A simulation model of island reef morphology: the effects of sea level fluctuations,growth, subsidence and erosion

Glacioeustatic sea level fluctuations continually cover and expose reefs, alternately allowing growth or erosion to operate. In a simulation model we examine the simultaneous effects of sea level change, island subsidence, reef growth, subaerial erosion, marine backwearing, and fluvial erosion (from central highlands) on reef development. Using values obtained from the literature, we vary the rates of these processes and compare the reefs produced. Our results indicate that subaerial erosion, subsidence and growth are of comparable importance in determining reef morphology. Fore reef terraces, as developed by the model, are primarily drowned growth features; marine backwearing is of little importance in their development. Reef terraces form readily at depths that never had a stable sea stand, their depth is influenced by growth, subaerial erosion, and subsidence rates. Thus reef terraces often do not indicate former sea stands. We examine the causes of reef drowning and attribute it primarily to rapid subsidence and subaerial erosion, not to truncation through marine backwearing. We propose that reefs deeply submerged today are not necessarily drowned out, but may be vertically stable through many sea level cycles. Fluvial erosion is likely an important agent of lagoon formation on high islands in areas with high erosion rates.     

ECOLOGY AND MORPHOLOGY OF RECENT CORAL REEFS

1. The classical ‘coral reef problem’ concerned the geological relationships of reefs as major topographical features; modern coral studies consider reefs both as complex biological systems of high productivity and as geological structures forming a framework for and being modified by coral growth. 2. Deep borings in reefs have conclusively confirmed the general arguments of Darwin, that oceanic reefs developed by progressive subsidence of their foundations. Darwin failed to take account of Pleistocene changes in sea level and their effect on the present surface features of reefs. Daly's alternative ‘glacial control theory’ was based on false assumptions concerning marine erosion rates during glacial periods, but if sea level during the Holocene was higher than at present, as Daly also supposed, the effects on reef features would be profound. 3. Reefs are complex biological systems in tropical seas, dominated by scleractinian corals. Coral faunas are larger and more diverse in the Indo-Pacific than in the Atlantic. Hermatypic corals are restricted to shallow water by the light requirements of their symbiotic algae, but temperature is a major control of worldwide distributions. Temperature, salinity and sediment tolerances of corals are wider than formerly supposed, and corals can survive brief emersion except when it coincides with heavy rainfall. Water turbulence is an important ecological control, but difficult to measure. 4. The trophic status of corals is still unclear, but in spite of their anatomical and physiological specialization as carnivores it is likely that they derive some nutrient substances from zooxanthellae. Suggestions that filamentous algae in coral heads play a major part in the economy of the corals have not been supported by later work, but biomass pyramids constructed on the basis by Odum and Odum remain the only ones available. Most reefs are apparently autotrophic, with 1500–3500 g. Carbon being fixed per m.² per year. 5. Few animals eat corals, which may account for their success. Important predators are fish and the echinoderm Acanthaster. Quantitative estimates of biogenic erosion of organic skeletons on reefs are high. Fish affect not only corals but other invertebrates, algae and marine phanerogams. 6. Corals may be killed by ‘dark water’, intense rain or river floodwaters, earth movements, human interference and especially hurricanes. Reef recovery after hurricanes may take 10–20 years. 7. In addition to fringing, barrier and atoll reefs, intermediate types are recognised. The main types may consist of linear reefs or faros. Smaller lagoon reefs include pinnacles, patches and platforms, and submerged knolls. Complex cellular or mesh reef patterns are also found. 8. Reefs are conspicuously zoned, both laterally in response to changing exposure to waves to form windward and leeward reefs, and transversely, as a result of steep environmental gradients across reef flats from sea to lagoon. Topographic and ecological zones may be characterized by particular coral species, but these vary widely from reef to reef. A major distinction can be made between reefs with and without algal ridges, which are common on open-ocean trade-wind reefs, in the Indo-Pacific, but are absent on Caribbean reefs and on Indo-Pacific reefs in more sheltered waters. gorgonians are common on Caribbean reefs, alcyonaceans in the Indo-Pacific. 9. Much of the difficulty in comparing reefs stems from the lack of uniformity in surveying methods. Problems of describing the complex three-dimensional patterns of organisms on reefs have yet to be solved, and hence little progress has been made in explanation of these patterns. Explanation in terms of simple environmental controls is inadequate. 10. Understanding the distribution of corals is made more difficult both by taxo-nomic problems and by the plasticity of growth form in different situations. 11. Growth of corals and reefs may be estimated by measuring the growth of individual colonies, measuring rates of calcium carbonate deposition in the skeleton, measuring topographic change on the reef and deducing net rates of reef growth from geological evidence. Massive corals may increase in diameter by 1 cm./year, branches of branching corals may increase in length by 10 cm./year. Study of deposition rates shows variation within colonies, between species, in light and dark, and seasonally. Rates of reef growth extrapolated from colony measurements reach 2–5 cm./year, and contrast with figures as low as 0–02 cm/year averaged over 70 million years from borehole data. Both colony growth rates and geological data suggest worldwide variations in rates of reef growth. 12. In spite of clear evidence of long-continued subsidence, present surface features of reefs, often only thinly veneered by modern corals, have been much affected by recent sea level fluctuations. Many slightly raised reefs at 2–10 m. above sea level date at 90–160 thousand years B.P.; there is evidence for a sea level at about the present level at 30–35 thousand years B.P.; and controversy continues over whether sea level has stood higher than the present at any time since the last sea level rise began about 20,000 years ago. Evidence from many reefs suggests a slightly higher sea level in the last 4000 years, but on other reefs such evidence is lacking. 13. Several reef features (submerged terraces, groove-spur systems, algal ridge, reef flat, reef blocks and reef islands) have been interpreted either as relict features dating from a higher sea level in the last 5000 years, or contemporary features developed in response to present processes. In some cases the evidence is equivocal; in others it is clear that diverse features are being grouped together under the same name. If such features are referable to a higher sea level, this may have been of last Interglacial or even Interstadial age rather than Holocene. 14. A reef consists of a rigid framework defining several major depositional environments within and around it. Sediments are of biological, mainly skeletal origin, except in unusual environments such as the Bahama Banks. The characteristics of sediments derived from organisms depend partly on the breakdown patterns of particular skeletons, partly on transportation and sorting processes. Fine sediments may be either detrital, or physicochemical precipitates. 15. Organisms affect sediments after deposition, by disturbance, transportation and probably comminution. Fish and holothurians have been studied in detail. 16. While new theories of coral reefs are proposed from time to time, the need is less for new theories than for standardised procedures to ensure comparability of reef studies and the identification of variations in reefs both on local and regional scales. While reefs as biological systems adjust relatively rapidly to changes, reefs as geological systems adjust much more slowly. Because of the magnitude and recency of Pleistocene fluctuations in sea level, many biological features of reefs are out of phase with inherited geological features, and this had led to much controversy.     

