Discovery Bay, Jamaica 6 Phillip Dustan and Judith C. Lang

Abstract Keywords Mesophotic coral ecosystems occur on fore-reef slopes Mesophotic coral ecosystems · Discovery Bay · Jamaica · (~25 to 55 m) and near-vertical deep fore reefs (~55 to Fore-reef slope · Deep fore reef ~122 m) off the north Jamaican coast. In the 1960s, T.F. Goreau and colleagues found corals and calcareous green algae (Halimeda) thriving to depths of ~70 m and coralline sponges even deeper. Boring sponges and calcar- 6.1 Introduction eous algae produced prodigious quantities of carbonate muds and sands, some becoming lithified into reef rock Thomas F. (Tom) Goreau pioneered the scientific exploration and the remainder draining into deeper water. Low-light of subsurface tropical reef environments in the 1950s. His flattening of corals facilitated sediment shedding and sta- two classic papers on Jamaican reef ecology (Goreau 1959a) bilized dislodged colonies. Huge pinnacle reefs with frame and the physiology of coral skeleton formation (Goreau work-building Orbicella dominated the upper fore-reef 1959b) are seminal contributions to our understanding of slope (~25 to 45 m) at Discovery Bay. Foliaceous agarici- coral reefs. In 1965, he established the Discovery Bay Marine ids monopolized some lower slopes and the upper deep Laboratory (DBML), a small field station (Fig. 6.1a) located fore-reef. Submersible-based explorations of the deep in Discovery Bay (DB). DBML attracted so many visitors fore-reef cliff in 1972 confirmed active framework con- that funding was soon secured for a larger facility (Fig. 6.1b). struction by corals at ~55 to 70 m and by coralline sponges When Goreau died unexpectedly and prematurely in 1970, and crustose coralline algae at ~70 to 105 m. Species of shortly after opening his new lab, exploration of the nearby deep reef fishes occurred below ~60 to 90 m. The base of reefs to the absolute physiological limits of scuba diving had the cliff at ~91 to 145 m consisted of lithified sediments given DBML an international reputation for innovative coral and debris. Non-coralline sponges and algae were the reef research. dominant benthos from 53 to 120 m in a 1984 submersible This chapter is centered on the DB area of the Jamaican study. Fore-reef slope corals have declined greatly since north coast (Fig. 6.2). Enormous ecological changes have the 1980s, and their skeletons are largely overgrown with occurred in the decades since its deep reefs were first algae. Contributing factors include enhanced herbivore explored with scuba and submersible. Therefore, all but the losses from overharvesting (fish) or disease(Diadema) , concluding sections are partitioned into temporal periods, of repeated bleaching-mortality events, increased trapping of which the first (1960s–1970s) predates the widespread deg- sediments, degraded water quality, hurricanes, and coral radation of benthic reef structure and function that began in diseases. Apart from identified sponges collected with the 1980s and continues to the present. open circuit trimix scuba in the 1990s, the post-1984 con- dition of the deep fore reef is unknown. 6.1.1 Research History P. Dustan (*) Department of Biology, College of Charleston, Charleston, SC, USA 1960s–1970s Goreau (1959a) described the species com- e-mail: [email protected] position and zonation patterns of Jamaican corals at what J. C. Lang are now considered “shallow-intermediate” (<15 m) depths. Atlantic and Gulf Rapid Reef Assessment, Ophelia, VA, USA He then began to investigate deeper reefs off the north coast e-mail: [email protected]

© Springer Nature Switzerland AG 2019 85 Y. Loya et al. (eds.), Mesophotic Coral Ecosystems, Coral Reefs of the World 12, https://doi.org/10.1007/978-3-319-92735-0_6 86 P. Dustan and J. C. Lang

Fig. 6.1 DB Marine Laboratory: (a) original and (b) current. (Photo credits: P. Dustan) where a rich and diverse benthic biota extending over the The Buoy, adding qualitative observations to 75 m. Jackson edge of a narrow shelf (Fig. 6.3) was readily accessible to and colleagues (e.g., Jackson et al. 1971; Jackson and Buss divers in small boats (Goreau and Wells 1967). Unfamiliar 1976) investigated benthic organisms beneath flattened cor- plants and animals were collected, previously named spe- als at 30–67 m near DB and at 10–21 m on a vertical cliff cies were rediscovered or given range extensions, and some formed by a drowned river canyon at West Rio Bueno (WRB). new species were identified. Collaborative investigations of Dustan (1975) surveyed the collective cover of frame-­ the deep reefs at DB were well underway in 1970, and building Orbicella faveolata and O. franksi (formerly both Goreau was a posthumous co-author on many publications. Montastraea annularis, hereafter called Orbicella) to 45 m In 1973, two issues of the Bulletin of Marine Science (Vol. at Dancing Lady Reef (DLR). Porter et al. (1981) established 23, Nos. 1 and 2) were dedicated in his memory to research quadrats in 30–47 m at Monitor (Mon) in 1976 for biennial that had originated at DBML. photography. Hughes initiated population dynamics and life Early efforts to quantify the deep benthos at DB were history studies with approximately annual photoquadrats of focused on its seaward West Fore Reef (Fig. 6.2 shows the foliose agariciids and flattened-massive corals at 35 m on locations of the italicized reef sites named in the text). Kinzie Pinnacle One (Pinn1) and at the WRB cliff site (Hughes and (1973) described alcyonacean and coral zonation to 40 m at Jackson 1985). Benthic cover was first measured with line 6 Discovery Bay, Jamaica 87

Fig. 6.2 Discovery Bay (DB), Jamaica, showing offshore bathymetry, inset of reef bathymetry and sites: M Monitor (Mon), L Long-Term Sampling (LTS), D Dancing Lady Reef (DLR), Z Zingorro (Zin), P Pinnacle One (Pinn1), MI Mooring One (Moor1), B Buoy (The Buoy), WB West Bull (West Bull), DBL Dairy Bull (Dairy Bull). The West Rio Bueno (WRB) cliff site is about 6 km west of DB (Image: Google Earth; reef bathym- etry based on Moore et al. 1976)

transects in 30 m at Zingorro (Zin) in 1977 (Ohlhorst, pers. 1980–2017 A Perry PC-8B submersible that was stationed comm.). at DBML from 1984 to 1985 (RoseSmyth 1985) allowed In 1971, R. Ginsburg demonstrated in Belize (then British Liddell and Ohlhorst (1988) to extend their deep-reef sur- Honduras) that General Oceanographic’s two-person Nekton veys to 120 m below Zin and Mooring One (Moor1). About Beta submersible was easily adapted to investigate reef mar- the same time, a maximum limit of 30 m for “routine” gins (Ginsburg and James 1973; James and Ginsburg 1979). SCUBA diving was initiated at the lab (Gayle, pers. comm.); L. Land then secured funding to bring Nekton Gamma to most subsequent teaching and research has been conducted examine the organisms, framework, and sediments occurring at shallower (<25 m) depths. The Mon quadrats were photo- “below the depth of active reef growth” at DB in August 1972 graphed biennially until 1986 and in 2016 (Porter, pers. (Lang 1974). Nekton averaged 7 h underwater daily for comm.); those on Pinn1 and at the WRB cliff were photo- 28 days with a small lab boat as a tender. As 29 researchers graphed approximately annually until 1993 and in 2013 sequentially descended with a pilot to its maximum operating (Hughes 1996; Charpentier, pers. comm.). Andres and depth of 306 m and performed a few simple experiments, one Witman (1995) surveyed benthic cover data to 30 m at the of us (JCL) developed the camera film and a student tran- Long-Term Sampling (LTS) site in 1992. In 1999, Leichter scribed the taped notes. When submerged at a pressure of one and Genovese (2006) measured temperature between 10 and atmosphere inside the submersible, those among us who had 55 m at LTS for a year. Qualified technical divers used trimix thought our diving on the deep reefs was carefully executed to ~107 m for several years in the 1990s, collecting sponges were astonished to discover the inaccuracy of our scuba-based for taxonomic identification and biomedical assays (Lehnert scientific perceptions, thus ending the era of reef scientists and Fischer 1999; Gogineni and Hamann 2018). Mon was conducting research below 60 m on compressed air at DBML. surveyed to 46 m in 2005, and corals that later bleached at 88 P. Dustan and J. C. Lang

Fore-Reef Terrace Fore-Reef Escarpment Reef 10 a Lagoon Barren Crest Fore-Reef Slope Zone Inner 20 Outer b 40 Moat

60

80 Deep Fore- 100 Reef 120

140

Island Depth(m) Slope

N

apprx 100m

Fig. 6.3 (a) Profile of DB reef tract, with pinnacle reefs on the FRS and the drop-off at ~55 m (Based on Lang et al.1975 ); (b) deep reef sketch by T.F. Goreau

