冷水珊瑚礁研究进展与评述

doi:10.13284/j.cnki.rddl.002802

摘要/Abstract

摘要： 相比浅海暖水珊瑚礁，对深海的冷水珊瑚礁研究相对薄弱。近年来随着水下机器人和深潜器技术的进步，国际上兴起了深海珊瑚礁研究热潮。文章综述了：1）冷水石珊瑚的物种多样性及格局分布特征；2）冷水石珊瑚的生长特征及测量方法；3）冷水珊瑚礁的类型、形成机制与演化过程；4）海洋酸化对冷水珊瑚礁的影响；5）冷水石珊瑚对海洋古环境的记录等。最后提出中国南海冷水珊瑚礁研究展望，建议开展探测南海冷水珊瑚礁的分布，监测珊瑚礁区的生态环境，分析冷水石珊瑚群落特征，探讨冷水珊瑚礁对海洋酸化的响应机制，并利用南海冷水石珊瑚重建南海冷水珊瑚礁的发育历史及所记录的古海洋环境等研究工作。

关键词: 冷水珊瑚礁, 冷水石珊瑚, 海洋酸化, 环境控制

Abstract: Coral reefs are generally associated with shallow tropical seas; however, there are widespread and diverse coral ecosystems in deep waters on continental shelves, slopes, seamounts and ridges around the world. Cold-water corals have been known since the 18th century, but much fewer studies have been performed on deep-sea coral reefs compared with on tropical coral reefs. Along with the applications of modern submersibles and remotely operated vehicles (ROVs) to deep ocean exploration activities, more and more studies have been conducted and deep-sea coral reefs have become a research hotspot recently. This review includes: 1) the diversity and biogeographical distribution of cold-water corals; 2) the growth characteristics of corals and related environmental controls; 3) the initiation and development of deep-sea coral reefs; 4) the response of corals to ocean acidification; 5) corals as valuable archives for paleoceanographic reconstructions. It also highlights the future research directions of cold-water coral reefs in the South China Sea.

Key words: cold-water coral reefs, cold-water coral, ocean acidification, environmental control

赵美霞，余克服．. 冷水珊瑚礁研究进展与评述[J]. 热带地理, 2016, 36(1): 94-100.

ZHAO Meixia，YU Kefu．. A Review of Recent Research on Cold-Water Coral Reefs[J]. TROPICAL GEOGRAPHY, 2016, 36(1): 94-100.

参考文献

[1] ROBERTS J M，WHEELER A J，FREIWALD A．Reefs of the deep：the biology and geology of cold-water coral ecosystems[J]．Science，2006，312：543-547．

[2] ROBERTS J M，WHEELER A，FREIWALD A，CAIRNS S．Cold water corals：the biology and geology of deep-sea coral habitats [M]．Cambridge：Cambridge University Press，2009．

[3] RODER C，BERUMEN M L，BOUWMEESTER J，PAPATHANASSIOU E，ALSUWAILEM A，VOOLSTRA C R．First biological measurements of deep-sea corals from the Red Sea[J]．Scientific Reports，2013，3：2802．

[4] HENRY L A，ROBERTS J M．Biodiversity and ecological composition of macrobenthos on cold-water coral mounds and adjacent off-mound habitat in the bathyal Porcupine Seabight，NE Atlantic[J]．Deep-Sea Research Part I，2007，54：654-672．

[5] AUSTER P J．Are deep-water corals important habitats for fishes? [M]//FREIWALD A，ROBERTS J M．Cold-water Corals and Ecosystems．Berlin Heidelberg：Springer，2005：747-760．

[6] AUSTER P J．Linking deep-water corals and fish populations[J]．Bulletin of Marine Science，2007，81（S1）：93-99.

