Patterns and possible environmental controls of skeletogenesis of Porites lutea,South Thailand

Eleven fringing reef sites were investigated over a distance of about 50 km in the Phuket Region. There is a wide range in exposure to wave energy, and also water turbidity across the area. Annual increments of growth of shallow-water reef-front colonies of Porites lutea were calculated for the period November 1984 November 1986 using seasonal fluorescent banding (revealed with ultraviolet light) and Alizarin staining. Measurement of linear extension rate, skeletal bulk density, calcification rate, polyp numbers per unit area and colony surface morphology were made and compared. Linear extension rate and skeletal bulk density are inversely related within and between reef sites. Linear extension rate decreases and bulk density increases along a gradient of increasing hydraulic energy of the setting. Calcification (the product of linear extension rate and bulk density), although varying slightly from site to site, does not appear to relate to any obvious environmental inshore-offshore gradient. Skeletal bulk density is the most sensitive discriminator between reef sites, and we suggest that hydraulic energy of the setting is the main control on these spatial variations in skeletogenesis.     

Holocene coral reef accretion in Hawaii: a function of wave exposure and sea level history

 In the high Hawaiian Islands, significant accretion due to coral reef growth is limited by wave exposure and sea level. Holocene coral growth and reef accretion was measured at four stations off Oahu, Hawaii, chosen along a gradient in wave energy from minimum to maximum exposures. The results show that coral growth of living colonies (linear extension) at optimal depths is comparable at all stations (7.7–10.1 mm/y), but significant reef accretion occurs only at wave sheltered stations. At wave sheltered stations in Hanauma Bay and Kaneohe Bay, rates of long term reef accretion are about 2.0 mm/y. At wave exposed stations, off Mamala Bay and Sunset Beach, reef accretion rates are virtually zero in both shallow (1 m) and deeper (optimal) depths (12 m). At wave sheltered stations, such as Kaneohe Bay and Hanauma Bay, Holocene reef accretion is on the order of 10–15 m thick. At wave exposed stations, Holocene accretion is represented by only a thin veneer of living corals resting on antecedent Pleistocene limestone foundations. Modern coral communities in wave exposed environments undergo constant turnover associated with mortality and recruitment or re-growth of fragmented colonies and are rarely thicker than a single living colony. Breakage, scour, and abrasion of living corals during high wave events appears to be the major source of mortality and ultimately limits accretion to wave sheltered environments. Depth is particularly important as a modulator of wave energy. The lack of coral reef accretion along shallow open ocean coastlines may explain the absence of mature barrier reefs in the high Hawaiian Islands. Accepted: 14 May 1998     

Phenotypic plasticity for skeletal growth,density and calcification of <Emphasis Type="Italic">Porites lobata</Emphasis> in response to habitat type

A reciprocal transplant experiment (RTE) of the reef-building coral Porites lobata between shallow (1.5 m at low tide) back reef and forereef habitats on Ofu and Olosega Islands, American Samoa, resulted in phenotypic plasticity for skeletal characteristics. Transplants from each source population (back reef and forereef) had higher skeletal growth rates, lower bulk densities, and higher calcification rates on the back reef than on the forereef. Mean annual skeletal extension rates, mean bulk densities, and mean annual calcification rates of RTE groups were 2.6–9.8 mm year⁻¹, 1.41–1.44 g cm⁻³, and 0.37–1.39 g cm⁻² year⁻¹ on the back reef, and 1.2–4.2 mm year⁻¹, 1.49–1.53 g cm⁻³, and 0.19–0.63 g cm⁻² year⁻¹ on the forereef, respectively. Bulk densities were especially responsive to habitat type, with densities of transplants increasing on the high energy forereef, and decreasing on the low energy back reef. Skeletal growth and calcification rates were also influenced by source population, even though zooxanthella genotype of source colonies did not vary between sites, and there was a transplant site x source population interaction for upward linear extension. Genetic differentiation may explain the source population effects, or the experiment may have been too brief for phenotypic plasticity of all skeletal characteristics to be fully expressed. Phenotypic plasticity for skeletal characteristics likely enables P. lobata colonies to assume the most suitable shape and density for a wide range of coral reef habitats.     

