Rt'~t'arch [ .Iournal of Coastal ~_ _"_____"__ 67-12 __l Fort Lauderdale, ___"__ Winter 1997 ~

Patterns of Damage to the Branching palmata Following : Damage and Survivorship of Hurricane-generated Asexual Recruits

Diego Lirman and Peggy Fong]

Rosenstiel Schoo! of Marine and Atmospheric Science MBF Division University of Miami 4600 Rickenbacker Causeway Miami, I<'L :3:3149 ABSTRACT _

URMAN, D. and FONG, P., 1997. Patterns of damage to the branching coral Acropora palmata following Hurricane ,tll',IIII:. Andrew: Damage and survivorship of hurricane-generated asexual recruits. Journal oj' Coastal Research, 1311 1,67­ ~ 72. Fort Lauderdale (Florida I. ISSN 0749-020H. Hurricane Andrew caused widespread damage to the Acropora palmata population on a patch on the euus~ ~ d Tract. After the storm, more than 50'/r of the A. palmata cover in the rubble and reef-flat zones was comprised of live -+ &-- fragments. Other species of coral were minimally damaged. Most fragments were distributed within or adjacent to the remaining patches of standing elkhorn colonies. Neither distribution nor mortality rate of fragments was depen­ dent on initial fragment size. However. rate of stabilization of fragments was related to substrate type and distance from a patch of mature colonies, suggesting that standing colonies may protect regenerating fragments from romoval from the reef. Differences in the substrate type Ihard 1's. unconsolidated rubble I where fragments landed, affected romoval , lotal mortality. and part.ial mortality rates of hurricane-generated fragments. Rubble substrate favored sta­ bilization and survival of hurricane-generated asexual recruits.

ADDITIONAL INDEX WORDS: Hurricane damoe« coustttl storm, .

INTRODUCTION physical IPmnEH. et al., 1982; DALLMEYER ('I al .. 1982; Dt .s­ TAN. 198.51, chemical IBAK, 19871 and biological IBAK and Elkhorn coral (Acropora palmata 1 is a branching, tree-like C\{IENS, 1981; GT.ATlFELTEH, 1982: JAAP, 1984 I stresses, Ac­ coral that forms densely aggregated patches on the shallow ropora palnuita persists as a dominant component of shallow, portions of reefs in the Florida Reef Tract and throughout the high energy reef areas. Many scientists believe that local IGlJl{!-;AI;, 1959: GEI:-iTEl{, 1977; GLADFELTER et dominance of this species is achieved by fast growth rates al, 1978; ROm;H:-i ct al.. 1982). Although it is adapted to high 147-H.~) crnyr ,) and high regeneration potential In.>\K, 1976; wave and surge zones of reefs ISCIlI1IlMA< 'IlEI{ and PLEWKA, GLADFELTER et ol.. 1971-\: HICIlSMl'I'll, 1982; HpSTO!', 19851. 1981), it is very susceptible to the intensity of physical forces Additionally, A. palmata has several morphological adapta­ generated by a hurricane I\VO(uu.icv cl al.. 1981\. Significant tions to wave stress. A thick stem composed of skeletal ma­ dislodgment and Iragmontution of this species were reported terial with unusual structural strength anchors the colonies after Hurricanes Edith (GLYNK ('I al.. EJ64 I, Gertn (HICIl­ to the reef ISCHl iHMAl 'IIEf{ and PLEWKA, 191-\11. Each colony :-iMITIl et al.. 191-\01, Allen l\VoOIJLI';Y ('I al.. 19811, David and aligns branches parallel to the prevailing water flow presum­ Frederic IROl;EI{:-i ct al .. 191-\21, Hugo IGLAllFEI/mll, 1991 I, ably to reduce the force exerted on them by waves and cur­ and Andrew IFONl; and LIIII\L\!\, 1994 I. In some cases, up to rents IGnAl's et al .. 1977: S(,II[fIlMN'II~;!{ and Pr,EWK,\, 9.5'1. of the existing A. palmula cover was lost I\V(H)J)I.EY v! 19S1I. 01., 1981: GLADF!-;LTE\{, 19911. This susceptibility has been Unlike acroporid species in the Indo-Pacific Province that related to A. polmato« high surface-urea-to-volume ratio that exhibit high sexual recruitment rat(>s IWALL\"E, 19851, the results from the upright branching form. as massive and en­ main propagation mode of A. polmuta in the Caribbean is by crusting with a low profile seem to be more resistant means of colony fragmentation ISAK and E;-':<;EL, i979: to damage IW, HI!ILI';Y ct al., UJH1: ROI;I-:II:-i ('I al.. 1~)82, HJ8:3: HI(;IISl\1IT1l, 1982; RYLAAI{:-iIlAl\l, 191-\:3: ,JoIUJAl'i-DA!IL!;ln;;-.:, ST(IIJIlAIH, 198.5; EIJ:'>1l1ND:-i and WITMAN, 1991; BYTIlELLi'I 19~j21, enahling this species to re-cover and re-populate areas al.. 199:31. disturbed by intense physical disturbances IHI(:I 1:-i;VIITII et al., Although highly susceptible to storms and many other 1980: R()(;E!{:-i d al. I~J82: LIH:YIAN and FON(;, llllPllhl/s/wd data I. Some resoarchc-rs believe the ability to produce asexual 942·11 /'(-('(-",,<1 18 -lu nuurv I,'},'}:i; acc/'pt"d !II /'(/S/II/l 28 Murch I.'I.'IS recruits readily from wave generated fragments is also an , Current address: P,-ggy Fong, P"cilic ~:stuar tH- Resl'arl'h Lahora­ tory, Dopurtmvnt of Biologv, San Dipgo State l niversitv. San Diogo, adaptation to wave stress IHH:II:-iMITII, 19821. Fast growth CA 921K2, U.S.A. rates, the ability to regenerate lesions IRo(;,;J(:-i />1 al.. 1982: 68 Lirma n an d Fong

