Effect of acclimatization to low temperature and reduced light on the response of reef corals to elevated temperature

This study tested the effects of acclimatization on the response of corals to elevated temperature, using juvenile massive Porites spp. and branching P. irregularis from Moorea (W149°50′, S17°30′). During April and May 2006, corals were acclimatized for 15 days to cool (25.7°C) or ambient (27.7°C) temperature, under shaded (352 μmol photons m⁻² s⁻¹) or ambient (554 μmol photons m⁻² s⁻¹) natural light, and then incubated for 7 days at ambient or high temperature (31.1°C), under ambient light (659 μmol photons m⁻² s⁻¹). The response to acclimatization was assessed as biomass, maximum dark-adapted quantum yield of PSII (F _v/F _m), and growth, and the effect of the subsequent treatment was assessed as F _v/F _m and growth. Relative to the controls (i.e., ambient temperature/ambient light), massive Porites spp. responded to acclimatization through increases in biomass under ambient temperature/shade, and low temperature/ambient light, whereas P. irregularis responded through reduced growth under ambient temperature/shade, and low temperature/ambient light. Acclimatization affected the response to thermal stress for massive Porites spp. (but not P. irregularis), with an interaction between the acclimatization and subsequent treatments for growth. This interaction resulted from a lessening of the negative effects of high temperature after acclimatizing to ambient temperature/shade, but an accentuation of the effect after acclimatizing to low temperature/shade. It is possible that changes in biomass for massive Porites spp. are important in modulating the response to high temperature, with the taxonomic variation in this effect potentially resulting from differences in morphology. These results demonstrate that corals can acclimatize during short exposures to downward excursions in temperature and light, which subsequently affects their response to thermal stress. Moreover, even con-generic taxa differ in this capacity, which could affect coral community structure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effects of temperature on the growth of juvenile scleractinian corals

Tropical reef corals are well known for their sensitivity to rising temperature, yet surprisingly little is known of the mechanisms through which temperature acts on intact coral colonies. One such mechanism recently has been suggested by the association between the growth of juvenile corals and seawater temperature in the Caribbean, which suggests that temperature causes a transition between isometric and allometric growth scaling in warmer versus cooler years, respectively (Edmunds in Proc R Soc B 273:2275–2281, 2006). Here, this correlative association is tested experimentally for a cause-and-effect relationship. During April and May 2006, juvenile colonies (8–35 mm diameter) of massive Porites spp. from Moorea, French Polynesia, were incubated at warm (27.8°C) and cool (25.7°C) temperatures for 15 days, and their response assessed through the scaling of growth (change in weight) with colony size. The results reveal that the scaling of colony-specific growth (mg colony⁻¹ day⁻¹) was unaffected by temperature, although growth absolutely was greater at the cool compared to the warm temperature, regardless of colony size. This outcome was caused by contrasting scaling relationships for area-specific growth (mg cm⁻² day⁻¹) that were negatively allometric under warm conditions, but independent of size under cool conditions. In April 2007, a 22 days field experiment confirmed that the scaling of area-specific growth in juvenile Porites spp. is negatively allometric at a warm temperature of 29.5°C. Based on strong allometry for tissue thickness, biomass, and Symbiodinium density in freshly collected Porites spp., it is hypothesized that the temperature-dependency of growth scaling in these small corals is mediated by the interaction of temperature with biomass.     