30–37 m were tagged for follow-up photography (Gayle ments (Greenaway and Gorgon-Smith 2006; Gayle, pers. 2009). In 2013, the DLR Orbicella was recensused to 45 m comm.). Terrigenous sediments, nutrients, and freshwater by Dustan (pers. obs.). accumulate offshore of streams and rivers, impacting nearby reefs during the rainy seasons and after storms. On reef slopes, fine-grained sediments drain into deeper water, unless 6.2 Environmental Setting constrained by coral formations (Goreau and Hartman 1963; Goreau and Goreau 1973). Some of the calcium carbonate Jamaica is situated ~145 km south of Cuba and ~190 km precipitated in situ by marine organisms on seaward reefs is west of Hispaniola in the West Indies. At 10,990 km2, it is the trapped within the coral framework and becomes lithified as third largest of the Greater Antilles and the largest island reef rock (Land and Goreau 1970). entirely surrounded by the Caribbean Sea. Northeasterly trade winds alternate with nocturnal offshore land breezes 1960s–1970s Water masses at DLR sometimes moved slowly unless interrupted by “Northers” (fronts from North America) eastward as countercurrents to their predominantly westerly in colder months. Jamaica’s north coast is bathed in surface flow direction, or upslope if deeper than 20 m, suggestive of waters from the Caribbean and Antilles Currents. The pre- upwelling (Dustan and Lang, pers. obs.). Near the mouth of dominant flow direction is westward but strong seasonal and the DB Channel, Kinzie (1973) recorded continuously chang- eddy variability lead to eastward and southward flow at times ing currents that were not obviously correlated with the tides (Mariano, pers. comm.). Wave fetch is limited and seas are at depths >28 m at The Buoy. At this site, J.C. Lang and usually moderate. Tides are mixed, ranging from 15 to 60 cm Y. Neumann were once swept downslope from 25 to 50 m by (Gayle and Woodley 1998). a strong current that dislodged loose corals and, given an No permanent rivers flow into DB, but freshwater enters unusually “dead slack” sea surface (von Arx 1962), was attrib- as surface runoff after rain. Groundwater originating inland uted to a large internal wave breaking on the island shelf. of DB continuously enters the bay via submarine springs, During the Nekton dives, an upslope current was encountered rocky shoreline fissures, and seepage through bottom sedi- at 110 m, contour currents were experienced in either an east- 6 Discovery Bay, Jamaica 89 erly or a westerly direction from 120 to 150 m, and downslope sonal variability, with a mean value of 28.1 °C, minimum of currents were felt below 150 m (Land and Moore 1977). 26.5 °C, and maximum of 30.1 °C (Leichter and Genovese Light fluctuates as the optical properties of seawater are 2006). Rapid cooling relative to the depth-averaged daily affected by changes in cloud cover, rain, waves, suspended mean temperatures was frequently recorded between May sediments, and phytoplankton and, in shallow water, by the and September being most pronounced near the edge of the optical properties of the reef (Dustan 1982, 1985). The DB island shelf at 45 and 55 m (Fig. 6.3). Thus (small) internal reefs face northward (Fig. 6.2), and the sun’s path illumi- waves may routinely oscillate the thermocline vertically over nates all but the deepest (>55 m) habitats for most of the day, the DB fore reefs, especially in summer (Leichter and with higher light levels for longer duration in summer. The Genovese 2006). Unusually warm seawater temperatures in submarine light field was a diffuse blue-green in the 1970s. the late summer-fall have been linked to bleaching, and Light levels decreased exponentially to about 2% of surface sometimes mortality, of DB corals in 1987, 1989, 1990, values, or approximately 1019 quantum scalar irradiance 1991, 1995, 1998, 1999, 2005, 2010 (Woodley 1988; Goreau (400–700 nm)/cm2/day, by 60 m at DLR in spring 1978 1992; Sebens 1994; Woodley et al. 1997, 2000; Gayle 2009), (Dustan 1982). and 2017 (Charpentier, pers. comm.). Divers commonly noted “mini-thermoclines” on the deeper fore reefs, especially after winter storms (Lang and Dustan, pers. obs.). Intermittent temperature measurements 6.3 Habitat Description at 15 m ranged between 29.2 °C in June 1969 and 26.0 °C in February 1970, and a maximum change of <2 °C was found Active tectonism (block faulting, uplift, and east-west dis- between depths of 12 m and 73 m (Kinzie 1973, pers. placement) and Quaternary glacioeustatic sea-level fluctua- comm.). The depth of the mixed (isothermal, surface) seawa- tions have created a series of Pleistocene reef terraces of ter layer off DB, as observed from Nekton, varied from ~50 locally varying size and elevation along the Jamaican north to 105 m in August 1972. Density schlieren (conspicuous coast. Modern reefs on the island shelf are presumed to over- layering of the upper thermocline layer) were frequently lay Pleistocene bedrock of variable topography (Goreau and recorded below ~90 m (Land and Moore 1977). Wells 1967; Hartman and Goreau 1970). Two distinctive habitats that correspond to “upper” and “lower” mesophotic 1980–2017 Hurricane Allen, the first violent storm since zones (Loya et al. 2016) are defined by abrupt changes in 1944 to strike DB, caused extensive damage to its shallower slope at their landward margins in 25–35 m and at a “drop-­ reefs in 1980 (Woodley et al. 1981; Gayle and Woodley off” in 55–70 m, respectively (Fig. 6.3). 1998). When Hurricane Gilbert again scoured them in 1988, When first explored by divers, Jamaican reefs had long rotting alcyonaceans (sea fans and sea whips) and sponges been artisanally overfished. The echinoid Diadema antilla- piled up in deeper channels and chutes. Both hurricanes dis- rum, a major reef herbivore, was abundant on shallow reefs placed vast quantities of sediment downslope over the deep (Jackson 1997). We can scarcely imagine how the deep ben- reefs (Woodley 1980, 1989). thos might have appeared had large predators and herbi- Terrigenous materials have accumulated in DB Harbour vores—finfishes, sharks, rays, turtles, manatees, and the from the bauxite port (Macdonald and Perry 2003; now-extinct Caribbean monk seal—still plied these waters. Fig. 6.2). Changes to upland drainage, local population growth, coastal development, and increased runoff from highway “improvements” now produce elevated turbidity 6.3.1 Fore-Reef Slope levels above the fore reefs that can persist for days or even weeks after heavy rain events (Gayle and Charpentier, 1960s–1970s The fore-reef slope (FRS) was a muddy-sand pers. comm.). By 2013, the underwater spectral light field plain of highly variable geomorphology and coral cover dip- had shifted toward green. Horizontal visibility at 30–50 m ping seaward at angles that steepen with increasing depth, on DLR had been reduced from ~20 to 30 m in the usually from ~20–30o at 25–35 m to ~60o at 50 m (Fig. 6.3). 1960s–1970s to ~10 to 20 m. The diffuse, conspicuous Sediments slowly flowed downslope around any coral for- “clouds” of particulate organic matter and gelatinous zoo- mations (Fig. 6.4a–c) or spread out as broad fans beneath the plankton in the upper 2–3 m of the water column in the channels or canyons draining the shallower fore reefs. Fleshy 1970s had become whiter and denser, with more fine par- and calcareous algae and seagrasses inhabited the sediments ticulates, less “organic snow,” and almost no gelatinous along with benthic foraminifera, Cassiopea, cerianthids, plankton (Dustan, pers. obs.). pennatulids, and garden eels. Goreau and Goreau (1973) Thermistors recording average temperature values at described the varied geomorphology and predominant coral 16-min intervals at 10, 20, 30, 45 and 55 m on LTS from cover of diverse Jamaican FRS sites. Some had huge talus January 1999 to January 2000 revealed relatively little sea- blocks, or pinnacle reefs (Fig. 6.5a), or immense foliose 90 P. Dustan and J. C. Lang

Fig. 6.5 FRS reefs: (a) Goreau at Pinn1, 25 m, 1966; (b) Agaricia undata “super colony,” West Bull, ~45 m, 1967. (Photo credits: E. Graham)

line algae, ascidians, foraminifera,­ filamentous algae, cyano- bacteria, and other cryptic biota grew under reduced light conditions beneath flattened corals on the FRS and in 10–20 m on the WRB cliff (Jackson and Winston 1982). (Cryptic habitats and their associated biota also occurred in Fig. 6.4 FRS sediments and (a) Penicillus, Udotea, and Rhipocephalus on the upper FRS, off Eaton Hall, Runaway Bay, 1967; (b) Halimeda the cavernous frameworks of some Jamaican reefs at <30 m and agariciid corals, The Buoy, 46 m, 1966; (c) corals and alcyonaceans depth (Jackson et al. 1971).) at the drop-off, Pinn1, 58 m, 1971. (Photo credits: E. Graham (a, b), Enormous (up to 10-m high) pinnacles (Fig. 6.6a) were P. Dustan (c)) capped with large, overlapping “shingles” of framework-­ constructing Orbicella between depths of ~25 and 45 m on the upper west FRS off DB (Goreau and Land 1974). In agariciid “supercolonies” (Fig. 6.5b). In areas of large sedi- 1977, live corals covered approximately half of the sub- ment input, corals were more abundant on the deeper FRS, stratum at 30 m on Zin and at 35 m on Pinn1 (Table 6.1). sometimes forming sill reefs, small pinnacles, or buttress- Foliose agariciids and flattened-massive Montastraea like mounds above the drop-off (Goreau and Goreau 1973; ­cavernosa, other benthic animals, and algae were inter- Goreau and Land 1974). Exposed rock surfaces were gener- spersed between patches of sediment from ~45 m to the ally covered with scleractinian corals, sponges, algae, alcyo- drop-off at ~55 m on the lower west FRS (Figs. 6.4c, 6.6b, naceans, antipatharians, fire coral (Millepora alcicornis, and 6.8a). Although plentiful on the shallower fore reefs, which also overgrew stalked alcyonaceans), and other sessile Diadema were naturally scarce on the FRS and generally organisms (Fig. 6.4c). Sponges, bryozoans, crustose coral- found no deeper than ~40 m. 6 Discovery Bay, Jamaica 91

