Ecology of the Symbiotic Coral Cladocora caespitosa (L.) (Faviidae, Scleractinia) in the Bay of Piran (Adriatic Sea): II. Energy Budget

Abstract. The temperate, symbiotic coral Cladocora caespitosa occurs naturally with variable numbers of zooxanthellae. This allows us to study differences in the physiology of symbiotic and "non-symbiotic" polyps and to correlate them with zooxanthellae activities.
Zooxanthellae density in "dark" (normal-colored) polyps was 50 times higher than in "white" (bleached) polyps.
The chlorophyll a content per zooxanthella was 3 times, and Chi c was 13 times higher in white polyps than in dark ones. O₂-flux experiments were conducted with colonies in situ and with single polyps - dark and white - under laboratory conditions. Two approaches were used to evaluate the contribution of zooxanthellae translocation products to animal respiration. Both revealed that the animal tissues derive a higher benefit from the zooxanthellae during periods of low water temperature than during warm periods.     

Climate change and coral reef bleaching: An ecological assessment of long-term impacts,recovery trends and future outlook

Since the early 1980s, episodes of coral reef bleaching and mortality, due primarily to climate-induced ocean warming, have occurred almost annually in one or more of the world's tropical or subtropical seas. Bleaching is episodic, with the most severe events typically accompanying coupled ocean–atmosphere phenomena, such as the El Niño-Southern Oscillation (ENSO), which result in sustained regional elevations of ocean temperature. Using this extended dataset (25+ years), we review the short- and long-term ecological impacts of coral bleaching on reef ecosystems, and quantitatively synthesize recovery data worldwide. Bleaching episodes have resulted in catastrophic loss of coral cover in some locations, and have changed coral community structure in many others, with a potentially critical influence on the maintenance of biodiversity in the marine tropics. Bleaching has also set the stage for other declines in reef health, such as increases in coral diseases, the breakdown of reef framework by bioeroders, and the loss of critical habitat for associated reef fishes and other biota. Secondary ecological effects, such as the concentration of predators on remnant surviving coral populations, have also accelerated the pace of decline in some areas. Although bleaching severity and recovery have been variable across all spatial scales, some reefs have experienced relatively rapid recovery from severe bleaching impacts. There has been a significant overall recovery of coral cover in the Indian Ocean, where many reefs were devastated by a single large bleaching event in 1998. In contrast, coral cover on western Atlantic reefs has generally continued to decline in response to multiple smaller bleaching events and a diverse set of chronic secondary stressors. No clear trends are apparent in the eastern Pacific, the central-southern-western Pacific or the Arabian Gulf, where some reefs are recovering and others are not. The majority of survivors and new recruits on regenerating and recovering coral reefs have originated from broadcast spawning taxa with a potential for asexual growth, relatively long distance dispersal, successful settlement, rapid growth and a capacity for framework construction. Whether or not affected reefs can continue to function as before will depend on: (1) how much coral cover is lost, and which species are locally extirpated; (2) the ability of remnant and recovering coral communities to adapt or acclimatize to higher temperatures and other climatic factors such as reductions in aragonite saturation state; (3) the changing balance between reef accumulation and bioerosion; and (4) our ability to maintain ecosystem resilience by restoring healthy levels of herbivory, macroalgal cover, and coral recruitment. Bleaching disturbances are likely to become a chronic stress in many reef areas in the coming decades, and coral communities, if they cannot recover quickly enough, are likely to be reduced to their most hardy or adaptable constituents. Some degraded reefs may already be approaching this ecological asymptote, although to date there have not been any global extinctions of individual coral species as a result of bleaching events. Since human populations inhabiting tropical coastal areas derive great value from coral reefs, the degradation of these ecosystems as a result of coral bleaching and its associated impacts is of considerable societal, as well as biological concern. Coral reef conservation strategies now recognize climate change as a principal threat, and are engaged in efforts to allocate conservation activity according to geographic-, taxonomic-, and habitat-specific priorities to maximize coral reef survival. Efforts to forecast and monitor bleaching, involving both remote sensed observations and coupled ocean–atmosphere climate models, are also underway. In addition to these efforts, attempts to minimize and mitigate bleaching impacts on reefs are immediately required. If significant reductions in greenhouse gas emissions can be achieved within the next two to three decades, maximizing coral survivorship during this time may be critical to ensuring healthy reefs can recover in the long term.     