Effects of temperature on photosynthesis and respiration in hermatypic corals

Photosynthesis and respiration rates of the reef corals Pocillopora damicornis (Linn.), Montipora verrucosa (Lamarck), Porites compressa Dana and Fungia scutaria Lamarck were measured under controlled temperatures. Results indicate that coral metabolism is closely adapted to ambient temperature conditions. Tropical corals measured at Enewetak, Marshall Islands, showed greater primary production compared to maintenance requirements at elevated temperatures than did subtropical varieties of the same species in Hawaii. Photosynthesis: respiration (P:R) ratios were significantly and negatively related with temperature between 18° and 31°C for all Hawaiian corals, whereas at Enewetak this ratio generally showed a curvilinear relationship for this temperature range. Extrapolations of P:R regressions on temperatures to a value of 2.0 (estimated as a minimum required for long-term functional autotrophy) coincide for Hawaiian specimens with published upper lethal temperatures. Extrapolation of P:R regressions for Enewetak specimens at temperatures above 25°C suggests lethal temperatures for these corals to be 2 to 5 C° higher than for Hawaiian corals, in good agreement with recent experimental findings. Interspecific differences in P:R temperature regressions for Hawaiian corals correlating with upper lethal temperature tolerances are described.Contribution No. 505 of the Hawaii Institute of Marine Biology.     

Effects of sheltering fish on growth of their host corals

Stony corals are the foundation species of tropical reefs, and their structures can harbor a diverse range of mutualist taxa that can confer important benefits, including provision of nutrients. Prominent among the associates of branching coral in the genus Pocillopora are groups of zooplanktivorous damselfishes that take refuge in the coral to avoid their predators. In field and laboratory experiments, we explored the effects of colonies of resident damselfishes on growth of their host corals. Laboratory studies revealed a positive relationship between biomass of fish and output of ammonium. In the field, levels of ammonium were significantly elevated in the water surrounding the branches of Pocillopora occupied by colonies of damselfish, particularly in time periods following active feeding by the fish. Experimental manipulation of the presence of fish on host corals during a month-long field experiment revealed that corals hosting fish grew significantly more than those that lacked fish, and coral growth was positively correlated with the biomass of resident fish. The Pocillopora colonies in the field experiment varied in the degree of openness of their branching structure, and dye studies indicated that this affected their ability to retain waterborne nutrients. Together with biomass of resident fish, colony openness explained 76% of the variation in coral growth rate during the experiment. Corals can exhibit considerable morphological variability, and mutualistic fish respond to colony architecture during habitat selection, with some species preferring more open-branched forms. This makes it likely that corals may face tradeoffs in attracting resident fish and in retaining the nutrients they provide.     

Turnover of photosynthetically fixed carbon in reef corals

Siderastrea siderea and Montastrea annularis were labeled in situ with NaH¹⁴CO₃. The corals were sampled over a period of 11 days and the radioactivity remaining in the ethanolsoluble and ammonia-soluble fractions measured. Total radioactivity in the corals fell to about 1/3 after one night and then to about 1/3 in the next 10 days. The ethanol-soluble radioactivity is probably converted to the less soluble, ammonia-extractable, material in the dark.Contribution No. 1514 from the University of Miami, Rosenstiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

Influence of UV radiation on the survival of larvae from broadcast-spawning reef corals

Effects of ambient ultraviolet light on the survivorship of eggs and planulae larvae was investigated for three species of broadcast-spawning reef corals, Acropora palmata, Montastraea annularis, and M. franksi. Eggs and larvae from these corals contain high concentrations of lipids (60–70% by weight) and float in surface waters for 3–4 days following spawning. Larvae originating from colonies living at deeper sites on the reef exhibited significantly lower survivorship than conspecifics originating from parents in shallow water when experimentally exposed for up to 4 days to ambient surface levels of ultraviolet radiation (UVR). Concentrations of the UVR-protective compounds correlated positively with survival and matched concentrations found in parent colonies, implying that higher concentrations of ultraviolet B protective compounds are responsible for greater survival of eggs and larvae from shallow compared to deeper-dwelling parents. Ultraviolet B appears to be responsible for most of the observed differences in larval survivorship with ultraviolet A playing a minor or insignificant role. Data presented here indicate that coral recruits on Caribbean reefs and elsewhere may originate primarily from adult colonies dwelling in shallow water.Communicated by P.W. Sammarco, Chauvin     