Reef coral diversity and global change

Regional anthropogenic processes such as pollution, dredging, and overfishing on coral reefs currently threaten the biodiversity of stony corals and other reef-associated organisms. Global climate change may interact with anthropogenic processes to create additional impacts on coral diversity in the near future. In order to predict these changes, it is necessary to understand the magnitude and causes of variation in scleractinian coral diversity throughout their 240 million year history. The fossil record documents long periods of speciation in corals, interrupted repeatedly by events of mass extinction. Some of these events relate clearly to changes in global climate. Diversity in reef corals has increased since their last period of extinction at the end of the Cretaceous (65 My bp ), and is still rising. During the last 8 million years, the fragmentation of the once pantropical Tethys Sea separated corals into two major biogeographical provinces. Periods of glaciation also have caused major changes in sea level and temperature. Accumulated evidence supports the theory that relative habitat area and changing patterns of oceanic circulation are mainly responsible for the two observed centres of recent coral diversity at the western tropical margins of the Atlantic and Pacific oceans. At predicted rates of climate change in the near future, coral reefs are likely to survive as an ecosystem. Increases in sea level may actually benefit corals and lead to regional increases in diversity if new habitat area on back reefs is opened to increased water circulation and thus coral dispersal. Rising temperature may cause higher rates of coral mortality and even local extinction in isolated, small populations such as those on oceanic islands. The effects of increases in ultraviolet radiation (UV) are largely unknown, but likely to be negative. UV may damage planktonic coral propagules in oceanic surface waters and thus decrease rates of gene flow between coral populations. This may result in increased local extinctions, again with the strongest impact on widely separated reefs with small coral populations. The largest threats to coral diversity are regional anthropogenic impacts, which may interact with global climate change to exacerbate rates of local species extinctions. Centres of high reef coral diversity coincide with human population centres in south-east Asia and the Caribbean, and thus the greatest potential for species loss lies in these geographical areas.     

