On Some Octocorallia (Alcyonacea) from Hong Kong, with Description of a New Species, Paraminabea rubeusa1

Y. Benayahu2,4 and K. Fabricius3

Abstract: Octocorals from Hong Kong were studied at 18 sites down to a depth of 25 m in 1999. The collection of @90 specimens yielded nine species distrib- uted in seven genera of the families Alcyoniidae, Nephtheidae, and Xeniidae (all are new zoogeographical records for Hong Kong), plus ca. 70 samples of azoo- xanthellate octocorals of the genera Dendronephthya (family Nephtheidae), Chiro- nephthya, and Nephthyigorgia (family Nidaliidae), which were not identified to species level. The collection included Paraminabea rubeusa Benayahu & Fabri- cius, n. sp., which is described here. The impoverished nature of the zooxan- thellate octocorals is reflected in the low number of species found in the families Alcyoniidae and Xeniidae (seven and one, respectively), families that typically contribute a high proportion of species in the Indo-Pacific region. It is crucial to implement effective conservation policies in Hong Kong to preserve its remaining zooxanthellate octocoral species and thereby prevent the local ex- tinction of these species, including the newly described Sarcophyton tumulosum Benayahu & Ofwegen, 2009, and Lobophytum mortoni Benayahu & Ofwegen, 2009, which may be endemic to the region.

Hong Kong (latitude 22 N) represents a Kong have been studied extensively and are marginal environment for corals within the considered subtropical formations (e.g., Clark South China Sea (Morton and Blackmore 1998, McCorry 2002, Lam et al. 2007, and 2001). Its coral communities are nonreef- references therein). Notably, coral cover has building communities, due to winter seawater declined to low levels in several areas, appar- temperature minima averaging 17C (with ently due to impaired coral recruitment occasional brief drops to 14–16C in surface (McCorry 2002). waters), and freshwater intrusions frequently Taxonomic studies of the Hong Kong Oc- reducing salinity. Furthermore, the reefs of tocorallia have recently been reviewed by Hong Kong are affected by a variety of Lam and Morton (2008). Those studies were anthropogenic perturbations that have led initiated in 1854 during the North Pacific Ex- to their gradual deterioration (e.g., Morton ploring Expedition by Stimpson (1855), who 1994, 2002). The stony corals of Hong described the new species Nephthya coccinea, which was placed in Echinomuricea by Nutting (1910). They were followed by Verrill 1 Manuscript accepted 1 February 2009. (1865a,b), who identified and described a 2 Department of Zoology, George S. Wise Faculty of further six new species from the specimens Life Sciences, Tel Aviv University, Ramat Aviv 69978, collected earlier by Stimpson. Later, Shen Tel Aviv, P.O.B 39040, Israel. (1940) described Dendronephthya sp. and 3 Australian Institute of Marine Science (AIMS), Nephthya sp. from Hong Kong. After a gap PMB No. 3, Townsville MC, Queensland 4810, Austra- lia. of 40 yr in the study of octocorals from the 4 Corresponding author: (phone: 972-3-6409090; fax: area, Zou and Scott (1982) surveyed the gor- 972-3-6409403; e-mail: [email protected]). gonians from the northeastern part of Hong Kong, recording a total of 26 species belong- ing to 15 genera and six families: Antho- Pacific Science (2010), vol. 64, no. 2:285–296 doi: 10.2984/64.2.285 thelidae (one species), Subergorgiidae (two : 2010 by University of Hawai‘i Press species), Acanthogorgiidae (three species), All rights reserved Paramuriceidae (12 species), Plexauridae (five

