Pacific Science (1989), vol. 43, no. 2 © 1989 by University of Hawaii Press. All rights reserved
Coral Reef Recovery Subsequent to the Freshwater Kill of 1965 in Kaneohe Bay, Oahu, Hawaii!
PAUL F. HOLTHUS,2 JAMES E. MARAGOS,3 AND CHRISTOPHER W. EVANS4
ABSTRACT: The reef coral community on the landward side of a patch reef near Kahaluu in Kaneohe Bay, Oahu, Hawaii was resurveyed 18 yr after all live coral was killed by a thick lens of freshwater runoff from a flashflood in 1965. The initial phase of recovery of the reef was documented from 1968 to 1973. A resurvey of the reef was conducted in 1983, using the same methods as the 1973 study. Species, abundance, and distribution of corals on the patch reef were measured and recorded along a series of 10 transects. Results show large in creases in size and numbers ofcolonies, area, and depth range covered by corals. Greatest coral abundance was reported in the upper 5 m, but community diversity did not increase because the fast-growing finger coral, Porites compressa, became more dominant. The pattern of coral community succession at this sheltered location was similar to that observed at other environments in the Hawaiian Archipelago. Recovery appears to be rapid in protected, low-wave energy environments such as Kaneohe Bay, which are infrequently affected by -major-disfurhances.Almost 20yi afteia -majordishirhance ~theKahariiupatcn reef slope coral community is approaching the climax conditions of other reef slope communities in Kaneohe Bay not disturbed by the 1965 flashflood.
CORAL REEF communities are subject to a tored several permanent meter quadrats on variety of natural or man-made disturbances reef flats since 1962. He has reported on their (Stoddart 1969a, Johannes 1975). Relatively recovery from cyclone damage, but the dy few instances ofcoral reefrecovery have been namics of reef flat coral communities may documented (Endean 1976), especially in the differ from those ofreefslopes, where optimum quantitative assessment of coral community coral development often occurs (Sheppard dynamics during recovery. Stoddart (1963, 1982). The recovery of coral communities on 1969b, 1974) observed the damage and re Guam following Acanthaster infestations was covery of British Honduras reefs following documented by Randall (1973a,b,c) . Acan cyclone Hattie in 1961. On the Great Barrier thaster destruction of coral reefs and subse Reef, Connell (1973, 1976, 1978) has moni- quent recovery were broadly investigated by Endean and Stablum (1973a,b) and in detail by Pearson (1981). The recovery ofcoral reef assemblages disrupted by natural disturbances 1 This paper is a product of the 1983 Hawaii Institute of Marine Biology Summer Program in Marine Science are considered by Loya (1976) to be mainly a - sponsored by the-Edwin-W.-Pauley-Foundation,-the - . function.of.time. Pearson.(1981) defined reef University of Hawaii Foundation, and Sea Grant recovery as the restoration of a coral assem Program Project No. F 238-F-582-B-315; under grant blage to a degree comparable to its original number NA8IAA-D-00070. Manuscript accepted May state. In reef slope habitats with high coral 1988. 2South Pacific Regional Environment Programme, cover, the recovery process may require several South Pacific Commission, B.P. D 5, Noumea, New decades following major natural disturbances Caledonia. (Pearson 1981, Sheppard 1982). 3 Environmental Resources Section, U.S. Army Engi- In Hawaii, the reestablishment ofcoral reef neer Division, Pacific Ocean, Bldg T-I, Fort Shafter, Hawaii96858-5440. communities was studied on submerged lava 4Department of Geography, University of Hawaii at flows of known age by Grigg and Maragos Manoa, Honolulu, Hawaii 96822. (1974), who proposed a pattern of long-term 122 Coral ReefRecovery after Freshwater Kill-HoLTHUS, MARAGOS, AND EVANS 123 succession of coral assemblages in Hawaii. spaced along the southern and western Coral colonization and succession in a newly (stream-facing) slopes of the reef, closely created harbor habitat were monitored at sev approximating the sites used in earlier studies eral times over an ll-yr period at Honokohau, (Figure 1). The stations were positioned per Hawaii (Maragos 1983). In Kaneohe Bay, pendicular to the reef slope at progressively Oahu, reef slope communities throughout the greater distances from