Coral reef recovery subsequent to the fresh water kil.l of 196'.

paul y .'>olthns9"partm4nt of Geography~rnivers'ty af hawaiir'onolulu, Hawaii 96822 OSA

Clrrr=tar har u. BVaasrr'niv . sit y of Calif ormia"ant: Cruz, California 9506V vsA

,lamer ". sara< as=nviror mental Resources sectionV. S. rrey yngineer prvrsron«o ' ~ha.'terr Bavaii 96856 VSA

' pres ar.t address: 628 11th Street, ltanhattan Peach, Calif amia 90266 USA

Ak stract

The recovery of the coral reef coaaunity on a patch reef near Kahaluu in'.aneolre ra; was studied. This reef vas denuded of live coral by a "fresh vaterkil'" .'n 1465. The present Study [1983! iS a resurVey nSing rmethads Of an ear-1 i r �o 31 s!udy. The species, abundance, and distribution of corals on thepatch reof ir. 1983 vere measured and recorded along a series of transects.f esults show large increases in the size and numbers of colonies and coral areacoverage. Analysis of di stribution reveals an extersion of coral coverage f ar-t her lawn the reef slope, but with highest abundance in the upper 5 nr.Community di vcrsi ty decreased slightly as the fast growing pyrites ~~regna be-came more dominant. The pattern of succession at this sheltered site appears tole cor sistent with results from other investigations in the HawaiianArchipelago..'.ecovery appears to he rapid in protected, lov vave energy. iafre-guentlv disturbed environments of Baneohe Bay. Almost 20 years after a aa jarreef kill, the Rahaluu patch reef slope coral coasunity appears to be approach-ing it» p. e-disturbance r:orrdition.

I n 't rod action

Pv ca use of the corn plexit! and longe vity of coral reef s yst eas, f ew long terr.vtudio ~ have lean made cf coral community dynamics, especially in the guantita-'. ' va 4::4< sneer,t nf coral reef recovery from natural or aan-sade disturbances.'.4 1" ', Kanmohe pay, Hawaii experienced heavy rains in conjunction with lov, hrs killed many nearshore cora 1 communities [llanner, 1968! . In the'. ~ lrv � . sector of the ha I, a patch reef slope f acin g th e entrance 'o Kah al uu

was completely denuded of live corals. The initial recovery of this cor-a e ' was monitored from 1 468 to 1973 haragos, 1 97g! . The present study is ar.sur vr y of the rahaluu patri. reef lope coral community to determine the extento i;s r ~ rovery 18 yearS after the original disturbanCe.

reef communities are su! ject to a variety of destructi ve events : mr 1!nrt, 1 at "; Jahannes, 1975! . pelatively yeW instanCeS Of COral reef reCOv-r -y 1 v 'een documented vndean, 1976! . pearson f1 9 ST! defined reef recovery.estorntion 4 a coral arsemhlage to a degree comparable to its orioinals ~ ate ~~ 1 maggested t hat this may require several decades following major natu-ra 1 ! r' =1 urbances..he reestabli haent of coral reef coaaunities vas studied oa

! at.ed 1<va flows, and long term succession of coral asseahlages vas docuaented gri" r and "aragas, 197rrI . The patterns of coral colonization and succession inn" 4 1! crea ted habitat. vere f o.'lowed in detail orrer an 11 year period"oaor "I du, Vawaii aaragos, 19gg! ~

ooral r slope comm«r.ities throughout yaneohe Bay were resurveyed in!or um~ nt t! eir ecoverv fi ve to six years af'ter sevage input vas divert-e! +4 ~u',side he bay yvars ot al., this publication! . In other areas, theover ! -.. ree a seablaoe. di srupted bv natural drsturbanCes are consideredra rr,' y a fore t ' on of r ime rloya, 1476! . The reSpnnae af coral reefa tO bath

arrl natural pert urba tie'ns rn a va.iet y of locations is reviewed and'v . ear sari f19p1! .

dr 6

In '. he present "tu "y, a series of t rausects are used '.o replicate earl zer in-vestioa ti ons and provide co mpara t ive in'or nation on t!.e size. a hund ance, an'dis rihut iOn Of cOralS alOng t>e Kahaluu patch reef slo!e.

Materials and methods

In duly, 19P'., a resur vey oi the coral. community on rahaluu patch reef,xaneohe '!ay, was carried out. Ten survey stations vere evenly spaced along thesouthern and western [stream facing! slopes of the patch reef, very closely ap-proxima .ing the sites surveyed in earlier studies Fig. 1! . At each station,coral colony ize, species composition, and number of species vas obtaired usirgthe exact methods of naragos �974! . A 1 4 frame vas posi.tioned on the reef

lat at the slope edge. 'fhe maximum diameter and species of each coral colonylying at less' 50 witkin the quadrat vere recorded. Tl e frame vas then moveddown tke slope in a series of consecuti~e 1 mv units. Information vas gatheredin each uccessive quadrat until no corals vere encountered. T'hus, a transectm ride beginning at the patch reef crest and extending to 13 m dovn the reefslope vas surveyed at each station.

Fig- 1. location of the pate! reef near the l.ahaluu .'tream where a aerie - ofsurveys vere conducted along a reef face denu 'ed of corals durxng thefreshvater kill from Waragos,

67

1 r reef fs at edge at this site is a feature of fairly uni form dept h composi-P ion and rel ief in relation to patch reef slopes in Kaneohe Bay Boy, 197D! .

re f cr-est th us served as a reasonable ref erence point in relocating the.",tart nf each slo pe transect. In guadrats of high per cent coral cover, colo-nies mav abutt and grov together, foraing aggregate heads of mote than one colo-Ily. ' n thi. S Sit uaoion, diSCernable colur and mOrphol.ogical dif f e renCeS enabledindividual colonies to be counted separately. A fe v larger colonies had dead

nt ral. and ur per portions. 'These vere measured as vhole live colonies unlesstighl fragmer.ted, 'n vhich case the distinct live portion vere aeasured. lnaddi p ion tO Specif ic data, photoS Vere taken and general observatiunS regardingvs ter iuality, substrate composition, anrl other organisms present vere noted

ir ld data vere compiled and compared to data from the ear' ier survey ofmaragos �?Ta! . This alloved us to determine the extent of recovery. Speciescomposit~on vas exaa'ned in terms of total numbers of colonies and total areaAve r . +o ca lcu 1 a t e area coverage, co lony di a meters vere con vs r ted using the

for vula xrea=y. 10 rz vhere r= 1/2 the diameter. This conversion formula may behiasrd in favor of flat, encrusting corals over multi-diaensional branching col-onios, Also, pet' cene. coverage per ouadrat mv may be distorted. por this rea-son absnlute amounts of coral cover are used in making comparisons and not percent cover per guadrat.

rara y distribution vas analyzed both vr rtically and horiczontally along thepatch reef slope Horizontal zonation vas examined by plotting coral abundance,as expressed in numbers of colonies and total coral area coverage for each traa-sect station. Vertical zona.ion vas displayed by plotting both the number ofcoloni-s and tctal coral area coverage per quadrat meter measured dovn the reefslopo. The maximum grovth rate for each species vas calculated by averaging thediamety rS Of the largest five COlunieS for eaCh sPecieS, in bnth 1973 and 1983.Coral community diversity vas measured and compared by applyiryg theShannon-n'ener Index to the information collected.

Results

Cpn ra' aluu patCh reef in «aneuhe Bay, betveen 1973 and 1983, Coral abundanCeincrease! in colony numbers and and area coverage [Table 1! . Coral cove" age in-creas<1 by omar 660% chile the total numbers of colonies rose about 1SO'e. The

u *r .y pe l your' tes ~cmyess, so~et yo y r u ~poc o ort~la i~or nis, a n d r~Bha~ct e~ o~c~li na ! const.i tote soar ly al I of the coverage andcoTnnirs .ncounvored. ize freguency distribution reveals the shift tovardsgreater numbers of larger colonieS far poritey co~a~asm g~ont' IyOra ~ve ~uC~apand og~ci |~os ~dmgc~r~n' s Table 2! . In 1973, no colonies greater than 30 csverv found. ry 1963, over 25y of the colonies measured vere larger than that.,iz". In contrast crf+naslrga oceQgjna disrlayed a marked increase in the nua-i er ryf smaller colonies io 19Bv, v hie!, aav be as a result of its response to de-creased sevagc ir. I'.aneohe Bay Tvans et al., this publication! .

.i charging distribution of coral abundance vas examined horizontally alongthe rvef face and vertically alorg the patch reef slope Ro distir.ct patternsre]atrd o distance from the surviving reef slopa community vere obvious in theae.sor ment o total coral coverage or number of colonies ITable 3, Table g!.Vert ically lovn the rerf slope, in 1473 a zone of high coral coverage and colonynumber. occur in th upper five guadrat meters By 19B3, the coral coverage int hi.- t an ha expan ied greatly vhile the total numbers of colonies has not in-creasy" much ', Tat le 5, Table 6! . Although the extent dovrslope at vhict: coralsre n"o rt ered had searle doubled ty 1983, the main zone of h' gh coral abun-

dance -xsr nds on' y slightly further dovnslope.

information for each species revealed po~l~s coapressa and QQt~orayf.rufosa to he grovrng rapidly. gocillopora gamico~nis shoved a gradual in-cr~as~, vhi le Cvphystrr h ocellinh grovth leveled off a fter 1973 Table 7! . Thesllif e .' n coral community composition iS apparent in 1 he inCreasing dominance Of"~r iten co~nrcssa Table 1! As the. teef slope coral assemblage returns to itsfor mer coy dition, the number of colonies per mz decreases gradually as coveraoe

m y increa" es, e:pecially in the case of ipor jtes ~mressa Table B! . This"har,-e i re" coed in a slioht decline in community diversity, ftom 0 $0 to

sva ured hy t! e ha anon-riener Index of commuxtity diversity-Discussion

study of co a' reef recoloni.zation and recovery in the Havaiian Islandsis '. cil: t atedy by the relat ave lv small number of coral genera present and the

Suv nary of txar,seCt Bata for the tao Surveyn.able 1.

Area Coloof

mv! .ot al Ho.Area

o' cmv! Totalspec ies

97.87 v. 1 358, 06634686. 3

3o 796

4, 433

9.611.410.2

67 14 23. 20 13930.3

7!or.t~ioza gatnlg

9 o~zt~s ~ic hen

1. 45, 792

0.1�. 130 07 09 10091 00%407, 88361, 256 100" 4'73 10oz

otals

Table 2. Size frequency distzibut ion foz coabin ed transect data. Ta ble valuesare num! er of coral colonies for each size class.

COlony siZe Class [di ameter in Cm!0-10 11-2D 21-3D 31 � 40 41-5n 51-60 61-70 71-80 81-90 91-'100

Pohi~tS COmlIZesSa1963 114 1121973 166 128

charge -52 -14

620

+62

00 00 +7

10

vl

290

9 29

117 9052 0

+65 v'90

t~rfrro o~ere aa1983 23 191'973 33 '19

change -10 0

10

+1O 0 00 0 04

0+5

30

+3

30

+3

019vg f,2 5 0 0

change 0 ~2 +3 0

0 0 00 0 0 00 0 0 0

0

0 00 0 0CV Ola1es -~ea pc~cling

06 0

+35 0

0 0 0 00

0 0 0 0 0 00 0 G0 01983I 4 13

chango

0 00 00 G

lie/en00

pozites0 00 00 0

mont il!gra pat!!~a,2 00 0

+2 0

Ot.hez19831'9v3

change

0 0 0D 0 00+1

TOT!'.v19631o73

chn Iqr

33650

v65

24 2 138 12kl 95267 152 54 0

5 -14 m600+2v7e32

e

o:'r.=. c~ore aa52, 882

run ti no ra ver rue os a6,259

P oc~lln oz a ~mi c~or ' s1,963

Cvoh astzea no~el inB

1 9'73 SU" VTY17uubPZ OvColonies

ofIln. Tot a!