膨胀蔷薇珊瑚与壮实鹿角珊瑚的胚胎和幼虫发育

造礁石珊瑚的有性繁殖是修复珊瑚礁生态系统的有效途径之一.文章为国内首次报道造礁石珊瑚有性繁殖和幼体发育过程,为利用有性繁殖技术恢复珊瑚礁生态系统提供了发育生物学上的理论基础.海南省三亚市鹿回头海域的大部分珊瑚在2009年3月底至4月中发生有性繁殖行为.2009年4月11日晚采集膨胀蔷薇珊瑚Montipora turgescens和壮实鹿角珊瑚Acropora robusta自然排放的受精卵,研究其胚胎及幼虫发育的过程.通过连续观察和显微拍摄记录了2种珊瑚的受精卵发育过程,结果显示,受精卵经过分裂变为桑葚胚;桑葚胚经过进一步发育变形为虾片状,进而发育成盘状幼体;盘状幼体由四周向内弯曲形成碗状的结构,进而出现原肠胚及纤毛,发育成为椭圆型或圆形的浮浪幼虫;浮浪幼虫发生附着变态,长出触手,发育成珊瑚幼体.通过观察还发现,膨胀蔷薇珊瑚的卵母细胞中本身就含有黄褐色虫黄藻,壮实鹿角珊瑚的卵母细胞中没有虫黄藻,其共生的虫黄藻足在发育过程中从周围环境获得的.     

The evolution of modern corals and their early history

Scleractinians are a group of calcified anthozoan corals, many of which populate shallow-water tropical to subtropical reefs. Most of these corals calcify rapidly and their success on reefs is related to a symbiotic association with zooxanthellae. These one-celled algal symbionts live in the endodermal tissues of their coral host and are thought responsible for promoting rapid calcification. The evolutionary significance of this symbiosis and the implications it holds for explaining the success of corals is of paramount importance. Scleractinia stands out as one of the few orders of calcified metazoans that arose in Triassic time, long after a greater proliferation of calcified metazoan orders in the Paleozoic. The origin of this coral group, so important in reefs of today, has remained an unsolved problem in paleontology. The idea that Scleractinia evolved from older Paleozoic rugose corals that somehow survived the Permian mass extinction persists among some schools of thought. Paleozoic scleractiniamorphs also have been presented as possible ancestors. The paleontological record shows the first appearance of fossils currently classified within the order Scleractinia to be in the Middle Triassic. These earliest Scleractinia provide a picture of unexpectedly robust taxonomic diversity and high colony integration. Results from molecular biology support a polyphyletic evolution for living Scleractinia and the molecular clock, calibrated against the fossil record, suggests that two major groups of ancestors could extend back to late Paleozoic time. The idea that Scleractinia were derived from soft-bodied, “anemone-like” ancestors that survived the Permian mass extinction, has become a widely considered hypothesis. The 14-million year Mesozoic coral gap stands as a fundamental obstacle to verification of many of these ideas. However, this obstacle is not a barrier for derivation of scleractinians from anemone-like, soft-bodied ancestors. The hypothesis of the ephemeral, “naked coral”, presents the greatest potential for solution of the enigma of the origin of scleractinians. It states that different groups of soft-bodied, unrelated “anemone-like” anthozoans gave rise to various calcified scleractinian-like corals through aragonitic biomineralization. Although there is evidence for this phenomenon being more universal in the mid-Triassic interval, following a lengthy Early Triassic post-extinction perturbation, it appears to have occurred at least three other times prior to this interval. This idea suggests that, because of ephemeral characteristics, the skeleton does not represent a clade of zoantharian evolution but instead represents a grade of organization. In the fossil record, skeletons may have appeared and disappeared at different times as some clades reverted to soft-bodied existence and these phenomena could account for notable gaps in the taxonomic and fossil record. A fuller understanding and possible solution to the problem of the origin of modern corals may be forthcoming. However, it will require synthesis of diverse kinds of data and an integration of findings from paleobiology, stratigraphy, molecular biology, carbonate geochemistry, biochemistry and invertebrate physiology.     