the coral areas less bay were resurveyed in 1983to document their damaged by the 1965flood. Stations 1 and 10 recovery 5 to 6 yr after sewage input was were positioned on undamaged corals and diverted to outside the bay (Maragos et al. were within 5 m of the damaged/undamaged 1985, Evans et al. 1986). reef boundary while stations 5 and 6 were over In the present study, a patch reef at 100 m from the boundary. At each station, Kahaluu, in Kaneohe Bay, was resurveyed to data on coral colony size,speciescomposition, document coral community composition 18 and number ofspecies were obtained using the yr after a freshwater "kill" (see Banner 1968). methods of Maragos (1974). A frame (1 m") From 2 to 8 May 1965, Kaneohe Bay water was positioned on the reef flat at the upper sheds experienced extremely heavy rains and slope edge. Water depth varied from 0.5 to flood conditions. Up to 17.16 in. (43.59em) of 1.0 m. The maximum diameter and species of rain fell in a single day and many streams each coral colony lying at least 50% within the equaled or exceeded previous maximum quadrat were recorded. The frame was then heights and discharge rates (Banner 1968). moved down the slope in a series of consecu 2 The runoff resulted in a freshwater layer, tive units 1m . The slope angle varied from 30 which formed over more dense marine waters to 90°_from the horizontal but was pre and coincided with verylow tide-s, subjecting dominantly within the 45 to 60° range . In nearshore coral communities to considerable formation was gathered in each successive salinity stress. General surveys showed coral quadrat until no corals were encountered or kills to depths of 5 ft on the inshore reefs in the observed beyond the last quadrat along the central sector of the bay (Banner 1968). transect line. Thus, a transect 1 m wide begin The reef slope on the landward-facing por ning at the top of the patch reef slope and tions ofa patch reef near Kahaluu Stream was extending a distance of 6 to 10m down slope completely denuded of living coral (Banner was surveyed at each station. Water depths at 1968).It is thought that floodwaters entering the base ofthe patch reef varied from 4 to 6 m. the embayment fronting Kahaluu Stream, In quadrats of high coral cover, colonies which is constricted by fringing reef on either may make contact and sometimes fuse to side, were blocked by the patch reef (Figure 1). gether, forming aggregate coral heads of ir This trapped the fresh water, extending the regular shape . Discernable discontinuities in zone of damage further down the reef slope on color and morphological features enabled the stream-facing side, resulting in more coral individual colonies to be distinguished and destruction here than elsewhere in Kaneohe counted separately. A few larger colonies had Bay. The coral communities on the opposite dead central and upper portions. These were side of the patch reef were seriously damaged measured as whole live colonies unless highly only in the upper 2 m (Maragos 1972). The fragmented, in which case the distinct livepor- initial phase of recovery ofthis reef slope was . tions.were measured separately. monitored from 1968to 1973(Maragos 1974). The 1983resurvey and the earlier survey by In July 1983, the Kahaluu reef slope was re Maragos (1974) provided comparative data surveyed at 10 transect sites replicating the on the size, abundance, and distribution of earlier investigation. corals on the patch reef 8 and 18 yr after the freshwater kill. Species composition was examined in terms of total numbers of colo nies and total areal coverage of each colony of MATERIALS AND METHODS each species. Colony diameters were con The coral communities of Kahaluu patch verted to areal coverage using the formula: reef were resurveyed at 10 stations evenly area = 3.14 ,2, where r = 1/2 the measured 124 PACIFIC SCIENCE, Volume 43, April 1989
oo Patc h reefs o & o o o o o o 2--'·l°':::':uo,...... 3---;" • 0-•0 0_ <1/,t, ,/10 6 7 9 Kaneohe Boy 0 Lagoon
o
Kahaluu
- Reef edge Shorel in e I I Highway o 300 600 Boundary of serious 1965 SCALE : m flood damage