1983 SDFVBYSumter of

nies~ ' ofTotal

530 74. 9

61 8.6

72 1D.2

41 5.8

3 0

-.ahle . 11orizor tal zonation: Area coveraoe caz! o' coral in ali guadrats ateach station.

station5 6 10

1983 53754 3'1 353 3404 0'1 973 5881 4129 5552

change +47'07 +24886 v26753

41688 49 50 50727 34098 42536 42884 270516281 5785 3169 4574 4921 7915 4674

+28o16 <24541 +40238 +29217 +35549 >29417 v 17462

aontiqora ~ve racogB19>3 36 1 19801'9 ~ 3 699 2'92

change -338 + 1688

5F35 17c155 72601270 694 10'9 5

r4365 +16761 +6165

29 1 986 3304 243529 0 '126 79

-230 +986 + 3'178 +164

go g i alloy or 4 gBa jc o~n j,s19<3 242 3581973 80 523

Chanc~ r 162 -165

~p't 98~83 ~ocei inB1983 fia 3 131973 0 3 0

ch angr +r4 P el 3

137

Qo tiger a patula, goO c! 00 0 00 0

983qv3 0 2054

P 00 +2054an 3~

053754 31353 34040 41688 49250 50727 34098 42536 42884 27051666 1 4q48 7049 7649 5813 4955 5152 5105 8078 4846

+470" 3 + 26405 >26991 +34039 442437 +45772 +28946 +37431 >34806 +222o5cha hcr

1

ch

-.a 1.9n3q7 1

168214aa4198

41

+28

942 145098 314

+644 +1136

110 "P0 20

+ 110 0

rites ~cen!3 P0 o

r 0

52570

-51a

8122

-11 4

0 00

1657

-41

0 0D

D 0 0

ap184

-164

600

+60

'I 8 030

+150

832q3

+739

lo30

+103

37370

+3737

Tat I e u. HOriZOntal ZOnati Oa: II saber Of COLOn ieS in all. g O aar a l S at enon St. a-t io n.

R E CCRC RSStation

2 3 4 5 6 7 R 9 '10

75 40 61 4617 24 20 45

+58 a 16 +41 sf

4025

~ 15

1983 541973 57

change -3

oorites

2 74 D 20 +2

31

v 2

610

1993 3I 'l7 3 12

change

Ifgn t~iararucosa

I I 72 4 3

� I -3 +4

1610+6

PgcjllofIora 1983 6 !ha jcornjs I'973 9

change -3

I 'I0

vlf

19831973 0

change + I

0 100 00 +10 0

~c!lh asthmaa~ 3

-1

I0

+I

'l98 3 0lo73 0

change 0

0 0 30 0 00 0 +3

Other l0 0

7136

435

675E

45 74 6430 24 5115 v50 v13

foe 3 64I 973 78

change -14

9855

TOTAL32

+11 v 56

~ llont~iora pats/ I. LQW~ LMhfn ~

Tatle 5 Vertical Zonation= Ilean Coral COver Can! per aa guadrat.

Distance �! dovn reef slops f ron crest2 3 3 4 4-5 5 6 6-7 7-8 8 9 9- I D0 � I

Pf�;j&eg goaera~a1983 26'96 8349 7165 66521973 97 762 2807 1426

change 42599 +7587 +4358 45226

96 15486 0

90 v 1'548

8520

+852

5026191

+4875 +21

il o 4 t. h Qtggh v~~ j3 c sf1903 8 I 5851973 0 33

change +81 + 552

270

4 27

1870

v 187

70 1270137 439-67 +831

96818

v 950 43

pccillonara Baaigornis19P3 48 2331973

change +." 7 v 'I 8 00

+40 0 02 025

61-36 o 15

CY2hastrRR M~Jkn31I'983 15 101973 I 0

change 4 100 0 0 0 0 011 3

104 I -1

pgh! tes lie>en!0 0 00 0 00 0 0

10390

+ 1 D.l9

2560'9 177 727 1 7"96848 3D15 1925

v8329 +4256 +6071

5998211

v 5787 i 2554

Ot.her Ilonti~ara1983 2C'51973 0

change +205

'TOTAL1903 304519v3 119

change v 2926

rontl17 4000

59 3944 42

+15 -3

211

+5 -9

8 24 4

-I -2

'I0

0 vl

0 00 00 0

71 4458 57

+19 -'l3

6637

429

158

-7

100

+10

10

vl

D

0 0

4237v5

156

+9

812-4

21

41

83

+5

39

-6

620

62

0 0

00 0

3640

v 364

I '9120

1 '912

12430

'1243

60

v6

0 0 0

3740

v 374

16270

v1627

0 0 0

Pl0

v81

0 0 0

cail ~ 6. VertiCal zona tiOnt I!ann nunber of Coral COl onion per nv gun drat.2 2 Cs 4 SSS 2 4 ~ ates CSC~ 4 scc 44 t'4 4

Distance [n! down reef slope frow creat12 23 34 45 56 67 79 09 9-10p-1

cp~r' Zgg ~onPre~sa19a3 8.4 tZ.3

4.0 9.1change t3-6 «3 ~ 2

2.9 1.2 0,30.0 O,D 0.0

«2 9 41.2 40. 3

2 ~ 20.0

«2.2

02875412. 9 6.5 1. 2-4. 1 42.2 au. 2

PODZZR [X9 vgZLPH?Bd191!3 P. s 0,9T9v3 0.0 0 6

change «0 5 «0. 30.5 0 0

0.0 O.p O.p«0-4 40 5 0 0

0-6O.D

«0 6

0.5 I.tl 1 !1.5 3.4 0.1

-'1,0 -?,4 «1 P

t.PPkllDPDXD hddkgtffpld'1993 1,3 1.7T973 li 5 2.9 l. 4

rhange -0.2 - 1.2 0.00.4 00 0 0P.P 0.0 0. 0

«0 4 0.0 0.0Ct 4 P h Opt f ed Modil M 91'7l}3 1. 5 1 1 0.8 0 5 0.0tgv3 D.2 0.0 0.2 0.2 O. 0

change 4 T. 3 «1. 'I «D.6 tp. 3 0. 00.0 0 0 0 00 ' 0 P.p 0.0D.O 0.0 0.0

other !9PZlpo~d Pa did. fgfltfg j~!1903 0.1 0? 0.1 P 0 0 D

p D D.D 0 0 p 0 0.0change «0.1 a0.2 «0.1 0.0 0.00.0 0.1 0.00.0 0. 0 0. 00 0 40 1 0.0

11. 7 16,2 10.6 11. 3 7-212.6 16.0 11.3 1.4

t4.2 «3 6 -5.4 0.0 45.02 e0.0

«2.8

3 7 1 7 D 30.0 O.O 0-D

+3.7 «1,7 «0.3

4 2 I css 2 4444 44� 4 2 4 2 ~

Tabfs 7. Ilariau ~ rural growth rates for each Specien. Ilaned oo average Oflargrst 5 colony dianetern [ca! .~ 4 ~ 4 '4 4 - 4 sc *est

4 sc 2 ct 2 2 ~ ss ~10 yr.1965 1973 1903 rate ca/yr!

ZPZlf 0 RKPfe~ngl 0 27 0 74 2 4. 1yto~to~r ~vvc~d 0 19 6 65 6 3. 6Zfygil~ogfd I!dg,'~n'a 0 13 6 22.4 1. 25 v p[t ~st 5-8 6.4 0 4

~ 4 4 ~ 42 2 4' tt 4 ct 2 4 ~ ss ss

Tt! !r !'. ConIIurt sot of nueher of colonies nnd nean coverage fOt the two sur-rey

~ « I � 92 4 .CC -t � 92 � csc 2 4ac*n no. of ColOniea

per sv gundrat lleun cover eve!per ~ 4 guadrat

T97$ 19!13 73 1983nor.tippy 4 vfrr~unsd3 opl!iopIortt >dfrj cor~ndCf [hatt r sd octo«ha

Total133'l.l 5P35. 6

cr«tsct 2 Cc 92� etc 2' '!clif op?a Ial Jld, ZOOPltes+i~he .

72

goy at19931973

chanae

1.1 P,7'I 2 0 ~ I

-0,! «0 6

'T. 61.21.5p

2. 60 3

«2. 3

0. 50. 0

«p s

p. '!D. 0

«0. 1

0. D0.00. 0

0-00. 00.0

3 20.3

«2.9

6,4D.70.80 5P 1

0.00,00- 0

0.00.00 0

0 ~ 00 00.0

'l 149 6136. I42 73 3O.p

4420. 6403. 954. 7~ .9

7'! 5

lov speciation within genera Naragos, 1977!. The occurrence of 30 coral spec-ies in the Xnneohe may area waters was studied by Naragas �972!, vho f ouodabout ha' f of these to be co~sonly distributed in the lagoon reefs, r iog andHaragus �97m! propOsed a patteZn of reef COral SuCCession af ter examini..g Carelcommunity composition, distribution, and structure or. a series known-age sub-merged lava flovs and on reference reefs ad!scent to the flows on the island ofHawaii. The number of coral species present, ds nsi+ I, per cent cover, and div-ersity vere studied in relation to age of the substr ate and degree of exposureto sea and swell at each station. Although all of these sites vere coze exposedto destructive physical disturbances than Vahaluu patch reef inside Kaneobe Bay,siailari'ties of coral assemblage devel.opaent are evident.

At ! l-,e most s? eltered stations, grigg and Wzragos found that coral cover ishigh and that the amount of cover is negativelv r~Ia ted to exposure. y urther,vhere coca' cover is high, diversity is lov, indicating doainance since thenumber o species present was relatively constant. In less exposed situations,where physical processes are not constantly interrupting succession, di versityrises initially as more and more corals colonize the available substrate. Asspace becomes limiting and interspecific interactions increase, the competitive-ly successful species becoae dominant and diversity decreases. In the shelteredenvironment of Xaneohe Bay, this pattern of less interrupted succession is ap-par nt The amount of cazal cover cortinues to increase vhile the number ofcolonies per 'unit area decreases. One species, rorite~ con oressa, assumes anincreasing dominance in the reef slope coamunity as diversity gradually decreas-es. ",zigg �983! considers this coral to le competitively superior and that 'tvill eventually doe.inate a community vhere wave st ess is not an import.act fac-tor.

ln a 'etailed study on the island of "avaii, Hazagos �983! systeaa' icallyfollowed the development of coral comvunities after the construction ofHonohohaa Harbor in 1970. Although the protected harbor envizonmert vas ~ uchmore -avorable to coral colonization and growth, successional patterrs s milerto those on lava f love vere found. he more rapid coral community developmenti.n the harbor is thougl-t to be due to greater protection from vave expo ure andthe stability and suitability of .he substrate, conditions comparable to thesituation. at Faha luu. In Fonokohau p arbor, the sean fr egue mr y of cora I colonynumners reached a plateau six years after the substrate became available whilecoral abundance, as estimated from per cent coverage, continues to rise elevenyears after construction. Presumably, colony numbers vould begin to decline inthe near future as space becomes limiting, similar to what is occurring in theeighteen year old recovering reef community at Yahaluu. An enczusting species.'varites lobate becaee rore and more dominant over time or the boa't harbor sub-st rates.

Growth rate information is similar from both the Honokohau and Xaha' uu situ-ations. Common species of the genera Zorites and. '!~out' or a sl-,ov initial andcontinued fast growth. Small encrust ing corals such as Closest rea ~c~el ~na ex-hibit persistent slow growth.,he dominarce of ~os i tes in t eras of size, fre-quency, aod surface coverage becomes established at advanced stages of communitysuccession in both study azeas. n the lava flow and Honokohau studies, he lovcommunity diversity of early stages rises rapidly to an 'ntermediate peak asmore corals become established and continue to grov. Hovever, with t' he increas-ing dominance of sorites, communit I' diversity at ll one kohao decreases markedlyseven years following the creation of the available habitat. The lack of inter-mediate time period data from Xahaluu prevents de+ermining vbether the sane pat-tern emerged as explicitly from this study.

However, as in other cases documented on Hawaii, diversity of the coral com-munity at Kahaluu is seen to be decreasing due to i~creased dominance ofPorites. In both the lava flov and Honokohau st.udice, ~o' 1~a~or meandrinm isa major component of the communi+y. ~Pocil opera ~man~dine is absent from areasother tl.an the kiah eave energy regions of Xaneohe bay Haragos, 1973l ..he re-covery of the reef slope community at Kahaluu probably also has been influencedby efi'ects of sewage vhich vas discharged into xaneohe aay until 197P Isee fvanset al., this publi,cationl .