Metapopulation vicariance explains old endemics on young volcanic islands

Terrestrial plants and animals on oceanic islands occupy zones of volcanism found at intraplate localities and along island arcs at subduction zones. The organisms often survive as metapopulations, or populations of separate sub‐populations connected by dispersal. Although the individual islands and their local subpopulations are ephemeral and unstable, the ecosystem dynamism enables metapopulations to persist in a region, more or less in situ, for periods of up to tens of millions of years. As well as surviving on systems of young volcanic islands, metapopulations can also evolve there; tectonic changes can break up widespread insular metapopulations and produce endemics restricted to fewer islands or even a single island. These processes explain the presence of old endemic clades on young islands, which is often reported in molecular clock studies, and the many distribution patterns in island life that are spatially correlated with tectonic features. Metapopulations can be ruptured by sea floor subsidence, and this occurs with volcanic loading in zones of active volcanism and with sea floor cooling following its production at mid‐ocean ridges. Metapopulation vicariance will also result if an active zone of volcanism is rifted apart. This can be caused by the migration of an arc (by slab rollback) away from a continent or from another subduction zone, by the offset of an arc at transform faults and by sea floor spreading at mid‐ocean ridges. These mechanisms are illustrated with examples from islands in the Caribbean and the Pacific. Endemism on oceanic islands has usually been attributed to chance, long‐distance dispersal, but the processes discussed here will generate endemism on young volcanic islands by vicariance.     

Model of fringing reef development in response to progressive sea level fall over the last 7000 years – (Kikai-jima, Ryukyu Islands, Japan)

 Kikai-jima in the central Ryukyu Islands of Japan is fringed by exposed terraces of Holocene reefs, which formed as a result of periodic local tectonic uplift associated with subduction/collision. The terraces form four topographically distinct features (TI-IV) around the island and represent reefs that grew to sea level at 9000–6065 y BP, 6065–3390 y BP, 3790–2630 y BP, and 2870 to 1550 y BP. The modern reef terrace has been growing since approximately 1550 y BP. The reef terraces were uplifted sequentially around 6050 y BP (4 m), 3390–3790 y BP (2.5 m), 2630–2870 y BP (1 m) and 1550 y BP (2.5 m). Five sites were studied to define reef development in response to periodic relative sea level fall and different stillstand recovery periods. Thirty coral genera and 70 species were recorded from four distinct shallow reef flat to upper reef slope and one deeper reef slope coral assemblage. Significant lateral variations in total coral abundance, genera number, diversity, and the coverage density of Acropora spp. and Faviids occur both within and between the terraces. Stratigraphically, drill core and outcrop data recorded shallowing upward sequences characterised by tabulate Acropora spp. overlying massive Porites sp. and Faviids. The biological variations may represent growth strategies responding to initial colonisation, episodic perturbation (relative sea level fall) and differing recovery times during stillstands, and indicate a reef ecosystem stable and strong enough to recover after substantial perturbations. However, this study suggests that relatively small geological changes have had substantial biological effects, and modelling indicates that such changes would have been more profound had a third factor, such as substrate angle, varied more dramatically. In such a case, the drowning growth strategy exhibited in the drill core transect may have been more prevalent, and reefs would be struggling to grow around Kikai-jima today. Accepted: 27 May 1998     

DO GEOLOGICAL OR CLIMATIC PROCESSES DRIVE SPECIATION IN DYNAMIC ARCHIPELAGOS? THE TEMPO AND MODE OF DIVERSIFICATION IN SOUTHEAST ASIAN SHREWS

Geological and climatic processes potentially alter speciation rates by generating and modifying barriers to dispersal. In Southeast Asia, two processes have substantially altered the distribution of land. Volcanic uplift produced many new islands during the Miocene–Pliocene and repeated sea level fluctuations during the Pleistocene resulted in intermittent land connections among islands. Each process represents a potential driver of diversification. We use a phylogenetic analysis of a group of Southeast Asian shrews ( Crocidura ) to examine geographic and temporal processes of diversification. In general, diversification has taken place in allopatry following the colonization of new areas. Sulawesi provides an exception, where we cannot reject within-island speciation for a clade of eight sympatric and syntopic species. We find only weak support for temporally declining diversification rates, implying that neither volcanic uplift nor sea level fluctuations had a strong effect on diversification rates. We suggest that dynamic archipelagos continually offer new opportunities for allopatric diversification, thereby sustaining high speciation rates over long periods of time, or Southeast Asian shrews represent an immature radiation on a density-dependent trajectory that has yet to fill geographic and ecological space.     