Gender-related differences in the apparent timing of skeletal density bands in the reef-building coral Siderastrea siderea

Density banding in skeletons of reef-building corals is a valuable source of proxy environmental data. However, skeletal growth strategy has a significant impact on the apparent timing of density-band formation. Some corals employ a strategy where the tissue occupies previously formed skeleton during as the new band forms, which leads to differences between the actual and apparent band timing. To investigate this effect, we collected cores from female and male colonies of Siderastrea siderea and report tissue thicknesses and density-related growth parameters over a 17-yr interval. Correlating these results with monthly sea surface temperature (SST) shows that maximum skeletal density in the female coincides with low winter SSTs, whereas in the male, it coincides with high summer SSTs. Furthermore, maximum skeletal densities in the female coincide with peak Sr/Ca values, whereas in the male, they coincide with low Sr/Ca values. Both results indicate a 6-month difference in the apparent timing of density-band formation between genders. Examination of skeletal extension rates also show that the male has thicker tissue and extends faster, whereas the female has thinner tissue and a denser skeleton—but both calcify at the same rate. The correlation between extension and calcification, combined with the fact that density banding arises from thickening of the skeleton throughout the depth reached by the tissue layer, implies that S. siderea has the same growth strategy as massive Porites, investing its calcification resources into linear extension. In addition, differences in tissue thicknesses suggest that females offset the greater energy requirements of gamete production by generating less tissue, resulting in differences in the apparent timing of density-band formation. Such gender-related offsets may be common in other corals and require that environmental reconstructions be made from sexed colonies and that, in fossil corals where sex cannot be determined, reconstructions must be duplicated in different colonies.     

Variability in skeletal bulk densities of common hard corals in Southeast Asia

Coral Reefs - Skeletal density, a measure of a coral’s investment in its structure and a known proxy for climate history, can potentially be used to enhance the precision of reef accretion...     

Effects of monsoon-driven wave action on coral reefs of Guam and implications for coral recruitment

Benthic cyanobacteria can respond rapidly to favorable environmental conditions, overgrow a variety of reef organisms, and dominate benthic marine communities; however, little is known about the dynamics and consequences of such cyanobacterial blooms in coral reef ecosystems. In this study, the benthic community was quantified at the time of coral spawnings in Guam to assess the substrate that coral larvae would encounter when attempting settlement. Transects at 9, 18, and 25-m depths were surveyed at two reef sites before and after heavy wave action driven by westerly monsoon winds. Communities differed significantly between sites and depths, but major changes in benthic community structure were associated with wave action driven by monsoon winds. A shift from cyanobacteria to crustose coralline algae (CCA) accounted for 44% of this change. Coral recruitment on Guam may be limited by substrate availability if cyanobacteria cover large areas of the reef at the time of settlement, and consequently recruitment may in part depend upon wave action from annual monsoon winds and tropical storms which remove cyanobacteria, thereby exposing underlying CCA and other substrate suitable for coral settlement.     

Skeletal adaptations during mammalian reproduction

Remarkable changes occur in the mammalian skeleton prior to, during and after the reproductive cycle. Skeletal changes occur with ovarian maturation and initiation of menses and estrus in adolescence, which may result in a greater accumulation of skeletal mineral in the female vs the male skeleton. There is also some evidence to suggest an excess skeletal mass in young female experimental animals. In early pregnancy, growth, modeling and perhaps suppressed remodeling promote the accumulation of calcium. Some changes may also occur with the transition from pituitary to placental control of the pregnancy. In later pregnancy, an increase in bone turnover appears to coincide with fetal skeletal mineralization. Rapid and important changes occur in the skeleton and mineral metabolism in the transition from pregnancy to lactation as the mammary gland rather than the uterus draws on the maternal calcium stores. Lactational demands are met at least partially by a temporary demineralization of the skeleton, which is associated with increased bone modeling and remodeling. Endochondral growth almost ceases during lactation, but envelope-specific bone modeling and remodeling are greatly increased. This is generally associated with a loss of skeletal mass and density, more apparent at sites with less of a mechanical role (e.g. central metaphysis regions and the endocortical envelope). The post-lactational period is profoundly anabolic with substantial increases in bone formation, but blunted resorption at almost all skeletal envelopes. Skeletal mass is increased during this period and it is associated with improved skeletal mechanical properties. There are several important observations. 1) The nulliparous animal appears to have an excess skeletal mass to perhaps compensate for maternal metabolic inefficiency of the first reproductive cycle. 2) Changes in growth, modeling and remodeling occur at different times and at different skeletal envelopes during the reproductive cycle. These site-specific, temporal changes appear to be adaptations that facilitate the use of skeletal mineral while preserving mechanical competence. 3) After the first reproductive cycle, modeling and remodeling optimize the existing skeletal mass into a structure that better accommodates the prevailing mechanical environment. 4) The post-lactational period is profoundly anabolic and may provide new strategies for preservation of skeletal mass when reproductive capacity ceases.     