285 286 PACIFIC SCIENCE . April 2010 species), and Ellisellidae (three species). Based materials and methods on a collection conducted in Hong Kong dur- ing 1980, Li (1986) recorded the families Al- Collection of material was conducted during cyoniidae (six species) and Nephtheidae (four November 1999 and comprised the following species), all of which were new records from 18 sites in Hong Kong: Bokhara Rocks (22 China. Based on a collection carried out in 13 0 25.71 00 N, 114 16 0 45.0900 E); Tai Tau 1995, Clark (1997) listed several octocorals Chau (Shek O) (22 13 0 56.63 00 N, 114 15 0 in the Cape d’Aguilar Marine Reserve (Hong 10.1900 E); Sham Wan (22 23 0 05.5900 N, Kong) in the families Paramuriceidae (nine 114 08 0 30.49 00 E); Pak Shau (Round Island) species), Telestidae (one species), Plexauridae (22 32 0 25.4500 N, 114 19 0 51.5800 E); Tsung (three species), Ellisellidae (two species), and O Wan ( Junk Bay) (22 16 0 39.4800 N, 114 Alcyoniidae (two species), including two new 15 0 12.45 00 E); Kung Chau (22 28 0 56.9100 N, records for the region. Based on a survey con- 114 220 14.1800 E); Ping Chao (22 32 0 22.30 00 ducted in 1999 on the shallow reefs of Hong N, 114 23 0 31.32 00 E); Breakers Rock (22 28 0 Kong, Fabricius and McCorry (2006) listed 39.5200 N, 114 50 13.72 00 E); Lo Chau Mun 42 octocoral species distributed among 23 (Po Toi Channel) (22 11 0 23.31 00 N, 114 genera of the families Alcyoniidae (six spe- 15 0 26.85 00 E); Waglan (22 11 0 02.6100 N, cies), Nephtheidae (six species), Nidaliidae 114 18 0 19.1600 E); Pak Kwo Chau (North (two species), Xeniidae (one species); Acan- Ninepin) (22 15 0 26.2300 N, 114 21 0 00.26 00 thogorgiidae (four species), Plexauridae (18 E); Nam Kwo Chau (South Ninepin) (22 species), and Ellisellidae (three species). Most 35 0 15.0500 N, 114 20 0 49.40 00 E); Fo Shek of the taxa mentioned in the study were not Chau (Basalt Island) (22 18 0 51.08 00 N, 114 identified to species level and are referred to 22 0 01.20 00 E); Long Ke Wan (22 190 12.83 00 as sp. (see table 2 of Fabricius and McCorry N, 114 22 0 58.25 00 E); Mat Chau (Po Toi) 2006). That survey presented results on the (22 10 0 12.67 00 N, 114 15 0 31.0200 E); Nam abundance of zooxanthellate and azooxan- Kok Tsui (Po Toi South) (22 12 0 12.29 00 N, thellate octocorals along the main spatial and 114 15 0 47.62 00 E); Lo Chau Pak Pai (Castle water quality gradients in Hong Kong, and Rock) (22 10 0 30.1500 N, 114 14 0 13.67 00 E); the authors concluded that the former are Luk Chau (Lamma) (22 13 0 42.43 00 N, 114 strongly affected by water clarity and that 07 0 28.4200 E) (Figure 1). their disappearance is associated with a dra- These sites were reached by boat, and a matic reduction in stony coral communities. careful examination of a variety of niches Recently, Lam and Morton (2008) reported was carried out by scuba diving down to 25 22 octocoral species from two submarine m. Approximately 90 samples were collected, caves in the oceanic southeastern waters of encompassing the variety of octocoral taxa Hong Kong, including one new species, Para- found at the various sites. Before collection minabea hongkongensis Lam & Morton, and an most of the colonies were photographed in additional 11 species that were new records situ. Samples were fixed in 4% formalin in for the region. In addition, that study pre- seawater overnight, then rinsed in freshwater, sented a list of 60 octocorals previously re- and transferred to 70% alcohol. Sclerites corded from Hong Kong. were obtained by dissolving the tissues in The study reported here deals with octo- 10% sodium hypochlorite and prepared for corals of the families Alyconiidae, Nephthei- scanning electron microscopy as follows: the dae, and Xeniidae from Hong Kong collected sclerites were carefully rinsed with double- by us in 1999. We provide a systematic list of distilled water, dried at room temperature, the species and describe a new species, Para- coated with gold, and then examined with an minabea rubeusa Benayahu & Fabricius. Nota- electron microscope ( Jeol 840A) operated bly, this collection has recently yielded two at 25 kV. Identification of species was in part other new species, Lobophytum mortoni Be- facilitated by comparisons with permanent nayahu & Ofwegen, 2009, and Sarcophyton sclerite preparations of type material kept tumulosum Benayahu & Ofwegen, 2009. in the Zoological Museum, Department of Octocorallia from Hong Kong . Benayahu and Fabricius 287

Figure 1. Map of Hong Kong showing the study sites. 1, Bokhara Rocks; 2, Tai Tau Chau (Shek O); 3, Sham Wan; 4, Pak Shau (Round Island); 5, Tsung O Wan ( Junk Bay); 6, Kung Chau; 7, Ping Chao; 8, Breakers Rock; 9, Lo Chau Mun (Po Toi Channel); 10, Waglan; 11, Pak Kwo Chau (North Ninepin); 12, Nam Kwo Chau (South Ninepin); 13, Fo Shek Chau (Basalt Island); 14, Long Ke Wan; 15, Mat Chau (Po Toi); 16, Nam Kok Tsui (Po Toi South); 17, Lo Chau Pak Pai (Castle Rock); 18, Luk Chau (Lamma).