[7] HUSEB? A，N?TTESTAD L，FOSSA J H，FUREVIK D M，J?RGENSEN S B．Distribution and abundance of fish in deep-sea coral habitats[J]．Hydrobiologia，2002，471：91-99．

[8] ROBINSON L F，ADKINS J F，SCHEIRER D S，FERNANDEZ D P，GAGNON A．WALLER R G．Deep-sea scleractinian coral age and depth distributions in the northwest Atlantic for the last 225000 years [J]．Bulletin of Marine Science，2007，81（3）：371-391．

[9] ROBINSON L F，ADKINS J F，FRANK N，GAGNON A C，PROUTY N G，ROARK E B，VAN de Flierdt T．The geochemistry of deep-sea coral skeletons：a review of vital effects and applications for palaeoceanography [J]．Deep Sea Research Part II：Topical Studies in Oceanography，2014，99：184-198．

[10] RADDATZ J，LIEBETRAU V，RüGGEBERG A，HATHORNE E，KRABBENH?FT A，EISENHAUER A，B?HM F，VOLLSTAEDT H，FIETZKE J，LóPEZ CORREA M，FREIWALD A，DULLO W．Stable Sr-isotope，Sr/Ca，Mg/Ca，Li/Ca and Mg/Li ratios in the scleractinian cold-water coral Lophelia pertusa[J]．Chemical Geology，2013，352：143-152．

[11] FOSSA J H，MORTENSEN P B，FUREVIK D M．The deep-water coral Lophelia pertusa in Norwegian waters：distribution and fishery impacts [J]．Hydrobiologia，2002，471：1-12．

[12] HALLSPENCER J，ALLAIN V，FOSSA J H．Trawling damage to Northeast Atlantic ancient coral reefs[J]．Proceedings of the Royal Society of London Series B-Biological Sciences，2002，269：507-511．

[13] CLARK M R，ROWDEN A A．Effect of deep water trawling on the macro-invertebrate assemblages of seamounts on the Chatham Rise，New Zealand[J]．Deep-Sea Research，2009，156：1540-1554．

[14] ROBERTS J M．Full effects of oil rigs on corals are not yet known [J]．Nature，2000，403：242．

[15] HALFAR J，FUJITA R M．Danger of deep-sea mining[J]．Science，2007，316：987．

[16] GUINOTTE J M，ORR J，CAIRNS S．FREIWALD A，MORGAN L，GEORGE R．Will human-induced changes in seawater chemistry alter the distribution of deep-sea scleractinian corals?[J]．Frontiers in Ecology and the Environment，2006，4（3）：41-146．

[17] TURLEY C M，ROBERTS J M，GUINOTTE J M．Corals in deep-water：will the unseen hand of ocean acidification destroy cold-water ecosystems? [J]．Coral Reefs，2007，26：445-448．

[18] DAVIES A J，ROBERTS J M，HALL Spencer J．Preserving deep-sea natural heritage：emerging issues in offshore conservation and management [J]．Biological Conservation，2007，138：299-312．

[19] VAUGHAN T W，WELLS J W．Revision of the suborders，families，and genera of the Scleractinia[J]．Geological Society of America Special Papers，1943，44：1-363．

[20] KELLER N B．The deep-sea madreporarian corals of the genus Fungiacyathus from the Kurile-Kamchatka，Aleutian Trenches and other regions of the world oceans[J]．Trudy Institut Okeanologii，1976，99：31-44．

[21] DAVIES A J，WISSHAK M，ORR J C，ROBERTS J M．Predicting suitable habitat for the cold-water reef framework-forming coral Lophelia pertusa（Scleractinia）[J]．Deep Sea Research Part I：Oceanographic Research Papers，2008，55：1048-1062．

[22] HOWELL K L，HOLT R D，PULIDO E I，STEWART H．When the species is also a habitat：Comparing the predictively modeled distributions of Lophelia pertusa and the reef habitat it forms [J]．Biological Conservation，2011，44：2656-2665．

[23] GASS S E，ROBERTS J M．Growth and branching patterns of Lophelia pertusa（Scleractinia）from the North Sea[J]．Journal of the Marine Biological Association UK，2011，91：831?835．