Estimation of photosynthesis and calcification rates at a fringing reef by accounting for diurnal variations and the zonation of coral reef communities on reef flat and slope: a case study for the Shiraho reef, Ishigaki Island, southwest Japan

Seven coral reef communities were defined on Shiraho fringing reef, Ishigaki Island, Japan. Net photosynthesis and calcification rates were measured by in situ incubations at 10 sites that included six of the defined communities, and which occupied most of the area on the reef flat and slope. Net photosynthesis on the reef flat was positive overall, but the reef flat acts as a source for atmospheric CO₂, because the measured calcification/photosynthesis ratio of 2.5 is greater than the critical ratio of 1.67. Net photosynthesis on the reef slope was negative. Almost all excess organic production from the reef flat is expected to be effused to the outer reef and consumed by the communities there. Therefore, the total net organic production of the whole reef system is probably almost zero and the whole reef system also acts as a source for atmospheric CO₂. Net calcification rates of the reef slope corals were much lower than those of the branching corals. The accumulation rate of the former was approximately 0.5 m kyr⁻¹ and of the latter was ~0.7–5 m kyr⁻¹. Consequently, reef slope corals could not grow fast enough to keep up with or catch up to rising sea levels during the Holocene. On the other hand, the branching corals grow fast enough to keep up with this rising sea level. Therefore, a transition between early Holocene and present-day reef communities is expected. Branching coral communities would have dominated while reef growth kept pace with sea level rise, and the reef was constructed with a branching coral framework. Then, the outside of this framework was covered and built up by reef slope corals and present-day reefs were constructed.     

Tectonic stability since the last interglacial offsets the Glorieuses Islands from the nearby Comoros archipelago

The four fossil terraces of the Glorieuses Islands are described, and new dates are provided to resolve their stratigraphy, depositional setting, and tectonic behavior. Most outcrops consist of a single sedimentary unit that represents the remains of an extensive reef flat dominated by Isopora palifera corals. At Lys Island, this unit is locally overlain by dipping layered beds composed of Halimeda segments, tentatively interpreted as storm overwash. Reliable U/Th dates obtained from corals sampled from the fossil outcrops mostly fall between 127 and 123 kyr, suggesting that these reefs formed exclusively during the first MIS-5e sea-level highstand, when sea level reached +3 m. The mean elevation of these terraces being +4.5 m, an uplift of 0.012 ± 0.002 mm year^?1 is inferred. This relative tectonic stability contrasts with the subsidence reported from Mayotte Island, suggesting a different geologic setting for the nearby Comoros and Glorieuses archipelagoes.     

Shoreline changes in a rising sea level context: The example of Grande Glorieuse,Scattered Islands,Western Indian Ocean

This paper provides baseline data on absolute and relative sea level variations and shoreline changes in the Scattered Islands region of the Indian Ocean, based on aerial image analysis, satellite altimetry and field observations and in situ measurements from the 2009 and 2011 TAAF scientific expeditions. The analysis shows the importance of regular observations and monitoring of these islands to better understand reef island responses to climate stressors. We show that Grande Glorieuse Island has increased in area by 7.5 ha between 1989 and 2003, predominantly as a result of shoreline accretion: accretion occurred over 47% of shoreline length, whereas 26% was stable and 28% was eroded. Topographic transects and field observations show that the accretion is due to sediment transfer from the reef outer slopes to the reef flat and then to the beach. This accretion occurred in a context of sea level rise: sea level has risen by about 6 cm in the last twenty years and the island height is probably stable or very slowly subsiding. This island expansion during a period of rising sea level demonstrates that sea level rise is not the primary factor controlling the shoreline changes. This paper highlights the key role of non-climate factors in changes in island area, especially sediment availability and transport. We also evidence rotation of the island, underscoring the highly dynamic nature of reef islands.     