Zoology, Tel Aviv University, Israel (zmtau). results The specimens are deposited in zmtau and in Nationaal Natuurhistorisch Museum, Leiden, The examined material yielded nine species The Netherlands (rmnh). Members of the in the three families Alcyoniidae, Nephthei- families Nephtheidae and Nidaliidae are still dae, and Xeniidae (Table 1), of which three being examined. are new species (Benayahu and Ofwegen 288 PACIFIC SCIENCE . April 2010

TABLE 1 description: The holotype is an arbo- List of Species of Octocorallia of the Order Alcyonacea rescent colony, 60 mm long and 55 mm wide (Lamouroux, 1816) from Hong Kong, with Museum (Figure 2a). Its has a low base, ca. 3–6 mm Inventory Numbers (zmtau Co), RMNH Coel high, that gives rise to a number of branching primary lobes. Lobes and lobules are finger- Family Alcyoniidae Lamouroux, 1912 like, stout with tapering ends, and compose Genus Cladiella Gray, 1869 most of the total length of the colony. Lobes Cladiella digitulata (Klunzinger, 1887) zmtau Co 30503 and lobules are clustered together, with some Cladiella pachyclados (Klunzinger, 1887) partially bent. The colony is dimorphic. zmtau Co 30555 Some of the autozooids with their white Genus Lobophytum von Marenzeller, 1886 anthocodiae protrude from polyp apertures. Lobophytum mortoni Benayahu & Ofwegen, 2009 Tentacles have 7–12 pinnules each, arranged zmtau Co 30508, 30517, 30546, 30550 Genus Paraminabea Williams & Alderslade, 1999 in one row along each edge. Siphonozooids Paraminabea rubeusa Benayahu & Fabricius, n. sp. minute (>1 mm in diameter) and hardly visi- zmtau Co 30536, 30545, 30536, 30562, RMNH ble with the naked eye. There are no sclerites Coel 38997 in the polyps. Genus Sarcophyton Lesson, 1834 Sarcophyton tumulosum Benayahu & Ofwegen, 2009 Surface of lobes with numerous six- and zmtau Co 30530, 30556 eight-radiates 0.05–0.08 mm long (Figure 3). Genus Sinularia May, 1898 Interior of lobes with two types of sclerites, Sinularia brassica May, 1898 including some radiates similar to those of zmtau Co 30516, 30532, 30549, 30553, 30564, the surface of the lobes, as well as elongated 34221 Sinularia inexplicita Tixier-Durivault, 1970 spindle like sclerites, 0.070–0.095 mm long, zmtau Co 30507 with thorns, either pointed or truncated, and Family Xeniidae Ehrenberg, 1828 a distinct median waist (Figure 4). Uppermost Genus Sansibia Alderslade, 2000 surface of base with similar radiates as on sur- Sansibia flava (May, 1898) zmtau Co 30513, 30514, 30515 face of lobes. Interior of base with numerous Family Nephtheidae Gray, 1862 barrels and a few radiates 0.05–0.09 mm long Genus Scleronephthya Studer, 1887 (Figure 5). Individual sclerites tinted reddish; Scleronephthya gracillimum (Ku¨kenthal, 1906) when packed densely on surface of lobes, zmtau Co 30497, 30509, 30510 30523, 30524, surface of base, and its interior, they provide 30541, 30552, 30556 colony with its red color; interior of lobes is white-beige. 2009; this study) and the other six are new color: Preserved colony is red with zoogeographical records for Hong Kong. white polyps; all sclerites reddish. No zooxan- Underwater photographs of some of the oc- thellae were found in the tissue. tocorals collected in this study are presented. living features: Red color; expanded autozooids white (Figure 6a). Colonies grow Paraminabea rubeusa Benayahu & Fabricius, in silt with base anchored to rocky substrate. n. sp. variation: The paratypes differ in size Figures 2–5, 6a (Figure 2c–g) but all have closely set lobes. There is variation in the degree of polyp re- type material: Holotype and four mi- traction: Co 30545 (Figure 2b–d) has numer- croscope slides (zmtau Co 34221), Waglan, ous protruding polyps, whereas Co 30562 Hong Kong, 16–18 m, 10 November 1999. (Figure 1e) and Co 30536 (Figure 1f,g) have Six paratypes: zmtau Co 30545 (two colo- retracted ones. All paratypes possess the nies), rmnh Coel 38997 (one colony), same same type of sclerites as the holotype. data as holotype; zmtau Co 30562, Bokhara etymology: The species name is derived Rocks, Hong Kong, submerged reef, 17 m, from Latin, rubeusa (red), in reference to the 12 November 1999 (one colony); zmtau Co color of the colonies. 30536, Mat Chau (Po Toi), Hong Kong, 18 remarks: The genus Paraminabea was m, 11 November 1999 (two colonies). established by Williams and Alderslade Figure 2. Paraminabea rubeusa Benayahu & Fabricius, n. sp.: a, holotype (zmtau Co 30530); b–d, paratypes: zmtau Co 30545; e, zmtau Co 30562; f, g, zmtau Co 30536. Scale bar 10 mm. 290 PACIFIC SCIENCE . April 2010