FIGURE 1. Location of the patch reef near Kahaluu Stream with position of 10 tran sect stations surveyed in 1973 (forty-six 1_m2 quadrats) and 1983 (eighty-one 1-m2 quadrats) (after Ma ragos 1974)0 Coral Reef Recovery after Freshwater Kill-HoLTHUS, MARAGOS, AND EVANS 125 diameter. Because coral colonies are multi mean coverage per I_m2 quadrat nearly quad dimensional forms that often overlap, areal rupled from 11.28% in 1973 to 43.59% in coverage was calculated in absolute amounts 1983 (Table 1). During that same period, the of coral cover. From this information, the mean number of P. compressa colonies per mean percentage cover of each species per quadrat declined slightly (7.56 in 1973 versus I-m 2 quadrat was calculated. The mean num 6.44 in 1983) while mean colony size almost ber ofcoral colonies ofeach species per quad doubled (Table 1). Data on size frequency rat and the mean colony size of each species distribution illustrate this shift toward greater were also determined. numbers of larger colonies for P. compressa, Coral distribution was analyzed both as a as well as for M. verrucosa and Pocillopora function of distance from the surviving coral damicornis (Figure 2). In 1973, no colonies community and depth. Horizontal zonation greater than 30 em in diameter were reported. was examined in numbers of colonies and By 1983, over 25% of all measured colonies total coral areal coverage at each transect sta were larger than that size. tion . Vertical zonation was plotted both as the There was no distinct pattern of coral re number of colonies and total coral area colonization in relation to distance from the coverage per quadrat along each transect less-damaged reef community on the opposite down the reef slope. Coral community diver side of the patch reef (Table 2). Vertically on sity was measured for the 1973 and 1983 the patch reef slope, high coral coverage and Kahaluu reef coral assemblages and com colony numbers occurred in the upper five pared using the Shannon-Wiener Index l-m quadrats in 1973. By 1983, the zone of (Pielou 1966). high coral coverage had extended downslope somewhat, although the total numbers of colonies had not increased much (Figures 3, 4). The total depth range of corals along the RESULTS reef slope nearly doubled by 1983. On the Kahaluu patch reef between 1973 Porites compressa thus showed a substan and 1983, coral abundance increased in terms tial increase in substrate coverage, mean per ofcolony numbers, areal coverage, and colo centage cover per quadrat, number of colo ny size (Table 1). Overall coral coverage in nies, and mean colony size between 1973 and creased by 660% while the total numbers of 1983 (Table 1). At the same time, the mean colonies increased by about 150%. However, number ofcoral colonies per quadrat, includ large variations occurred between quadrats in ing P. compressa, decreased, indicating a shift the amount of coral coverage, number ofcolo in coral community composition. The greater nies, and size ofcolonies, as evidenced by the dominance of P. compressa is manifested in a standard deviations (Table 1). The same four slight decline in community diversity between coral species reported in 1973 [Porites com 1973and 1983, from 0.50 to 0.45, respectively, pressa Dana, M ontipora verrucosa (Lamarck), on the Shannon-Wiener Index of community Pocillopora damicornis (Linnaeus), and Cyph diversity. Although the decrease is not strik astrea oce/lina (Dana)] contributed nearly all ing, the results show that diversity has not of the coverage and colonies encountered in increased in the 10 yr between surveys. 1983.-Two additional species (Porites lichen Dana and Montipora patula Verrill) were re ported in 1983, but were minor constituents. DISCUSSION Porites compressa and M ontipora verrucosa together composed 96.5% of the total coral Kahaluu patch reefis similar in structure to cover in 1973and 97.5% in 1983. P. compressa other patch reefs in Kaneohe Bay (Maragos alone accounted for 86.3% and 87.9% of 1972, Evans et al. 1986, Holthus 1986). The coral coverage in 1973and 1983, respectively. seaward side of Kahaluu patch reef, which In contrast to the relatively constant propor was not extensively affected by the freshwater tion of areal coverage by P. compressa, its runoff event of 1965, is similar in its species TABLE I
S UMMARY OF THE TRANSECT DATA* FOR THE 1973 AND 1983 SURVEYS
CORAL NUMBER OF COLONY COVER COLONIES StZE
' TOTAL : CORAL MEAN% TOTAL MEAN MEAN CORAL I COVER % OF COVER/ NO. % OF NO./ DIAMETER SPECIES : (m") TOTAL QUADRAT S.D. COLONIES TOTAL QUADRAT S.D . (em) S.D.