BAK, 1983; FONG and LrRMAN , 1994), and the ability of bro­ ken branches to cement to the substra tum (TUNN ICLIFFE, ". 1983 1, are all characteristics that en hance the probabi lity of

fragment survival following mechanical disturbances . &r'05", Th e recovery of mecha nically dam aged reefs is highly de­ t pendent on the continued growth of those colonies that re­ -, N mai ned upright after a disturban ce, coloniz ation by new ly I produced coral planulae , and the stabi liza tion and growth of 1 FOWEY J coral fragm ents (ENDEAN , 1976; PEARSON, 1981). Due to th e nocss J BISCAYNE I low re cruitm ent success of sexually produced planulae in this NATlONAL PARK , species (DUSTAN, 1977; RYLAARSDAM , 1983; ROSESMYTH, ,I 1984), the survivorship of dam aged ad ults and asexua l re­ I cruitment by coral fragment s may play significant roles in ".,.,-+--!;1 I f I' I ~""" th e recovery process.In this pa per, we accomplish three ob­ • ,/mIUMf'HREEF BISCAYNE BA Y jectives . First, we docum ent spatial pattern s of hurrica ne­ ' ./lLONG REEF generated damage to A. palmata by recording the dist ribu tion 195k'>llh of damaged adu lt colonies an d fra gments. Second , we mea­ 25"5 l ~iA I sure rates of early su rvivorship of new re cruits derived from ' / AJAX REEF I I fragments. Third , several hypotheses about factors affecting · / PACIF'C AEEF I survivorship of Acropora palmala fragmen ts are tested, in­ AUN A'SRHF I cludin g the effect of dista nce from an existing A. palmata * I 1.Ur I ',*~~:N I patch, initial size of each fragment, and substrate character­ I 21n~ istics of the area where a fragment landed. I BALL BUOY I J REEF ~ I , 1 2'0,,",," METHODS 'I 25~ 1 5 T'1: F ~ Study Site ..-" o 1 2 3 Elkhorn Reef (25°21.766 'N , 800 0.9.961'W) is an isolated reef in th e northern region of the Florida Reef Tract (Figure 1). It is ap proximate ly 4.5 hectare s and su rrounded by sand Figure 1. Map of South Florid a including the path of Hurricane Andrew with demarcations indi cating bands of approximate maximum susta ined plai ns and seagrass beds. Altho ugh it is a mid-s helf formation winds, the boundaries of Biscayne Nat iona l Park, and several of the larg­ located 2.5 km from the outer reef, it is afford ed little pro­ er reefs of northern Florida Reef Tr act. The star sh ows the location of the tection from wave action beca use to the E and S-E, the di­ study site . El khorn Reef. rection of the prevailing wind, th e fore-reef is discontinuous (SHINN et al., 1989). The long axis of th e reef is oriented N and S an d there is no typical reef crest formation; ra ther surveyed a belt transect that crossed all three ree f zones . Pla ­ there is a relatively wide and shallow ree f flat. A Pri ncipal nar percentages of cover by each species were estimated in Component Ana lysis perfor med on th e a bundance of all coral situ with SCUBA in March 1993 by placing a 1 m" quad ra t species on Elkhorn Reef (LIRMAN an d FONG, 1995) revealed a long a tran sect bisecting the reef from W to E. Along with that the cora l communi ty can be statistically divided into cover, we estimated the percentage of colonies of each species th ree major zones: rubble zone (mean depth = 1.7 m), ree f­ tha t had sustained any physical da mage during the hu rr i­ flat (1.2 m), and fore-reef (3.1 m), cane. Acropora palmata was divided into standing colonies and live fragmen ts th at were brok en from colonie s and de­ Stor m Description posited on th e benthos . We defined as sta nding A. palmata those colonies that remained up righ t an d had a visible at­ Hurricane Andrew was a compact and quickly moving tac hmen t to the bottom through the characteristic stalk storm that crossed South Florida on August 24, 1992 (Figure (SCHUHMAC HER an d PLEWKA, 1981). We defined as frag­ 1). Maximum sustained winds of 225 kmhr -, wit h gusts of ments th ose broken branches of A. palmata covere d by live up to 285 km-hr ' ' were measured 15 km N of our study site tissue lying on the substratum and lacking an obvious stalk. (POWE LL and HOUSTON, 1993; RAPPAPORT and SHEETS, Fragments become new asexual recruits when t hey attach to 1993). This storm ranked as a Ca tegory 4 sto rm on the Sa ffir­ the botto m. It was not possibl e to estimate the da mage that Simpson hurricane damage-potential scale (AHRE NS, 1993), might have occurred to Millepora spp ., beca use it was not making Hurricane Andrew th e strongest storm to hit South always ap paren t when fine bra nches may have been re­ Florid a in at least 50 yr . moved. Altho ugh we found ske leta l remains of fine branches in th e rub ble field, we saw few obvious sca rs whe re rem oval Field Methods occurred. It is possib le th at these small wound s were already Visual sur veys of th e reef comm unity on Elkhorn Reefwere healed by our sampling time as Mill epora spp . is often very perform ed 3 wk after Hurricane Andrew. To quantify the rel­ fas t-growing. ati ve dam age to different cora l species among reef zones, we To determine the large-scale distribution patterns of both