Figure 3. Paraminabea rubeusa Benayahu & Fabricius, n. sp.: holotype (zmtau Co 34221), six- and eight-radiates of surface of the lobes. Scale 0.02 mm.

(1999), referring to alcyoniid soft corals with these two species. Paraminabea rubeusa, n. sp., dimorphic colonies commonly unbranched has a more robust morphology, with closely and dome-shaped or hemispherical to digiti- set lobes (Figure 2). It also differs in its inte- form or, rarely, digitate/lobate with a few rior lobes possessing spindles with a distinct branching processes. Sclerites of surface or waist, whereas P. arborea Williams & Alder- interior of colony are densely packed: mostly slade, 1999, has mostly barrels and six- or barrels and six- or eight-radiates; spindlelike eight-radiates. Paraminabea rubeusa, n. sp., forms derived from radiates and tuberculate has barrels in the base interior, and P. arborea spheroids may also be present. Paraminabea Williams & Alderslade, 1999, has similar rubeusa, n. sp., fits the diagnostic features of sclerites that may resemble spheroids. It Paraminabea Williams & Alderslade, 1999, should be noted too that colonies of P. indica and thus raises to nine the number of species (Thomson & Henderson, 1905) can also be in this genus. Before this study, P. arborea lobate, with thick short lobes arising from Williams & Alderslade, 1999, was the only a distinct stalk that occupies as much as half digitate/branched species described in the of the colony’s total length (see Williams genus. There are several differences between 1992), thus differing from P. rubeusa, n. sp. Octocorallia from Hong Kong . Benayahu and Fabricius 291

Figure 4. Paraminabea rubeusa Benayahu & Fabricius, n. sp.: holotype (zmtau Co 34221), spindle like sclerites of in- terior of lobes. Scale 0.02 mm.

In addition, these two species have differently noted that our collection also yielded ca. 70 shaped sclerites. Paraminabea rubeusa, n. sp., samples of azooxanthellate octocorals of the differs too from the recently described P. family Nephtheidae and Nidaliidae, indicat- honkongensis Lam & Morton, 2008, both in ing their high abundance at the study sites colony morphology and sclerites. Therefore, (see also Fabricius and McCorry 2006). The based on the above comparisons, it is evident latter included Scleronephthya gracillimum that the examined specimens (Figure 2) have (Ku¨kenthal, 1906) (Figure 6b), and others a unique morphology and sclerites and differ not identified to species level from the genera from all previously described congenerics. Dendronephthya (family Nephtheidae, Figure The recent findings of Lam and Morton 6c), Chironephthya, and Nephthyigorgia (fam- (2008), along with ours reported here, further ily Nidaliidae, Figure 6d,e). It is interesting indicate a presumably high degree of ende- that no zooxanthellate nephtheid taxa were mism of Paraminabea species in the western found in our surveys. Moreover, the zooxan- Pacific, as was noted by Williams and Alder- thellate species of the families Alcyoniidae slade (1999). and Xeniidae had a rather limited distribution on the studied reefs, and they constitute only discussion a small part of the collection. This is demon- strated by the rarity on the reefs of Cladiella The overall objective of this survey was to in- pachyclados (Klunzinger, 1887) and Cladiella vestigate the octocoral fauna of the families digitulata (Klunzinger, 1887) (Figure 6f ). In Alcyoniidae, Nephtheidae, and Xeniidae in addition, this observation is further demon- Hong Kong. Our findings indicate an impov- strated by our finding that Sinularia brassica erished diversity of these three families May, 1898 (Figure 6g), and S. inexplicita (Table 1), which comprised only eight species Tixier-Durivault, 1970, were the only species distributed in seven genera. It should be of their genus found in Hong Kong (Table Figure 5. Paraminabea rubeusa Benayahu & Fabricius, n. sp.: holotype (zmtau Co 34221), barrels and radiates of in- terior of the base. Scale 0.02 mm. Figure 6. Underwater photographs of Hong Kong octocorals: a, Paraminabea rubeusa Benayahu & Fabricius, n. sp., with expanded autozooids; b, Scleronephthya gracillimum (Ku¨kenthal, 1906); c, Dendronephthya sp.; d, Nephthyigorgia sp. in part covered with sediment; e, Nephthyigorgia sp., closeup of branches; f, Cladiella digitulata (Klunzinger, 1887); g, Sinularia brassica May, 1898, and Sansibia flava (May, 1898) (bluish turquoise polyps). 294 PACIFIC SCIENCE . April 2010