Rafting of reef corals and other organisms at Kwajalein Atoll

Studies conducted at Kwajalein Atoll (9°N; 168°E) in early 1988 reveal that marine organisms are commonly rafted into the area on drift pumice, drift wood and other flotsam. Coralskeletons on pumice provide the most useful quantitative data because they persist after rafted motile organisms have departed and rafted sessile forms have decomposed or been scavenged. The estimated minimum number of pumice fragments carrying corals into Kwajalein Atoll during its geological history is on the order of 10⁹, with a more realistic estimate in excess of 10¹¹. The estimated number of coral colonies rafted into the atoll would probably have to be increased several-fold if rafting on floating organic materials such as wood, charcoal, nuts and seeds could be determined. In the present study, a typical sample of beached pumice from Kwajalein Atoll contained 10³ coral colonies per m³ of bulk pumice fragments. Major pumice rafting episodes frequently result from volcanic eruptions. Estimates based on the observed K wajalein coral fouling rate and literature values for drift pumice production suggest that 10⁵ colonies could be rafted through the tropics by the smallest reported pumice-producing events and up to 10¹² colonies for large events. In the present study, most of the rafted corals that were recovered belong to species of the genus Pocillopora, but speccics of Porites and Millepora were also collected. Analysis of surface-current data and reported drift patterns for pumice, logs, drift bottles and wrecks suggests that movement of rafted corals and drifting larvae is predominantly from peripheral areas of low coral-species diversity into centers of high coral diversity. Coral diversity centers might be viewed as areas of coral species accumulation rather than centers of coral species origin.     

Effects of kinship on growth of a live-bearing reef fish

The influence of genetic relatedness on the individual performance (e.g. growth, development) of animals is often tied to agonistic or cooperative behaviors among conspecifics, and studies of the effects of kinship have produced mixed results. To explore genetic relatedness independent of these behaviors, we investigated the effects of kinship on the growth of the kelp perch Brachyistius frenatus, a live-bearing, planktivorous marine reef fish that is capable of only limited dispersal. Although juveniles occur in aggregations and compete for food resources, they do not exhibit overt aggressive or cooperative behavioral interactions. We hypothesized that under competition and in the absence of these behaviors, sibling and non-sibling groups of juvenile B. frenatus raised at the same densities in the field would not differ in average growth, but that siblings would exhibit lower variation in growth, simply due to genetic similarities in inherent growth rates. Pregnant, female kelp perch were collected and placed in cages until parturition was complete. Groups of young, recently born from the same mother or from different mothers, were then raised in the field for 9 wk. Our results revealed that average growth rates were similar between sibling and non-sibling treatments. While variation in growth increased initially in non-siblings, siblings showed little such variation. This divergence, however, was not consistent over the duration of the experiment, and variation in the growth of siblings ultimately converged with that of non-siblings. Effects of genetic relatedness would be most likely to manifest themselves early after birth, before environmental factors exert their influence, and this may explain the initial separation but eventual convergence in variation in growth between sibling and non-sibling treatments. For B. frenatus and other organisms that will encounter relatives and compete for resources without overt behavioral interactions, the degree of kinship may play a minor role in the demography of local populations. Received: 26 January 1998 / Accepted: 30 September 1998     

Experimental tests of suspension feeding in Atlantic reef corals

The ability of 15 species of Atlantic reef corals to act as suspension feeders was demonstrated by their removal of suspended particles from sea water in culture vessels. Mean clearance rates varied from 16.6 to 145.5 ml water cleared/h/cm² of live coral tissue. The lowest rates was found in Porites porites which is primarily a tentacle feeder, and the highest in Diploria clivosa which acts as both a tentacle feeder and suspension feeder. Rates of particle clearance in Agaricia agaricites, which is primarily a suspension feeder, were influenced by current velocity and type of food.     