Journal of Coastal Resea rch, Vol. 13. No. 1, 1997 Hurri ca ne Dam age an d Earl y Survivorship 69 sta nding colonies and fragm ents of A. palmata, we surveye d Table 1. Percent coral cover ( ::!:S .E.! and percentage ofcoral colonies th at three line-intercept transects along th e E-W axis of th e reef was damaged within each major zone on Elkhorn Reef Ma ssive corals include species in the genera Montastraea , Siderastrea , and Diploria . nd in April and May 1993. These transects were 10 m apa rt, = no da ta auailab le. We were un able to quantify damage to colonies of running par all el to each other. The cover of fragm en ts a nd Millepora spp. th at may have had finely bra nch ing sections removed. standing colonies was estimated at every meter-interval by calculating th e fraction of th e length of the line th ese inter ­ Fore-reef Reef-flat Rubble zone cepted (LoYA, 1978). ( n = 39) ( n = 29) (n = 28 1 The distribution and abundance of fragm ents in re lation to Acropora palm ata a patch of damaged Acropora palmata colonies that may have Sta nding colonies 0.0 (00 18.2 (4 0) 54 ( 1.7 ) been th e fragmen t source was measured. Five parall el belt Percent dam aged > 90 > 90 Live fragment s 00 (0.0) 17.9 (2.8) 11.1 (3.2) transects 5 m apart were surveyed. These tran sects bisected Porites astre oides 4.8 (1.0) 11.3 n .7) 2.6 (1.2 ) th e largest fragme nt field on the reef, starting at the edge of Percent dam aged 0 < 5 0 a large standi ng A. palmata patch. Along each tra nse ct, a 1 Porite s porites 20.5 13.31 2.1 (1.3) 0.3 (0.2) rn- qu adrat was flipped from E to W until no more live frag­ Percen t damaged 0 0 0 5 9 (1.3) (2 2) 1.9(0.7) ments were found. The number of qu adrats along each tran­ Millepora spp. 5 5 Percen t da maged nd nd nd sect ra nged from 8 to 12. Within each qu adrat, we coun ted Ag aricia spp. 01 (0.02) 0 05 ro. oi: 0.1 10.03) the nu mber of fra gments, mea sured the size of each fragme nt Percent damaged 0 0 0 as longest length and widest width , and noted whether th e Mas sive cora ls 8.9 ( 1.5) 1.2 (0.5) 2.3 (09) fragment was loose or cemented to the substrate. This survey Percent damaged 0 0 < 5 was done in May 1993, 9 mo after th e storm. Two hy potheses about th e spa tia l patte rns of fragment abundance were test­ ed. The first hypoth esis was tha t lar ger fragm ents tr aveled missin g branches easily dete cted by the pre senc e of a circu lar shorte r distances. To tes t th is, we compared average size of to oval area of exposed white skeleton. Standing A. palmata fragments in the closest (1-3 m) an d furthes t (10-12 m) qu ad­ cover ed over 18% of the reef-flat, and an equa l area was cov­ rats from the edge of th e A. palmata pa tch . The second hy­ ered by A. palmata fragmen ts . The oth er domin ant species of pothesi s was th at more unattached fra gments were closer to cora l in this zone, Porites astreoides, was almost undam aged, the source patch. Thi s was based on the assumption that only a few colonies showing signs of surface scour ing. None proximity to an A. palma ta patch redu ced wave action and of th e other species or species groups were damaged in this removal of fragments. To test this , we compared the mean zone. Total cover of hard corals was low in the rub ble zone, distance from the