There are many variables in the factors governing tho recovery of co-alcommunities. Pe arson lgsl! discusses three important areas of contention whichcloud the issue. First, it i ~ r ot clear what con ' i. ales a climax covmuni y.It miglt dif fer f roe one localitv to tie next Also, the gues ion of .in~ scalr

73

present st mdy suggests that 'the patch reef at Kahaluu is not fully recov-ered -jghteen yeats after d .struc ion. Coral community succession appears to beProc< ling ra!idly in 4 pattern suggested bp previous investigations in Hawaii ~Recause o! the leng th of time reguired f or complete coral reef recovery, futraresv udice "hould continoe to f ollov the reestablishment of coral communities inthese 'ypes o situations vhere initial surveys have been made after ma jor dis-t ur na ness.

Literature Cited

rrann r, ',. K. roF8. K freshvatr r kill on the coral reefs in Hawaii. Harvaiir,. tir ute nf "arine Fioloqy -.echnical peport r '. University of Hawaii,rronulul u.

andean, S. r "~e. D tructron anrl recovery of coral reef communities. pages2'l'-234 in Uiology and Geology of Coral Peefs Vol. 3. 0 ir Jones and R-'.rr d e err, e dn. a c a 1 e 4 ic Pr a ss, o n d on.

Gri I;, S. l:. and J. ", Haragos. 7g74. recolonization of heraatypiC corals on: u! nerged lava flows in !lavaii. Ecology 5S;387-395.r,rre ' .. e. 'lgp3.9, . Comeunaty structure, succession and development of coralre~!s in Uavaii. aarine ECology � progresS SerieS IIt1-'le.rol af

P r err st 1!IJ! l.,

lny'. pollution and dc gralatron of coral reef comaunities.1!- l'. in Trol xc"ai ".arine pollution. Elsevier Oceanography Series Uo-Ferqeson-eood and n. E. Johannes, eds. el sevier Scientificnev york.

y r, v. l ovr . Peroloni=ation of Eed Sea coral s af feCta d by natural Cat aStrO-aan-vade pr rt uri.ations. Ecology 57r 278-289.

ea . 4,,1 ! $%vr a study of t,udy o. he ecol ogy of Hawaiian peef corals. ph.fr.-r r ar ic r., Pr l r, o oceanography, nniversi ty of yravaii, Honolulu. 2gO'r 47 r. Vier ur it ut ion and abundance of reef cnr als P s 37-egrn r t! a., of Ka neohe ea": A.! .,: .e ay: Reef Ecosystem Under Stress. S. v. Smith, K-a . agesav ~ and " o rrae eds. Drrl "I-syaGEAHT-TR-v2-01. sr' av ~ ". ", -.. - � 2-0l. Sea rant Collegev r i. y o, ava i, Ronolulu.

r y rn74 o create preserve, andCo« ! t ravsplan tati on: a method tree ", Vrriversity of Hawaii Sea G rant ad visory Report-7, Vonol lrl rr-

in vol vv d. Secondly, erterna l dist urbances aay alter succession, prew, »tingt h»' ' ai r men' of any form of a cli maa community. Thi. dly, coral set tleaeat isa~! r f >ed by hotl.. the successional process and anv disturbance, as are the fac-tors ir. luercino coral settlement and survival. Lova >g76I concludes that tbadif Sonance between aan-made polluting activities and natural catastrophes oncora; rc efs is the possibility that the human perturbed errvironrvent siLL not re-r,urn ro it= former configuration, while reconstitution of reef areas denuded byna'ural disturber ces is earnly a unction of tin o. Grigg and !Saragos �974! es-t irat complete recovery of reef corals on lava flow- off the island of Harraiimew rrrr ur vitf 'n 15 to 50 yea s del ending on environmental factors. The resultsfrnv l r vious rorve ys in raneohe Fay by ".aragos 'r g741 indicate that protected:nvr ronaental con]itions and other factors may result in complete reef slope re-cclonirar ion vithin 30 years.

'Us?agon, J. y. 1o7 CrderSclel aCtinia, slory coral .. "ages 1=8-241 1n Feeiand 8! ore au< a of P avaii. D. <. pevan y ar d . 8 Fldredge, e s. cornice

"ishnp .'use us special Publication f a t1! .

Ra:agos, '.. E. 1'983. The status c f reef coral populations in Bonokohau SmallPoat !iarzor 1971-1881. pages 3:- sg in R Deca "e of vcological Studies'Fol losing <onst uct ion of 8onotoh au Sma1 I Roar ''-.ar bor 8 osa, Basaii ", ". ]traySnqineer District, Honolu lu.

PearSO4, R. G. 198 1. neCOVery arrl recoluni at i nn O COral res f s ".!arZne'cology progress series IU:1c5-122.

Foy, 8.,1. 1878. Change in l.athy metric con igura! ior Raneohe kayg Clah1882-1869 Dn' v. Ramaii. Revs' i Inst. nSeophysics Rept. 70 � 15, 22f

StoVart, 8.. 1<88 Ecology and morpholoey ov recent coral reefs. Eiol.Re v. u uvu '33 � a38.

25

eef cora» in xaneohe Bay. Six years before aud after termination of sevagedi chary«�ahu, Hawaiian archipelago!

pent. nf 11iplogIh I 3 " I 6 I t '3 p f C a I i f Or II i aSa tt Cruz, Cl 95060

3ah'- 'araaoshnli~r'.hhe.. vestal Pesources section

I r ty 1'nq}neer Iliv ~'.. 3" after, III 9685'

paul IIoItI3ue9 rar . ahnt of ceogra ph 1'Ilni1 et sit 1 of Havaiiyorola lu, Ilf 96822

Itbs tract

aa,e.sheds sur roundosr he fare 1939. IIevers f:o ected vroa thtart dredving and fillred, vsperiall,y in anseaagh discharges autolarge savage outealls

Ira neoef corale open ocing, res'd around

the 1 agoin the

the saic ntif ic con a unit y, including Itaragos lg t 2!, peg an torn '. h e 1 a goon. 1 t v as spec u 1 a ted t ha t eut roph ica t ion an d sedi aen-result O urbaniratian and COnstructron, vere the cauee of an ob-

e ir. lagoon coral communities in the south laqoon and explosaveqreen algae gjc~t~os hae.ia cavernosa, vhich vas smothering coral,lagoon. Reef corals in tie bay's northwest laaoon remained abun-

ared unaf fected.

� 3'sure f roa the publrc and scientif ic coaaunj.ty compelled t he local govern-3enI »d eilitarv to terminate latge Sevaqe diSChargeS in the SoutheaSt lagOOn

Iao» only a minor amount of sewage is discharged in t.he nor thvest la-OOOII .

yed the lagoon and coral transects srtes o f Haragos �972!These surveys revealed a remarkable recovery of corals,

pressa and eoat12o~a ~v~rc~jl. in the southern aad middleigh coral abundance in the northern lagoon at inor coralamicornis and C~fthastrea ocellina, also vere more abundantntra.t, tictvosnbaeria declined g eatl y ex cept for a aiuorrn lagoon.

Ih lghl ve re-surVeasia; 'oe save methodsesoecially porg+es coalaophh and COntir.Oed bspent<, Paga llopora dio t.'e laonon. fn coinr:aha r rr, t he r.ar the

"tidy and other recent ir.vestiga!i ons corroborate that sevage vas a aa-

'rene "-tudies indicate that the detrimental ef fects of sewage on coralsa e s.rrraily aequi f ied in combined cuba yaents vith restricted circulation.

Io tr od pc t j,on

response of lagoon reef coral popula tions in Xaneoheqe discharge into Iaaneohe Bay by com paring the abun-rals before and after the as~age reaova1. The ora.g-1970-71 at 14 lagoon stations by Haragos �972! andpublished>. ln late 1977 to aid 3978, the tvo majorbarge into the lagoon vere eliai.sated. Tn 1983 ve

at the same 15 lagoon stations using the same tech-

.'ic stadt document s thereduction of seva

'imtrat utiOn Of COioal:a'.3 ver e co 1 lect ed in

station by tokiel funhoist sources of sevage disrresirt~yid rural r,opulationsnay!ho

I..". hr lgh5study caaioestat' or., assertr1 de lragrnvt.h O' the

th~ haddiedent aad anne

he Ital vere dominated by rural and agriculturalC OIh anni tieS f 1 aura shed on latroen reef Slapem andean by a large barrier reef. hf ter 193'9, aili-dential development, and populati on grovth occur-the conf ined southeast lay. as population gree,on increased, culminating in !be constroctaou ofutheast bay by 1963.

Fancohe Bay, located oo the vindvard nor the ast coast of Oa hu, is ttavaii 'slargest embayment< stretching 13 ka along the coast and 4 ka of shore 4 lar ge4 km-long barrier reef occurs 1-2 km of f shore f rom the center of the nay. Thehar'rier and adjacent fringing reefs protect the lagoon from hear! save actionan d restrict va ter circulation and erch ange, especially in ' he isola ted southernend of the lagoon pig. 1!. The barrier reef is deeper in tbe north end of thebay, contrrbuting to more vigorous vav action and vater circulation there- ksa consequence the lagoon can be divided into three physiographic sectors thenorthvest seai-erposed, the central, and the isolated southeast sectors. Thevhole system of lagoon vater" and associated reefs in the bay is unigue toBavaii Kaueohe Bay has been extensively studied because of the presence of theHavaii Institote of Harine Biology since 1951 at coconut Island in the .southernlagoon. prolif ic coral coaaunities have developed in t.be pr otected lagoon . nvi-ronaent of the bay, especially on the steep lagoon slopes and shallov out er mar-gins o f the harrier, patch and fringing reefs. Although most ree coral speciesreported froa Hasaii are found in haneohe Bay< only a fev pecies dominate la-goon habitats ~ vith the finger coral oor ates co~mr~ssa comprising over POC ofthe tOtal live Coral cover Haragos, 1972! . The only other abundant SpecieS iSHongtocra v~ruooea. Other COnmoo Species inClude pOCillognrtl damiCOrniS,p yond at'ldl, cyyyh~e me~thy, Fo~n' cutat', a d noato ta yat r .kbout a dozen other coral species have been reported from the laooon, includinga ft tal yo t~ho ~f'c< a d th ah typ c al n tf cctHistorical Background

Prior to vestern contact, the Havaiian population in the Faneohe vatershedvas principally involved vith taro and fish culture Devaney et al, 19761. Thebay vaters vere regularly fished, and over 30 rock-valled fishpond- aanagedalong the i.nner reef a nd aud f lats. The 19th century sav the decline of theHawaiian colture and the development of ranching and laroe-scale agriculture inthe bay, including rice, pineapple and sugarcarle. Hovever, by the end of thecentury, aucb of the bay vatershed had reverted tack to small scale farming,rangeland. and rural use.

Beginning in 1931, the bay undervent major changes to accommodate a' litaryand reaidentia1 develOpment, espeCially in the south. 6 ma jor deCade long n i li-tary dredging and filling operation, concentrating on lagoo~ teefe. resulted inexpansion of land area and the construction of a military air station at thesouthern headland of the bay Hokapu!; clearing of reefs and other hazards inthe south lagoon to establish a seaplane runvay; and dredging of a ship naviga-tion Channel f rOa the south laoOOn to the nor th end of t he b ay. ra re dged mat er i-als not used for landfilling vere dumped back into tie lagoon. The 194O's ust.�ered in a era of ma!or residertial development in the soot bern bay vith rurallands cleared and many f ish ponds f i l led for housiog tracts The land dr stu r-bance caused soil erosion and sedimentation in the lagoon from stream runoffduring periods of. heavy rainfall. duct of the sediment set tied in the de<do la.�goon { Boy, 1 9701 Betveen 1920 and 1 9p D, the popul at ior, of t he bay g ev f roeless than 5,OOO to over 60<OOO-

Sevaoe discharges into the lagoon increased in response to populatiorf grovthThe 'Ha rine Corps Air Station at ttokapu began to dischar ge sevaoe into the southlagoon in 1951 Site "A"< 1'ig. I]. Bet veen t951-1972, that sevage sas suh!a ctedonly to brief settling of the major solids Lf tervards an additional step ofbiological. decomposition vas added to the treatment process !n 1963 a largeCity and County aunicipal outfall legan discharging secondar y treated seve geinto the south lagoon Site "B". Fig. t!. 6 smaller secondary sesage treataentplant and outfall vas established at Ahuiaanu in 1970 to service the light ruraland reSidential population Of the ~orth 1 ay {Sate "C", pig 1!. This sehrage en-tered hhuiaanu Stream and eventually the northern lagoon vaters of the bay. By'1977 the coatbined effluents Of the three Sevage plants an d Out falls tot alledover 7.5 million gallons per day �0,00D as day!, «ith 95'L beirfg dis~ha~ged intothe south lagoon Smith et al., 19tf1!