Patterns of recruitment of coral reef fishes in a monsoonal environment

The abundance of newly settled recruits of coral reef fishes was monitored at a total of 11 sites at two islands and two coastal locations in the central Philippines for a 20-month period (February 2008 to September 2009) that included two monsoon cycles. Recruitment occurred throughout the year. Most of the abundant species exhibited protracted recruitment seasons. This confirms the expectation of extended breeding of reef fishes at lower latitudes. The annual pattern of recruitment of reef fishes as a group was predictable. Annual fluctuations of sea surface temperature and wind strength largely explained the pattern. Rainfall, however, did not significantly influence the pattern of recruitment. Peaks in density and species richness of recruits occurred during the southwest monsoon and the second inter-monsoonal period of the year (July to October) when temperatures were highest and when most of the sites were sheltered from winds or when winds were weak. Conversely, lowest density and species richness were observed during the northeast monsoon (November to March) when temperatures were lowest and most sites were exposed to winds. The same pattern could also be seen in the recruitment of both damselfishes (Pomacentridae) and wrasses (Labridae), notwithstanding a tenfold difference in abundance of recruits between the two families. The pattern was fairly consistent across most sites, among most of the species that were examined, and between the 2 years that were sampled. This study is one of the few to provide insights into the influence of environmental factors on the recruitment patterns of fishes on Indo-Pacific coral reefs situated at lower latitudes.     

Skeletal development in Acropora cervicornis

Monthly linear extension and calcium carbonate accretion were measured over a year in the Caribbean staghorn coral, Acropora cervicornis. X-radiographs were made of cross sections of branches to analyze radial growth. Correlations were made between parameters of skeletal growth and four environmental parameters monitored over the same sampling periods: temperature, daylight hours, sun hours, plankton abundance. The results indicate that linear extension does not change during the year with the possible exception of April. It is suggested that temperatures outside an optimal range (ca. 26°–29°C for staghorn Acroporas) might cause a decrease in linear extension, however. Specific accretion (mg. mm^-1) does show significant variations through the year. Calcium carbonate accretion (mean specific accretion times mean linear extension, mg. tip^-1) is most strongly correlated with number of sun hours. A comparison is made between diel patterns of extension and accretion and longer term measurements. It is suggested that the accretion process is probably most influenced by some activity influenced by light. There are no annual growth bands in X-radiographs of cross-sections of the branches of A. cervicornis. This may result from secondary infilling in the skeleton.     

Skeletal growth of a spongiose radiolarian Dictyocoryne truncatum in laboratory culture

The stages of skeletal growth of the radiolarian Dictyocoryne truncatum were observed using scanning electron and light microscopy. Four growth stages were recognized through comparison of laboratory-grown individuals with various sized skeletons which were collected by plankton tows near Barbados. Skeletal size is expressed as test height, which is the largest value among the three dimensions between a base line and arm apex of a triangular skeleton. The early stage (stage 1), with a maximum height of less than approximately 120 μm, is a uniformly spongiose, triangular skeleton lacking arms and a patagium. The next stage (stage 2) is a triangular skeleton with slightly concave sides of ca. 120–200 μm size with three arms and a lace-like patagium. Further accretion of silica on the patagium conceals its lace-like structure, making a skeleton (stage 3; ca. 200–280 μm) with a more uniformly spongiose patagium. The mature stage (stage 4) with a maximum height more than ca. 280 μm is a triangular skeleton with convex sides, resulting from an outward growth of the patagium. Additional maturation produces an accretion of spongiose silica on the central part of the skeleton, forming a thickly biconvex skeleton in transverse section. One hundred and one individuals of D. truncatum were cultured in the laboratory at 28°C, 35‰ salinity and 165 μE/m²/s light intensity, with maximum and mean longevity of 37 and 6.4 days, respectively. The mean growth of 57 specimens which grew in the laboratory culture was 24.7 μm, with a maximum of 95 μm. The initial size of cultured specimens ranged from 103 to 325 μm, covering all of the four growth stages. Some specimens grew from one stage to the next stage changing skeletal morphology. The mean growth rate of four specimens which grew from stage 3 to 4 in laboratory culture was 5.4 μm/day (range 4.5–6.5 μm/day). Sporadic growth patterns exhibiting periods of rapid growth (15–20 μm per day) punctuated by intervals of little or no growth were observed in some cultured specimens. The sporadic growth may be related to a physiological rhythm of the organisms rather than environmental factors. The silica-depositing capacity is related largely to the vitality of the organisms. Discolored specimens and specimens with weakly extended axopodia showed no skeletal growth.     