The importance of historical ecology for interpreting evolutionary processes in plants of oceanic islands

Oceanic islands and archipelagos are natural laboratories for investigating patterns and processes of evolution. Islands change with the course of time, resulting in a dynamic ontogeny over millions of years. The combined forces of tectonic plate subsidence and erosion from waves, wind, and rainwater bring about substantial geomorphological change over millions of years, until islands eventually disappear under the sea. Added to these long‐term natural changes to the environment of the islands are the changes caused by human activities in recent centuries. After humans reach a previously unpopulated island, they utilize the natural resources for their own survival, cutting forests for making houses, boats, and firewood. The size of the human population and the length of time on the island determine the degree of environmental impact. Evolutionary processes in plants of oceanic islands take place during ontogeny of the islands, resulting in population divergence, speciation, and hybridization. Due to the dramatic alterations suffered by many islands over millions of years, the present patterns of distribution and ecology of species within endemic groups may have little to do with the patterns when the species originated. Understanding these environmental changes is fundamental to infer a founder effect, reasons for levels of genetic variation within and among populations, and modes of speciation. Special caution must be exercised while making comparisons between groups located on islands of different geological ages and that have endured differing environmental modifications from humans. Examples are provided from the Juan Fernández Archipelago and Lord Howe Island.     

The impact of the Mid-Pleistocene Transition on the composition of submerged reefs of the Maui Nui Complex,Hawaii

The submarine reef terraces (L1–L12) of the Maui Nui Complex (MNC—the islands of Lanai, Molokai, Maui and Kahoolawe) in Hawaii provide a unique opportunity to investigate the impact of climate and sea level change on coral reef growth by examining changes in reef development through the Mid-Pleistocene Transition (900–800 ka). We present an analysis of the biological and sedimentary composition of the reefs that builds directly on recently published chronological and morphological data. We define nine distinct limestone facies and place them in a spatial and stratigraphic context within 12 reef terraces using ROV and submersible observations. These include oolitic, two coral reef, two coralline algal nodule, algal crust, hemi-pelagic mud, bioclastic and peloidal mud facies. These facies characterise environments from high energy shallow water coral reef crests to low energy non-reefal deep-water settings. Combining the bottom observations and sedimentary facies data, we report a shift in the observed sedimentary facies across the submerged reefs of the MNC from dominant shallow coral reef facies on the deep reefs to coralline algae dominated exposed outcrop morphology on the shallower reefs. We argue that this shift is a reflection of the change in period and amplitude of glacioeustatic sea level cycles (41 kyr and 60–70 m to 100 kyr and 120 m) during the Mid-Pleistocene Transition (MPT, ~ 800 ka), coupled with a slowing in the subsidence rate of the complex. The growth of stratigraphically thick coral reef units on the deep Pre-MPT reefs was due to the rapid subsidence of the substrate and the shorter, smaller amplitude sea level cycles allowing re-occupation and coral growth on successive cycle low-stands. Longer, larger amplitude sea level cycles after the MPT combined with greater vertical stability at this time produced conditions conducive to deep-water coralline algae growth which veneered the shallower terraces. Additionally, we compare reef development both within the MNC, and between the MNC and Hawaii. Finally we suggest that climatic forcings such as sea-surface temperature and oceanographic currents may also have influenced the distribution of coral species within the sample suite, e.g., the disappearance of the Acropora genus from the Maui Nui Complex in the Middle Pleistocene.     

Cycles of coral reef ‘turn‐on’, rapid growth and ‘turn‐off’ over the past 8500 years: a context for understanding modern ecological states and trajectories

Human activities threaten reef ecosystems globally, forcing ecological change at rates and scales regarded as unprecedented in the Holocene. These changes are so profound that a cessation of reef accretion (reef ‘turn‐off’) and net erosion of reef structures is argued by many as the ultimate and imminent trajectory. Here, we use a regional scale reef growth dataset, based on 76 core records (constrained by 211 radiometric dates) from 22 reefs along and across the inner‐shelf of the Great Barrier Reef, Australia, to examine the timing of different phases of reef initiation (‘turn‐on’), growth and ‘turn‐off’ during the Holocene. This dataset delineates two temporally discrete episodes of reef‐building over the last 8500 years: the first associated with the Holocene transgression‐early highstand period [～8.5–5.5 k calibrated years bp (cal ybp )]; the second since ～2.3 k cal ybp . During both periods, reefs accreted rapidly to sea level before entering late evolutionary states – states naturally characterized by reduced coral cover and low accretion potential – and a clear hiatus occurs between these reef‐building episodes for which no records of reef initiation exist. These transitions mimic those projected under current environmental disturbance regimes, but have been driven entirely by natural forcing factors. Our results demonstrate that, even through the late Holocene, reef health and growth has fluctuated through cycles independent of anthropogenic forcing. Consequently, degraded reef states cannot de facto be considered to automatically reflect increased anthropogenic stress. Indeed, in many cases degraded or nonaccreting reef communities may reflect past reef growth histories (as dictated by reef growth–sea level interactions) as much as contemporary environmental change. Recognizing when changes in reef condition reflect these natural ‘turn‐on’– growth –‘turn‐off’ cycles and how they interact with on‐going human disturbance is critical for effective coral reef management and for understanding future reef ecological trajectories.     