1), compared, for example, with 32 Sinularia Henderson, 1905) from Cape d’Aguilar Ma- species in the South China Sea (Ofwegen rine Reserve. Due to the lack of an appropri- 2002), 30 in Taiwan (Benayahu et al. 2004), ate revision of the genus Dendronephthya, in and 15 in Singapore (Y.B., unpubl. data). the study reported here we could not identify The impoverished nature of the Hong Kong colonies of that genus with species in our zooxanthellate Alcyoniidae fauna is further collection or verify the appearance of D. gi- demonstrated by the low number of species gantea. Clark (1997) wrongly assigned D. gi- in the other recorded genera: Cladiella (two gantea (Verrill) to the family Alcyoniidae and species), Lobophytum (one), and Sarcophyton listed E. indica (Thomson & Henderson, (one), which typically contribute a much 1905), which is not a known taxon. It is possi- higher number of species in Indo-Pacific ble that the latter species was Paraminabea reefs (e.g., Ofwegen and Vennam 1994, Be- indica (Thomson & Henderson, 1905). nayahu 1997, 2002, Fabricius and Alderslade Fabricius and McCorry (2006) indicated 2001, Benayahu et al. 2004). Notably, our that the octocoral communities in Hong results are the first to report species of the Kong comprised 42 species in 23 genera. genera Sarcophyton and Sinularia for Hong Unfortunately Lobophytum depressum Tixier- Kong (see also Benayahu and Ofwegen Durivault, 1996, and Sarcophyton tenuispicula- 2009). Recently, Lam and Morton (2008) re- tum Thomson & Dean, 1931, were wrongly corded 20 octocoral species from submarine identified by Y.B., but their correct identifica- caves there and listed a total number of 60 tion is now presented in Table 1 (Lobophytum species already recorded from Hong Kong mortoni Benayahu & Ofwegen, 2009, and waters. Regrettably, that study did not in- Sarcophyton tumulosum Benayahu & Ofwegen, clude species of the families Alcyoniidae, 2009, respectively). Nephtheidae, and Xeniidae that had been re- Li (1986) identified 10 species from Hong corded earlier for Hong Kong by Fabricius Kong belonging to four genera of the fami- and McCorry (2006). Obviously, none of the lies Alcyoniidae [Cladiella humesi Verseveldt, species identified in the study reported here 1974; C. madagascarensis (Tixier-Durivault, (Table 1) appeared in Lam and Morton’s 1944); C. subtilis Tixier-Durivault, 1970; Lobo- (2008) list. The poor zooxanthellate octocoral phytum denticulatum Tixier-Durivault, 1956; fauna of Hong Kong is further indicated by L. depressum Tixier-Durivault, 1966; and L. the presence of Sansibia flava (May, 1898) venustum Tixier-Durivault, 1957] and Neph- (Figure 6g) as the only species found there theidae (Spongodes spinifera Holm, 1895; S. of the family Xeniidae (Table 1), whereas studeri Ridley, 1884; Scleronephthya corymbosa species of this family contribute substantially Verseveldt & Cohen, 1971; and S. pustulosa to the octocoral diversity in both the East and Wright & Studer, 1889). We found none of South China reef systems (e.g., Roxas 1933, these species, and only S. pustulosa Wright & Benayahu et al. 2004). It is suggested that it Studer, 1889, was also found in the recent is the environmental conditions prevailing in survey by Lam and Morton (2008). Hong Kong that have caused the paucity of It is striking that all of the species obtained Xeniidae species there, which require high- in our study are either new species or new clarity water (see also Fabricius and McCorry zoogeographical records for Hong Kong 2006; Y.B., pers. obs.). (Table 1). Species of the family Alcyoniidae Some of the previous studies on Hong are undoubtedly the best-known reef- Kong Octocorallia contain confusing mis- dwelling octocorals and are broadly distrib- takes. For example, Shen (1940) wrongly as- uted throughout the Indo-Pacific region signed Dendronephthya sp. and Nephthea