Measurement of luminescence in coral skeletons

Luminescent lines in skeletons of the massive coral Porites record periods when seawater was significantly diluted by land runoff. Records developed from such lines would be useful in a wide range of areas, including climatology, oceanography, civil engineering, agriculture, water resources and reef management. To realise this enormous potential, we built an instrument for routine, reliable recovery of luminescence information from coral skeletons. Skeletal slices were laid on a table that moved in 0.25 mm steps. The coral skeleton was illuminated with ultra-violet light (UV) at 390 nm and luminescent emissions at 490 nm were recorded. Light at 490 nm was then shone on the same 2 mm diameter point on the skeleton and the reflection of 490 nm light was recorded. Luminescent emissions from a point were then standardised by the reflectivity of that same point.Slices cut from three corals that grew at an inshore reef had many strong luminescent lines. Measurements of luminescence in these colonies were nearly identical, both for multiple tracks across one slice and for tracks across slices from the different corals. There was a clear link with discharge from a nearby river. Slices cut from two corals that grew at a reef 56 km offshore had occasional, weak luminescent lines that were also linked with river discharge. Tracks across these slices were similar but the weak luminescence due to river discharge was partially obscured by weak luminescence associated with the annual density banding pattern that characterises massive coral skeletons. The technique recovered excellent information about skeletal luminescence. However, there is a need to gain better understanding of the link between seawater dilution and luminescence and to develop procedures for data processing before the technique can be used to construct useful proxy environmental records.     

Coral reef growth in an era of rapidly rising sea level: predictions and suggestions for long-term research

Coral reef growth is intimately linked to sea level. It has been postulated that over the next century, sea level will rise at a probable average rate of 15 mm/year, in response to fossil fuel emissions, heating, and melting of the Antarctic ice cap. This predicted rate of sea level rise is five times the present modal rate of vertical accretion on coral reef flats and 50% greater than the maximum vertical accretion rates apparently attained by coral reefs. We use these predictions and observations to offer the following hypothesis for reef growth over the next century. The vertical accretion rates of protected reef flats will accelerate from the present modal rate up to the maximum rate, in response to the more rapidly rising sea level. This more rapid vertical accretion rate will be insufficient to keep up with sea level rise, if present predictions prove to be correct. Less protected reef flats will slow their rate of growth as they become inundated and subjected to erosion by progressively larger waves. This projected sea level rise and postulated reef response will provide an opportunity for long-term studies of the response of coral reef systems to a predictable and measurable forcing function. If the scientific benefits from this uncontrolled global experiment are to be maximized, it will be necessary to establish a permanent international coordinating body to assist with the identification and preservation of long-term study sites and to provide guidelines for baseline data surverys, methods selection and comparison, and other procedures and decisions.     

The zonation patterns of Caribbean coral reefs as controlled by wave and light energy input,bathymetric setting and reef morphology: computer simulation experiments