[24] SABATIER P，REYSS J L，HALLSPENCER J，COLIN C，FRANK N，TISNéRAT- Laborde N，BORDIER L，DOUVILLE E．210Pb-226Ra chronology reveals rapid growth rate of Madrepora oculata and Lophelia pertusa on world’s largest cold-water coral reef[J]．Biogeosciences，2012，9：1253-1265．

[25] DAVIES A J，GUINOTTE J M．Global habitat suitability for framework- forming cold-water corals[J]．PLoS One，2011，6：e18483．

[26] QURBAN M A，KRISHNAKUMAR P K，JOYDAS T V，MANIKANDAN K P，ASHRAF T T M，QUADRI S I，WAFAR M，QASEM A，CAIRNS S D．In-situ observation of deep water corals in the northern Red Sea waters of Saudi Arabia[J]．Deep Sea Research Part I：Oceanographic Research Papers，2014，89：35-43．

[27] WHITMIRE C E，CLARKE E．State of deep coral ecosystems of the US Pacific Coast：California to Washington[M]//LUMSDEN S E，HOURIGAN T F，BRUCKNER A W，DORR G．The State of Deep Coral Ecosystems of the United States．NOAA Technical Memorandum CRCP-3．2007：109-154．

[28] CAIRNS S D．Deep-water corals：an overview with special reference to diversity and distribution of deep-water scleractinian corals[J]．Bulletin of Marine Science，2007，81：311-322．

[29] TITTENSOR D P，BACO A R，BREWIN P E，CLARK M R，CONSALVEY M，HALLSPENCER J，ROWDEN A A，SCHLACHER T，STOCKS K I，ROGERS A D．Predicting global habitat suitability for stony corals on seamounts[J]．Journal of Biogeography，2009，36：1111- 1128．

[30] BOSTOCK H C，TRACEY D M，CURRIE K I，DUNBAR G B，HANDLER M R，MIKALOFF FLETCHER S E，SMITH A M，WILLIAMS M J M．The carbonate mineralogy and distribution of habitat- forming deep-sea corals in the southwest pacific region[J/OL]．Deep Sea Research Part I：Oceanographic Research Papers，2015，100：88- 104[2015-05-01]．http://www.sciencedirect.com/science/article/pii/S0967063715000400-aff0015．

[31] GUINOTTE J M，DAVIES A J．Predicted Deep-Sea Coral Habitat Suitability for the US West Coast[J]．PLoS One，2014，9（4）：e93918．

[32] RENGSTORF A，YESSON C，BROWN C，GREHAN A．Towards high resolution habitat suitability modelling of vulnerable marine ecosystems in the deep-sea：resolving terrain attribute dependencies[J]．Marine Geodesy，2012，35：343-361．

[33] HOWELL K L，HOLT R，ENDRINO I P，STEWART H．When the species is also a habitat：comparing the predictively modelled distributions of Lophelia pertusa and the reef habitat it forms [J]．Biological Conservation，2011，144（11）：2656-2665．

[34] OREJAS C，GORI A，GILI J M．Growth rates of live Lophelia pertusa and Madrepora oculata from the Mediterranean sea maintained in aquaria [J]．Coral Reefs，2008，27：255．

[35] OREJAS C，FERRIER-Pages C，REYNAUD S，TSOUNIS G，ALLEMAND D，GILI J M．Experimental comparison of skeletal growth rates in the cold-water coral Madrepora oculata Linnaeus，1758 and three tropical scleractinian corals[J]．Journal of Experimental Marine Biology and Ecology，2011，405（1）：1-5．

[36] MIKKELSEN N，ERLENKEUSER H，KILLINGLEY J S，BERGER W H．Norwegian corals：radiocarbon and stable isotopes in Lophelia pertusa [J]．Boreas，1982，11：163-171．

[37] MORTENSEN P B，RAPP H T．Oxygen and carbon isotope ratios related to growth line patterns in skeletons of Lophelia pertusa（L）（Anthozoa，Scleractinia）：implications for determining of linear extension rates [J]．Sarsia，1998，83：433-446．