cente r of a patch for cemented and unce­ an area dominated by Acropora pal ma ta forming discrete and mented fragments. den sely aggrega te d clumps. Here, cover of sta nding A. pa l­ In March 1993 (7 mo post-storm l, app roximatel y 80 Aero­ mata was lower th an cover of fragme nt s (Table 1). In thi s pora palmata fragm ents were marked in two different areas zone where massive corals are rare, a few small colonies « 30 of th e reef where fragments had accumulated after th e hur­ cm in diamete r) had been dislodged and overturned during ricane. The subst ratum in one of th ese areas, located W of the storm. No oth er types of coral were damaged in thi s zone. the lar gest patch of standing A. palmata, was covered by piec­ On th e deeper fore-reef, an area dominated by th e bra nching es of dead cora l rubble. The second set of fragments was lo­ cora l Porites porites, th ere was no app arent hurricane dam­ cated E of th e large A. palmata patch on a ha rd, consolidated age observed. substratum. Each fragm ent was individually labeled with a Alth ough the distribution of sta nding colonies and frag­ metal tag and photographed with a fixed fram er quadropod ments of Acropora palmata was patchy, th ere were three gen­ in June 1993. The fragm ent fields were surveyed again in eral patterns obse rved (Figu re 2). First, there were fewer May 1994, and th ose fragm ents bearing tags wer e re-photo­ fragmen ts and more sta nding colonies of A. pal mat a in the graphed using similar methods. The resulting photo graphs northe rn section of th e reef than in th e center or th e S, sug­ were digiti zed, and th e ar ea covered by live coral tissue was ges ting that the force of the hurricane may have been dim in­ estima ted for each fragment. The hypothesis that initial size ished in th e northern part of the reef. Second , patches of of fragments affected th e survivorship of cora l fragments was st anding colonies seemed to fall into two categories. Eith er tested with linear regression. The hypothesis th at substra te colonies within a patch were lightly dam aged and sur roun ded type affected fragm ent removal, rates of partial mortality, by few fragments (as in int erval 35-50 and 85- 100 in Figure and fragm ent stabilization (cementation to substrate) was 2a and 32-50 in Figure 2c), or there were only a few standing tested with t-tests. colonies remaining and most were reduced to fragmen ts (as in interval 15-40 a nd 85- 100 in Figure 2b). Last , man y of RESULTS the A. palmata fragm en ts th at survived through th e 7 mo afte r the hurrican e were distributed withi n or adjacent to the The percen ta ge of th e benthos of Elkhorn Reef covered by remaining patches of standing colonies. There are several corals was very pat chy, ranging from 0-100%. Mean coral possible explan ations for th is distribution, including differ­ cover for the whole reef was 42.3% (S.E. = 2.8). Cover was entia l tra ns port and depositi on during th e storm, differential highest on the reef-flat, wher e approximately 90% of th e Ac­ remo val after the storm, and differential surv ival after the ropora palmata that rema ined standing after the storm was storm . dam aged to some extent (Table 1). Most colonies had many To determine whe ther larger fragments were tr an sported