1983 Porites compressa 3,580.66 87.9 43.59 37.28 522 75.0 6.44 5.09 25.23 15.14 Mon tipora verrucosa I 39J.l6 9.6 4.84 13.19 59 8.5 0.73 1.35 20.38 17.74 Pocil/opora damicornis 39.33 1.0 0.49 1.77 67 9.6 0.86 1.79 6.90 8.09 Cyphastrea ocellina 3.93 0.1 0.05 0.15 43 6.2 0.53 1.56 8.05 8.17 57.95 4.71 0.7 0.06 0.29 31.50 33.73 M ontipora patula + 1.4 0.72 5 '"ti Porites lichen I > T o tal ~, 07 3 . 0 3 100.0 50.31 696 100.0 8.62 (J "I1 1973 - (J Porites compressa 528.82 86.3 11.28 12.70 348 73.0 7.56 5.27 12.66 15.48 - CIl Mon tipora verrucosa 62.59 10.2 1.36 2.78 53 IJ.l 1.15 2.07 10.94 6.18 (J Pocillopora damicornis 19.63 3.2 0.43 0.83 70 14.7 1.52 1.47 11.61 11.32 Cyphastrea oce/lina 1.52 0.3 0.03 0.15 6 J.2 0.13 0.40 5.17 2.56 -~ (J T otal 612.56 100.0 13.10 477 100.0 10.36 ..tIl ' Tota l number of tra ns eets surveyed : 1973 = 46,1983 = 8 1. <: a-0 S (l> ~ UJ
"0> 2: \0 00 \0 Coral Reef Recovery after Freshwater Kill-HoLTHUS, MARAGOS, AND EVA NS 127
300
fIJ 250 ...... Q) ~ 1983 o ...- 10 20 30 -- 40 50 - 60 ·· 70 80 90 100 Coral Colony Size (ern) 300 fIJ 250 ...... Q) ~ 1973 o ...- a a 10 20 30 40 50 60 70 80 90 100 Coral Colony Size (ern) F IGURE 2 . Size frequency distributi on for comb ined transect data, showing number of coral colonies (n) for each size class in 1973 (n = 477) and 1983 (n = 696). 128 PACIFIC SCIENCE, Volume 43, April 1989 \0 M M NNO composition and level of coral coverage to v-)~....;oo .-I \0 lr) r-. ;:: ('fiN-V) r- ('fj M 0'\ patch reef communities elsewhere in central M +- 00 - -'"+ Kaneohe Bay (Maragos 1972, Evans et al. 0'\ - 1986). The reefs in central Kaneohe Bay have ~ been subject to similar levels of stress from M N 0'\ M t- ' E;( '<1"00\0 \0 r-- 00 CIl Coral Recruitment fo< ~~oO"I:t OO("'.l\Otr) z\O '.r'lNN N 00 ('fj lr) r- ~ N NO +N $2 +- _ J 3e_caust: .n() living coral survived on the ~ E;( ::J landward reef side, the recolonization of CI !;; ...l Kahaluu patch reef was dependent upon the ...l <: "'-' ..--.. a S 1----4 . _ --.- . "-" - 0_. -'-'- ' I-'~ Q) p. 2 , .' 0 ,./ .-< (f) / ...... ;;>" Q) 4 Q) P::: _.-I • __ 0 -- ~ ...... I ;;: 6 / -' 0 ~ ( Q) C) ~ 8 \ (\j , +J .rn...... 1983 ~ - 10 a 10 20 30 40 50 60 70 80 90 100 Me a n % Coral Cover Per Quadrat ..--.. a S "-" -, Q) 1----'-.;-1 c, 2 0 f;'-;.--i .-< '- - , (f) , - ...... ~ Q) /' Po r ites comp ressa Q) 4 P::: Montip ora ver r u cosa ~ ;;: 6 Po cillopora damicor nis 0 ~ Cyphast r ea ocellina Montipora p a tula Q) Porites lich en C) ~ 8 (\j +J .rn...... 1973 - -- ~ - - 10' - a 10 20 30 40 50 60 70 80 90 100 Mean % Cor al Cover Per Qu adr at FIGURE 3, Vertical zonation: mean percentage coral cover per m- down reef slope. Number of quadrats at each distance down reef slope (I, 2, 3, etc.) was 10, 10, 10, 10, 10, 10,9, 7, 4, I, respectively, for 1983 and 10, 10, 10, 10, 5, I, respectively, for 1973. Total number of quadrats: 1983 = 81, 1973 = 46. 130 PACIFIC SCIENCE, Volume 43, April 1989 .--... 0 "-"s Q) c, 2 0 -' ...... -- ' 1 -' (/) \ -' ...... Q) 4 Q) -' 0::: -'-' ~ '-.-1 - ~ 6 I 0 ..... 1 Q <, Q) <, C) ./ ~ 8 ./ ...,(\j I ...... rJl 19 8 3 Q 10 o 2 4 6 8 10 12 14 Me an Number of Coral Colonies Per Quadrat .--... 0 S "-" -- .-...- .--. 