(Fig. 6.7), no changes to the benthos were initially apparent at 30 m on Zin (Liddell and Ohlhorst 1986). Noncrustose algae continued their expansion at 35 m at Pinn1, reaching ~45% when coral cover and larval recruitment by corals started decreasing in 1986 (Hughes 1996). Noncrustose algae began to increase on the WRB cliff within 6 months (at 10 and 15 m) to a year (at 20 m) of Diadema’s 1983 demise (Hughes et al. 1987) and were the predominant benthic cover by 1986 (Table 6.1). Large-scale bleaching of Jamaican corals, including many Agaricia lamarcki to depths of 35 m, first occurred in 1987 during a period of unusually warm (maximum of 30.5 °C off DB) summer-fall seawater temperatures (Fig. 6.7; Woodley 1988). Hurricane Gilbert temporarily “scrubbed” noncrus- tose algae from the reefs in 1988 and smashed flattened cor- als on the WRB cliff (Woodley 1989). Many A. lamarcki at ~30 m bleached when >30 °C seawater temperatures lasted longer than usual in 1989 and 1990, followed by partial or complete tissue mortality of some colonies, especially those that had previously bleached (Savina 1991; Sebens 1994; Precht, pers. comm.). By 1989, noncrustose algal cover had increased to ~45% at 30 m on Zin and coral cover had decreased to ~33% (Table 6.1). Grazing pressures on reef algae were further reduced Fig. 6.6 Coral communities on DLR FRS in 1971 dominated by (a) when artisanal fishing activities intensified during the 1990s Orbicella spp. with downslope “trails” of lighter sediments, 30 m; (b) (Fig. 6.7) due to greater economic duress and increased use agariciids, Pseudoplexaura (large alcyonacean below diver), small of spearguns (Woodley and Sary 2000; Ennis and Aiken alcyonaceans, sponges, and Stichopathes lutkeni (wire coral antipathar- ians), 45 m. (Photo credits: P. Dustan) 2014). In 1992, coral cover at 30 m on LTS was only half that of macroalgae (27% vs. 52%); by 1993, coral coverage at 35 m at Pinn1 had fallen to ~13% and noncrustose algae 1980–2017 In 1980, Hurricane Allen (Fig. 6.7) smashed, occupied ~76% of the benthos, while corals were <6% at toppled, or scoured more corals on the exposed East Fore 10–20 m on the WRB cliff (Table 6.1). Reef at DB than on the West Fore Reef. Shallow habitats Although data are lacking for its effects on FRS corals, were generally damaged more severely than deeper habitats bleaching was severe at DB in 1995 and milder in 1998 and (Porter et al. 1981; Woodley et al. 1981). A huge coral over- 1999 (Woodley et al. 1997, 2000). A major bleaching event hang at ~20 m was detached from the fore-reef escarpment occurred in 2005 (Fig. 6.7d; Jones et al. 2008) when tem- south of Pinn1, landing among a group of similar blocks at peratures reached 30.2 °C for several weeks during the top of the FRS (Lang, pers. obs.). Some foliose agariciids September-October (Gayle 2009). About 80% of the corals were damaged to a depth of 50 m further west on the west in phototransects on Mon at 30 m and ~65% of those at 37 m FRS. Coral cover immediately after the hurricane was bleached (Charpentier, pers. comm.), with some sustaining unchanged from pre-storm estimates of 64% at 33 m on Mon partial or complete mortality. After a minor bleaching event and of 71% at 10 m on the WRB cliff, where losses at both 15 in 2010, <5% of corals photographed on the upper FRS at and 20 m were only from 33% to 32% (Woodley et al. 1981). Mon showed any mortality (Charpentier, pers. comm.). Coral cover at 35 m on Pinn1 had declined by about 10% in The upper west FRS pinnacles still retain their “shingled” 1981 (Table 6.1), presumably from storm damage, and non- morphology (Fig. 6.7c). By 2013, some of the surviving crustose algae (= macroalgae + turf algae) had increased DLR Orbicella showed signs of disease, with sediment-­ from ~9% to 15% (Hughes 1996). In 1982, coral cover on trapping turf algae growing immediately adjacent to the Zin was greater at 30 m than deeper, whereas noncrustose affected tissues (Fig. 6.7h). Most of the “deepwater” Agaricia algae showed the opposite trend from ~12% at 30 m, where (Fig. 6.8a) on rocks in the sand channels and on mounds in the density of Diadema was ~0.1/m2, to ~32% at 45 and the lower FRS had either died or disappeared, and the reef’s 56 m (Table 6.1). three-dimensional structure just above the drop-off had When most Diadema antillarum in Jamaica perished in largely collapsed (Fig. 6.8b). Small (juvenile) corals, primar- summer 1983 during its Caribbean-wide mass mortality ily Agaricia and Mycetophyllia, were scarce; no Orbicella or 92 P. Dustan and J. C. Lang

Table 6.1 Mean percent composition data for corals, macroalgae, turf algae, and noncrustose algae at sites on the (a) west FRS (30–56 m) and (b) the WRB cliff (10–20 m) Depth (m) Site Date ∑ Orbicella ∑ Coral Macroalgaea Turf algaeb Noncrustose algaec References Discovery Bay West fore-reef slope sites 30 DLR 1972 ~53 1 2013 ~8 1 Zin 1977 45.5 51.3 5.4 3 ~8 2 1982 54.2 ± 29.3 58.9 ± 9.7 8.2 ± 7.9 4.1 ± 3.5 ~12 3 1989 ~33 ~45 4 LTS 1992 ~15 27 ± 7.6 52 ± 8.0 3.3 ± 1.5 ~55 5 35 DLR 1972 ~43 1 2013 ~10 1 Pinn1 1977 ~20 47.5 ± 4.1 7 6 1980 50.8 ± 5.5 6 1981 40.5 ± 5.3 6 1985 >40 26 ± 6 7 1993 12.6 ± 2.7 76 ± 4 7 45 DLR 1972 ~9 1 2013 ~1 1 Zin 1980 ~43 ~26 4 1982 19.1 ± 16.0 45.0 ± 12.4 26.5 ± 5.4 5.3 ± 4.8 ~32 3 1984 13.3 ± 8.3 43.1 ± 9.3 3.6 ± 2.9 ~47 8 M1 1984 21.5 ± 11.9 26.4 ± 21.0 3.8 ± 1.9 ~30 8 56 Zin 1982 0 38.5 ± 14.1 24.4 ± 21.0 7.6 ± 11.0 ~32 3 Cliff sites WRB 10 1977 <1 9 1983 ~43 <1 9 10 1986 23.9 ± 3.4 ~13 ~59 72.4 ± 2.8 9 10 1993 5.4 ± 1.2 10 WRB 15–20 1977 <2 9 1983 <2 9 1986 13.1 ± 7.5 ~15 ~66 80.9 ± 9.3 9 1993 5.6 ± 0.9 10 1 Dustan (1975) and Dustan (unpubl. data); line intercept transects 2 Liddell and Ohlhorst (1981) and Ohlhorst (pers. comm.); line point transects 3 Liddell et al. (1984); line point transects 4 Liddell and Ohlhorst (1992); line point transects in 1980, phototransects in 1989 5 Andres and Witman (1995); line point transects 6 Hughes (1996); marked photoquadrats 7 Hughes and Tanner (2000); marked photoquadrats. 8 Liddell and Ohlhorst (1988) and Liddell (pers. comm.); phototransects 9 Hughes et al. (1987); marked photoquadrats 10 Hughes (1994); line-intercept transects aAlso called fleshy algae bAlso called filamentous algae c= macroalgae + turf algae

M. cavernosa recruits were found (Dustan, pers. obs.). The drop-off and an island slope at ~122 m that dipped seaward prevalence and intensity of bleaching at 30 m on Mon in late to oceanic depths. Near-vertical-overhanging promontories 2017 resembled that observed in 2005 (Charpentier, pers. (Fig. 6.9), usually located beneath coral-covered areas of comm.). the lower FRS, alternated with reentrants positioned mostly below sand channels or chutes, creating a “vertically corru- gated” morphology. Irregular promontory surfaces had 6.3.2 Deep Fore Reef many projecting ledges (Fig. 6.10a) and numerous holes, caves, and crevices. Several large caverns of unknown ori- 1960s–1970s The deep fore reef (DFR) is a rugged cliff of gin were found at ~82 to 91 m. Sediments covered all the spectacular relief. At DB, it extended between the ~55 m horizontal and sloping surfaces (Fig. 6.10b) as they cas- 6 Discovery Bay, Jamaica 93

100

Overfishing intensifies

Water quality deteriorates 80 a

DLR b est

c 60

Orbicella ZIN %Cover (30m) DLR ZIN LTS d 40 ZIN

Algae e non-crustose %cover (30m)

20 Severe Chronic Diadema bleaching Overfishing mortality Lionfish Bleaching invasion events 0 1970 1980 1990 2000 2010 Hurricanes Allen Gilbert Algal overgrowth Algal overgrowth Yellow band begins continuing disease common fgh

Fig. 6.7 Ecological changes on upper FRS reefs at DB, 1972–2017, as Orbicella decreased and non-crustose algae increased, overgrowing corals and other benthos. Cover data in 30 m from DLR, Zin, and LTS (Table 6.1); est. = noncrustose algae estimated from photographs. Multiple stressors included chronic overfishing,Diadema mortality, bleaching events, hurricanes, land-based sources of pollution, and introduced lionfish. All photos except (d) on DLR. (a) Orbicella, ~35 m, 1972; (b) fish trap, ~32 m, 2014; c( ) Orbicella remnants overgrown by Halimeda, other noncrustose algae and invertebrates, 32 m, 2015; (d) severely bleached Orbicella on LTS, ~35 m, 2005; (e) lionfish(Pterois), 26 m, 2014; (f) turf algae, macroalgae, and cyanobacteria overgrowing H. cucullata and A. agaricites, 27 m, 2014; (g) Caulerpa overgrowing Orbicella and Agaricia, 35 m, 2013; and (h) Orbicella with yellow-band disease, 35 m, 2013. (Photo credits: P. Dustan (a–c, e–h), B. Charpentier (d))

caded down the promontory faces. Debris and sediments of their dead skeletons produced a reef framework with a shed from the shallower­ reef habitats scoured the reentrants, predominant growth direction that was laterally outward limiting ­epibenthic organisms primarily­ to crustose coral- with an upward component (Land and Moore 1977). Rock line algae (Lang 1974; Lang et al. 1975). samples excavated at ~65 m below Pinn1 had a horizontal The upward-facing surfaces of the promontory ledges accretion rate of approximately 0.22 m/1000 years (Land were dominated to depths of ~70 m by agariciids and 1974). Halimeda and fleshy green algae were also com- flattened-­massive corals (Figs. 6.10a and 6.11a) that were mon on the ledges (Fig. 6.10a, b). Near-vertical surfaces of tilted seaward, facilitating sediment drainage. Cementation the promontories on the upper DFR were covered with 94 P. Dustan and J. C. Lang