J ourn al of Coastal Research, Vol. 13, No.1, 1997 70 Lirman and Fong

I OO~ 1 I' 4OT, ------, 9 0 ~ Fragments flO- 70 o Standing Colonies " eo '0 ­ 30 40 ­

30 ­ Vl 25 20 § 10 · "ge 20 u, .0 o 15 ~ . 0 70 oo 10 SO 4 0 30 · 20 · ' ~ ~ S ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ $ ~ ~ ~ £ ~ g ~ 8 8 ~ ~ ~ ~ 9 ~ g ~ a ~ ~ ~ ~ ~ ~ ~ g ~ ~ ~ ~ ~ § ~ ~ 90 00 LENGTH INTERVAL (em) 70· 00 Figure 3. Size frequency distribution or fragments or Acropora palmata so in the largest fragment field. Size or each fragment was measured as the 40 longest length of living tissue JO 2iJ 10

ment size. Although there were few fragments with a longest

INTERVAL NUMBER (m) length less than 5 ern, a large proportion of fragments were within the next three smallest size classes (5-20 ern). This Figure 2. Distribution of standing Acropora palmata colonies and Irag­ ments along three parallel E-W transects bisecting Elkhorn Reef. Tran­ indicates that mortality may not be directly dependent on sects were surveyed with the line-intercept method, recording the length size above a 5 em threshold. In addition, there was no sig­ or each 1 m in terval that was covered by either standing or fragmented nificant relationship between initial fragment size and sur­ Apalmata. vivorship of the 80 marked fragments (r 2 = 0.048, P > 0.05). Differences in substrate type affected removal, total mor­ tality, and partial mortality rates of hurricane-generated shorter distances, we compared the average size of fragments fragments. In the rubble zone, only 34% of the fragments that close to and distant from the edge of a patch of standing col­ were marked were removed from the reef during the 11 mo onies. There was no difference in mean size of fragments in interval between sampling dates, while 57% of the fragments the closest (1-3 m) and the farthest (10-12 m ) intervals (t­ were removed from the hard-bottom area. Thus cementation, te st, p > 005), Thus, our data indicate that the distribution an important process in the transformation of a fragment to of fragments is not directly dependent on size One possible a recruit, may be slower or less efficient in the hard bottom explanation of this distribution is that fragmentation may area. Total mortality, or complete loss of live tissue was also have occurred in several stages, first from the hurricane and higher in the hard bottom (50%) than the rubble (4%) sub­ then through several winter storms. Alternatively, clusters of strate. In addition, many of the marked fragments (73%) lost large fragments away from standing colonies may represent tissue over the 11 mo of the study. Average partial mortality the shattered remnants of isolated colonies. rates of surviving fragments was significantly higher for frag­ Proximity to a patch of A. palmata may limit removal of ments in the hard bottom area (t-test, p < 0.05). Fragments fragments by reducing wave and current action. Ifloose frag­ in the hard bottom area lost 61'/0 of their live tissue area (8.E. ments are removed more rapidly away from a patch than next = 15.5, n = 23), whereas fragments in the rubble areas lost to it, then there should be fewer un cemented fragments close 31% of their live tissue area (8.E. = 6.9, n = 10). to a patch of standing colonies. To test whether there was Only a few fragments increased the area covered by tissue differential removal after the storm, mean distance from the (6 in the rubble zone and only 2 in the hard bottom area) over patch to uncemented and cemented fragments was compared. the 11 mo of the study; for these fragments mean increase in Mean distance to uncemented fragments was significantly tissue area was 15.5% (8.E. = 2.2). During the summer of less th an for cemented fragments (t-test, p < 0.05), suggest­ 1993, the marked fragments started developing vertical ing tha t standing colonies may protect uncemented frag­ growth features on their upper surfaces. The rapid upward men ts from removal from the reef. growth of these features (3.65 ± 0.04 cm/yr, FONG and LIR­ In May 1993, 218 fragments in 48-1 m' quadrats within MAN, unpublished data) sharply contrasts with the planar tis­ the hurricane generated field of fragments were measured sue loss shown by the fragments. This seems to indicate that (Figure 3), for a mean of 4.56 fragrnentsrn:" (8.E. = 0.54). after the fragments have re-cernented to the substratum, There are two lines of evidence that suggest that survivorship most of the energy allocated for growth is dedicated to the of fr agments may not have been dependent on initial frag- upward extension of these vertical growth features.