In the late I96O ' s, marine scientists became ala teed at the det eri or at ingcondition of the lagoon's coral coaaunities and suspect.ed d redoing aud fill inu,sevage pollutioa ~ and sedimentation f ron urbanization. to be t he major caus=s Boy 19701 Banner and Bailey, 1970: Waragos, 1972; Smith e ~ al edsl . 1913;~ ~ o.'tars gos, 19741 'Banner, 1969, 1914: Detfaney et al., 1976; and Rollet t, 1, 11} . In1968, ttaragos initiated coral grovth and mortal it y studies using tr an= i lan edcorals to monitor response to bav environment a l conditions. ht each oftransplant stations, many of shici. vere in the lanoon, h~ also conducted trati-77

47ae 45157 5'I 5032

Figure 1. Hap showing Kaneohe Bay andwatersheds. Numbers and circles depictcoral transect number and location inlagoOn. Letters depict locatiOn ofsewage outfalis. B01d line delineatesshoreline. Fine lines offshore depictislands. Very irregular lines offshoredepict upper margins of shallow reefs.HIaiB is the Hawaii 1nstitvte of piarineBiology.

31

r

'b

q30

30

NOr t1I

II/

2929

II

I/ I ~ ~2e

II% II

I //I //,I I / I' I I /

/III ///III////////////// /

~ I/I// IIIIIIII'II, /3///I, I//II / I/r

IIIII I II IIA rr

II

2727 I

5,

South

2on

2125

21

PJI I

II ~ I /////I I /~ /I/// /I r/ /I II ///I/II 'I / I I/

asa7ae15751 w J950

I,II I/I III // Ir/ I I/ I I~ II. IIIII '.'III/r I I/II I II//r/I'III/I... I/// r r..Ir/rrr. I ~'//I I

:I/I. /I//II /I III

//

qo p //I/

8 / I/Ir/III////II I I' / I I / IIII // I//I/I II I II I / I //I

' I II I // / / I I / I' //

//I/ II / I'/// I I'I / I / //I / // I' I /////

/Irrrrrrrr/I// I'///I //I/I/r///I / I'// I r////I'II/I I/////r//r/. I//rr

I//IIII /I//I 'I/I I///I ////I I // //r/ ////////I' ///t

sect guadrat Surveye in 1971 to estimate abundanCe and distr ibotion of reef cor-al populations. The scientific coaaunity collectively blamed tbe decl iae ofcorals in the south bay on pollution froe dredging and urban ration, and impli-e t e .e ge a tr t ti g the gr th of gr rg ~D' t~*h r' t th tsmothering corals in other parts of the lagoon. However, there vere never ar:yadequate baseline surveys of reef coral populations in the lagoon prior to baydredginq and filling operations, sedieentatior, and sewage pollution. Deug:e thedecline of coral populations vas never really documented hist orically nor thecausative factors clearly deliniated.

tv ever theless, considerable sci en t if ic and pu bli c concern ov er t he ve l f are othe hay led the county and federal governments to remove sewage discharges fromthe south lagoon, and by mid 1978 bath major sources vere complete'y diverted toa deep ocean outfall outside the bay. Smith et al ['f991!, anticipating thediversion project, studied and monitor ed bay ecosystem response ty measurinophysi" al, chemical, geological and biological characterist ics hef ore and af teractual sev age diversion. kl though this study succeeded in d ocu mer.ting the shorttera bay response to sevage diversion, i t vas not speci f ical ly designed to moni-tor reef coral popalations and the one year po t-diversion phase of the stud»vas insufficient to document the post-diversion response of lagoon reef coralpopulations. Corals typically respond ~ ore slovly than other organisms.

Be thuds

The ln tranSeot SurVeyS aCCOmplished by Naragos in lgf1 in tl. lagOon verereanrveyed by uS {Fig 1!. Nn additiOnal StagiOn {'8 on Fig. 1! located in thpmiddle laqooa and originally surveyed bv Paul Jokiel in late 1970 vas also re-surveyed because Jolrrel accomplished the survey using the same techni gues.During the earlier study, notes on locations vere recorded. a iso markers vereleft at the Haragos transects< vhich consisted of the oriqinal coral transplantplat forms. OnCe over eaCh Site, 1tar au os used sn oiheling gear to re loca t e theplatforms aad properly position the transect lines.

ht several sites the original platforms coald not be located because they hadmoved {85!, become overgrown with live coral e61, vere buried under natural ac-CueulatiOnS of reef Sediment t7! Or vere not present lad, 4 ln!. Fovever, fromeemory and notes, haragos vas able to position all but one of the resurvey sitesto vithin one to five meters of the original alignments. at anly one site n6

Fig. 1!, the original survey site could not be accurately relocated becausecoral grovth vithin the 12-year interval vas so prolific as to obliterate previ-ous bathymetric features and the 'seascape' as it. vas remembered ee estimatethat tbe resurveyed site vas still vithin 50 m of the original survey site. Inany case it vas not possible to exactly replicate the earlier transect studiesat any of the stations.

In addition to data variation attrit uted to changes in the coral pope»ti»sover time, the inability of replicating the earlier al ignments introduces datavariation attributed to spatial changes in abundance and distribution of coralsover small distance on a reef. To esti sate the magnitude of this variation or"error" ! and coapare that to the magnitude of variation attributed to the tiaefactor, ve accomplished tvo transects at each of the '15 stat. ions dnrir g our 1983resurveys. The transects vere aligned parallel. to one another separated bydistance of 3-5 w,. Data obtained from the "replicate" transects a eacl statiorvere la t er con pared to one a not he r in addi ti on to t.he co mpari s on s be t veen st a-tions

The survey data vere collected using tbe contiguous guadrat meth od originallyfollowed by Baragos �972! . h each station, SCUBh assisted divers laid out. a25 m long transect line from the top to the bottom of the eef slope. perpergdic-alar to the depth contours. at all stations except v6, th line vas long enough

In 1983, a summer course on coral reefs vas heHarine Biology and alloved us to collaborate on asect sites of Haragos �972! . Since our resurveyof sewage from the south lagoon, ve felt this intment changes, if any, to reef coral populatians atage. Hence ve used the saee techniques and sitesment the more recent condition of reef corals andearlier survey results accomplished at tle same siten sion.

ld at theres ur veyva- 5-6 yerval vastributedof the eato comp ares 6 year

Ha vari Inst1t ute ofof the la goon trao-

ears af ter d versionsu f fi cient t o docu-

to reduction of suv-rlier study to docu-e the results to thes before sevage div-

to encompass .e en iretl. t' coral community on the reef slope with the deep end oftbe lr ne on the noddy lagoon ooh dd l floor devoid of coral! aod t.he shallow eod nreef flat where coral grovt is noth is norwally less developed. At station nueber six.29 guadrats vere require . anI I at least 20 quadrats vero accoaplisbed at all.stat inns.

h one aeter square qua rat rased t frame divided with. vires into a grid of 100 squareof r qual area feeaserinq 10 cv. on a side! vas laid and centered over tbe hottoeend of the transect line. eThe diver then recorded on undervater vriting tabletsthe coral pecies, ot er o om1 = ' th r bottom types 1sand ~ eud, rubble. dead coral, coralline

d 1< r<ct <~h~r' < <~. < hh<v <.! «e«ac<square, The nin aue tesoTh inj,aue "tesolutionn of this technique vas half a grid sguare �

hcev! . Species ess . anl than t his si "e vere noted as being "present n. Data on deptnnd slop< angle vere a lso recorded for each quadrat. After coepleting the cen-sus 'fr>r the f irst quadrat posit.ion on the line, the quadrat frame vas then flip-

o<r< t t o t he next posit ion up r he s lope, on the line, and the consus repeat eth drat survey reach d the top of the line. This approach enabled a

htop- to-3!ot toe profile of the reef slope one ee.er vide to be surveyed for eactrnns rt.

The rav data froe the transects were later tr auscribed and converted to per-rent. r ovor estimates f or each spec'ies of coral or algae enumerated in tbe quad-rat. Later the quadrat data vere pooled to give percent live cover aod distri-butio<i estimates for each species reported on the transect

3resu 1 t. s

space did not pr.sit the present atior, of replicate transect data plots foreach i! the 35 stations. trovevs t, data in Table 1 shov that the vatiation be-tveen rr pl irate transect. dat.a vas in most cases much lr.ss than the variation be-tween the '1971 and 1983 transect I ata at the same stations. This is stroagll'evid en t for' plots of total live coral cover vs. depth at all except cost northbay sr .t ions aad stati ous u and 6 in the transition zone. This is also evidentfor p'3 t s of Oict vase!aeyja cover vs. rlepth at all except the south bay stations

St at inn. 12 and 13 in the tranvitiOn Zone. These resultS Suggs st that OVer-all c <ral corerage in the north lagoon did not increase by 1983. Coral coverage

a! r eadv hioh at 3 of rr StationS. IncreaseS in Coral coverage at StatiOn 13,and <o a ! es:er extent Station 12, vere attributed to rapid recovery froe afreshwater coral hill io 1965 lace trolthus, this voluee} a slight decrease incoral covr rage at station 15 aay ha ve been the result of recent encroacheent byVictvo-.nhacrria sl.ich increased dramatically at the site since 1971

O.ct vosr h aeriesout 3; l a y nr es usals not ~d ea. 1 ier,ent pop.lati ori coovas coot d< fferent"deCll n~" in coraldepth at Stat i or,ir croa < r- racked ly

vas nearly absent in both the 3971 and 1983 survevs in thebl 7 beca use cond i tions vere still unfavorable for the algae.coral surveys in 1983 at station 6 probablv involved a diffet-pared to the 1971 survey, and the bathymetry at the 1983 sitefrom the 1971 site. Tb is could explain the anomalous minorcoverage at station 6. Although coral coverage in shallov

not increase over tine, the deep vater populations didin co verso e.

',:ot. i 6 ~ t a ndr r g t he north bay exception, total live coral coverage in the la-goor. as a vhole increasr d dramatically betveen 1971 aod 1983. el cost doubling in

Th rav data from the 1911-72 and 1'983 surveys are presented in Table 1 foteach transect of the 15 stati ons. Transect det.a on to tal live coral. and+gtvo~chae~s coverage plotted against depth are summarized by region tnorth,eidd1<, and south be f3 in Fia. 2 A transitioo zone region vas added as a sub-division of t be eiddle bay region vbich includes the four aid bav transect sta-tions clo-est to the south region. The transition "zone" encoapasses the lagocnsector immediately south of the barrier reef but vitbin the influence of en-hanced vater circulation near Sawpau Channel and also vithin the influence ofsouth lagoon vaters eoving north past Coconut island. Transect Stations 1-3constr t eted the south bav sites, stations v-7 the tra nsition zone sites,Stations 8-11 the eid bay sites. and Stations 12-15 the north bay sites. Afount! south bay station vas also surveyed located adjacent to Kaneohertunicir<al outfall "A" on pig. 13 tut no living corals or 0+ctvos~haeiia were re-nortrd dur ing either the 1971 or 1983 survey. Table 2 presents distributionaldata ror individual species of corals encountered during the surveys ~ and Fig. 3navar rzes for the bay as a vhole the changes in vet tical abundance of

2r "tvpnlrhaerid and live corals bet veen t he earlier and later surveys.

pa[1 1 d* d ta O [' 1 d ~dr~to ha [* o dd Pr. *P d Prd-centaze of total hot to[[ coveraoe, at. one meter depth intervals on each transectand for all stations. Rlanks indicat e tbe transect did not estend to thatdepth. h dash indica tes absence 1 zero abundance! . h "p" indicates presence,bat in very lov aaovnt.s

I.lvs ccshl5

st.at ron 4

I 2 3 4 5 6 7 8 9 10 11 12 13 14 15

13 9.8 - 0.352 44 69 55 4240 5'I 57 45 42

3 5 1100

1 28

4.5 72 3398 48 1292 94 43

10 9 3 6'I 5467 . 62

10 55 67 98

3.5 6652 3437 55

6.5 2045 244 7

I 4 50 7P27 64 76 tp714 73 43 93

1.5 1 634 42 46

22 30

0 53340

P5 5

22 1. 50.44

0. 5 15 0.5 13ZB 17 39 6ZtD 4. 3 59 62

31029

1011

717 2. 112

147 70 6

44 33I 3 Zl41 29

542 5859 73

115

51226

2. 5 0.7

212

1

238 2825 70

I3426

293

2323.5

17.5I

10 10zo

1 0.54.5 130.2 1 3

0 328

5.59.3 D.S

1.52 4 3[I 5 5. 7 10 26

0.5 4 21 2 I 0.5

p. 520.5

1 P60.8 1

2. 3 1.5

81

DepthI tleter19711983a 3. 31983b 1. 6

2 set ers1971 0 11983a 7. 61983b B. 3

3 net ers19711983a 61983b 5. 5

set era1971'l983a 31983b 2

5 aetersI 9711983a 161983b 6

6 aeter s19711983a 0 51983b I

7 aeters19711983a1983b P

8 aeters19711983a1983b D. 5 6

9 veters1971198341983b 2

10 aeters19711983a1983b

11 seters1 o711983aI 983b

'I 2 seters1971198341983b

12144

463053

127774

12543

422616

1.1 2715 3612 'l6

2.7 4220 6828 65

1848 514H 76

1837 6D66 s6

148. 7

ll

72214

103515

646

8

I2922

21312

I 445 64Be 4O

0 5 1050 5246 55

0.537 2357 21

318 1220 16

3.5 211 722 33

5 22I II 6

22 37

897854

2260IP

115712

100 9396 271 ilo 49

99 799P 2098 54

68 6D[8 Ztd

32

25 1356 1532 26

I 2ID 1115 6.5

0 52 5

Ip

Table 1. [cot0 t itI u ed!