Fig. 6.10 Upper DFR promontory, Pinn1, 1971: (a) large overhang with corals (M. cavernosa, H. cucullata, M. aliciae, and M. reesi), Halimeda, sponges (including Agelas spectrum below the M. caver- nosa), Antillogorgia, 60 m; and (b) small ledges with corals, Halimeda Fig. 6.8 FRS just above the DLR drop-off, 55 m: (a) Agaricia undata patch, 1973; and (b) same location with algae, a few sponges, and sediment (above), and sponges (below), 63 m. (Photo credits: corals, and alcyonaceans, 2013. (Photo credits: P. Dustan (a), P. Dustan) L. Wheeler (b))

massive, tubular, and branching sponges and encrusting corals and other cnidarians (Figs. 6.10b and 6.11b). Numerous colorful organisms, including Ceratoporella nicholsoni and other coralline sponges (formerly called sclerosponges), crustose coralline algae, peyssonnelids, encrusting sponges, azooxanthellate corals, whip and fan antipatharians, hydroids, bryozoans, brachiopods, tube worms, crinoids, and ascidians, were restricted to areas protected from falling sediments under ledges or corals and in crevices and caverns (Jackson et al. 1971; Hartman 1973; Lang 1974). Zooxanthellate corals were scarce below depths of ~75 m, presumably from light limitation on a northward-facing cliff. Sponges and other sessile organisms dammed sediments and debris that fell from above onto the promontory ledges in the mid DFR. The most conspicuous calcifying organisms at 70–105 m were nodular Ceratoporella that, along with some crustose coralline algae, grew on vertical surfaces and on Fig. 6.9 Upper DFR: Goreau and agariciids at The Buoy, 55 m, 1967. downward-facing surfaces of ledges or cracks where they (Photo credit: E. Graham) 6 Discovery Bay, Jamaica 95

Fig. 6.11 DFR promontories, 1972: (a) flattenedM. cavernosa with A. spectrum at the drop-off, 58 m; (b) fleshy sponges shaded from sedi- mentation below an overhang. (Photo credits: Nekton Project: J. Woodley (a), P. Goreau (b))

were sheltered from sediments. Some Ceratoporella received enough light for endolithic green algae to grow in their outer skeletons (Fig. 6.12a), but the upper surfaces of any that grew in the open had died and were either overgrown by other organisms or covered with sediment (Fig. 6.12b). Rock samples collected from 85 to 105 m consisted of coralline sponges, lithified sediments, and crustose coralline algae that were creating a reef framework (Fig. 6.12b, c) oriented pre- Fig. 6.12 Mid DFR promontories, 1972, C. nicholsoni: (a) with green endolithic skeletal algae and partially under a ledge near azoo- dominantly laterally outward with a downward component xanthellate corals (Thalamophyllia riisei), hydroids, and sponges; (b) (Lang et al. 1975). with crustose coralline algae and encrusting sponges; (c) partially Below ~105 m, the ledges and inclined outer surfaces of overgrown with sponges, sediments, and algae, interspersed with wire the promontories were relatively barren and blanketed with coral antipatharians, and creating reef framework, vertical lines are ~3 m apart, 84 m. (Photo credits: Nekton Project: P. Goreau (a, b), mucus-rich, sediment-trapping mats of unknown microbial J. Woodley (c)) origin that were too fragile to collect with Nekton (Fig. 6.13a). C. nicholsoni and other sponges were still abundant on verti- cal surfaces (Fig. 6.13a, b), along with crustose coralline mechanically abraded by flowing sediments (Lang1974 ; algae. Lithified rock at the base of the DFR was composed Land and Moore 1977). primarily of fallen sediments and debris (Fig. 6.13c). Gullies Spiny lobsters, crabs, sea urchins, and brittle stars were in the vertical reentrants at these depths were being actively the primary motile scavengers and grazers observed on the bioeroded by endolithic green algae and probably also DFR. The large “primitive” gastropod, Entemnotrochus 96 P. Dustan and J. C. Lang

abundant and widespread, with ~16 to 28% cover at 53– 120 m. Turf algae were fairly common (~16 to 21%) below about 68 m. Zooxanthellate corals (~9 to 17%) were only found at 53 and 60 m. Macroalgae, averaging ~14%, reached a maximum depth of ~75 m. Crustose coralline algae aver- aged ~11% from 53 to 90 m, but only ~1% at 106 and 120 m. Coralline sponges were present in low abundance (~1 to 7%) from 60 to 90 m (Liddell and Ohlhorst 1988).

6.4 Biodiversity

Classification and taxonomy have undergone radical changes in recent decades. The scientific names for organisms given in the text or tables in earlier publications have been changed, as needed, from their original designations to conform to nomenclature accepted by the 2017 World Register of Marine Species (marinespecies.org). The apparent number of spe- cies recorded in surveys on the deep DB reefs has varied over time, both among investigators and in response to taxonomic changes. No attempt has been made to update earlier esti- mates of species diversity or evenness.

6.4.1 Macroalgae

Fore-reef slope. 1960s–1970s. Calcareous green and red macroalgae (Table 6.2) were important FRS sediment pro- ducers. Among the earliest of the deepwater Jamaican species to be described, Halimeda cryptica (Colinveaux and Graham 1964) and H. copiosa (Goreau and Graham 1967) were attached to coral skeletons and rocks, along with other Halimeda and red calcareous macroalgae. Diverse calcareous green algae (Penicillus, Udotea, Rhipocephalus, and some Halimeda) inhabited the fine- grained sediments on the slopes (Fig. 6.4a). Crustose cor- alline algae and peyssonnelids overgrew solid surfaces. Fleshy brown and red ­macroalgae and the lightly calcified brown Padina were common below ~40 m (i.e., Diadema’s maximum grazing depth) in some areas of the FRS Fig. 6.13 Lower DFR promontories, 1972: (a) sediments trapped (Goreau and Goreau 1973). by “mucous mats” on ledges and sponges in vertical areas, 116 m; (b) C. nicholsoni, other sponges and turf algae, 130 m; (c) fallen debris and sediment with white segments of Halimeda, 125 m. (Photo credits: 1980–2017 Macroalgae that proliferated after Diadema’s Nekton Project: J. Lang (a), H. Reiswig (b), N. James (c)) demise at 35 m on Pinn1 were mostly fleshyLobophora and deepwater Halimeda (Hughes 1996). Gayle (2009) found Halimeda and fleshy macroalgae (Lobophora, Cladophora, adansonianus, was first seen in its natural habitat below Dictyosphaeria, Dictyota, and Sargassum) at 28–42 m on depths of 115 m (Yonge 1973). Most fishes were small, and Monitor in 2005. In 2013, complex cyanobacteria-turf algal sightings of larger vertebrates (several groupers, a nurse assemblages and macroalgae were trapping sediments and shark, and a turtle) were rare (Lang 1974). copiously overtopping corals on DLR (Fig. 6.7c, f, g) at 25–40 m. Halimeda, Lobophora, and the fleshy Caulerpa 1980–2017 Benthic cover was quantified in 1984 (Liddell, were particularly conspicuous macroalgae along the drop-off pers. comm.) with phototransects from the PC-8B submers- at 50–55 m. Crustose coralline algae and peyssonnelids still ible at Zin and Moor1. Non-coralline sponges were the most covered rock surfaces (Dustan, pers. obs.). 6 Discovery Bay, Jamaica 97

Table 6.2 Distribution of calcareous FRS and DFR chlorophytes and down the promontory faces DFR (Fig. 6.13a, c) and a com- FRS rhodophytes near Discovery Bay, Jamaica, in 1973 ponent of the lithified rocks collected at the cliff base (Land Fore-reef and Moore 1997, 1980). slopea Deep fore reefb Chlorophyta Upper Lower Upper Mid Lower Halimeda copiosa x x x x 6.4.2 Anthozoans H. cryptica x x x x H. discoidea x x x H. gracilis var. opuntioides x x 6.4.2.1 Scleractinian Corals H. goreaui x x Early scientific scuba diving on deep Jamaican reefs H. incrassata x x increased the list of zooxanthellate scleractinians from 35 H. opuntia x species in 1959 to 50 in 1973 (examples in Fig. 6.14), one of H. simulans x x which, Madracis pharensis, also occurred in an azooxanthel- H. tuna x x late form. The number of azooxanthellate species (including H. tuna f. platydisca x x M. pharensis f. pharensis) rose from 5 to 13 (Goreau 1959a; Penicillus capitatus x Wells and Lang 1973). Wells (1973a) described five new spe- P. dumetosus x x cies and one new form of zooxanthellate corals. Mycetophyllia P. lamourouxii x ferox (which does not inhabit the deep reefs), M. reesi, and P. pyriformis x x M. aliciae (Fig. 6.14g, h) were distinguished by differences Rhipocephalus phoenix x x Udotea conglutinata x x in aggressive behavior and habitat distribution in addition to U. cyathiformis x x skeletal characters (Lang 1973). Wells (1973b) also described U. flabellum x Gardineria minor, a new, solitary azooxanthellate coral with U. spinulosa x a wide depth range that encompassed the deep reefs. U. wilsonii x x Fore-reef slope. 1960s–1970s. Forty-six species of zoo- Total 20 15 3 2 xanthellate corals were recognized on the upper FRS Rhodophyta (Table 6.3a). Fifteen species characteristic of shallower Amphiroa x reefs reached their lower depth limits here, and five other Dichotomaria obtusata x x (formerly Galaxaura obtusata species (A. grahamae, A. undata, M. pharensis f. luciphila, v. major) M. aliciae, and M. reesi) were at, or near, their upper depth Ganonema farinosum x limits. Two “deepwater” Agaricia (A. grahamae and A. (formerly Liagora farinosa) undata), Helioseris (formerly Leptoseris) cucullata, and M. G. pinnatum (formerly L. x cavernosa were proportionately more abundant on the pinnata) lower FRS, where ~31 zooxanthellate corals were recorded. Jania x In 1972, Orbicella cover on DLR varied from ~53% at 30 m Liagora ceranoides x Titanophycus validus x to 9% at 45 m and was ~46% at 30 m in 1977 on Zin (formerly Liagora valida) (Table 6.1). In 1977–1980, cover at 35 m on Pinn1 aver- Total 7 1 aged ~20% for Orbicella and ~25% for agariciids (A. Species names and distributions on the fore-reef slope based on Goreau lamarcki > H. cucullata > A. agaricites); abundance by and Goreau (1973) numbers of physiologically distinct colonies was A. agaric- Nomenclatural differences from Goreau and Goreau (1973) conform to ites > H. cucullata > Orbicella > A. lamarcki (Hughes and information online in the World Register of Marine Species (marinespe- cies.org) in October 2017 Jackson 1985). Goreau and Wells (1967) reported nine azo- aFore-reef slope, upper = ~25 to 45 m; lower = ~45 to 55 m oxanthellate coral species at upper FRS depths, five of bDeep fore reef, upper = ~55 to 70 m; mid = ~70 to 100 m; lower = ~100 which were found on the lower FRS along with to ~120 m Balanophyllia floridana (Table 6.3b).