1-1--="'-.-.;:-,------1 Q) ...... c, 2 - ...... 0 '-.,_ 0."",:::- (/) _0, ._ 0- °_. ...... p Q) 4 Q) -'-' 0::: Porites compressa ~ ~ 6 Montipora ver r u cos a 0 Q Pocillopora damicornis Q) Cyphastrea oc ellina <. C) ~ 8 Montipora patul a ...,(\j Porites lichen -: - ...... rJl------Q 10 1973 o 2 4 6 8 10 12 14 Mean Number of Coral Colonies Per Quadrat F IGURE 4. Vertical zonation: mean number of coral colonies per quadra t down reef slope. Coral ReefRecovery after FreshwaterKill-HoLTHUS, MARAGOS, AND EVANS 131 epifauna (Fitzhardinge 1985).The recoloniza able for coral colonization and growth. The tion of the Kahaluu patch reef was thus coral community developed more rapidly in probably initiated with mixed recruitment of the outer harbor than on the flows as a result planktonic coral planula of the common of the greater protection from wave exposure species found in the Kaneohe Bay lagoon. and the stability and suitability of the sub strate. The mean frequency of coral colonies Patterns ofSuccession in Hawaii per meter square unit area reached a plateau after 6 yr. Coral abundance, as estimated from Patterns of coral succession on Hawaiian percentage coverage, however, continued to reefs were first described by Grigg and rise II yr after harbor construction. An en Maragos (1974), who assessed coral com crusting species, Porites lobata, became more munities that had colonized submerged lava and more dominant over time. Correspond flows. Community composition, distribution, ing with the increasing dominance of Porites and structure were measured on a series of lobata, coral community diversity decreased known-age lava substrates 10 to 100 yr old following a peak 7 yr after the harbor was and on adjacent reference reefs on the island available for coral recruitment. of Hawaii. The number of coral species pres ent, density , percentage cover, and diversity were studied in relation to age ofsubstrate and Coral Community succession in Hawaii: degree of exposure to sea and swell at each Kahaluu Patch Reef station. At the most sheltered stations, Grigg and Maragos (1974) found that coral cover In an overview of coral reef community was high and the amount of cover was nega- structure and succession in Hawaii, Grigg tively related to wave exposure. In addition, (1983) suggested that most differences in where coral cover was high, diversity was low, community structure are those related to indicating dominance by one or two species. species adaptations. The species that even The total number of species present per sta- tually dominate coral communities in Hawaii tion was relatively constant. are usually either the most tolerant to wave In less exposed areas , where physical pro- stress (Porites lobata) or competitively supe cesses were not constantly interrupting succes- rior (Porites compressa). Grigg (1983) con sion, diversity rose initially as all coral species eluded that the primary mechanism control colonized the available substrate. As space ling diversity, community structure (domi became limiting and interspecific interactions nance and patchy distribution of coral increased , the competitively successful species species), and succession of