Depth analysis of fatty acids in two caribbean reef corals

Total fatty acid compositions of colonies of two hermatypic, reef-building corals collected during the day-time over a depth range of 21 m were determined to assess the effect of depth-related environmental factors upon the lipid content of these organisms. No systematic changes were found, suggesting a steady-state balance between algal and animal lipogenesis in these symbiotic partnerships. Stephanocoenia michelinii, a day and night feeder, contained lipids indicative of external dietary sources such as copepods, whereas Montastrea annularis, a night feeder, did not.     

Effects of eutrophication on reef-building corals

Fourteen environmental variables were monitored at seven locations along the west coast of Barbados on a weekly basis over a one-year period, 1981 to 1982. The physicochemical and biological data indicate that an environmental gradient exists as a result of increased eutrophication of coastal waters. Growth rates (linear extension) of Montastrea annularis (Ellis and Solander), measured along the environmental gradient, exhibit high correlation with a number of water quality variables. Concentration of suspended particulate matter is the best univariate estimator of M. annularis skeletal extension rates (r ²=0.79, P<0.0001). The results suggest that suspended particulate matter may be an energy source for reef corals, increasing growth up to a certain maximum concentration. After this, reduction of growth occurs due to smothering, reduced light levels and reduced zooxanthellae photosynthesis.     

Effects of delayed feeding and temperature on the age of irreversible starvation and on the rates of growth and mortality of Pacific herring larvae

The time periods from exhausion of the yolk to the age of irreversible starvation for Pacific herring Clupea harengus pallasi larvae were 8.5, 7.0 and 6.0 d at 6°, 8° and 10°C, respectively. These periods are within the range perviously measured for Atlantic herring larvae and other temperature zone fish species; they are long compared to the periods for tropical species. The variation in the length of this period is due almost entirely to temperature; the natural logarithm of the time period from fertilization to irreversible starvation is highly correlated (r=0.91) with the mean rearing temperature for 25 species of pelagic marine fish larvae. The rates of growth and mortality, measured for 26 experimental populations of Pacific herring larvae reared at 6°, 8° and 10°C and ten ages of delayed first feeding, decreased and increased, respectively with increasing age of first feeding and increasing temperature. These rates, adjusted for the effects of rearing conditions, were compared with the rates for natural populations of herring larvae. Growth is generally faster in the sea than in experimental enclosures. Two of the eleven estimates of natural mortality rate were high enough to indicate possible catastrophic mass starvation. This is consistent with Hjort's critical period concept of year class formation and it suggests that mass starvation occurs in 18 to 36% of the natural populations of first feeding herring larvae.     

温度对小叶海金沙生长的影响

用野外调查测定和室内模拟的方法研究了温度对小叶海金沙生长的影响.结果表明,小叶海金沙疯狂生长群落中的温度比正常生长群落高,且温度与其净光合速率呈极显著正相关,而这两种类型群落内大气相对湿度以及光照差异较小.此外,土壤养分和土壤pH与小叶海金沙的生长速度无明显的关联性.根据研究结果推测:温度可能是小叶海金沙疯狂生长原因之一.     

Influences of habitat and natural disturbances on contributions of massive Porites corals to reef communities

We compared densities, distributions and size frequencies of massive corals in the genus Porites on five relatively exposed, mid-shelf reefs (50 km offshore) in the central Great Barrier Reef with those on a sheltered inshore reef (10 km offshore). Data included various transect and mapping studies between 1984 and 1990, estimates of size-dependent damage from the crown-of-thorns starfish Acanthaster planci, estimated densities of herbivorous sea urchins (potential predators of juveniles), and observations of size-specific effects of tropical cyclones. Assemblages of Porites spp. on mid-shelf reefs were dominated by small colonies (2 to 10 cm diam) established either from planula larvae or from small tissue remnants that had survived A. planci predation in the early to mid-1980s. Large colonies (up to 10 m diam) were scarce, except for localized aggregations on terraces at the base of reef slopes (6 to 12 m deep). Extensive space suitable for settlement by coral larvae can be attributed to recurrent cyclones and A. planci outbreaks. Despite low sea urchin predation, the slowly growing Porites juveniles are likely to die from overgrowth by numerous, much faster growing corals. On the sheltered inshore reef, the coral community was dominated by very large (>5 m diam) Porites colonies, several centuries old; recruitment was mainly by fragmentation of large colonies; there was little space available for settlement, and probabilities of juvenile mortality from grazing urchins were high. Differences in settlement and early survival of Porites spp. are exacerbated by different regimes of storm damage. A model is proposed that links wave climate with the size and age reached by corals before dislodgement by storm waves, and which is consistent with observed densities and size-frequency distributions of Porites in sheltered and exposed areas.     