[38] LARTAUD F，PAREIGE S，DE RAFELIS M，FEUILLASSIER L，BIDEAU M，PERU E，ROMANS P，ALCALA F，BRIS N L．A new approach for assessing cold-water coral growth in situ using fluorescent calcein staining[J]．Aquatic Living Resources，2013，26（2）：187-196．

[39] RIDING R．Structure and composition of organic reefs and carbonate mud mounds：concepts and categories[J]．Earth-Science Reviews，2002，58：163-231．

[40] HOVLAND M．Do carbonate reefs form due to fluid seepage?[J]．Terra Nova，1990，2：8-18．

[41] CORDES E E，MCGINLEY M P，PODOWSKI E L．BECKER E L，LESSARD-PILON S，VIADA S T，FISHER C R．Coral communities of the deep Gulf of Mexico[J]．Deep-Sea Research Part I，2008，55：777-787．

[42] HUVENNE V A I，BAILEY W R，SHANNON P M，NAETH J，DI Primio R，HENRIET J P，HORSFIELD B，DE Haas H，WHEELER A．The Magellan mound province in the Porcupine Basin[J]．International Journal of Earth Sciences，2007，96：85-101．

[43] WILLIAMS T，KANO A，FERDELMAN T，HENRIET J P，ABE K，ANDRES M S，BJERAGER M，BROWNING E L，CRAGG B A，MOL B D，DORSCHEL B，FOUBERT A，FRANK T D，FUWA Y，GAILLOT P，GHARIB J J，GREGG J M，HUVENNE V A I，LéONIDE P，LI X，MANGELSDORF K，TANAKA A，MONTEYS X，NOVOSEL I，SAKAI S，SAMARKIN V A，SASAKI K，SPIVACK A J，TAKASHIMA C，TITSCHAK J．Cold-water coral mounds revealed[J]．EOS：Transactions American Geophysical Union，2006，87：525-526．

[44] CHENG H，ADKINS J，EDWARDS R L，BOYLE E A．U-Th dating of deep-sea corals[J]．Geochimica et Cosmochimica Acta，2000，64（14）：2401-2416．

[45] ELTGROTH S F，ADKINS J F，ROBINSON L F，SOUTHON J，KASHGARIAN M．A deep-sea coral record of North Atlantic radiocarbon through the Younger Dryas：Evidence for intermediate water/ deep water reorganization[J]．Paleoceanography，2006，21（4）：PA4207．

[46] CAIRNS SD，HAUSSERMANN V，FORSTERRA G．A review of the Scleractinia（Cnidaria：Anthozoa）of Chile，with the description of two new species[J]．Zootaxa，2005，1018：15-46．

[47] LINDBERG B，BERNDT C，MIENERT J．The Fugl?y Reef at 70_N：acoustic signature，geologic，geomorphologic and oceanographic setting [J]．International Journal of Earth Sciences，2007，96：201-213．

[48] DORSCHEL B，HEBBELN D，RUGGEBERG A，DULLO W C．Carbonate budget of a cold-water coral carbonate mound：Propeller Mound，Porcupine Seabight[J]．International Journal of Earth Sciences，2007，96：73-83．

[49] KANO A，FERDELMAN T G，WILLIAMS T．HENRIET J P，ISHIKAWA T，KAWAGOE N，TAKASHIMA C，KAKIZAKIY，ABE K，SAKAI S，BROWNING E L，LI X．Age constraints on the origin and growth history of a deep-water coral mound in NE Atlantic drilled during Integrated Ocean Drilling Program Expedition 307[J]．Geology，2007，35：1051-1054．

[50] REED J K，POMPONI S A，WEAVER D，PAULL C K，WRIGHT A E．Deep water sinkholes and bioherms of South Florida and the Pourtal_es Terrace：habitats and fauna[J]．Bulletin of Marine Science，2005，77：267-296．