coral reefs in (especially Porites lobata Dana and Porites Hawaii is disturbance. On exposed coasts, compressa) became dominant and diversity de- which dominate reef habitats in Hawaii, long creased. Grigg and Maragos (1974) estimated period swell and wave action are the major that complete recovery of reef coral commu- sources ofphysical disturbance (Dollar 1982), nities on lava flows off the island of Hawaii while in protected environments a lack of may occur within 15 to > 50 yr depending on disturbance may be equally or more impor degree ofexposure, with elimax achieved more tant. In areas sheltered from wave distur- _~ ~ ~ _ qu i ck l yjn _exp~o s ed . envi ronm en t s . ~ _ ~ ~_.__ ~. __bance.zsuch .as.Kaneohe.Bay. Iagoonv.coral. In a detailed 11-yr study on the island of communities can develop to a level at which Hawaii, Maragos (1983) followed the devel- control is exerted by biological factors (e.g., opment of coral communities at Honokohau competition for space) or less frequent physi Harbor after it was quarried from sterile ba- cal disturbances (e.g., major floods). salts inland from the shoreline and opened to The relatively small number ofcoral genera the sea in 1970. Successional patterns similar present in the Hawaiian Islands and the low to those on the "natural" lava flows were number of species within genera facilitate the encountered in the protected harbor environ- study of reef recolonization and recovery. Of ment, which was determined to be very favor- the approximately 45 species of hermatypic 132 PACIFIC SCIENCE, Volume 43, April 1989 corals identified in the Hawaiian Archipelago tions in Hawaii. However, the pattern ofcoral (Maragos 1977, Grigg 1983; Maragos, per reef succession in subtropical Hawaii may not sonal communication), 30 species have been hold for reef environments with a greater recorded in Kaneohe Bay. Halfofthese corals diversity ofcoral species and habitat. are commonly distributed on lagoon reefs in the bay, although community composition is dominated by Porites compressa (Maragos ACKNOWLEDGMENTS 1972). Porites lobata is not abundant inside the bay and is confined to outer Kaneohe Bay We thank the staffofthe summer course for reefs, which are subject to higher levels of their assistance. Special thanks are given to wave energy (Maragos 1972). P. Jokiel for support and encouragement, to At Kahaluu, in the protected central sector P. Alifio for fieldassistance, and to M. McGinn ofthe Kaneohe Bay lagoon, coral community for assistance with statistical analysis and succession proceeds uninterrupted by wave graphics. disturbance. The amount of coral cover con tinues to increase while the number of colo- nies per unit area decreases with time until LITERATURE CITED a biologically controlled climax community is achieved . Competitively superior Porites BANNER, A. H. 1968. A freshwater kill on the compressa assumes an increasing dominance coral reefs in Hawaii. Hawaii Inst. of Mar. of the reef slope community as diversity grad- BioI., Tech. Rep. 15. Univ. of Hawaii, ually decre~s~s . _I-:Io_w~y~r , l~e J~ag ~le_ cqlQfl)' Honolulu. form of P. compressa renders it less suited to -----.-i974.