Joumal or Coast,,1 Research, Vol. 13. No.1, 1997 Hurricarw lIamag« and Early Survivorship 71

DISCUSSION ricane-generated fragments can be influenced by substrate type and distance from standing coral colonies. Hurricane damage to Elkhorn Reef was extremely patchy. However, there were three general patterns observed. First, CONCLUSIONS almost all the coral damage that occurred was to colonies of Tropical storms are undoubtedly major shaping forces act­ Acropora palmata. Damage to this species was extensive; we ing on coral community structure. The damage caused by found very few undamaged colonies of A. palmata in all of Hurricane Andrew, although significant, was not as devas­ our surveys. Destruction of A. palmata colonies is commonly tating as may have been predicted given the magnitude of reported after hurricanes (WOOIJLEY et al., 1981; and many the storm. The Acropora palmata population on Elkhorn Reef others), and may be attributed to colony shape (GJ{AUS et al .. exhibited remarkable recovery capabilities even though most 1977) and the location of most colonies in high energy reef of the colonies experienced some degree of fragmentation due zones (GoREAU, 1959; GEISTER, 1977; ROBEKrs et al., 1977). to waves and coral projectiles. This coral species, with limited Second, damage was variable among patches of A. palnuita. sexual recruitment capabilities, was able to increase its dis­ We attribute this difference to a downcurrent "domino effect", tribution due to the asexual recruitment of hurricane-gener­ where if one colony is shattered, its fragments can cause a ated fragments. The survival of these fragments was shown cascade of damage to nearby colonies before coming to rest to be size-independent, but affected by substrate type and on the benthos. Thus, the probability of sustaining damage distance from standing colonies. On Iy the continued study of increases with the amount of damage to neighbours. Third, the survivorship and growth of these recruits as well as the many surviving fragments were retained within or adjacent healing of standing colonies will enable us to fully evaluate to standing A. palnuita colonies. Fragment retention within the long-term effects of Hurricane Andrew on this coral com­ patches was also observed after Hurricane Gerta in munity. (HIGHSMITH et al., 1980). One explanation may be that con­ tact with standing colonies of the same species enhances the ACK.~OWLEDGEMENTS re-cementation process and therefore decreases the probabil­ ity of a fragment being removed from the reef by subsequent This research was funded, in part, by National Science wave action. BAK and CJ{IENS (1981) found that A. palmato Foundation Grant # OCE-9-00649 to the University of Miami. fragments fuse to other elkhorn fragments faster than to any Additional support was provided by the Elizabeth Ordway other type of substrate. Additionally, proximity to standing Dunn Foundation and the Key Biscayne Rotary Club. Bis­ A. palmata colonies may produce a baffling effect on waves cayne National Park provided boat and dive support. We and currents that prevents or delays the removal of unce­ thank Richard Curry, Science Director for BNP, as well as men ted fragments from the reef. our volunteer divers, Juan Mate. Guillermo Diaz, Maia The most important factor affecting the ability of frag­ McGuire. and many others. Our appreciation