1980–2017 Although much diminished since the early 1980s Deep fore reef. 1960s–1970s. Halimeda cryptica and H. (Table 6.1), the skeletons of Orbicella still dominate the copiosa were prominent to ~100 m on the promontories upper FRS (Fig. 6.7c). In 2013, when its cover at DLR was (Fig. 6.10a, b), with cover estimated at ~10% at ~61–91 m. ~8% in the area formerly of highest (~53%) cover at 30 m and H. cryptica was the more abundant, locally occupying up to ~1% at 45 m, only 59 of the 210 colonies surveyed between ~60% of the substratum (Moore et al. 1976). Crustose coral- 27 and 45 m in 1972 were alive (Dustan unpubl. data). line algal cover estimates ranged from 5% to 25% (Hartman Deep fore reef. 1960s–1970s. Less was known about the 1973). White, disarticulated Halimeda segments were visu- composition and distribution of corals, especially the azoo- ally conspicuous on the light-colored sediments that flowed xanthellates, below the drop-off. Foliose agariciids, 98 P. Dustan and J. C. Lang

Fig. 6.14 FRS zooxanthellate corals at DB: (a) flattenedM. cavernosa, 60 m, 1971; (b) flattenedOrbicella , 28 m, 1971; (c) Eusmilia fastigiata f. flabellata, 1968; (d) Agaricia lamarcki, 50 m, 2013; (e) Agaricia grahamae and Orbicella, 33 m, 2013; (f) Helioseris cucullata, 30 m, 2013; (g) Mycetophyllia reesi, 28 m, 1968; (h) Mycetophyllia aliciae, 27 m, 2013; (i) Madracis formosa, 1966. (Photo credits: P. Dustan (a, b, d–f, h), E.A Graham (c, g, i))

flattened-­encrusting M. cavernosa, Mycetophyllia, and 6.4.2.2 Alcyonaceans Madracis dominated the 21–22 zooxanthellates in the upper DFR (Table 6.3a; Figs. 6.9 and 6.10a). Twelve of these spe- 1960s–1970s Nicella goreaui (Fig. 6.15a) was described cies were noted at mid depths; the only in situ zooxanthel- from specimens first collected by Goreau at DB (Bayer1973 ). late coral seen with Nekton on the lower DFR was an Kinzie (1973) found 24 zooxanthellate holaxonians and two Agaricia at 99 m. Eight species of azooxanthellate corals scleraxonians on the upper FRS at The Buoy; 12 of the former were present on the upper DFR (Table 6.3b). Thalamophyllia also occurred on the lower FRS (Table 6.4a). Species richness (formerly Desmophyllum) riisei (Fig. 6.12a) and other azoo- at 24–40 m was correlated to the diversity of the substratum xanthellates too small to identify from inside the submers- (live coral, sponges, sand, dead coral, and rock), whereas col- ible grew abundantly on the undersurfaces of foliose or ony density was greatest in areas of reduced coral cover and flattened-­massive corals below overhangs and in holes or with more exposed hard surfaces (Kinzie 1973). Although caves (Lang 1974). Eunicea and Pseudoplexaura were more conspicuous, numer- 6 Discovery Bay, Jamaica 99

Table 6.3a Distribution of FRS and DFR zooxanthellate scleractini- Table 6.3a (continued) ans near Discovery Bay, Jamaica, in 1973 Zooxanthellate Fore-reef slopea Deep fore reefb a Zooxanthellate Fore-reef slope Deep fore reefb scleractinians Upper Lower Upper Mid Lower scleractinians Upper Lower Upper Mid Lower Siderastrea radians x Acropora cervicornis x x S. siderea x x x A. prolifera x Solenastrea bournoni x Agaricia agaricites x x x x Stephanocoenia intersepta x x x x A. fragilis x x x x x (formerly michelinii) A. grahamae x x x x Total 46 31 21–22 12 1 A. lamarcki x x x x Species names and distributions primarily based on Goreau and Wells A. undata x x x x (1967), Lang (1971), Goreau and Goreau (1973), Pang (1973a), Wells Colpophyllia natans x x (1973a), Wells and Lang (1973), Gayle (2006), and Charpentier (pers. Dendrogyra cylindrus x comm.) Dichocoenia stokesii x x x Nomenclatural differences from Wells and Lang (1973) conform to Diploria labyrinthiformis x information online in the World Register of Marine Species (marinespe- cies.org) in October 2017 Eusmilia fastigiata x x x aFore-reef slope, upper = ~25 to 45 m; lower = ~45 to 55 m Favia fragum x bDeep fore reef, upper = ~55 to 70 m; mid = ~70 to 100 m; lower = ~100 Helioseris cucullata x x x x to ~120 m Isophyllia rigida x Madracis auretenra x x x? (formerly mirabilis) M. decactis x x x Table 6.3b Distribution of FRS and DFR azooxanthellate scleractini- M. formosa x x ans near Discovery Bay, Jamaica, in 1973 M. pharensis f. luciphila x x x x Azooxanthellate Fore-reef slopea Deep fore reefb Manicina areolata x x scleractinians Upper Lower Upper Mid Lower Meandrina danae (formerly x x x Astrangia solitaria x meandrites f. danai) Balanophyllia floridana x x Meandrina jacksoni x x (formerly meandrites) Colangia immersa x x Meandrina meandrites x x Caryophyllia sp. cf. x x x x C. antillarum Montastraea cavernosa x x x x Gardineria minor x x x x Mussa angulosa x x Guynia annulata x x Mycetophyllia aliciae x x x Madracis pharensis f. x x x x M. danaana x pharensis M. ferox x Paracyathus pulchellus x x x x ? M lamarckiana x x (formerly dephilippi) M. reesi x x x x Phacelocyathus (formerly x Oculina diffusa x Caryophyllia) flos Orbicella (formerly x Rhizosmilia (formerly x Montastraea) annularis Caryophyllia) maculata O. faveolata (formerly M. x x x Thalamophyllia (formerly x x x x ? annularis) Desmophyllum) riisei O. franksi (formerly M. x x x Tubastraea coccinea x annularis) Total 9 6 8 7 ?2 Porites astreoides x x x x Species names and distributions based on Goreau and Wells (1967), P. divaricata x Wells and Lang (1973), and Wells (1973b) P. furcata x x Nomenclatural differences from Goreau and Wells (1967) conform to P. porites x information online in the World Register of Marine Species (marinespe- Pseudodiploria (formerly x cies.org) in October 2017 and from Cairns (pers. comm. for Diploria) clivosa Phacelocyathus flos and Rhizosmilia maculata) P. (formerly D.) strigosa x aFore-reef slope: upper = ~25 to 45 m; lower = ~45 to 55 m Scolymia cubensis x x x x bDeep fore reef, upper = ~55 to 70 m; mid = ~70 to 100 m; lower = ~100 S. lacera x x x to ~120 m (continued) 100 P. Dustan and J. C. Lang

Fig. 6.15 DFR organisms: (a) orange Nicella goreauii with cryptic Analcidometra armata, DB, 1972; (b) Stichopathes lutkeni close-up behind Davidaster discoidea, DB, 128 m, 1972; (c) C. nicholsoni close-up showing excurrent canals increasing in size and converging at oscular openings, Pear Tree Bottom cavern, Runaway Bay, 25 m, 1973; (d) spotted moray eel, Gymnothorax moringa, DLR, 55 m drop-off, 2013. (Photo credits: Nekton Project: J. Woodley (a), N. James (b); Other: P. Dustan (c, d)) ical dominants were Antillogorgia (formerly Antipathes (A. atlantica and A. gracilis) are also common. Pseudopterogorgia) bipinnata at ~24 to 33 m and A. (formerly Two commercially important species have tree shapes, one P. ) elisabethae at ~33 to 60 m. The number of zooxanthellate of which (Plumapathes pennacea) is rare; the other species decreased with increasing depth, while azooxanthellae (A. caribbeana) is small (<1 m tall) and often intermingles species, such as Ellisella elongata (formerly barbadensis), with large Iciligorgia on cliff edges exposed to the longshore Iciligorgia schrammi, and Diodogorgia nodulifera, were current. The sprawling A. rubusiformis clings by multiple increasingly common below ~45 m (Table 6.4b). Five azoo- holdfasts to the undersides of overhangs in 30 m at Rio xanthellates were observed to depths of at least 100 m, of Bueno, and the cave-dwelling A. umbratica was once found which N. goreaui and Hypnogorgia pendula (?) were the two inside a crevice at DB (Warner 2005, pers. comm.). most common on ledges in the upper DFR (Kinzie 1973).