Fishing degrades size structure of coral reef fish communities

Fishing pressure on coral reef ecosystems has been frequently linked to reductions of large fishes and reef fish biomass. Associated impacts on overall community structure are, however, less clear. In size‐structured aquatic ecosystems, fishing impacts are commonly quantified using size spectra, which describe the distribution of individual body sizes within a community. We examined the size spectra and biomass of coral reef fish communities at 38 US‐affiliated Pacific islands that ranged in human presence from near pristine to human population centers. Size spectra ‘steepened’ steadily with increasing human population and proximity to market due to a reduction in the relative biomass of large fishes and an increase in the dominance of small fishes. Reef fish biomass was substantially lower on inhabited islands than uninhabited ones, even at inhabited islands with the lowest levels of human presence. We found that on populated islands size spectra exponents decreased (analogous to size spectra steepening) linearly with declining biomass, whereas on uninhabited islands there was no relationship. Size spectra were steeper in regions of low sea surface temperature but were insensitive to variation in other environmental and geomorphic covariates. In contrast, reef fish biomass was highly sensitive to oceanographic conditions, being influenced by both oceanic productivity and sea surface temperature. Our results suggest that community size structure may be a more robust indicator than fish biomass to increasing human presence and that size spectra are reliable indicators of exploitation impacts across regions of different fish community compositions, environmental drivers, and fisheries types. Size‐based approaches that link directly to functional properties of fish communities, and are relatively insensitive to abiotic variation across biogeographic regions, offer great potential for developing our understanding of fishing impacts in coral reef ecosystems.     

Implications of reef ecosystem change for the stability and maintenance of coral reef islands

Coral reef islands are among the most vulnerable environments on Earth to climate change because they are low lying and largely constructed from unconsolidated sediments that can be readily reworked by waves and currents. These sediments derive entirely from surrounding coral reef and reef flat environments and are thus highly sensitive to ecological transitions that may modify reef community composition and productivity. How such modifications – driven by anthropogenic disturbances and on‐going and projected climatic and environmental change – will impact reef island sediment supply and geomorphic stability remains a critical but poorly resolved question. Here, we review the unique ecological–geomorphological linkages that underpin this question and, using different scenarios of environmental change for which reef sediment production responses can be projected, explore the likely resilience of different island types. In general, sand‐dominated islands are likely to be less resilient than those dominated by rubble grade material. However, because different islands typically have different dominant sediment constituents (usually either coral, benthic foraminifera or Halimeda) and because these respond differently to individual ecological disturbances, island resilience is likely to be highly variable. Islands composed of coral sands are likely to undergo major morphological change under most near‐future ecological change scenarios, while those dominated by Halimeda may be more resilient. Islands composed predominantly of benthic foraminifera (a common state through the Pacific region) are likely to exhibit varying degrees of resilience depending upon the precise combination of ecological disturbances faced. The study demonstrates the critical need for further research bridging the ecological–geomorphological divide to understand: (1) sediment production responses to different ecological and environmental change scenarios; and (2) dependant landform vulnerability.     