Effects of feeding history on the growth characteristics of a reef fish at settlement

A feeding experiment was conducted on the pelagic stages of the tropical goatfish Upeneus tragula (family Mullidae) to examine how food quantity affects growth characteristics and potential success at settlement. Pelagic goatfish were collected from aggregation rafts 3 nautical miles west of Lizard Island on the northern Great Barrier Reef during December 1990. Three tanks in each of four feeding regimes were stocked with 25 pelagic goatfish between 20 and 23 mm standard length (SL). The four feeding regimes were: fed ad lib. (fed): fed once per day (f1pd); starved every second day (stld); starved for 3 d and re-fed (st3d). Fishes were fed 36 to 48 h old Artemia sp. nauplii (Ocean Star strain). Fish were removed from tanks when they underwent metamorphosis, changed pigmentation and settled to the bottom of the tanks. Morphology, muscle development, time taken to settle and biochemical condition were examined. Growth attributes of the treated fish were compared to fish which settled within 24 h of capture (field). All attributes examined were significantly influenced by the feeding treatments. Fish within the fed and st3d treatments were significantly larger and heavier than fish in the less well-fed treatments (f1pd, st1d). Similarly, concentrations of total lipid, carbohydrate and protein in the settled fish were significantly higher in the fed and st3d treatments compared with the st1d and f1pd treatments. For all these morphological and biochemical attributes the st1d and f1pd fish did not significantly differ from the field fishes, but did differ from the fed and st3d fishes. Water content was significantly higher in the f1pd, st1d and field fishes compared with the fed and st3d fish. Furthermore, the average time taken to settle followed the pattern: fed (14d)U. tragula are physiologically well suited to exploiting a patchy food source, and that food availability within the pelagic stages can have a major influence on the growth characteristics of this reef fish at settlement. The ramifications of this finding are discussed in relation to survival and success once fish have recruited to the reef population.     

The relationship between bleaching and mortality of common corals

Reef corals are likely to have many subtle but four gross responses to anomalous warm water. These are (1) not bleach and live (mortality <10%), (2) not bleach and die (mortality >20%), (3) bleach and live, and (4) bleach and die. The frequency of these four possible gross responses was determined for 18 common coral taxa over an exceptionally warm 1998 El Niño where intense bleaching was observed, and mortality determined from line transects averaged 41.2±34.7 (±SD). Field studies included (1) recording the loss of color (bleaching) and observing recently dead individuals among 6,803 colonies during five sampling periods and (2) estimating mortality based on 180 m of line-intercept transects completed 4 months before and near the end of the bleaching episode. There was no clear relationship between the loss of color and either direct observation or transect-based estimates of mortality for the 18 taxa. The morphology of the taxa did not influence color loss but branching and encrusting taxa had higher mortality than massive and submassive taxa. Loss of color and mortality are the most common responses to warm water as only Pavona did not lose color or die and only two taxa, Cyphastrea and Millepora, did not significantly lose color but died. Of the 15 taxa that lost color, five taxa, Astreopora, Favia, Favites, Goniopora, and Leptoria, did not die. These taxa are those most likely to have reduced potential mortality by the loss of pigments and associated algal symbionts. Death of the branching taxa was detected reasonably by direct field observation but some taxa were underestimated when compared with mortality estimates based on line transects. Death of encrusting and massive taxa including Echinopora, Galaxea, Hydnophora, Montipora, Platygyra, and massive Porites was poorly detected from direct observations but they proved to have modest to high mortality (20–80%) based on line transects. There was no single response of these common corals to warm water but these data, collected during an extreme warm-water anomaly, indicate that the loss of color is most frequently a sign of morbidity, particularly for branching and encrusting taxa.Communicated by P.W. Sammarco, Chauvin     