0 Std t to~10 acr i a

2 3 4 5 6 7 8 9 19 I 1 12 13 14

51 8.9 41 18 1142 14 5 97 147.8 8.tt 4.3 9 5G. 5

77 102. 6 4. 5 265. 9 3.5 27

0.20.3

9 15 612 3,5 112 5 1 8

733

19 1.5 856

15 13

48 2. 5 12 3. 3 69 400.

2 47 40 3.56 63.7 15

92 371.5

27 656

3.5 28 850.5

20

18 98l. 5

10 2.5

5. 5

7 c

8'0

1 4Iet er197 Ilt!83a 0.1983b 0.

2 aIe> + rs19'1 119834 0.198 3ti

3 aI~tr rs197 I1983a19 83b

~ rt mrs197 1198 341983..

5 aeters'l97 I1983a198 3I0

6 aetrrs197 1193 34198 3h

7 eetr rs197119834198 3b

8 III e t r r s197 1198 taI '04

9 ai t~rsI I7119934108 3 10

IC er trr14711o8 Ia19P. 11

11 p19711>83419831

1! a ~ tr -s19711 a83a1>Ala

tIt-t< rS1 a! 11 "8 3a198 310

12 141'I 4.3

4.2 21

40 265.7 223 5 20

el116

741911

0.53. 56.5

522.25

55

25

35 l. 81. 5

351338

151421

98 6.5I

90140.5

0.4052750.4 S.S 14

2 162 29

30

53529

I IO

28 36

0 52018

HORTH AY / e ~ .a in co ea oo aoco ~

~ D la la Iaoa ff~ iI

z

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CORAL

MID BAYO Fa

CO44O Fa ea la SO COca ~~ D 'lo la taa

' ~ED FoID ~

EPDAEPIA

ICT R AH SIT IOIE o C..e,~ D OD la IDD

CeD Fa 4O SD OD OaD FD

7.r/ e ~e!

IE ~ro1AL LivE

COP A i.o r0 ~

DiC'YOSPSFAEnil

IO

Ie

SOUTH SAY D rea ro ec cc o./, Fee

AC eo OD OeC Fc

FOSf

OTAL LiVECORAL

DiCTDOEDHAERIA

F tao itr. F

I'ig. 2. lrire Coverage of coral and IIi-toos~haeria in 192'I and 1D93 plottpiaqaiaSt depth for f onr regions in KaneOhe lagaon Data fr on al] st ations e/ztn AneaCh r egion pOOled. The dOts represent 1971 data and the "X's" represent 19p3data

83

E es F

Ocf ~

sE

'iOFD ~

E 4

cca ~

t oie .. D stria ti o d t iot species or eeet co:ai d patios aaeria stered during the 1'971 aud 1983 transeCt surveyS at the 15 laaOOn statiOnS.Hunbers indicate total nuuber of tines each species vas encountered in theSquares of the quadrat along eaCh transeot. The 1983 nueberS repreaent the~ cans of the replicate transect data at each station. 4 dash indicates thepe i., a eot. pr t o ta t * t. ep � ~t'dra ~ato a, pd

g~o~ ~ae n rona, and pS � L33l~e~a ~st l~t

LOCA TI 0' RRiddle 84ySOut h Bay Transition Zone Rortl. 8ay TOrALS

Sta tion No8 9 10 111 2 3 4 5 12 13 14 15

3 10

26 172

7 158 67 588

1926 i 9 57 18 1 1028

20 12 34 10 100 27 328~ht 42tyea ooe~lna

191 1

19832 2 9 '10 61

~yng+a ~sc tria'197 1

1 12 11 14198 1

14

~P vo /Id 3~ann1971

1983

36ptther orals

1 d? 7 1

2H P -- 3 hP 3?dp 88P 1 PS 171PR 1PS 2 71P 1PS

0i = t y~:Ltha er2 a197 1

cayernosa

520 327 44R 907 991 610 630 25 134 48351983

8 8 76 225 74 160 40 10 118 30 326 1246

84

rpti' " ~coa essa197 1

16 237 50 9431'98 319 n 313 220 351 374

«o8t in2Za ~ver uCOSa19713 -- 10 43 7 254

199358 22 208 78 172 138

ddc gl~t ~ds co1971

119836 e 8» 18 30

15 365 129 114 376 612 29 711 564 4168

438 1006 353 730 620 872 504 922 424 7154

CymarD Bl jO v ID

e

Fig. 3. Live coverage of coral and Dzctgosghaeria in 1971 and 1993 lottedagainst depti . Data f rom all stations pooled.

cor al. coverage vas sub=tantially hi aber in11993 at all depths a< south< transition, and aiddle bay regions con pared to 197

levels. The abunrlance of coral in 1983 at 'the ai d bay sta tions had nearly ap-prOached t he hi qhe st levd le in the lagoon reported in the nOr th lagOOn. Tbetransition zone and patticularly tbe south lagoon coral populations did not yetrear e ig a un ancerh the high abundance levels reported sore to the narth. Despite spectaca arircreases in south lagoon coral coaeunities by 1983, overall abundance is stn i llonly a fraction of those of other regions and it aay t.ake decades for the coaaff-niti es to recover coapletely [Fig. 2! . Notable coral recover y on old dredgedsur f aces vas also reported at Station 2 and observed elsevh ere in the south la-goon. In contrast, coral recovery on deteriorated reef surf aces covered vit!fsediaent had not yet begun at one south bay stat»n possibly due to non-suit-abili< y af the substrate for coral settleaent.

tell, <ictd~sha c' ~ . r ' th i o '.tha er ileh cetaispecies and its abundance vas equivalent to all coral species cosbined Table2! . I!oveve r by 1903 0iotygsphaeria shaved draaatiC declines in COVerage at~ ost oay stations vith it.s recent overall le~el being only 29% of its earlierlevel Table 2; F ig. 3! . Spectacular decreases vere reported at all transitionaod aid bay stations vhere Eiglyoslthaeria had earlier erhibited its peak abus-da rice ir, the ba y Fig. 2! . ~Dr~tospa he~id declines in deeper eater appeared tobe I arget than those af shel lov vater although declines vere generally substan-tial at all depths yiqs 2,3; Table 1! llthough ~b~tos~haeria continues tabe nearly absent in the south tay, it shoved a eoderate increase in sost northbav stations Table 1,2; rig. 2! in 1963 cospared to 1971 levelsInterestingly, the alga sha~ed a draaatic inCrease at Station 13, the AOrth baystat ion closest to the ahuiaanu Sevage outfal.l vhich ras still discharging sev-a9e at the tive of the 1983 survey.

ftraakatic increaees in the abundance and diStribution Of indiVidual speCies Ofcora!a vere also reported Table 2! . The aost abundant species ~or l~tsCoaitgfgye near ly doubled its abundance Since fgy 1 and reCOrded aa !or inCreaeeSat all stations ezcept tvo lo and f15: Table 2! . la~res vas coapletely absentat statiori 1 in fgy'I but present in 1983. The grovth of porit<.s vas so iapres-sive at soae stations �0-�! that tie platforas vere heavily colonized andnear ly hidden b! coral gtovth. up to 50 ce of grovth vas reported over theplat ora at Station 12 and the olatfOrn cotILd no<. be relOCat ed at all at Station6 h:: *r i ca th -.:tc c. ted t h ed it. L k 's it~oct ra eer-fucg.a nearly dani led its ovetall abundance in tbe lagoon shaving increases at12 nl. +ha 1 S station Table 2! . The species vas absent f ron Stations 2 andin 1'171 but present at those stations in 1 9R3.

v ral of the les . coaaon pe< ies re por ted at the st ations sl oved reaarkaoleincr ea.-< s in thea r distribution and abundance Table 2! . hlthoug h goci~l~oo~rdaaicorn~ van ab. < nt !roe all south bav and one transition zone stations init vae present at a ll 1 s stat~onS and Shoved increaSed a! fu stationS. isi ef lar but nore iapressive tt end vas recorded far ~C+astrea oce~ll 'na Tablenuring the fey 1 ..urvey, the speCies vas reported at only 3 statianS � sid-dle bay and one north bay station! . Ity 1983 Cyphastrea vas reported at 12 ofthe 1'9 lageon stat ions and shou<'.d increased abundaaCe at all of these stations.I r Iidl'9 at nen! only at tvo soot h bay and ooe traaaition zone st ation. !A can-trast, <.hc sof t coral goantbus !!acjgca vas observed to be present at all south!.ay, on< traosit ion and one aid bay station in 1971 but vas absent at all sta-tionn lfuring t ha 19R3 survey. The other rarer specxeS Of Cur ale vere not en--oust ~ 9 ed n'.tr n enough during the transeCt surveys to den Onst rate any trendsov< ~ 9 iee.

frise u, .ion

vnd study of "..attn et al. !981! is the only other to docusent ecologicalcord d t tons in Raneohe both hef are and after the CeSSatian of Sevage diSChargeStn thea -ou<h laqoon. They vieved the bay as a Single nutrient SubSidiZed eca--y '. "t rlr i or to ',.i ver Sior, and pl aced e Vkphauis On SOnitOring biOeaSS and prOduC-tiv t v of aalor croani a groups rather t.ban the progress of j.ndividual species.Ttr v irtcuet r ted a rapid decline in phytoplankton, zoaplankton and benthos pri-ratil, susrensi or feeders! i n the south bay and a oecline in f!icl~os ha~eja inh p ai L! 3 I e ba y fc' In vi rkq diversi or. and t he eliaanation of se vage nutrient subsi-st ver t hale ~ - 't he st ody vas not abye to dacnaent recovery by reef corals.Hov -v--., Frock an 1 af t h � gn 3! <loni<ared cryptofauna1 coen unities on reefs be-fore a:id after diversion and noted increased bioaass of hard-bat toned cryptofau-ra1 ~crt ur it ice duri no r va ge load ing and declines after diversion.

I n contrast, our St udy dOCumented an eS sent ia I ly Opposite pat tei rl arid r a t.e Ofresponse by reef coral comauriities. Coral populations vere at depressed levelsiu the south to middle lagoon before diversion Although respand ing slowly atfirst to the termination of sevage, reef corals are nov dra mat ical ly recoveringtheir abundance< distribution< and diversity throughout all regnons of the la-goon pre viouslv af fectvd. surprisinoly these responses vere not li mi ted merelyta south lagoon corals earlier affected directly by sevage or transition andnidd le bay coral populations, pri.sari ly garites a nd ~mont i era, the t vere previ-ously af fected by aassive grovths of Dict os haeria. Even some less cosmos cor-al g. ' i clsdi g rggdrrs ra hd c~htr sh d sshst ti Iabundance and distribution throughout. t.he entire la goon vith the six year peri adfolloving sewage diversion. Althovgn it is difficult to esplatn hov the terai-nation of sevage discharges could stimulate measurable coral recovery to ro 10Ra to the north, the exhibited patterns of coral recovery cannot be dismissed asa coincidence nor are other explanations for the recovery plausible -.ndeed rfthere is a relationship, then some of the coral species responding to the reduc-tion of sewage stress uay serve as ex remely sensir.ive indicators of sewage pol-lution.