Mid-oceanic carbonate platforms as oceanic dipsticks: examples from the Pacific

Framebuilders of Cenozoic coral reefs are limited by their photic requirements to the contemporaneous sea-level, and therefore shallow water reef facies are reliable paleo sea-level indicators. Sea-level lowstands leave no record on coral reefs in areas subject to tectonic uplift, such as the Huon Peninsula, New Guinea, but are recorded by coral reefs in areas subject to tectonic subsidence. A eustatic sea-level fall which exceeds the rate of subsidence subaeriallyexposes the upper section of the reef complex, creating a meteoric ground water system whose diagenetic imprint on the reef carbonates offers a good indicator of a sea-level stillstand. Cenozoic reef platforms thus may contain records of sea-level fluctuations, whether eustatic and global, or tectonic and local. Those reef platforms which developed on seamounts formed in mid-oceanic plate settings are particularly useful for the study of eustatic sea-level changes because their subsidence history is relatively simple, and the tectonic factor can be accounted for when estimating the eustatic sea-level component. Conventional petrographic and biostratigraphic methods used to delineate erosional unconformities in Cenozoic carbonate sections are often deficient. We demonstrate here that stable oxygen and carbon isotopes of the carbonates can reveal the location of both the exposure surface and the paleo water table with greater confidence on account of the specific imprint of meteoric diagenesis. In addition, the⁸⁷Sr/⁸⁶Sr isotope technique offers a promising dating tool of disconformities linked to sea-level lowstands with a resolution superior to the conventional biostratigraphic techniques. Although oxygen, carbon, and strontium isotopes monitor different aspects of global sea-level changes, when used in conjunction they provide deeper insights into the past than either one could achieve alone. Examples from previous and ongoing studies of Pacific mid-oceanic carbonate platforms illustrate the potential of the isotope techniques to unravel sea-level changes. At Midway Atoll, stable carbon and oxygen isotopes along with lithologic and biostratigraphic data suggest a sharp eustatic sealevel fall during the Early Miocene and a series of rapid, brief eustatic fluctuations during the Pliocene-Quaternary. The frequency and timing of the latter is supported by sea-level data from Enewetak Atoll obtained on the basis of detailed strontium isotopes and lithology. The Enewetak data also indicate a series of rapid, brief eustatic fluctuations around the Early-Middle Miocene boundary. At Niue, a carbonate platform about 500 km south of Samoa, oxygen, carbon, and strontium isotope records cover the critical interval of the Miocene-Pliocene boundary and show two distinct disconformities. The mid-oceanic carbonate platforms offer a testing ground of Vail-Haq type eustatic sea-level curves derived primarily from sections along passive continental margins and continental interiors. We show that Neogene sea-level data obtained from Midway, Enewetak, and Niue differ from Vail and Hardenbol's contemporaneous sea-level curve and support Haq et al.'s version.     

热带珊瑚礁区海参的生境选择与生态作用

珊瑚礁生态系统是一个高生产力、高生物多样性的特殊海洋生态系统,具有为生物提供栖息地、参与生物地球化学循环、防浪护岸、指示水体污染程度等生态功能。珊瑚礁生态系统的突出特点是其生境异质性很高,各种各样的生境斑块为种类繁多、习性各异的游泳和底栖生物提供栖息场所,这些礁栖生物通过参与各项生态过程而形成各种特定的功能群,共同完成重要的生态功能。在热带珊瑚礁生态系统中,海参是大型底栖动物区系的重要一员。种类繁多的海参具有各自不同的生境选择特征,通过摄食、运动等行为活动发挥着改良底质、促进有机物矿化和营养盐再生等生态作用。近几年来,全球热带海参受人类过度捕捞和珊瑚礁退化的影响而面临资源衰退、物种多样性丧失等问题,深入认识其生态学功能、加强热带海参资源保护迫在眉睫。综述了国内外热带珊瑚礁海参的基础生态学研究进展：海参对珊瑚礁生境斑块呈现显著的偏好选择特征以及种间差异和季节变动,不同生境斑块的食物质量、底质类型和水动力条件是影响海参生境偏好的重要因素;海参通过生物扰动可以改变珊瑚礁生境沉积物的含水量、渗透性、颗粒组成、再矿化率、无机营养物质释放速率以及孔隙水的化学梯度,并增加沉积物中的溶氧浓度、促进溶解...     