goes to Mark ments to become new recruits was the substrate type where Harwell, Peter Glynn, Susan Akana, Maia McGuire and two the fragment landed after the storm. To become a recruit, a anonymous reviewers for their critical evaluation of the manuscript and to Laura Kaplan for help with the figures. fragment must preserve live tissue area t survive ), stabilize itself on the reef by cementation, and begin to grow upward so that it can compete for light. Fragments on rubble sub­ LITERATURE CITED strate suffered less partial or total mortality, and more frag­ AHltEN:-i, C.O.. 199:3. Essentials o] Mrtcrolog». Minneapolis: W. Pub­ ments cemented to the bottom than those on hard, consoli­ lishing Co. dated substrate. It will be important to determine if these BAK, RP.M., 1970. The growth of coral colonies and the importance of crustoso > and burrowing sponges in relation with differences are complemented by differences in rates of up­ carbonate accumulation. Netherlands Journal 0( Sea Research. 10. ward growth through time. The first signs of upward growth 2S5-:3:37 on hurricane-generated Irugment.s were observed in the sum­ BAK, RP.M.. 19S:3. Ne-oplasia. regenenltion and growth in the reef­ mer of 1993. building coral Acroporu palniat.a, Marine lIiologv. 77.221-227. BAK. RP.M., 19S7. Effects of chronic oil pollution on a Caribbean Due to the lack of success of sexual recruitment in Acropora coral reef. Marine Pollution Bulletin, lS, 5:34-5:3~l. palmata. the recovery of palmata-dominated reefs after me­ IlAK. RP.M. and CHIENS, S.R.. 191'1. Survival after fr.urmcntut ion chanical disturbances (e.,/{., hurricanes, groundings) will be of colonies of Madroci« mirubili«. Acropora polmota. and A. err highly dependent on the survivorship and growth of asexual ricorni» I Sch-rr-ctinia I and the subsequent impact of a coral dis­ ease. Procvcdinu» Fourth International Coral Reel.'ivmposiulII, 2, recruits. The effects of initial size on the survivorship of hur­ 221-22S. ricane-genornted Acropora palmata fragments has been stud­ BAK, RP.M. and ENeEI., M.S., 1979. Distribution, nhundance and ied previously with conflicting results. HIC;IISMITIl ct of. survival of juvenile hermatypic corals .Sch-rectinin : and the' im­ 119801 found a significant positive correlation between frag­ portance of life history stratehrjps in tho parent coral comrnunitv. Marine RlOlogv. ,,4,341-:3.';2. ment size and survivorship after Hurricane Gerta. In con­ B,THEI.I., J.C.. ; GI.AIlFI·:I.TI·;H E.H.. and BVTHEI.I., M., 199:3. Chronic trast, RO(;EI{s e! al. (19821 found that fragments that died and catastrophic natural mortality of throe common Caribbean were larger than frugments that survived after Hurricanes corul«. Coral Reels, 12, 14:'3-152. S~IITH. David and Frederic. In our study, we did not find a significant D,\I.I.MI':Yi':H, O.G.; Pown:H. J.W.. and G.•1.. 19S2. EIl',·c!s of peat on the behavior and physiology ofthe .Iamaican reef building relationship between fragment size and survivorship. Nev­ coral Montastrra annulnris. Marine Biologv, oS, 229-2:3:3. ertheless, we determined that the early survivorship of hur- Ol:STAN, P., 1977. Vitality of reef coral populations off Key Largo.

.Journal of Coastal R,'search. Vol. I:i, No. I. Im17 7~ Lirrnun and Fong

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.Iournal of Coastal [{psearch, Vol. 1:3, No. I, 1997