[51] ORR J C．Recent and future changes in ocean carbonate chemistry [M]//GATTUSO J P，HANSSON L．Ocean Acidification．Oxford：Oxford University Press，2011：41-66．

[52] MAIER C，HEGEMAN J，WEINBAUER M G．Calcification of the cold water coral Lophelia pertusa under ambient and reduced pH [J]．Biogeosciences，2009，6：1671-1680．

[53] FORM A U，RIEBESELL U．Acclimation to ocean acidification during long-term CO2 exposure in the cold-water coral Lophelia pertusa [J]．Global Change Biology，2012，8：843-853．

[54] MAIER C，WATREMEZ P，TAVIANI M，WEINBAUER M G，GATTUSO J P．Calcification rates and the effect of ocean acidification on Mediterranean cold-water corals[J]．Proceedings of the Royal Society of London B：Biological Sciences，2012，279（1734）：1716-1723．

[55] MOVILLA J，OREJAS C，CALVO E，GORI A，LóPEZ-Sanz A，GRINYó J，DOMíNGUEZ-Carrió C，PELEJERO C．Differential response of two Mediterranean cold-water coral species to ocean acidification[J]．Coral Reefs，2014，33（3）：675-686．

[56] MAIER C，SCHUBERT A，BERZUNZA Sa′nchez M M，WEINBAUER M G，WATREMEZ P，GATTUSO J P．End of the century pCO2 levels do not impact calcification in Mediterranean cold-water corals[J]．PLoS One，2013，8（4）：e62655．

[57] EMILIANI C，HUDSON J H，SHINN E A，GEORGE R Y，LIDZ B．Oxygen and carbon isotopic growth record in a reef coral from the Florida Keys and a deep-sea coral from Blake Plateau[J]．Science，1978，202：627-629．

[58] SMITH J E，SCHWARCZ H P，RISK M J，MCCONNAUGHEY T A，KELLER N．Paleotemperatures from deep-sea corals：overcoming ‘vital effects’[J]．Palaios，2000，15：25-32．

[59] COHEN A L，LAYNE G D，HART S R，LOBEL P S．Kinetic control of skeletal Sr/Ca in asymbiotic coral：implications for the paleotemperature proxy[J]．Paleoceanography，2001，20（1）：20-26．

[60] CASE D L，ROBINSON L F，AURO M E，GAGNON A．Environmental and biological controls on Mg and Li in deep-sea scleractinian corals [J]．Earth and Planetary Science Letters，2010，300：215-225．

[61] BURKE A，ROBINSON L F．The southern ocean’s role in carbon exchange during the last deglaciation[J]．Science，2012，335：557-561．

[62] MANGINI A，LOMITSCHKA M，EICHSTADTER R，FRANK N，VOGLER S，BONANI G，HAJDAS I，PATZOLD J．Coral provides way to age deep water[J]．Nature，1998，392：347-348．

[63] COLIN C，FRANK N，COPARD K，DOUVILLE E．Neodymium isotopic composition of deep-sea corals from the NE Atlantic：implications for past hydrological changes during the Holocene[J]．Quaternary Science Reviews，2010，29（19）：2509-2517．

[64] COPARD K，COLIN C，HENDERSON G M，SCHOLTEN J，DOUVILLE E，SICRE M A，FRANK N．Late Holocene intermediate water variability in the northeastern Atlantic as recorded by deep-sea corals[J]．Earth and Planetary Science Letters，2012，313：34-44．

[65] MONTERO-Serrano J C，FRANK N，COLIN C，WIENBERG C，EISELE M．The climate influence on the mid-depth Northeast Atlantic gyres viewed by cold-water corals[J]．Geophysical Research Letters，2011，38：L19604．

[66] 张旭东．“蛟龙”号超额完成试验性应用南海航段任务[N/ OL]．[2013-07-10]．新华网，http://www.chinanews.com/gn/2013/07- 10/5027386.shtml．