Our 19B3 documentation of Dictfospbaeria declrnes are corsistent with theearlier measurements of Saith et. al �9B1! . ue determined that the aloae de-clined to one fifth of its prediversion abundance overall and noted major popu-lation coll.apses at seven out of the eight. stations in the transition and middlela go on z egi on s. The rapid decline following di version suggests thatDict~o~sha~ia was previously stimulated by nutrient subsidies from the sewage.Furthermore, Smith et. al { 19B 1! explained the nor them shi f t o Dict vosldhae~ripopulations between 1970-1977 to be t' he conseguences of incroa sad sevaoe dis-charges over the period additionally, the initiation of sevage discharges inthe north ha v froth the Ahuima nu Sewage Treatment plant in 197 0 could explainboth this shift as veil as the moderate increases in Dirtyosnhaeria coverage veobserved at the north bay transects

The continued absence of Dic~to~shaeria from the south lagooa is more diffi-cult to explain. Smith et. al �981! concluded that the algae's grovth vas lim-ited by light due to 1 educed light pe net ration in t he va ter colu un f rom suspend-ed particulates. Although light aust be limiting to some degree, at does notappear to be the sole explanation for its absence in the south lagoon.,"a ragos{1972! repOrted huge populatianS Of the algae in deeper water to 9-11 m! duringthe nid bay and transition xone transect surveys in 1971 and dur ~ ng gualitativeobservations i,n the bay between 1970-1974. Part her nore, he vas constant l y re-moving grovths of the algae from his deeper platforms to prevent the algae fromsmotheti ng the coral transplants. h7ur 1983 surveys shov th at most of theQct~shaeri a decline occorred on !he deeper por tion o f the re. f slope seeFigs- 2,3! during a time interval when light penetration vas expected tocrease. Clearly light was not i+siting the algae on ni d lagoon reef slopes andOther factors aay be responsible f or its contin ued absence in t.he south lagoon.Substratum instabili.ty on the steeper deteriorated reef slopes see Kinsey,1979! and residual toxicity of the bottom sediments i n the south laooon frompreViauS se wane diScharges may render bot tord habitats there inboSpitable toDzctXo~shaeria colonixation and orovth perhaps the inhibitory factors are srm-ply the same as those which have discouraged devel o peen t of ot her ti pica 1 reefbenthos including corals in the sauth lagoon over many years

Until nov it has been dif f icult to di fferentiate tht net cont ri1'utior. of ear-ly major dredge and fill operations from that af later sevaqe discharaes to thedestruction of reef coral populations in the soutu lagoor.. our 1983 south baytransect results appear to support the hypothesis that savage discharges hadprevented or inhibited the recolonixati on of cor.als on dredged surfaces in thesouth lauoon. our r ecent studies indrcate corals h ave begun to rapidly recolon-

ixe dredged surfaces there nov that sewage has been diverted. Furthermore,Haragos �972, 197m! earlier observed greater coral recovery on dredged surface�outside of the soeth lagoon. The more recent estimates of military dredging andfilling see Devaney et al., 1976! indicate that 280 acres of reefs vere f illedat HOkapu and Over 15d 000,00D Cunio yardS of reef material vas dredged betveen1936 and 19rrS ~ most of this in the south lagoon. Dredged ma terials not used forfill vere discharged into the lagoon adjacent to the reefs. Although the docu-mentation of the ef fects of this activity on coral reef cov mani ties vas rot. ac-complished at the tiae, it is certain that dredging and filling had a cata-strophic i epact on south lagoon corals. However, Raragos �972! also indicatedth t tr ssl hted corals, p ' ill ~orite d ~Plls *ta o . d d g

poor ly an tbe south laooon during the period of act.ive sevage discharge. Hence,coral populations in the soutt lagoon may have been destined f or ma jor damageaad destructior by sevage stress, even in the absence of the earlier aassivedredge and fill operations.

In rrtrospect, the earlier decision to eliminate sevage discharges from thesouth lagoon resulted ia major benefit to reef coral populations in Kaneohe la-tyoon. I siarlmr diversion of the ahuiaanu sevage should result in additionalrecovery of reef corals and bettefit to the reef ecosystea i n the bay. Bguallyiaportant is the fact that the diversion of this sevage to a deep ocean outfallof fshore f rom nokapu Peninsula and outside the bay has not resul ted in notice-able adverse impact to the exposed reef communities adjacent ot the outfall. Iofact, reef fish populatioo are much greater in the vicinity of the outfallpl uses.

at first it aay pose as an enigma that the saae amount of sevage that causedcatastrophic damage to corals inside the hay vould have only negligible impactto reefs outside the bay. Tie ocean outfall site is exposed to strong currents,vaves, and vater circulation and va ter residence tiaes measured on the order ofveeks see 5 ~ ith et. al, 1981!. The rapid turnover of vaters ootside the lagoonprevents a buildup of nutrients greater biomass of phytoplankton and the eu-tropt,ic vaters that characterized the south lagoon during the period of sevagedischa roe. In retrospect, the location of the outfalls in the south lagoon vasperhaps the vorst of all alternat.ives and points to the need to site sevage out-falls in veil flushed vatera to avoid adverse ef facts to coral reefs.

Co nc la i one

The results of ecological studies completed af ter diversion of sevage fromthe lagoon of Itaneohe Bay enablm os to reconstruct more accurately tbe recenthistor y of corals and aajor ecosysteas iu the bay. Beef cora ls vhich ooraallyflourish om the reef slopes in the lagoon of It'aneohe Itsy vere subjected todredging and filling and urbar, pollutiOn after 1938. Dredge and fill OperatiOuSvere concentrated in the south lagoon and destroyed or degraded many reefs.Sedimentation from dredged material disposal and vroa land sources led to tvometers of deposition and shoaling of the lagoon floor vhich buried coral habitatat the hase of the reef slopes I freshvater flood in 1965. probably a naturalevent, led to mass mortalrty and injury to shallov rater corals and other reeflife above depths of 2 meters.

Betveati 1950 and 1978, sevage dischar qe into the isolated southern lagoonprovrded nutrients vhich subsidized high levels of plankton biomass in the ea-ters and Iaroe populations of beothic suspensioa feeders in the south oay.Coral coamunrties, dominated by a single species sensitive to sevage gouttescgmnressal ~ dt-.clioed in the south bay or vere displaced by the suspension feed-ers. In the arddle lagoon, the savage nutrients stiau la ted tbe grovth of at:enthic alga, gjc~tosphge~ia cavernosa vticb overgre v and smothered many coralson t he reef slopes. Even some rare species of corals op to 10 Lm avay froa theoutf al ls exhibi ted decreased levels during the period of sevage discharge. In1970 additional sevage began to be discharged into the uorthern lagoon, result-ing in moderatr grovth of Di~ct ~os h~ae ia there and possible displacement of somecoral.

Tr tarnation of tbe southern lagoon savage discharges reversed the earliertr end > ovards eutrophicatron and led to rapid declines of plankton, sooth hay

;p ' f 8 r.".. f yp'. fai . d .'d I ric~to a ~t' p p lat onsrcora is toot longer to respond but are nov beginning to recolonize all previousLydegt alt a lagoon coral habitats. Althouoh coral recovery appears to be nearingcorpl- <ion in the middle lagoon, at least one and tvo decades respectively mayice r ego ~ red f o rc cover y to pea It t.he transition and south l.agoon. Tbe plannedd t rersion of t'h e reeai ning sevage discharges from the north lagoon should accel-

he diversion of tbe sevaoe to a deep open oceaa outf all. outside the bay, ap-parent ly ha rot resol ted in ma jor adverse impacts to coral reef s. vapid flush-ing of ocean vaters prevent, present levels of sevage discharge from leading toeutropl icatiOn and associated adverse effects. In retronpeCt, the preelous din-charge of sevage into 1 he poorly flushed sottth lagoon eagni fied adverse effectson cor als ao l-oth the south and ~ iddle lagoons. Zxperi ence vitb Itaneobe maysu goes ts t hat, in qe nera l, =eva ge out f alla should not be placed ia protectedcoral laoooos and emha pments with restricted vatr r exchange

The response of reef corals to the introduction and then he elimination oflagoon sevage sffluents did not correspond to those of the domina nt ecollogica1components of the vater column, and particularly components directly or indi-rect ly subsidized by the sen a ge nut r ie at s. Cor al reef s ac hie ve t hei r greatestdevelopment in clear, nutrient poor! vaters and are not subsidized by large ex-ternal sources of nutrients. This plus the fact that coral reefs are able togrov and flourish in rare exposed environaents, creating its ovn habitat in theprocess. are tvo principal reasons vhy coral reefs succeed and have a competi-tive edge in some tropical aarine environments othervise unfarorable to majordevelopment by other kinds of ecosy stems. 1n the case of K aneohe Bay, the in-troductior of large ext'ernal sources of sevage nutrient into the reef systea en-abled other communities to compete successf ally against and event uall y displacecorals rn thi respect coral.s serve as useful indicators of serene pollution.and several species studied by us appear to be extremely sensiti ve to pollrtion.

Bystees approaches and aodels primarily emphasizing the transfer of nutrientsand energy through systems nay not be as accurate in predicting the responses ofcoral reef ecosysteas to perturba tions. Is the 1I aneohe Bay era mple has sho wnus, the coral responses to sevage fluxes vere different, slover, less predicta-ble, and more complicated than the gros s responses of the other ra jor ecosystemcomponents. Factors such as suhstrati suitability ecological succession, taro-nomic variability, physiological stress, and competition for space probanly hadmajor influences on the responses of corals.

Ac k nov le dg caen ts

Qe thank the University of Bavaii !nstitute of t1arine Biolooy BIBB! for. in-viting us to participate in the 1983 summer course on coral reefs, inclu'ing theorganizers paul Jokiel ard Bobert Kinzie, !TI. ve also thank porfirio Ilino forassisting in the field surveys. BIBB also prorided boat, logistical, cov puterand typing support. The U. 6. hr ay Cor ps of Engineers, pac ific Ocean Bi ri sion<provided vriting tiae, draftino, and typinq support. The earlier phase o! thestudy vas supported by Bational Oceanographic and Ataosoheric Admini.,trationgrantS tn the UniVerSity of pavaii Sea Grant progran. The 19B3 Summer courseras funded by riniversity of Havaii Sea Grant proqraa:, university of HavaiiFoundation. and the Pdvin Pauley poundation.

literature Cited

Banner, A. R. 1968. 4 fr as heater "kill" on Bava ii an cora 1 reef s. Havaii Inst.Pl ar. Biol. Tech. pep. 15, Univ. of I1a ma i i. 29 pg s.

Banner h. II. 1'974. Kaneohe Bay< Havaii: Urban po I I ation and a coral reef eco-svstem. proc. 2nd 1nt Coral neef Syap: Vol. 2, creat Barr ior BeefCoaait tee, 6risha ne.

Banner A. B. and J. 8 Bailey 1970. The effects o f ur tan pol lut ion uponCoral reef systeat a preliminary report. Bavaii !n t. Sar Biol. Tech. Rep25. Univ. of Havai i< Honolulu. 66 pqs.

BrOCk, R. p. and B. V. Sai!h. 1983. Response of coral re< I cryptof annal COanu-nities to food and space. Cora l Bee s. 1: 179-16 3.

Devaney, n B., ". Eel lr, p. J. lee, and l. s sot teler. 1976. Kareohe:Histor'y of Change 1776-195U! . Bernice p. Bishop susen a, Honolulu. 271 pgs.

Hollett r. J. 1977. Shoalinq of vaneohe Bay, Oahu, Havaii, in the period 1927to 197h, based upon bathymetric, sedinentologica I, and geographical stu dieB. S. thesis, Univ. of Bavaii, Honolulu. 145 pgs.

kinsey, D. Q. 1979. Carbon turnover and accumulat! on by coral reef s. Ph. Othesis, Univ. of Ravaii, Honolulu 284 pgs.