6.4.2.3 Antipatharians 6.4.3 Sponges

1980–2017 Warner (2005, pers. comm.) has observed seven Ceratoporella nicholsoni was initially thought be a tabulate species of antipatharians since 1998 at upper FRS depths off coral when a skeleton was dredged from 200 m off Cuba in DB and on the WRB cliff. Stichopathes lutkeni, an unbranched the nineteenth century, then a bryozoan, and eventually “wire coral” (Figs. 6.6b, 6.12c, and 6.15b), is the most abun- described as an octocoral by Hickson (1911). Finding this dant, its holdfasts being restricted to cryptic locations below enigmatic creature abundantly lining the walls of caves, crev- projecting pieces of coral or within crevices. Two fan-shaped ices, and tunnels in cryptic Jamaican reef habitats was a 6 Discovery Bay, Jamaica 101

Table 6.4a Distribution of FRS zooxanthellate alcyonaceans near Table 6.4b Distribution of FRS and DFR azooxanthellate alcyona- Discovery Bay, Jamaica, in 1973 ceans near Discovery Bay, Jamaica, in 1973 Fore-reef slopea Fore-reef Zooxanthellate alcyonaceans Upper Lower slopea Deep fore reefb Suborder Holaxonia Azooxanthellate alcyonaceans Upper Lower Upper Mid Lower Antillogorgia (formerly Pseudopterogorgia) x Suborder Calcaxonia acerosa Ellisella elongata x x A. (formerly P. ) americana x Nicella goreaui (formerly x x x x x A. (formerly P. ) bipinnata x x spp.) A. (formerly P. ) elisabethae x x Suborder Holaxonia Eunicea clavigera x x Hypnogorgia pendula (?) x x x E. (formerly Plexaura) flexuosa x x Lignella richardi x x x E. fusca x Swiftia exserta x x x E. laciniata x Suborder Scleraxonia E. pinta x x Diodogorgia nodulifera x x Gorgonia flabellum x Iciligorgia schrammi x x x x x G. mariae x Total 4 4 5 5 5 G. ventalina x Species names and distributions based on Kinzie (1973) and Bayer Muricea laxa x x (1973) Muriceopsis flavida x Nomenclatural differences from Kinzie (1973) based on information M. petila x x online in the World Register of Marine Species (marinespecies.org) in Plexaura homomalla x Oct. 2017 and Robert Kinzie (pers. comm.) a Plexaura kukenthali (formerly homomalla f. x x Fore-reef slope, upper = ~25 to 45 m; lower = ~45 to 55 m b kukenthali) Deep fore reef, upper = ~55 to 70 m; mid = ~70 to 100 m; lower = ~100 to ~120 m Plexaurella dichotoma x x P. grisea x P. nutans x x Pseudoplexaura flagellosa x x living or dead coral skeletons on the FRS at DB. Four other P. porosa x x clionaids (formerly called clionids) C. caribbaea (as C. P. wagenaari x langae), C. janitrix, C. schmidti, and C. (formerly Pterogorgia guadalupensis x Anthosigmella) ­varians were also collected in this habitat Suborder Scleraxonia (Pang 1973b). Reiswig (1973) observed that at least 100 Briareum asbestinum x species of non-coralline sponges could be found on the Erythropodium caribaeorum x FRS, many perhaps attaining their greatest biomass per unit Total 26 12 surface area in the FRS. Species names and distributions based on Kinzie (1973) Nomenclatural differences from Kinzie (1973) based on information online in the World Register of Marine Species (marinespecies.org) in 1980–2017 Agelas, Ircinia, Mycale, Neofibularia, Oct. 2017 and Robert Kinzie (pers. comm.) Spheciospongia, and Verongula (formerly Verongia) were aFore-reef slope, upper = ~25 to 45 m; lower = ~45 to 55 m the most abundant genera at 35 m on Pinn1 from 1977 to 1993 (Hughes 1996). When the “shallow-water” (to 60 m) momentous discovery (Hartman and Goreau 1970). C. nich- Jamaican demosponges were listed in 1998, ~41 species olsoni is a massive, nodular coralline sponge (Figs. 6.12, inhabited FRS habitats in the general area of DB. One of 6.13b, and 6.15c) with a compound skeleton of crystalline these, Plakina jamaicensis, was a new species, and another, aragonite, siliceous spicules, and organic fibers closely Mycale jamaicensis, had been previously described by resembling the stromatoporoids that were important reef con- Pulitzer-Finale in 1986 (Lehnert and van Soest 1998). structors during the early Paleozoic and late Mesozoic Deep fore reef. 1960s–1970s. Although M. normani was (Hartman and Goreau 1970; Jackson et al. 1971; Wood 1990). only found at 83 and 91 m, the other coralline sponges were Further scuba searches provided Hartman (1969) with speci- widespread on the DFR. As in shallower habitats, C. nichol- mens for describing four new coralline species (Goreauiella soni was the largest, growing to >1 m in diameter in caves auriculata, Hispidopetra miniana, Stromatospongia norae, where protected from falling sediments. It was also the most and S. vermicola) and recording a rare Merlia sp. that is now abundant (Fig. 6.12), except at depths of ~60 to 70 m, where classified withM. normani (van Soest 1984). S. vermicola was more common (Lang et al. 1975). The non- Fore-reef slope. 1960s–1970s. All six Jamaican species coralline sponge fauna on the upper DFR was similar to that of coralline sponges grew on the undersides of FRS corals of the lower FRS, with the addition of enormous (nearly 1 × and talus. Pang (1973b) described three new species of bor- 2 m) individuals of Agelas flabelliformis. Hartman (1973) ing (cavity-excavating) sponges (Cliona aprica, C. pepona- estimated that sponge cover on the promontories increased cea, and Siphonodictyon brevitubulatum) that inhabited from ~40% to 50% near the top of the DFR to ~60 to 80% at 102 P. Dustan and J. C. Lang their bases. Although not visible from Nekton, boring and Table 6.5 Distribution of FRS and DFR finfishes near Discovery Bay, cavity-dwelling sponges were present in all rock samples Jamaica in 1974 collected from 72 to 120 m (Lang 1974). Many of the corals Fore-reef slopea Deep fore reefb that accumulated on the lower DFR were thought to have Finfishes Upper Lower Upper Mid Lower fallen after boring sponges weakened the bases of their skel- Apogon lachneri x x x x etal attachments to the substratum (Hartman 1973). Apsilus dentatus juvc juvc x x Bodianus pulchellus x x x Canthigaster rostrata x x x x 1980–2017 During four trimix dives at 70–90 m on LTS in Caranx ruber x x x 1993, Lehnert gathered specimens for the first taxonomic Cephalopholis cruentata x x x x study of Caribbean DFR demosponges. Ten of the 27 described (formerly Epinephelus species were new (Lehnert and van Soest 1996). Further tri- cruentatus) mix dives in 1994 extended the depth range of the collections Chaetodon guyanensis x from 60 to 107 m and the number of sponge species to 60. Chromis cyanea x x x Twenty-three of these species were new, 24 also occurred in C. insolata (formerly x x x insulatus) shallower reef habitats, and 36 were restricted to the DFR C. scotti x x x x x (Lehnert and Fischer (1999). Species found on less inclined C. enchrysurus x x x surfaces differed from those with more vertical orientations or Clepticus parrae x x x under flattened corals and overhangs where they would have Coryphopterus personatus x x been protected from sedimentation. Lehnert and Fischer Cookeolus boops x (1999) noted that the inventory of DFR sponges was still far Elacatinus evelynae x x from complete. E. (formerly Gobiosoma) x x x x louisae Emmelichthys ruber x x Epinephelus mystacinus x 6.4.4 Fishes Equetus lanceolatus x x x x E. punctatus x x x x x Fore-reef slope. 1960s–1970s. As elsewhere along the nar- Ginglymostoma cirritum x x x row north Jamaican coast, artisanal overfishing had already Gonioplectrus hispanus x x depleted larger-sized reef fishes (grouper, snapper, and par- Gramma melacara x x x x x rotfish). Young individuals of small-sized fishes dominated Gramma linki x x the fishery (Woodley and Sary 2000) and the deep DB reefs. Gymnothorax moringa x x x x x Fish traps were frequently placed on sand in the upper FRS, Haemulon striatum x x x x x where “ghost traps” slowly moved downslope via surface Halichoeres garnoti x x x x creep (Goreau and Land 1974) unless retrieved by divers for Holacanthus tricolor x x x x x Holanthius martinicensis x x the fishers. Holocentrus adscensionis x x x x x FRS dives in the 1960s often attracted a great barracuda H. marianus x x x x x (Sphyraena barracuda) to the flash bulbs then used with H. rufus x x x x x underwater cameras (Lang, pers. obs.). Bicolor damselfish Hypoplectrus puella x x x x (Stegastes partitus) were the most abundant pomacentrid Ioglossus helenae x x x x x below 30 m in 1971, and herbivorous fishes were scarce Lipogramma evides x (Colin 1971, pers. comm.). Colin (1978) studied a group of L. klayi x x several hundred, small striped parrotfish(Scarus iseri, L. trilineata x x x x ­formerly croicensis) that moved daily from their shallower Lipoproma aberrans x L. mowbrayi x x x x x feeding areas to spawn above the DLR pinnacle, occasionally Lucayablennius zingaro x x x x attracting predatory mackerel (Scomberomorus sp.) or liz- Lutjanus analis x x x x x ardfish (Synodus). Some of the released eggs were quickly L. buccanella juvc juvc x x x consumed by groups of blue chromis (Chromis cyanea, for- Mulloidichthys martinicus x x x x x merly cyaneus) and creole wrasse (Clepticus parrae). Pristigenys alta x Table 6.5 lists the 40–41 species observed during the Nekton Prognathodes aculeatus x x x x x dives in 1972 (Colin 1974). Ptereleotris helenae x x x x Rypticus saponaceus x x x x x Schultzea beta x x x x 1980–2017 In 1984, Itzkowitz et al. (1991) surveyed DB fish populations with the PC-8B. The most common of the Seriola dumierli x x x x x 43 species encountered at 50 m were the blackcap basslet (continued) 6 Discovery Bay, Jamaica 103

