures in assemblages of the South American Titanosauria during the Late Cretaceous. Estimates of species richness were made utilizing a nonparametric estimator and null models of species co-occurrences and overlapping body sizes were applied to determine the occurrence of structuring in this assemblages. The high estimate of species richness (n = 57) may have been influenced by ecological processes associated with extinction events of sauropod groups and with the structures of the habitats that provided abundant support to the maintenance of large numbers of species. The pseudocommunity analysis did not differ from that expected by chance, indicating the lack of structure in these assemblages. It is possible that these processes originated from phylogenetic inertia, associated with the occurrence of stabilized selection. Additionally, stochastic extinction events and historical factors may also have influenced the formation of the titanosaurian assemblages, in detriment to ecological factors during the Late Cretaceous. However, diagenetic and biostratinomic processes, influenced by the nature of the sedimentary paleoenvironment, could have rendered a random arrangement that would make assemblage structure undetectable.     

Patterns of Species Richness at Varying Scales in Western Kenya: Planning for Agroecosystem Diversification

Agroforestry tree domestication research is geared at promoting diversification of on-farm tree species composition. A survey was conducted in western Kenya with the objective of exploring possibilities for diversification for a particular agroecosystem, involving a complete tree census, tree measurement and collection of ethnobotanical information in 201 small-scale farms. Various approaches to landscape diversification were explored, including random distribution of trees to increase alpha richness and species richness at higher scales in the landscape, and random distribution of species composition over villages to increase the average richness of villages. The results showed that random distribution would result in increments of average species richness in the landscape, without requiring increments of total and average abundance. A new, fast and exact method of calculating site-based species accumulation curves was presented. The method yielded results that were extremely close to classical algorithms using 10,000 randomisations. Four use-groups (beverage, fodder, charcoal and soil fertility enhancement) were identified as use-groups with alpha richness smaller than one species, but only beverage and fodder had lowest richness at all scales (fruit and construction wood joined the four use-groups of lowest average species richness at higher scales in the landscape). The novel approaches used in this study could be used in future biodiversity studies on species accumulation patterns, or on spatial distribution patterns of species richness in a landscape.