'.1ara go, J. E. 1972. a stud v of the ecology of Havaiian reef COralS. pb. D.thesis, Uo jv. of Havaj i, Hooolula. 290 pgs.

Hara go-,, J. K. 1974. coral transplantation. a sethod to create, preser ve, aod.manage coral reefs. Uoiv. of Hamaii Sea Grant. Pub. UHIHZ-SEAGH hNT LR-74-0330 gqs

Roy, K. J. 1970. change io bathymetric configuration, Kaneohe Bay ~ Oabu,1982-1969. Univ. of Havala, Havaii Inst. Geophys Rep. 70-15. 26 pgs.

Setto, S. V., V. E. Chave, and 0. T. O. Kam. 1973. atlas of Kanenhe Rayreof ecosystem under stress. Univ of Havaii Sea Grant P ub. TR � 72-01. 128pgs.

smith, s. v. ~ H. J. Ki amerer, E. a. Lave, R. E. Brac%, and T. R. Knish. 198!Kaneahe Bay sevage diversion erperisent: perspectives oa ecosystes responsesto notratiooal perturbation Pacif. Sci. 35 Q! t 299-395.

A comparison o coral community structu e on reef lats in Kaneohe hay, navaii

"orf'rio N. hlrKoUnieerSity Of the philippines Harine Sciences CenterUniversity of the philippines 'nilisan!>uezon City, philippines

Abstract

Coral colony size- freguenc; distribution, species composition and tota ' coralcoverage vas esti mated for various reef f 'let. station .. Tl ese data provrde addx-t ional evidence tl.at reef recovery !,ad proct eded rapidly in the five years fol-' ovino termination of sevage discharge ir to san cobe Pay. proximity ro streamsouths is an important factor controlling reef coral conmunity structure, sug-gesting that oc asioral "fresh va ter k. lls" are an important factor xn this re-giov.

Introduction

The characteristics of coral community st;ucture can provxde insights intosome aspects o mast environmental conditions. Loya �972! and i einberg �981!s'u9 gested that size distribution pat terns o t! e cora 1 popo la 1.iona could reflectage and grovty modes. size measurements can be useful in discerning patterrsand overall dynamics of a community. The use of size measurements e I diase-ter or lengt'h! has limitations in determining ab olute aoe due ta coapl' cationssuch as colon!' regeneration fragmentation! and fusi on. The usef ulness o f tvie"size-freguency" approach has been demonstrated by origg and paragon �974! intheir stud} of coral colonization on Havaiian lava f love. Dodge and vaisnvs�977! employed C'Oral age and SiZe ClasS populati On patterns using X-radxologyas a measure of coral responses to sedxsentation and turbi" ity associa ed vi hdredging. Connell's �978! assessment of tropical rain forests and coral reefsalso sl:oved tte importarce of the relative aoe cospositior. of the comiriant spec-ies in "climax" communities so as to in er t!e relevance o. diversit t iri rela-tion to pert.ubations

yicsults of studies on cora reef 8 estructi on and recover y have been revievedbv,lohannes �975!, endean �976! and I'earsor. �981! . In raneohe Bay, rnitialstud'es by Danner �968! docusented some aspects of a coral community "vill"caused by freshvater runoff. his studies vere later extended and relared o ur-ban pollution Farmer and Bailey, 1973: 'panner, 197rr! . nore extensive and de-tailed vork vas later conducted by naragos �972!, vith emphasis or. coral commu-nity structure. Integrated approaches relating to both the structural andfunCt.ional aspects of the !raneohe Bay eCOsystem have been made by Sm.tb �977!and 'Smith et al. �'973 and 1981! . These studies vere spec' fically related tothe effects of increased nutrier t loading nitrogen and phosphorus! accompanyingsevage discharged into the bay. Saith's �977! delineation of the bay irrto the!rorthvest. Central., and southeast sectors is relevant 92o this st udy. sevage vasdiverted from the bay in 1978. Smith et al. �9 81! pornted out that the bentnrccospositional response vas sloe and could not be ascertained at tfat time.Comparison of patterns of coral cosmunitv structure from reef flats t' rougbontthe three regions tooether vith other studies [e. g. Evans et al., thrs volume!vould broaden the data base in assessing t' he impact of these events on coralreefs, It may 'be possil le to deduce staoes of se ial successior. by tal; ng ir toconsideration the t heoretical concepts of Gonnel 1 �o78! and Grig; �963'I-!5aterial and fret!rods

The reef lats sampled vere iv, the saut!ieast, the "crt al and northvestSectorS o f the lagOCn area at 8 aneOhe Bay see pig. 1! . These si tes sere se-l ected so that depth, distance from edge of reef f la., proxxmity to streamsor ths, orientation to tradevind induced vaves vater circulation arrd turbl-lenCe! and ot!er faators vere simzlar for each ser'es o 3 stationsSampling methods

7 vO a m plots vere sampled at each statiOr,. Tl.eSe vere at a dist ric~ Of 38from each other. Each sample vas situated approxinatel! tvo to three m srore-

30'

1 ' r. 1. 1 oont inn map nf samplinr stations at Kaneohe Bar, 'stations 1-3 for the SoutheastVec irn; S; ni in»a 4-6 fc r the Renter Region; and 'Stat irma 7-9 for the 5orthMeetern Regian.i 2 sf t ~ I froai the Il.5. : i <;.S. Chart. 4134 and aerial photographs.

97

Results

and similarities in the coral"Average colo~y size" totalcoral coverage increased froms see Table 1! . Though thehave direct hiolo;ical mean-rara ge d ist ribu t r ona l p atter nS

tterns of differencesreef flats stitdied.of colonies! and tot alZorthvest study statione" may not necessarilycomposit iona 1 and si ze2 to 4].

There vere discernible pacommunity structure of thelive coral cover/ total no.the sriutheast sec 'tor to t.hei de a of "ave=a ge col on y sizing, there vere particularat each station see Figs

Table 1. Data summary for the 9 stations.

Station

5 6 7 8 9

1.12 0.64 014 8 0

pon~ti ora verrucosa89 1 07 3 59 n 17 5 94

142 d'9 33 31 220. 10

28Total Co ve r ca v !I'o. of Colonies

Porit. co~area-aTot.al Cover cm'!;to. o f Colonies

0 0.89 C. 47 7.52 14. 9 19.7 25. 8 40. 0 on. 69 25 12 66 63 50 101 136 81

p pc i ll~oora da mic or nis.otal Cover crt! 0. 11mo. o Color ies 14

'. 98154

6. 3o 6.43 0 19112 95

1. 88IPo

5 06 1.26 9.4369 86 'I 69

C~haSt r ea OCel1 iI, aTotal Cover cm'!IIo. of Colonies

P P 12 0 5o 0 3o 0.12 P 2P 1P5 0,57 0.500 21 52 25 o I 8 49 Io 13

Funoia scutariaTotal over cm'!ItO. of Colonies

0 C 0 C 0 II l.a5 l.s5 0 460 0 0 0 0 0 14 14 5

TOtal CoralTotal Cover cmv!Ito o f Co 1onieas

35. 2 35. 6 5t. 0 85. 7263 293 2. 2 102

4. 69 10. 1 'lb. 3 27. 5297 307 I9' 189

P. 2651

Dict vosohaeria cavernosa0 0 2.12 3.37 1.?S 19 2 3.3, 0.25 7.87rota' Cover cmv!

coralline algaeiotal Cover cm'! 0 0 1.37 0 18.7 16 ? 6 50 16. 0

1. Pl C.B7 C'. 72 . 081.00 1 o2 1.16 C.70

0-98 l. 13 1 021. 13 1. 30 l. 1 I

1. 141, 12

IIC 'I]I ra '

1. 06C. 99

1 67 1. 25 1. II 2l. 66 l. 89 1-65 1- 1 6

1.89 '.63 1.88 1 7Cl. 85 l. 88 2. 16 1. 94

2.372.21

Jcidn I

average Colony areacmz Colony 0 0 1 0.02 0.03 0.08 0- 4 0.14 0.12 C

vord from t] e reef edge The 2 m x ? m sguare plots � mz! vere subdrvzded intomz areaa ar d Ir eaSured uaing a I mz guadr at f rane Strung vxth line z ntO 'ICC

subdivisions each 10 cm x 1C cm! . Subsamples proved to he relatively homoge-nous at. each sl.at,ion ard vere combined. This !ave a sample size of 9 ma perstati on tvo samples of 4 m' located 30 m frot each other! . All ver e n vaterdepths of 1 to 1. 3 m. ln these sample areas, the maximum diameter lengths ofall t'hs scleract ini an coral s prese ot vere measured vit h a xeterstx ok to thenearest ca. The data oathered vere used to plot a size fregxency distr xtutzon

tl.e r.oral species in the area see Figs. 2-4! . Tstiaates of li ve coral coverare based on an assumed circular hot tom cover e timate II 3.14 rz! . RelativebOttOm cOver estimates Of Sub trats types pr esent vere made based on the numberof gride �0 X 10 cm! COvered Vithin the 1 mz 9oadrat. The estimates of coVerand numbers of indxviduals ver e used to calculate haonon-sexier zndex o dxver-sity 8' and equitability

D

V Q

~ J

lh

0C'7

Vlrd

0 VII<O

M 0

a Lt+ N

V g 4v'fL'4 I

N 04-

CVVl

V 0Cv

'i+

Fig. 3. Size- regueacy distritati ops of compels in he Lentzal Secto~4 to 6!.

95

p e rce n tag eI re quenc y

I0

4'

10

10

14 I

'4 ~

II

14

, II

~ 0 41SiZK Diameter in ce

96

0 l'l.l 0' ' ' 0!l,. ~ . 0l -F. II''Jej rv distr�'tutiOn of coralS io the Horth44eSt SeCtor [Sta-

the Southeast sector sta tions 1 to 3! the "sizz" of pt~ct~ora ver~rue sa see fig. 2! vas obviously skewed «ith a maximum in he 0 to 5 cm range .'tostof these colonies seemed to have oziginated from planula settlement with onlyabout 3 to 5% icing derived from fragmentation based on crass morophology and'x r a liogr aph v ar deer ibed by zs guive 1 this volume! . ft n increasing ab unde nce of.

gJJolpora damigoznis in coverage and number of colonies vas observed. ~Patesgpm~r~ssa and ~Cpp tst-ea oceggina vere less common and occupied a smaller por-tion of l.ottom cover. This sector had the highest proportion of sand vit h siltsubstratum coral rubble estimated at approxinbrtely 30 to 50"! .

T' he Central Sector neaZ heeia stations 4 to <! appeared to hav~ seen influ-enced by exposure at extreme lov tide auditor freshvater runoff as evidrnced bydead ur per port ons of coral colonies and presences of terri genous silt. T'h' sregion shows a less skeved SiZe pat td rn porites ~Coe re<sa had the broadestrane, ' llo ed rf f~so~r de~cot l, n ~n' a a~usa nd ~r~hastocellina in order of decreasin" size. Po jtf s co~~re sa occop'ed the greate "tCural cover '1 to 1 8..! iO all the Stations v th an inCrea ing bo t Ox' cove r romStations m tO 6. Poci11O~Ora ~m~COZriS shoved the hi~heSt numoer Of Colamiesin th' s region though its coverage {5 to "u! vas relatively less than P.com~ressa. f. verrucosa had only 3 to 6, cover and c. oc~e~iay had the lowesthottoa cover and number of colonies. This region had a more developed reeframe than the southeast region and the presence of coralline algae ra s more no-+ ic�osa b' e.

The "ozthvest region Stations 7 to u! vas characterized by the increasingbund e of ". rloubrle a d th eppes c of,u dfd a~cuter' . lh. s re ra u

Of g. Compressa tended to be less Skeved tovards the h!Orth with the gzeateStku tosis at t? e northernaost station. P. da~mcornis and v,. verzucosa seemed tobaca dcreaduabtd toad tbothr ~ stat'.lone.~cadrelatively high cover O.S tn 1.0T! as cotapared to the othter sectors. Though~vrucosa vas not present in t.he sample guadrats, a fev colonies were sightedoutside the sample plots. The reef fraaeworl. in this sector vas relatively moredeveloped and co"el line al.gae vere more abundant � to 16T! .