Table 6.5 (continued) over 100 m, verifying that the depth range and distribution of Fore-reef slopea Deep fore reefb some gobies are invariably tied to the distribution of their Finfishes Upper Lower Upper Mid Lower host sponges (generally tubular aplysinids). About 60% of Serranus luciopercanus x x the 51 species of observed or collected DFR fishes were Sparisoma atomarium x x x x shallow-­water species with maximum depth ranges of 120– Sphyraena barracuda x x x 150 m; a true “deep reef” fauna, including Epinephelus Urolophus jamaicensis x x x x x ­mystacinus, Gonioplectrus hispanus, Holanthias marticen- Xanthichthys ringens x x x x x sis, Lipogramma klayi, and Serranus luciopercanus, Total 40 41 39 40 33 appeared below ~60 to 90 m. Juvenile snappers (Apsilus den- Species names and distributions based on Colin (1974) and Colin (pers. tatus and Lutjanus buccanella) were seen in <30 m although comm. for E. evelynae) Nomenclatural changes from Colin (1974) based on information online adults were absent shallower than 70–80 m. Transects at in the World Register of Marine Species (marinespecies.org) in Oct. depths of 90, 105, and 120 m indicated that fish densities 2017 and with Patrick Colin (pers. comm. for Gramma linki) decreased with increasing depth (Colin 1974). a Fore-reef slope, upper = ~25 to 45 m; lower = ~45 to 55 m Five species of Grammatidae (basslets) were present on bDeep fore reef, upper = ~55 to 70 m; mid = ~70 to 100 m; lower = ~100 to ~120 m the deep DB reefs, and their importance at mesophotic cJuveniles only Caribbean depths was first revealed duringNekton dives. The blackcap basslet (Gramma melacara) occupied cervices and caves in vertical areas of the DFR, while the bicolor basslet (Gramma melacara), sunshine fish(Chromis insolata, for- (Lipogramma klayi) ranged from 90 to 125 m on steeply merly insolatus), blue chromis, and creole wrasse. Some sloping rock shelves that were dammed or covered by sedi- non-territorial cocoa damselfish(Stegastes variabilis) ment (Colin 1974). The Jamaica work was pivotal in the roamed over large areas without visibly affecting small eventual realization (Starck and Colin 1978) that G. melac- groups of striped parrotfish and carnivorous spotted goatfish ara, arguably the most abundant mesophotic reef fish of the (Pseudupeneus maculatus). western/central Caribbean, is absent from the eastern Invasive Indo-Pacific lionfish (Pterois spp.), first reported Caribbean, with the break in its range at the Mona Channel a few km east of DB in 2008, were soon found throughout in the north and the Peninsular de Guajira in the south. Jamaican waters (Chin et al. 2016). Fishing has continued unabated on the FRS and fish traps are still in use (Fig. 6.7b). The most common of the 18 species recorded in a short sur- 6.4.5 Other Biotic Components vey by Wheeler (pers. obs.) at 30 m on DLR in 2013 were tiny bluehead wrasse (Thalassoma bifasciatum), blue 6.4.5.1 Comatulid Crinoids chromis, bicolor damselfish, and small princess parrotfish (Scarus taeniopterus). Red hind (Epinephelus guttatus) were 1960s–1970s Davidaster discoidea (formerly Nemaster; the largest grouper present and no snappers were seen. A few see Hoggett and Rowe 1986) was the most common crinoid larger predators were also noted on the DLR FRS in 2013– on both the FRS and DFR off DB in June-July 1968, where 2015: lionfishes to depths of ~35 m (Fig. 6.7e), an adult it clung beneath corals or crevices exposing only its extended green moray (Gymnothorax funebris), smaller spotted arms for feeding (Fig. 6.15b). Thirteen individuals of the morays (G. moringa; Fig. 6.15d), an eagle ray (Aetobatus characteristically shallower-occurring D. (formerly N.) narinari), and southern stingrays (Hypanus americanus; rubiginosa were repeatedly found along with six D. discoi- Dustan, pers. obs.). dea on a large outcrop on the upper FRS near The Buoy Deep fore reef. 1960s–1970s. Colin (1974) employed (Meyer 1973a). Analcidometra armata (formerly carib- deep scuba diving, deepwater trapping, and Nekton in the baea, Meyer et al. 1978) clasped the stalks or branches of first comprehensive effort to characterize DFR fish commu- Antillogorgia spp. and other FRS holaxonians, spreading its nities (Table 6.5). One submersible-assisted collection arms in a filtration fan plane parallel to its perch (Meyer method allowed otherwise unobtainable specimens to be 1973a). On the DFR, A. armata grasped Nicella goreaui picked up with a dip net attached to a manipulator that was (Fig. 6.15a) and Stichopathes lutkeni with arms generally “muscle-powered” by the collector immediately after the extended normal to the prevailing current. Crinoid species geologists had gathered rock samples. A second technique, richness and population densities were generally higher on involving ichthyocides in a hot-water bottle, proved less suc- continental Colombian and Panamanian reefs than in cessful. Two new DFR basslets, Gramma linki (Starck and Jamaica when Meyer (1973b) suggested that overall avail- Colin 1978) and Lipogramma evides (Robins and Colin ability of their suspended food particles was greater in con- 1979) were collected. The sponge-dwelling spotlight goby tinental areas of higher or more diversified primary (Elacatinus (formerly Gobiosoma) louisae) was collected to productivity. 104 P. Dustan and J. C. Lang

6.4.5.2 Ascidians >20 species were found growing abundantly to 70 m in the DB area. Individual colonies of some species attained large 1960s–1970s Descriptions by Millar and Goodbody (1974) sizes and, in some places, corals covered very large areas of of new Jamaican ascidians included four species (Ascidia the FRS (Figs. 6.5 and 6.6) and upper DFR (Goreau and xamaycana, Bathypera goreaui, Clavelina puertosecensis, Wells 1967). Reef framework construction was initially pre- and Halocynthia microspinosa) that had been collected on sumed to be limited to “shallow, well-illuminated, turbulent the DB FRS and/or DFR. B. goreaui was only found attached environments above wave base” (Goreau 1963). Subsequent to the undersides of overhanging coral rocks; specific habitat research at DB showed that benthic organisms grow on a details were not provided for the remainder. framework of modern corals (to ~70 m) or of coralline sponges and crustose coralline algae (at ~70 to 105 m) that lithify in situ with reef sediments by processes which are 6.4.5.3 Brachiopods likely to be microbially mediated (Land and Goreau 1970). The morphology and species composition of deep 1960s–1970s Jackson et al. (1971) described the distribu- Jamaican reefs emerges in part from the physiological/evolu- tion and abundance patterns of three brachiopod species tionary responses of the zooxanthellate corals and other ben- encrusting the undersurfaces of flattened corals from 30, thic organisms to decreasing light energy. Kawaguti (1937) 45, to 60 m at The Buoy and from 30, 45, 60, to 67 m at described the flattening of massive corals in shallow, shaded West Bull (see Fig. 6.2). Argyrothea johnsoni was the larg- habitats in Palau. Massive zooxanthellate corals on the est and most widespread brachiopod on the FRS and rela- Jamaican FRS and DFR have similarly flattened morpholo- tively more abundant away from the growing edges of large gies (Goreau and Hartman 1963; Fig. 6.14a, b, e, h). In situ corals. Its maximum densities were at 60 m and greater at and transplant experiments with Orbicella on DLR revealed The Buoy, which is less exposed to sedimentation, than at that under conditions of low light intensity, and presumably West Bull (~23 m−2 vs. ~16 m−2, respectively). The smaller low rates of light-enhanced calcification, a flattened mor- Thecidellina barretti was more common below ~60 to phology allows corals to increase in surface area by adding 80 m, and Crania pourtalesii was the least abundant spe- new polyps at their growing edges, with substantially less cies (Jackson et al. 1971). increase of skeletal volume than occurs when massive corals grow in unshaded, shallow habitats (Dustan 1975, 1979, 1982). The small caves which are gradually created below 6.4.5.4 Bryozoans flattened Orbicella in ~25 to 45 m as it grows can extend >2–3 m out from the substratum, enhancing rugosity on the 1960s–1970s Steginoporella sp. nov., Reptadeonella “pla- upper FRS (Fig. 6.6a). giopora” (formerly violacea), Stylopoma spongites, and Erect corals, sponges, alcyonaceans, and antipatharians Parasmittina sp. were the most abundant of 15 cheilostome increase the external geometric complexity of reefs, whereas species encrusting the undersurfaces of flattened corals col- burrowing organisms (fungi, algae, sponges, worms, and lected in 1977 at 11–20 m on the WRB cliff (Jackson 1979). bivalves) weaken coral skeletons, rendering them more sus- In 1978, bryozoans were usually more common under corals ceptible to mechanical destruction during storms. Clionaid taken at 10 m than at 21 m, and near the outer edges of the sponges, the major bioeroders of Jamaican fore reefs, release corals than toward their bases. Abundances varied greatly, small carbonate chips identifiable in the silt-sized sediment especially for Steginoporella and Stylopoma, and the compo- components that flow down the FRS and over the DFR sition of the edge-zone bryozoan communities, which now (Moore et al. 1976). Thin foliose Agaricia undata and A. totaled 46 species, was found to vary significantly with coral grahamae found on deep reefs (especially >50 m) are heav- size (Jackson and Winston 1982; Jackson 1984). ily bored at their bases and, unless held in place by large encrusting sponges, are easily detached by their own weight or that of any attached organisms or from diver-generated turbulence. Skeletal flattening provides a broad area of 6.5 Ecology ­contact for snagging on surface irregularities that may pre- vent dislodged foliose corals from overturning and/or sliding Shallow-water Jamaican reefs (<15 m) are shaped by envi- downslope (Goreau and Hartman 1963). The cryptic habitats ronmental factors that vary seasonally and geographically created by flattening shelter diverse, sediment-intolerant and by the biological attributes of their constituent coral spe- encrusting organisms (Jackson et al. 1971; Jackson and cies (Goreau 1959a). Earlier perceptions of the depths to Winston 1982). which zooxanthellate corals occur were overturned when 6 Discovery Bay, Jamaica 105