-K.aneohe Bay, Hawaii: Urban the rigorous shallow water conditions char- pollution and a coral reef ecosystem. Proc. acterizing exposed reefs in Hawaii. Conse- 2nd Int. Coral Reef Symp. 2: 685-702. quently, Porites lobata and , possibly, Pocil- CONNELL, J. H. 1973. Population ecology of lopora meandrina Dana are instead the domi- reef-building corals . Pages 205-245 in O. A. nant forms in those environments. Jones and R. Endean, eds. Biology and In the lava flow and Honokohau studies, ecology of coral reefs. Vol. 2. Academic the low coral community diversity of early Press, London. stages rises rapidly to an intermediate peak as ---. 1976. Competitive interactions and more species become established. Although the species diversity ofcorals. Pages 51-58 the present study at Kahaluu reveals the in- in G. O. Mackie, ed. Coelenterate ecology creased dominance of Porites compressa and and behavior. Plenum , New York. nearly the same community diversity index - - - . 1978. Diversity in tropical rain forests values after 18 yr, a pattern of intermediate and coral reefs. Science, N.Y. 199: 1302- higher diversity cannot be determined because 1310. only two surveys, 10yr apart, were conducted. DOLLAR, S. J. 1982. Wave stress and coral The overall pattern of coral community community structure in Hawaii. Coral succession at Kahaluu does seem to follow Reefs 1:71-81. that proposed for Hawaiian coral reefs (Grigg ENDEAN, R. 1976. Destruction and recovery and Maragos 1974, Grigg 1983). Results from _ofcoral reef communities. Pages 215-::254 in previous surveys in Kaneohe Bay by Maragos O. A. Jones and R. Endean, eds. Biology (1974) indicate that protected environmental and geology of coral reefs. Vol. 3. Academic conditions and other factors may result in Press, London. complete reef slope recolonization within 30 ENDEAN, R., and W. STABLUM. 1973a. A study yr. The present study shows that, 18 yr after of some aspects of the crown-of-thorns destruction, the coral community on Kahaluu starfish (Acanthaster planci) infestations of patch reef is proceeding rapidly in a pattern of Australia's Great Barrier Reef. Atoll Res. succession suggested by previous investiga- Bull. 167:1-62. Coral Reef Recovery after Freshwater Kill-HOLTHUS, MARAGOS, AND EVANS 133 - --. 1973b. The apparent extent of re - --. 1974. Coral transplantation: a meth covery of reefs ofAustralia's Great Barrier od to create, preserve, and manage coral Reef devastated by crown-of-thorns star reefs. Univ. ofHawaii Sea Grant Advisory fish (Acanthaster planci). Atoll Res. Bull. Rep. UNIHI-SEAGRANT-AR-74-03. Ho 168: 1-26. nolulu. EVANS, C. W., J. E. MARAGOS, and P. F. --- . 1977. Order Scleractinia, stony HOLTHUS. 1986. Reef corals in Kaneohe corals. Pages 158-241 in D. M. Devaney Bay. Six years before and after termination and L. G. Eldredge, eds. Reef and shore of sewage discharges (Oahu, Hawaiian fauna of Hawaii. Bernice P. Bishop Mus. Archipelago). Pages 76-90 in P. I. Jokiel, R. Spec. Publ. 64(1). H. Richmond, and R. A. Rogers, eds. Coral ---. 