砗蚝(Hippopus hippopus)的人工繁育

于2016年4—9月开展了砗蚝(Hippopus hippopus)人工繁育技术研究。采用五羟色胺进行催产、促使配子排放;精卵分别收集,进行异体间受精以避免自交;受精卵经过30h孵化,选出600万D形幼虫进行培育。砗蚝的早期生活史与砗磲(Tridacna spp.)相似,经历前期面盘幼虫、中期面盘幼虫、后期面盘幼虫、足面盘幼虫、单水管稚贝、双水管稚贝、外套膜触手稚贝、幼贝等阶段。与砗磲不同的是,砗蚝怀卵量较少,但卵径较大,D型幼虫也较大,幼虫趋光性更强,壳长2.0mm以后外套膜不伸出壳缘外,幼贝贝壳形态也不同于砗磲。同砗磲幼虫一样,砗蚝幼虫需要构建虫黄藻系统之后,才能出现鳃、次生壳等,从而完成变态成为稚贝。砗蚝幼虫变态率较低,仅为1.4%。中间育成期间,丝状藻和锥形螺是稚贝培育的主要敌害,需及时清理才能确保稚幼贝正常生长发育。经过120d的精心饲育,培养出平均壳长6.3mm幼贝500余个。本研究为进一步开展砗蚝人工繁育、中间育成、增殖放流、资源修复及移植保育提供了参考。     

Trophic Potential and Photoecology of Endolithic Algae Living within Coral Skeletons

Abstract. The biomass of the endolithic algae Ostreobium quekettii Phyllosiphoniaceae living within skeletons of the scleractinans Mycedium elephantotus and Leptoseris fragilis averages 300 μg protein. cm^-2. This represents approximately 7% of the protein of the zooxanthekie-containing tissue of M. elephantotus and approximately 38% of that of L. fragilis. Oxygen production Pmax_net of 0. querkettii in bare skeletons of M. elephantotus averaged 0.7 μg O₂.cm^-2· h^-1 measured in large skeletal fragments. This amount is approximately 6% of the productivity of the zooxanthellae Symbiodinium microadriaticum living in the same scleractinian species at the same depth Pmax_net 11 μg O₂· Cm^-2· h^-1. Light compensation of O. quekettii - within skeletons - was reached at approximately 10 and saturation at 35 40 μE·m^-2· s^-1. Algae within the M. elephantotus skeletons receive a maximum of 4–6% of the ambient irradiance, which is approximately 0.9 μE · m^-2· s^-1 approximately 0.04% surface irradiance at a depth of 88 m. In L. fragilis at a depth of 145 m, the photon flux decreases to 0.3 μE·m^-2· s^-1, which is less than 0.004% of surface intensity. With increasing depth, the ratio of Chl b to Chl a increased in endolithic algae colonizing L. fragilis, indicating improvement of light harvesting under low light conditions. In free-living O. quekettii cultured at irradiance levels from 0.5–60 μE·m^-2· s^-1, the concentrations of chlorophylls increased and that of siphonein and β-carotene decreased with decreasing photon flux.     

Experimental bleaching of a tropical sea anemone in situ

Bleaching (whitening) of cnidarians such as corals and sea anemones has caused widespread degradation of coral reefs around the world and is therefore an urgent issue for coral reef science and conservation. Although cnidarians often bleach in aquaria, methods for experimental induction of bleaching in wild cnidarians are lacking, which impedes scientists’ ability to understand the ultimate effects of bleaching on the broader ecosystem. In this study, we investigated the utility of an in situ method for experimental induction of bleaching in the tropical sea anemone Heteractis crispa. Healthy, wild anemones were covered with opaque black plastic sheets, mesh cages or left undisturbed (controls) and tentacle colour and body size were monitored with a colour reference card and flexible tape, respectively, every 1–3 days for 15 days. Caged and control anemones remained unchanged for the duration of the experiment, but covered anemones commenced whitening after 4–6 days and were completely white after 7–14 days (mean time to bleaching ± SE = 10.1 ± 0.7 days). Experimental bleaching occurred without reduction in anemone body size and was visibly similar to natural bleaching seen previously in H. crispa. We hypothesize that light‐deprivation, reduced water flow, physical contact or some combination of these factors caused the bleaching. This study provides the basis for a simple and rapid method of inducing bleaching in situ, which releases scientists’ dependence on sporadic natural bleaching events or artificial aquarium experiments, and provides a means to investigate the effects of bleaching on other ecosystem components such as fishes.     