Flattened corals encounter other organisms primarily at had only decreased from ~59% to 33% between 1982 and their growing margins. Hierarchical spatial competition 1989 (Table 6.1). Corals were also more abundant at 30 m among corals was first noticed on the FRS at DB when rela- than in 5–20 m on LTS in 1992 (Andres and Witman 1995). tively small or slowly growing mussids deployed digestive Coral cover at 10 m in the Port Royal Cays (Fig. 6.2) and mesenterial filaments as a defense against direct overgrowth along the north coast dropped from ~52% at 10 sites before by more rapidly expanding branching and foliose species, Hurricane Allen to ~3% at 15 sites in 1990–1993, while mac- both in situ and in the wet laboratory (Lang 1973). Jackson roalgae increased from ~4% to ~92% (Hughes 1994). and Buss (1976) described nontransitive competitive net- These initial differences have been obscured by the simi- works among cryptic sponges and ascidians encrusting the larities of their phase shifts: loss of reef architectural struc- undersides of Agaricia at 15–60 m using tissue homogenates ture and complexity from collapse of fragile corals in aquaria. Complex nontransitive networks of competitive (Acropora spp. in shallow habitats and Agaricia spp. on the overgrowth abilities that occurred naturally under flattened lower FRS), extensive partial mortality of surviving massive corals were found to be more common between, rather than corals, macroalgal overgrowth of reef framework builders within, species of different major taxonomic groups. Spatial (especially Orbicella) and crustose coralline algae, and relationships (e.g., for cheilostomes—encounter angles and diminished recruitment of coral larvae on substrata covered active vs. inactive zooids), relative growth rates, and redi- with noncrustose algae. However, Edmunds and Bruno rected growth were recognized as important determinants of (1996) described large (kilometer-scale), horizontal varia- benthic distribution patterns, delaying resource monopoliza- tions in coral and macroalgal cover, coral species richness, tion by any single crustose species on space-limited hard and juvenile coral densities at depths of 10 m between DB substrata (Buss and Jackson 1979; Jackson 1979). and Runaway Bay (Fig. 6.2). Early descriptions and photo- Known interactions among corals soon expanded to graphs also reveal enormous spatial heterogeneity in the include diverse mechanisms of direct tissue contact (e.g., geomorphology and ecology of Jamaica’s FRS habitats sweeper tentacles on the FRS Montastraea cavernosa at DB; (Goreau and Hartman 1963; Goreau and Goreau 1973). Richardson et al. 1979) or indirectly at a distance. Outcomes Regrettably, our understanding of the condition of FRS varied from unilateral exclusion to temporal dominance reefs, both before and after Hurricane Allen in the 1980s, is reversals (Chornesky 1989) or apparently benign “stand-­ limited to the area around DB. offs.” Complex, natural competitive networks resulted Positive changes may also occur more quickly in shal- from variations in the morphology, size, growth rates, growth lower water. Although some tissues died, the Orbicella and plasticity, health, and motile associates of competing corals, other FRT corals that bleached in 1987 recovered zooxan- along with spatial and temporal effects (Lang and Chornesky thellae faster than the discolored Agaricia lamarcki in 1990). As their abundance on the FRS has declined, corals 15–35 m (Woodley 1988). Patches of Diadema appeared on encounter other corals less frequently and are generally los- the shallow FRT in the DB area in the early-mid 1990s ers when competing with algae and cyanobacteria (Figs. 6.7 (Woodley 1999). Within 3 years, Diadema populations in and 6.8). 4–6 m on LTS had increased by an order of magnitude, and a Mesophotic habitats have been proposed to be more sta- combination of crustose coralline algae, turf algae, and “bare ble and environmentally more predictable than those at shal- space” had replaced macroalgae as the predominant benthos lower depths, potentially serving as a refugia and/or source (Aronson and Precht 2000). In 1997, “opportunistic” Porites of propagules to repopulate shallower and more-easily astreoides, P. porites, Agaricia agaricites, and other corals degraded reefs after disturbances (Bongaerts et al. 2010; but averaged ~15 to 16% of the benthos at 5–10 m and ~11% at see Smith et al. 2016). The “lesson” from the west FRS (and, 15 m in 27 sites along 10 km of coastline around DB (Cho more broadly, Jamaica) implies rather that changes occur and Woodley 2000). A phase-shift reversal at Dairy Bull more slowly in deeper habitats, where hurricane wave ener- coincident with the appearance of Diadema was apparent gies are diminished and less light is routinely available for after macroalgae in 6–8 m fell from ~45% in 1995 to ~6% in photosynthesis, than in shallower water. When most of the 2004 and coral cover rose from ~23% to ~54% (Idjadi et al. Jamaican Diadema died in 1983, noncrustose algae expanded 2006). Other surveys at 5–8.5 m on Dairy Bull tracked a more quickly on the shallower fore-reef terrace (FRT) above sharp decline (from ~48% to ~13%), in coral cover immedi- the WRB cliff and at Zin than in deeper habitats (Hughes ately after the major bleaching event of 2005, followed by a et al. 1985, 1987; Liddell and Ohlhorst 1986). Bleaching in subsequent rebound to ~28% in 2012 (Crabbe 2014). 1987 affected more corals on the FRT than the deeper FRS Diadema is naturally limited to < ~40 m depths. As long as (Woodley 1988). Coral cover at 15 m on Zin was ~33% Jamaica’s herbivorous fish populations are overharvested, before Hurricane Allen, but it had declined and partially we are unlikely to learn how quickly, easily, or even if a recovered several times before ending the decade at only phase reversal could restore the former framework-building ~2% (Liddell and Ohlhorst 1992), whereas corals at 30 m role of its FRS corals. 106 P. Dustan and J. C. Lang

6.6 Threats and Conservation (Palmer et al. 2017). Photogrammetry is employed to gener- ate photomosaics and three-dimensional models of moni- Stressors impacting the Jamaican reefs have been obvious tored reef condition at DB (Charpentier et al. 2017; Porter, for decades, especially local overfishing and declining pers. comm.). Nevertheless, tropical islanders do not hold water quality from land-based sources of pollution, regional the solutions to all the problems that are killing reef organ- diseases, and global warming (Fig. 6.7). Impacts from isms because we all share responsibility for curbing excesses Hurricane Allen and the mass mortality of Diadema were of overpopulation, overdevelopment, and overconsumption well described, particularly in shallower reef habitats (e.g., (especially fossil fuels). Hughes 1994; Jackson et al. 2001). The effects of mortality The Natural History Museum in London, under from bleaching and diseases, reduced three-dimensional K. Johnson’s leadership, has responded to the need to pre- reef structure, and disrupted food webs have been relatively serve early ecological baselines for coral reefs and related ignored, particularly on the deep reefs. (But see Jackson marine habitats. An open-access archive of historical imag- and Winston (1982) for the influence of predators of ery (Yarlett et al. 2016), established with negatives donated encrusting organisms under flattened corals at 10–21 m on by E. Graham, is attracting contributions from other research- the WRB cliff and Knowlton et al. (1990) for the influence ers. Similar efforts should be made to conserve all the imag- of predators on hurricane-fragmented Acropora cervicornis ery in this volume describing mesophotic coral ecosystems, at <13 m.) while accurate accompanying metadata are likely to still be Efforts at DB to actively engage with the local fishing available. communities and collaboratively reduce fishing pressures began in the mid-1980s. By 1991, funds had been raised to Acknowledgments We are grateful to all the dedicated staff of the support gear changes and create a DB fishermen’s associa- University of the West Indies, Mona Campus, including boatmen and tion. Agreement was reached in 1994 to establish a tiny, vol- diving assistants, who have kept DBML funded and operational since 1965. We especially appreciate the availability of imagery donated to untary, no-take fisheries reserve on the western side of the the Natural History Museum by E. Graham and to the Nekton project by bay; its boundaries were marked in 1995 and daily patrols P. Goreau, N. James, and J. Woodley. B. Charpentier and L. Wheeler began in 1996. The reserve was extended around the bay in also contributed photographs. We are particularly indebted to 1998 as fishery benefits became evident to fishers, but -com B. Charpentier, P. Colin, P. Gayle, R. Kinzie, W. Precht, G. Warner, and P. Willens for facilitating access to the literature and/or unpublished pliance declined after financial support was lost, and the last data and/or patiently responding to a multitude of queries. J. Bruno, rangers were laid off in 1999 (Woodley and Sary 2000). T.J. Goreau, S. Gun, J. Jackson, K. Johnson, B. Lapointe, L. Land, However, the reserve was further expanded in 2009 when H. Lasker, H. Lenhert, D. Liddell, D. Meyer, J. Ogden, S. Ohlhorst, DB was declared a fish sanctuary by the Ministry of S. Palmer, J. Porter, K. Sebens, L. Wheeler, J. Wulff, and S. Zea helped with answers to other questions. Reviewers are thanked for challenging Agriculture and Fisheries, and the local Alloa fishermen’s comments that improved the focus and accuracy of the manuscript. association was given its day-to-day management responsi- Financial support was provided by the University of the West Indies at bility. In 2012, the sanctuary status was revoked and replaced Mona, the Waitt Foundation, and the Department of Biology, College of with the declaration of the DB Special Fishery Conservation Charleston. This chapter is dedicated to the memory of the late Tom Goreau. We express our sincere gratitude to his late wife Nora and their Area to encourage the reproduction of fish populations to a three sons for accommodating so many “Goreau enthusiasts” in their level that would support a sustainable fishery. A 10-year family life during the early days of deep-reef exploration and discover- management plan, developed through community-based par- ies in Jamaica. This is DBML Publication number 791. ticipation, was finalized in 2015 (Ministry of Agriculture and Fisheries 2015). DBML and the fishermen’s association have Author Contributions since secured funding to collect coastal water quality data to The authors shared the conceptualization of this contribution. PD col- lected the DLR data, collated photographs, and created figures; JCL promote regulatory best practices and provide educational assembled the DB-area literature and drafted the text and tables. outreach for reducing local point sources of pollution. Skills training will be offered to select community members (par- ticularly women and youth) to increase their options for alternative local employment (Gayle, pers. comm.). References Young researchers in Jamaica are trying to reverse the degradation of the reefs off DB and in other managed areas Andres NG, Witman JD (1995) Trends in community structure on a using structural and biological restoration techniques. Jamaican reef. Mar Ecol Prog Ser 118:305–310 Aronson RB, Precht WF (2000) Herbivory and algal dynamics on the Included are developing three-dimensional models of small-­ coral reef at Discovery Bay, Jamaica. 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