Late Devonian carbon isotope stratigraphy and sea level fluctuations, Canning Basin, Western Australia

The Upper Devonian reef complexes of the Canning Basin contain some of the world’s best exposed, continuous stratigraphic sections through the Frasnian-Famennian boundary. The facies distribution and composition of these reef complexes record interactions among sea level changes, sediment supply, ocean chemistry, and paleoecology. Changes in relative sea level produced spatial shifts in reef platform development and regional changes in sediment supply that can be correlated across facies boundaries using a combination of sequence stratigraphy, biostratigraphy, and carbon isotope stratigraphy. During the lowstand interval below the Frasnian-Famennian boundary, the reef margin advanced down the reef slope in shallow-water environments, and siliciclastics locally dominated in the marginal slope environment. Compilation of a broad late Frasnian to early Famennian sequence stratigraphic framework for the Canning Basin demonstrates that transgressive intervals correlate to positive carbon isotopic excursions within the basin. These isotopic shifts also can be correlated to time-equivalent positive carbon isotopic excursions reported from transgressive intervals in Europe. Thus, the late Frasnian transgressions in the Canning Basin were primarily eustatic rather than tectonic in origin, and positive carbon isotopic signatures of the Kellwasser horizons are globally correlative.     

Fine scale site fidelity in sea kraits: implications for conservation

The shores of coral reef islands are major sites for biodiversity, but unfortunately they are also subject to strong anthropogenic disturbances. Indeed vast arrays of organisms live exclusively in these very narrow and well structured zones, many others depend on the rich and diverse micro-habitats for essential part of their life cycle (to reproduce, forage, etc.). Sea kraits are sea snakes that depend on the shore of coral islets; they forage at sea but digest, reproduce and rest on land. They have been killed in extremely large numbers in many places, causing local extinctions. In the current study we demonstrate through recapture and translocation studies that these snakes exhibit a strong and fine-scale fidelity for particular segments of the shore. Consequently, these specific areas should be under strong protection, as it the case for the breeding beaches used by marine mammals, birds or turtles.     

Evidence of exceptional oyster‐reef resilience to fluctuations in sea level

Ecosystems at the land–sea interface are vulnerable to rising sea level. Intertidal habitats must maintain their surface elevations with respect to sea level to persist via vertical growth or landward retreat, but projected rates of sea‐level rise may exceed the accretion rates of many biogenic habitats. While considerable attention is focused on climate change over centennial timescales, relative sea level also fluctuates dramatically (10–30 cm) over month‐to‐year timescales due to interacting oceanic and atmospheric processes. To assess the response of oyster‐reef (Crassostrea virginica) growth to interannual variations in mean sea level (MSL) and improve long‐term forecasts of reef response to rising seas, we monitored the morphology of constructed and natural intertidal reefs over 5 years using terrestrial lidar. Timing of reef scans created distinct periods of high and low relative water level for decade‐old reefs (n = 3) constructed in 1997 and 2000, young reefs (n = 11) constructed in 2011 and one natural reef (approximately 100 years old). Changes in surface elevation were related to MSL trends. Decade‐old reefs achieved 2 cm/year growth, which occurred along higher elevations when MSL increased. Young reefs experienced peak growth (6.7 cm/year) at a lower elevation that coincided with a drop in MSL. The natural reef exhibited considerable loss during the low MSL of the first time step but grew substantially during higher MSL through the second time step, with growth peaking (4.3 cm/year) at MSL, reoccupying the elevations previously lost. Oyster reefs appear to be in dynamic equilibrium with short‐term (month‐to‐year) fluctuations in sea level, evidencing notable resilience to future changes to sea level that surpasses other coastal biogenic habitat types. These growth patterns support the presence of a previously defined optimal growth zone that shifts correspondingly with changes in MSL, which can help guide oyster‐reef conservation and restoration.     

Holocene growth of a mid-Pacific atoll: Tarawa,Kiribati

Cores from ten holes, drilled to a maximum depth of 30 m, on Tarawa atoll in the central Pacific have been utilised in a study of the Holocene development of the atoll. Four dominant lithologies, in descending order, are cay rock, unconsolidated sediment, corals and leached limestone. Petrographic and radiometric age analyses indicate that the Holocene reef has developed on a previous (last interglacial) reef; the latter shows the effects of both vadose and phreatic freshwater diagenesis. Hydrological investigations beneath the present islands indicate the presence of freshwater lenses up to 29 m thick; the modern lenses are unrelated to freshwater diagenetic imprints preserved within the limestones. Vertical accretion rates of 5–8 m/1000 years for the Holocene reef section on Tarawa are significantly higher than rates measured for other Pacific atolls. The dated coral sequences suggest a more rapid rate of sea level rise during the early Holocene, and a relatively earlier stabilisation of sea level than has been suggested previously.