Effects of eutrophication and sedimentation on juvenile corals

Settlement of juvenile scleractinian corals was investigated from 1987 to 1990 on eutrophic and less eutrophic fringing reefs on the west coast of Barbados, West Indies. The number of coral recruits and number of recruiting coral species on cement blocks decreased with increasing eutrophication of the reefs. This may suggest lower settlement rates on eutrophic reefs, but could also liave resulted from higher post-settlement mortality, since blocks were examined only once after 3 yr of immersion. Coral settlement rates to artificial plates that were checked monthly were also lower on the more eutrophic reefs. This could result from lower local availability of larvae caused by fewer adult corals and/or lower reproductive rates of corals on eutrophic reefs. However, the ratio of coral recruits to adult coral abundance was considerably lower on eutrophic reefs, suggesting that local coral abundance alone can not explain lower settlement rates on eutrophic reefs. The lower rates on eutrophic reefs may result from a lower probability of coral larvae settlin when present, perhaps because of a limited availability of suitable settlement substrate. Colonization of settlement plates by non-coralline organisms was heavier on eutrophic reefs, and unoccupied space was lower, supporting the suggestion that suitable coral settlement substrate may be limiting on eutrophic reefs. Moreover, coralline algae, which facilitate metamorphosis and settlement of coral larvae, were less abundant on settlement plates on eutrophic reefs.     

The effects of partial mortality on the fecundity of three common Caribbean corals

The recent intensification of human disturbances in the Caribbean has increased the prevalence of partial mortality on coral colonies. Partial mortality can change colony size by directly shrinking colonies or by splitting colonies into fragments. A reduction in colony size can also adversely affect fecundity and fitness as internal resources shift away from reproduction toward colony maintenance. This study aimed to determine whether three Caribbean coral species, Siderastrea siderea, Montastraea faveolata, and Diploria strigosa, along the reef tract in Puerto Morelos, Mexico (20^o52′N, 86^o51′W), continued to dedicate resources to reproduction when colonies were fragmented to pre-maturation size. Contrary to expectations, eggs were found in colonies that were smaller than the maturation size and had been subjected to partial mortality. The continued dedication of resources toward reproduction, even in the smallest colonies, suggests that resource trade-offs away from reproduction are not as rigid as previously suggested in stressed corals.     

Effects of sand deposition on scleractinian and alcyonacean corals

The ability of corals to withstand experimental sand deposition was investigated for two experimental periods (17 h and 6 wk) in eight scleractinia (Favia favus, Favites pentagona, Platygyra daedalea, Gyrosmilia interrupta, Galaxea fascicularis, Cyphastrea chalcidicum, Favites abdita, Goniopora dijboutensis) and five alcyonacea (Lobophytum depressum, L. venustum, Sinularia dura, S. leptoclados, Sarcophyton glaucum) collected in 1992 from Natal, South Africa. Scleractinia were active sediment shedders, alcyonacea passive, relying on water motion and gravity. Short-term sand clearing efficiency was primarily dependent on corallum shape. Sand application led to hydrostatic inflation of polyps in scleractinia and the entire colony in alcyonacea as well as to increased tentacular action in the scleractinian Gyrosmilia interrupta. Under continuous sand application, inflation remained while other activities, such as tentacular motion, ceased completely. In scleractinia and alcyonacea, tissue necroses appeared after the first week of continuous sand application. Death of entire colonies and partial bleaching of continually sandcovered areas were observed in alcyonacea only. Different grain sizes of sand had no influence on clearing reaction or efficiencies.