The computer model COREEF was used to simulate variations in the zonation patterns of Caribbean reefs in relation to parameters that affect the magnitude and distribution of wave and light energy. We first developed a simulated standard reef by exposing a simplified profile of the reef at Discovery Bay, Jamaica, to the known wave and light energy conditions to establish a reference coralgal and sedimentological zonation pattern. We then varied 13 parameters related to the wave and light energy input, bathymetric setting, and gross morphology of this reef to determine the effects of each parameter on the zonation pattern. Analysis of the simulation results indicates that submerging the reef or altering the wave or light energy input to the reef produces the greatest modifications of the zonation pattern. Morphological structures that alter a reef's horizontal dimensions only minimally affect the zonation pattern, but those structures that alter a reef's vertical dimensions-particularly steep-sided, wave reflecting structures-can significantly modify the zonation of the structure itself and that of more leeward areas. The more seaward the location of a morphological structure, the more profoundly it can affect the overall reef zonation. If waves break at the reef crest, wave energy conditions in the back reef are greatly reduced and the bottom consists of lower wave energy zones than those found at the same depths in the fore reef. If waves do not break at the crest, the back reef is subjected to almost the same wave conditions that exist in the fore reef, and the zones tend to be similar. The zonation patterns of some existing reefs resemble those of our simulated reefs, but other zonation patterns cannot be reproduced accurately because our simulation experiments do not consider the interactions between multiple parameters found on many existing reefs.     

Sedimentation on Rasdhoo and Ari Atolls, Maldives, Indian Ocean

A first systematic study of composition, texture, and distribution of modern sediments in two Maldivian atolls reveals the predominance of skeletal carbonates. Fragments of corals, calcareous algae, mollusks, benthic foraminifera, and echinoderms are identified in the grain-size fraction >125 μm. Non-skeletal grains such as cemented fecal pellets and aggregate grains only occur in small percentages. Fragments of skeletal grains, aragonite needles, and nanograins (<1 μm) are found in the grain-size fraction <125 μm. Needles and nanograins are interpreted to be largely of skeletal origin. Five sedimentary facies are distinguished (1–5), for which the Dunham-classification is applied. Fore reef, reef, back reef, as well as lagoonal patch reef and faro areas in both atolls are characterized by the occurrence of coral grainstones (1), which also contain fragments of red coralline algae, the codiacean alga Halimeda, and mollusks. On reef islands, coral-rich sediment is cemented to form intertidal beachrock and supratidal cayrock. Skeletal grains in atoll-interior lagoons are mainly mollusks and foraminifera. The lagoon of Rasdhoo Atoll is covered in the west by mudstones (2), in the center by mollusk packstones (3) and mollusk wackestones (4), and by hard bottoms with corals in the east adjacent to channels through the atoll reef margin. The interior lagoon of Ari Atoll contains mollusk wackestones (4) in the center and mollusk-foraminifer packstones (5). Marginal lagoon areas are characterized by hard bottoms with corals. Facies distribution appears to be an expression of depositional energy, which decreases from the atoll margin towards the center in Ari Atoll, and towards the west in Rasdhoo Atoll. Predominant sediment mineralogies include aragonite and high-magnesium calcite. Mean aragonite content decreases from 90% in coral grainstone to 70–80% in mollusk packstone, mollusk wackestone, and mudstone, and to 50% in mollusk-foraminifer packstone. Stable isotopes of oxygen and carbon in bulk samples range from −3 to −1.5 (δ¹⁸O) and from +0.4 to +3.2 (δ¹³C). It is not possible to delineate facies based on O- and C-isotopes.     

Computerized tomography and skeletal density of coral skeletons

In this paper I describe and discuss the use of medical X-ray computerized tomography (CT) in the study of coral skeletons. CT generates X-ray images along freely chosen sections through the skeleton and offers, as well, the possibility of density measurements based on X-ray attenuation. This method has been applied to measure the skeletal density of the Caribbean reef-building coral Montastrea annularis, from Curaçao, Netherlands Antilles. The observed, non-linear increase of skeletal density with depth can be attributed to decreasing photo-synthetic rates with increasing water depth. A comparison with extension rate measurements shows the inverse relationship between extension rate and skeletal density. CT proves to be aquick and non-destructive method to reveal growth structures (density banding) since it measures skeletal density.     

The seasonal dynamics and persistence of stream macroinvertebrates in Nepal: do monsoon floods represent disturbance?