在细胞水平上对高温珊瑚白化的初步研究

全球变暖背景下的异常高温能够导致珊瑚及其虫黄藻组成的共生体系崩溃,虫黄藻大量损失,出现珊瑚白化,并可能进一步导致珊瑚礁生态系统退化.文章通过对6种造礁石珊瑚的急性高温胁迫实验,分析不同种属的石珊瑚虫黄藻共生体系对高温的耐受性差异,为全球变暖背景下珊瑚群落演替趋势提供理论依据.结果显示:1)在急性高温胁迫下,石珊瑚耐受的差异性与其形态有关,枝状珊瑚耐受性最低,在高温胁迫下最先白化、死亡,而叶片状和块状珊瑚对高温的耐受性较强,这与野外珊瑚礁白化的现场观测结果一致.2)在高温胁迫下,不同种属珊瑚共生虫黄藻损失的方式不同:珊瑚持续排出虫黄藻,如鹿角杯形珊瑚 Pocillopora damicornis;珊瑚先排出一定的共生藻,之后珊瑚组织携带大量虫黄藻与珊瑚骨骼分离,如风信子鹿角珊瑚 Acropora hyacinthus 和松枝鹿角珊瑚 Acropora brueggemanni;先排出部分虫黄藻后,虫黄藻以有丝分裂增殖的方式迅速补充其数量,如十字牡丹珊瑚 Pavona decussata;虫黄藻细胞直接坏死而损失虫黄藻,如澄黄滨珊瑚 Porites lutea.研究强调,预测珊瑚对全球变化的响应问题时,应当同时考虑珊瑚宿主和共生藻的作用.     

Differential effects of thermal and chemical stressors on tissue balls from scleractinian corals

Coral cell aggregates (tissue balls) from four species (Acropora muricata, Fungia repanda, Pavona cactus and Pocillopora damicornis) were used as an indicator to investigate the effects on the corals of thermal stress and of chemical extracts from three sponges (Adocia sp., Haliclona sp. and Lissodendoryx sp.) and one ascidian (Didemnum molle). The formation and disintegration of tissue balls were studied through exposure to a temperature range of 23–30 °C at time intervals of 0–90 min, and to sponge and ascidian crude extracts at concentrations of 50–200 µg ml^?1 at temperatures of 23 and 30 °C and at time intervals of 10, 60 and 120 min. The negative effect of temperature on overall tissue ball density (number per cm² of coral surface) was greatest at higher temperatures (28 and 30 °C) but varied among coral species. Tissue balls of P. damicornis were the most robust whereas those of A. muricata were the most sensitive. High concentrations of extracts of Adocia sp., Haliclona sp. and Lissodendoryx sp. generally inhibited the formation of tissue balls or caused their disintegration, or both, most markedly at 30 °C. Adocia sp. induced the least negative effects and Haliclona sp. the most. No tissue balls were formed in the presence of D. molle extracts (50 and 100 µg ml^?1), indicating a high level of interference with tissue ball formation. The differential susceptibility to thermal and chemical stressors exhibited by the corals under study have possible implications for the interactions of the corals with other sedentary reef organisms under climate change-driven ocean warming.     

珊瑚共生虫黄藻密度的季节变化及其与珊瑚白化的关系——以大亚湾石珊瑚为例

定量分析了2006年6月、2007年8月和2008年2月采自南海北部大亚湾海域的共8科、13属、23种170个石珊瑚样品的共生虫黄藻密度,探讨了石珊瑚共生虫黄藻密度的季节变化及其与珊瑚白化的关系.结果显示,所有珊瑚种属的共生虫黄藻密度都显示出明显的季节性波动,总体上夏季低、冬季高(约为夏季的2倍),是海表水温和太阳辐射协同作用的结果.夏季大规模的珊瑚白化(热白化)可能是珊瑚共生虫黄藻密度逐渐降低(排出)到一定阈值的外观表征,而非突发的生态现象;冬季珊瑚白化(冷白化)则可能是极端低温直接致珊瑚死亡,进而快速排出虫黄藻的突发现象;高共生虫黄藻密度对冬季低温乃至极端低温条件下的珊瑚生存起到一定的保护作用.