sp. (e.g., Fabricius and Alderslade 2001, Be- from Hong Kong to the family Alcyoniidae nayahu 2002, Ofwegen 2002). Members of and also misspelled some of the taxa (i.e., Oc- this family are poorly represented in the tocoralla, Alcyonidae, Nephthya). Later, Clark Hong Kong reefs, however, and the record (1997) listed Dendronephthya gigantea (Verrill, of surveys there (e.g., Li 1986, Lam and Mor- 1864) and Eleutherobia indica (Thomson & ton 2008; this study) may suggest a decline in Octocorallia from Hong Kong . Benayahu and Fabricius 295 their richness over time and even the possible ———. 2002. Soft corals (Octocorallia: extinction of certain species, as reflected in Alcyonacea) of the southern Ryukyu the findings of Li (1986) compared with this Archipelago: The families Tubiporidae, study. Seawater quality has severely deterio- Clavulariidae, Alcyoniidae and Briareidae. rated in Hong Kong in the last few decades Galaxea JSRS 4:1–21. (Morton 1994), and its marine environment Benayahu, Y., M.-S. Jeng, S. Perkol-Finkel, has been chronically polluted by sewage, and C.-F. Dai. 2004. Soft corals (Octocor- land reclamation projects, industrial waste, allia, Alcyonacea) from southern Taiwan. contaminated sludge, trace metals, pesticides, II. Species diversity and distributional pat- and PCBs (McCorry and Cumming 1999, terns. Zool. Stud. 43:548–560. Fabricius and McCorry 2006, and references Benayahu, Y., and L. P. van Ofwegen. 2009. therein). The recent octocoral surveys there New species of Sarcophyton and Lobophytum (Fabricius and McCorry 2006; this study) (Octocorallia: Alcyonacea) from Hong have indicated that Indo-Pacific typical Kong. Zool. Meded. (Leiden) 83:863–876. shallow-water zooxanthellate octocoral com- Clark, T. H. 1997. The distribution of aher- munities are on the verge of disappearance matypic corals at Cape d’Aguilar, Hong from the Hong Kong reefs. Certain resilient Kong. Pages 219–232 in B. Morton, ed. species can nonetheless still cope with the en- Proceedings, 8th International Marine Bi- vironmental deterioration, and new ones are ology Workshop: The Marine flora and even being discovered there (Lam and Mor- fauna of Hong Kong and southern China, ton 2008; this study). In light of this finding, Hong Kong, 1995. Hong Kong University it is imperative to implement in Hong Kong Press, Hong Kong. effective conservation policies that will pre- ———. 1998. The distribution of hermatypic serve the remaining octocoral fauna and pre- sceractinian corals at Cape d’Aguilar, vent the local extinction of species, some of Hong Kong. Pages 151–164 in B. Morton, which may be endemic to the region. ed. Marine biology of the South China Sea: Proceedings, 3rd Conference on the Marine Biology of the South China Sea, acknowledgments Hong Kong, 1996. Hong Kong University Press, Hong Kong. We thank L. P. van Ofwegen and two anony- Fabricius, K. F., and P. Alderslade. 2001. Soft mous reviewers for comments on the manu- corals and sea fans: A comprehensive guide script. We are particularly grateful to B. to the tropical shallow water genera of the Morton for initiating the octocoral survey central-west Pacific, the Indian Ocean and and inviting us to carry out this research. the Red Sea. Australian Institute of Marine We thank K. Lam and D. McCorry for ad- Science, Townsville. vice and help during the fieldwork and the Fabricius, K. F., and D. McCorry. 2006. Swire Institute of Marine Science, The Uni- Changes in octocoral communities and versity of Hong Kong, for providing facilities. benthic cover along a water quality gradi- We thank Y. Delaria for valuable assistance ent in the reefs of Hong Kong. Mar. with the scanning microscopy, A. Shoob for Pollut. Bull. 52:22–33. photography, V. Wexler for graphic assis- Lam, K., and B. Morton. 2008. Soft corals, tance, A. 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