1. The monsoon causes major flood events in some Himalayan streams, but their seasonal predictability might reduce the resulting disturbance. We assessed seasonal change in the benthos of 16 streams in central Nepal over a gradient of declining rainfall and increasing altitude from 600 to 3800 m. All sites were surveyed on four occasions, two in winter (November) and two pre-monsoon (June), with additional sampling during the monsoon (August) at four low altitude sites. Invertebrate abundance, taxon richness and persistence were assessed at all sites, and density and meso-habitat distribution at the four low altitude sites only.
2. Strong seasonal variation among invertebrates was confined primarily to streams at low altitude (600–800 m) where monsoon rainfall was greatest and catchments were dominated by terraced agriculture. At these sites, a significant reduction in benthic density (on average by 77%) and taxon richness (by 20%) occurred between the winter and pre-monsoon periods, so that invertebrate numbers were already low before the monsoon. A further significant decline occurred in all meso-habitats during the monsoon, but the change in density was small in absolute terms.
3. Persistence in rank abundance was equally low at all sites, but turnover in composition was significantly lower at sites in semi-natural forest than in catchments managed for terracing or alpine pasture.
4. These data provide no evidence that monsoonal floods represent major disturbance, instead supporting the view that the ecological response might reflect an adjustment to predictable flow pattern. However, catchment land use in the Himalaya appears to be a significant source of ecosystem instability, and confounds the simple interpretation of monsoon effects.     

Skeletal muscle: Energy metabolism, fiber types, fatigue and adaptability

Skeletal muscles cope with a large range of activities, from being able to support the body weight during long periods of upright standing to perform explosive movements in response to an unexpected threat. This requires systems for energy metabolism that can provide energy during long periods of moderately increased energy consumption as well as being able to rapidly increasing the rate of energy production more than 100-fold in response to explosive contractions. In this short review we discuss how muscles can deal with these divergent demands. We first outline the major energy metabolism pathways in skeletal muscle. Next we describe metabolic differences between different muscle fiber types. Contractile performance declines during intense activation, i.e. fatigue develops, and we discuss likely underlying mechanisms. Finally, we discuss the ability of muscle fibers to adapt to altered demands, and mechanisms behind these adaptations. The accumulated experimental evidence forces us to conclude that most aspects of energy metabolism involve multiple and overlapping signaling pathways, which indicates that the control of energy metabolism is too important to depend on one single molecule or mechanism.     

Extreme spatial heterogeneity in carbonate accretion potential on a Caribbean fringing reef linked to local human disturbance gradients

The capacity of coral reefs to maintain their structurally complex frameworks and to retain the potential for vertical accretion is vitally important to the persistence of their ecological functioning and the ecosystem services they sustain. However, datasets to support detailed along‐coast assessments of framework production rates and accretion potential do not presently exist. Here, we estimate, based on gross bioaccretion and bioerosion measures, the carbonate budgets and resultant estimated accretion rates (EAR) of the shallow reef zone of leeward Bonaire – between 5 and 12 m depth – at unique fine spatial resolution along this coast (115 sites). Whilst the fringing reef of Bonaire is often reported to be in a better ecological condition than most sites throughout the wider Caribbean region, our data show that the carbonate budgets of the reefs and derived EAR varied considerably across this ~58 km long fringing reef complex. Some areas, in particular the marine reserves, were indeed still dominated by structurally complex coral communities with high net carbonate production (>10 kg CaCO₃ m^?2 year^?1), high live coral cover and complex structural topography. The majority of the studied sites, however, were defined by relatively low budget states (<2 kg CaCO₃ m^?2 year^?1) or were in a state of net erosion. These data highlight the marked spatial heterogeneity that can occur in budget states, and thus in reef accretion potential, even between quite closely spaced areas of individual reef complexes. This heterogeneity is linked strongly to the degree of localized land‐based impacts along the coast, and resultant differences in the abundance of reef framework building coral species. The major impact of this variability is that those sections of reef defined by low‐accretion rates will have limited capacity to maintain their structural integrity and to keep pace with current projections of climate change induced sea‐level rise (SLR), thus posing a threat to reef functioning and biodiversity, potentially leading to trophic cascades. Since many Caribbean reefs are more severely degraded than those found around Bonaire, it is to be expected that the findings presented here are rather the rule than the exception, but the study also highlights the need for similar high spatial resolution (along‐coast) assessments of budget states and accretion rates to meaningfully explore increasing coastal risk at the country level. The findings also more generally underline the significance of reducing local anthropogenic disturbance and restoring framework building coral assemblages. Appropriately focussed local preservation efforts may aid in averting future large‐scale above reef water depth increases on Caribbean coral reefs and will limit the social and economic implications associated with the loss of reef goods and services.