1983. The status ofreefcoral popula reef population biology. HIMB Tech. Rep. tions in Honokohau Small Harbor 1971 No. 37. Sea Grant Coop. Rep. UNIHI 1981. Pages 32-59 in A decade ofecological SEAGRANT-CR-86-01. Honolulu. studies following construction of Honoko FITZHARDINGE, R. 1985. Spatial and temporal hau small boat harbor, Kona, Hawaii. U.S. variability in coral recruitment in Kaneohe Army Engineer District, Honolulu. Bay (Oahu, Hawaii). Proc. 5th Int. Coral MARAGOS, J. E., C. W. EVANS, and P. F. ReefCongr. 4 :373 -377. HOLTHUS. 1985. Reef corals in Kaneohe GRIGG, R. W. 1983. Community structure, Bay six years before and after termination succession and development of coral reefs of sewage discharges (Oahu, Hawaiian in Hawaii. Mar. Ecol. Prog. Ser. 11:1-14. Archipelago). Proc. 5th Int. Coral Reef GRIGG,R. W., and J. E.)vtARAGOS. 19].4. Re Congr. 4: 189-194. colonization of hermatypic corals on sub PEARSON, R. G. 1981. Recovery and recolo merged lava flows in Hawaii. Ecology 55: nization of coral reefs. Mar. Ecol. Prog. 387-395. Ser. 4 : 105-122. HODGSON, G. 1985. Abundance and distribu PIELOU, E. C. 1966. The measurement of di tion of planktonic coral larvae in Kaneohe versity in different types of biological col Bay, Oahu, Hawaii. Mar. Ecol. Prog. Ser. lections. J. Theor. BioI. 13: 131 -144. 26:61 -71. RANDALL, R. H. 1973a. Reef physiography HOLTHUS, P. H. 1986. Structural reefs of and distribution of corals at Tumon Bay, Kaneohe Bay: An overview. Pages 1-18 in Guam, before 'crown-of-thorns' starfish P. I. Jokiel, R. H. Richmond, and R. A. Acanthaster planci (L.) predation. Micro Rogers, eds. Coral reef population biology. nesica 9 : 119-158. HIMB Tech. Rep. No. 37. Sea Grant Coop. - --. 1973b. Distribution of corals after Rep. UNIHI-SEAGRANT-CR-86-01. Ho- Acanthaster planci (L.) infestation at Tan nolulu. '" guisson Point, Guam. Micronesica 9: 213 JOHANNES, R. E. 1975. Pollution and degrada 222. tion ofcoral reefcommunities. Pages 13-51 ---. 1973c. Coral reef recovery following in E. J. Ferguson-Wood and R. J. Johannes, extensi ve damage by the 'crown-of-thorns' eds. Tropical marine pollution. Elsevier starfish Acanthaster planci (L.). Publ. Seto Oceanography Series No. 12. Elsevier, New Mar. BioI. Lab. 20:469-489. York.- - - SHEPPARD, C. R. C.-1982. Coral-populations LoYA, Y. 1976. Recolonization of Red Sea on reef slopes and their major controls. corals affected by natural catastrophes and Mar. Ecol. Prog. Ser. 7:83-115. man-made perturbations. Ecology 57 :278 SMITH, S. V., K . E. CHAVE, and D. T. O. KAM. 289. 1973. Atlas of Kaneohe Bay: A reef eco MARAGOs, J . E. 1972. A study of the ecology system under stress. University of Hawaii ofHawaiian reefcorals. Ph.D . dissertation, Sea Grant. UNIHI-SEAGRANT-TR-72 Dept. of Oceanography, Univ. of Hawaii, 01. Honolulu. Honolulu. STODDART, D. R. 1963. Effects of Hurricane 134 PACIFIC SCIENCE, Volume 43, April 1989 Hattie on the British Honduras reefs and British Honduras reefs and cays: Re-survey cays, October 30-31 , 1961.Atoll Res. Bull. of 1965. Atoll Res. Bull. 131: 1-25. 95:1-142. - -- . 1974. Post-hurricane changes on the - - . 1969a. Ecology and morphology of British Honduras Reefs: Re-survey of 1972. recent coral reefs. BioI. Rev. 44: 433-498. Proc. 2nd Int. Coral Reef Symp. 2 :473 - - - . 1969b. Post-hurricane changes on the 483.