Trodr ' h ee tO f i ve Species of cleZactinian coral vr re obser ve 1 in 1 he reeflat stations. <he lowest num!er of species [3! was in Station 1 and the high-

est vas in Station 6 of the Central sector and Station 1 of the I'orthvestSector. shannon-l-einer ndex !a,'! and eguital ility J u! f or each s tat ion sho~edsome varying patterns. There does not seem to le much drf ference in ave.age H 'cfor the SOutheaSt �. 06 + 0. 07! i ut a sudderh decrease vas seen rn the '.'.or thveStregion � 56 + 0. Q5! even with t»e added presence ~. SCutaria. }! un for oll theregions vere <xuite variable. The sudden decrease in diversity is also seen inthe avera ge eguita bi lity va l ues; vith the la or t hvest regioru ha via g the lowesteguitabilitv dn = 1. 57 o 0, 3R and Jc = 0. 61 z 0. Sg! reflecting the ' ong domi-nance o f p. gifmppuussa in the region.

ftiscussion

C!azacteristics of coral communities are usually studied f". compari by d ffer-enCeS ir. S paCe and/or time. The high variance encO untered ir, sOme coral Zeefmeasurements does not necessarily exclude the presence of oatterns. patternscan be discerr.ed to uggest commonality of their po itions in a certain dimer.-sion..o prioritize the variables in a particular f.amevork is usuallv ehe goalfor the student assesSing community structure. In this stud!' the southeast tonorthwest sampling of some leeward reef t lat: xfd Ya neoho par show pat t -ms vhi c.cou'd he attributed to water circular�'on, substra+e cor post tior� i -oxrnrty tostream mouths, and their stage in serai succession.

're area rear station 1 had no previous signi= icaot coral pop u la tio;t "aragos72! . Smith et el. �981] implied that tl i a sf a vas possr fly under got rt 2 "de-

calcif ' cation" and biorrosion due to sevage and s' ltati on. This a; ea vas ofparticular interest because it shoved distinct signs o. recove v as evider.ced lythe skewed d ir tzi'hut ion and a laz ge nus ber of n ev cc.rais ir the si=e class be-tveen 1 dO 5 cm. This reCruitmpnt Could have occurreo ortly reCertt ly..",oSt oftide nev Coloniea vere leSs than 3 yearS of age oaSed ord polaClteCf,'s �978!grovth data foz !i. verrucgsa and p. damic~onis. Apparently nev recruitmer.t cidnot occur for the first fev years following sewage d' versior. Smith et al. I1981!. his is not surprising because the reef van covered vith organic mud andthe substrate vas unsuitable for cora' set tlemer,t. In time, be muds vere win-nowed rom the flats exposing !;ard substrate which vas later colorized.

97

.'.r hioh recruitment of i. vgr~ucc2sa and P. daarcorniy is coaparab'e to vhatar d raragos �97rr! and Baragos cited by Jokiel et al., 1983! considered

as f Jgitive species. Also this sector is the most protected Bathes 'l968I andthus as,tokiel �97tr! has suggested ". ~VZguCOSi and g. +~ai at'i',s aay be soreadapted for t!is condition. It is also interesting to note that the coapositionof most of thr color ies vere of planular settlement and only 3 to 51L' vere fromf rag ments. T> is seeras to coincide with vhat Jok iel et al. �983! reports; thatasexual re product i,on vas less j sports nt in 'f. vi',~rr gs1i being only 5%! than I .9 j'itat a based or. araft coapatibility. The vide spaces bet veen coral coloniesand sof t sediment.s does not yet seem to limit th e cxpa nsion of tie specres pres-ent. nnly a fev species of fish in lov abundance vere observed only sj,x spec-xes vere re;orted t'y Snith et al., 1981! . This does not represent a high preda-'.rorr p~assure at the present time. predators such as Chae~todo spp. see Cox,this volume! vere not seen in this area though some scarids vere observed.

«all scar ids ir. Havaii normally do not graze live coral.

Or.1' four species of corals vere encountered in the central region, but highHt ar d J' vere obtained due to the ever distribution of the spec'ies. The ab-sence of g. ~3rrig may be the result of sampling liaitations. usually thisspecies is found near the margins of reefs in this region fr aragos, 1972 . Eventhough there is a relatively well developed reef framework, lov tide exposurecompo rnded by fr> sh water runoff and siltation could have limited coral grovth inthis r" n'ral sec tor as Compared to the nOrt!rvest SeCtor Banner 1968, and Bannerand Pai lry 1'9 3! .

Th+ sor't h vt st sect or vh ich is supposedly the least inf lnenced by disturbanceshad the invest rti varsity, .he laraest colonies, and the highest coral cover re-flee ting a lorg period of undisturbed grovth. p. cr2m~es~s seems to outcoapete

ve~iLcoye and '. d amicornis. This could be related to the relative coapeti-tiv. ability of q. gyg2~egsd to this environment compared to g. dear~corn's vhichis more adapted to ex;loit environaerts that are disturbed occasionally. Thehigher fregoencv of g. scutaria in the samples suggests that this area has astrongor vater movement to vhich this species is sore adapted Jokie1 and"ovdin, 1976! .

are vary no decrees of differences in environraental regimesure to pollution, ronofi, siltation, etc.! in the three1 patterns siailar to those suggested by Grigg �983! and

'nferred. That is, that degrees of disturbances at tribut-ff ard siltation, complicated by sevage could be responsi-

di.versity of theme reefS. Io the mOSt nOrthern regianf conpetetive exclusion ty ". c~m~esa «hich are coapara-C~no~atrd forests of Conn ell �978! and Grigg's �983!

nce in Hawaii.

Acknowledges nts

vo ld li' e to t!danr r and a'vice 'npaul Po lthu~, and peg

-ummer cour er o. t wr nor ai ie.

itrrature Cited

< ~:, r, ',. 1 ~ ~ n. r, f rash mater "kill" on the coral reefs of Bavaii. Ravaxi. r.' '.- Gf n:r.. iol. Tech. Pep. 15: 1- 9.

h'aneoh Bal, Havaiir urban pollutiOn and a COral reef eC--.; -' r .; ~roc..'nd ln .. Coral Fee f symposium, Brisbane 2: 665-702

and,t. 8 Ba el. 1473Ba le.. 1 7 ..he ef acts of urban pollution upon aoct a1 rrr f ocosy ter . Bava' i Inst. of nar. Biol. I'ech ~ Rep. 25: 1-66.

: =. ~ her . -.. 3 r n< B. A dcscriptrve study of physical oceanography of Kaneohe! u, .'~av, i-. Ra vari In t. of sar. Biol Tech. Pep 1aa 7-353.

9S

Ever thoug.' there l. v. previous expossectors, successionaCouncil �978! can berd to frvshvater runo! le for the 'evel oft here are evidences obio to the "climax"

itr-" ggl atf. do nina

arik 1'au 1 Jokiel and Jim .'.a ragos for the'r' challenge, gui-the process of this study. I aa rndebted to Chris Evans,Bartlett vho assisted in f ield vork. and to all my frierdsof 1983 at Coconut Island vho have made my stay in Hawaii

Connell, J. 1978. Diversity in '.ropica I rain forests and c~r al reefs. 'cience1o9: 1302-1310.

Dev anei, D., I'. Eel ley, P. J. Lee, and L. S. vIot tele= . 197 6. Kane ohe: a hi s-tory of change �770-1'930! Bishop,.use un, ."onolulu. 271 pgs.

Dodge, P. '. and J. R. Vais. ys. 1977. Coral population and grovth pattezrs:responses to se 3inentation ann turbidity associa ted vi th dredging J .",ar.Res 1m u!:715-730

Endean, R. 1976. Destruction and recover v of coral reef coaniunitifs Pages215-254 ' n Biology and Geology of coral Reef s, Vol. I=I, Biol 2. 0, A.,1o..eS and m. Endean, eds. AcadenrC P rr ss, London.

Grigg, P. 19o3. Coaaunity stzuctuze, succession arid development of coral reefsin !Iauaii. sar ECol. PrOg. Ser. 11t 1-1u.

Griqu, P. and J. 8aragoS. 197II. ReColonisation o heraatypiC Cora ls On Sub-merceR lava flovs in Hawaii. Ecology 55:387-395-

Johannes, Fi ". 1975. Pollution and de gradation of coral reef coaiaunities.Pages 13-51 in Tropical Ear inc Pollution. E. J. Fergusson-Rood and R E.Jo'hanres, ede, Else VieZ OCeanOgraphy Seri< s ',I O. 12. E l aevi er Scxenti SicPubl., Nev Yo-k.

Jok in l, P. 'L. 19'7 8. Ef fects o f' v a ter motion on ree. cora ls J. Exp.Biol. Ecol. 35: 87-97.

1Okiel, P. L. and H, P. COadin. 1<76. PydromechaniCal adaptations in the SOli-tari free living coral Fungia ac~aria 9 tare Lordon 202.212-213.

Jokiel, P. L., I'. H. Hildemann< aud C. 8 igger. 1983. Cloual popnlat' onstructure of two sympatzic species of the reef coral 1g~t~ry. Bull. IIar.Sci. 33 �! t 181-187.

Loya, E. 1 "72. commuriity structure and species div.rsit v of hermatyprc coralsat v'ilat, Fed Sea. IIar Biol. 13;100-173.

Aaraaos, J. 1972. A study of Havaiian reef cora ls. pb, D. Dissertation, Dept.of Ciceanography, University of naeaii. 190 pcs.

Pearson F. ".. 1981. Recove y and recoloniratron o. coral reef s. sar Ecol.Prog. Ser. II. 105-'I 22.

polachecb, T. 1978. The population biologv of f our '.Ianna iran corals. 8. 5Thesis, Dept. of Zoology, 92lriiversity of !Iesaii. 15'1 pgs.

Smith, S. V. 1977 Kaneohe Bay: a preliminary report on the responses of acoral reef estuarv ecosystem to relaxation of seva9e stress. Proc. 3rd Lnt.Coral Reef syap., Miami 2:577-58u.

Smith v 5. V., K. F.. Chave and D. T. 0. K aa. 197 i. Atlas of Kaneohe Day = A ReefEcosystea Under 5tress- HEIDI-SEAGRAHT-. R-72-01. sea G" ant. Col.lege program,Univ. of Baca i i. 12n pgs

99

-"mi ', s. V., s. n. xismerer B. A. ha ms, B. 9. Brock . x. >alsh. 1991.,'anemhe sewage diversion experiment: perspect'ves on ecosystem responses tonutritional pert urhatians. Pac. Bc' . 35:270-395.

i'eon,'erg, B. 1981. z comparison of ree~ survey methods: Bij iragen totRcershunde 5 [2!: 199-216

100

Srnvth o the Pee Cor el por ries COSgressa on the Coconut I Sian d Re f, haneohev

pay ~

Paul h. Jox ie 1Havaaz T nstitute of .".a rane pioloc�P, 0, pox 1346, Eaneohe, BaVaii o670a USA

Abstract

pirect measurement of grovth in the reef co=el pot iten =ospressa over a fourveer orovth inter va 1. between 1979 and 1 9R3 yielf ed the f ollovinu value f orbranCh elongation: 1-5 Ca yr � ' in a shallov backvater lagoon, 2.u cm yr-i on tbevindvard reef crest, 3.5 cm vr � ~ at a depth of 3 m on the reef slope and 2 8 cm

a depth of 7 m. his translates into a maxr mum theoret zeal skeletalcarbonate aCCret' On rate Of 25 kg m v yr-' fOr the Skelr tal reef fr arne vork.y1ovs ver, th is value ca n be doubled i f me a d d i of r ll ing o f t h e r e ef mar gr n f ras e-vork vith sediment derived from the adjacent reef flat and other locat ron s.baring thi.s study Hurricane Iva damaged sections of the reef and revealed thepresence of a veakened layer of reef framevork that vas lard dovn during theperiod of sevage pollution circa 1960 to 1978.

untrod uctxon

The "finger cora 1" porites conpressa is the doainant reef- f orminq oor al i nKaneehe may, Oahu, Haeaii. This species accounts for mOSt of the living coralvithin t he bay. p. co~mressa often covers f rot 80% to 100% of the substrate onthe reef margins in the bay soithus, 1985; Bolthus et al.. 1985; Evans et al.,1985! . hiving tissue covers only the outermost fev ca of the branches. Thesebranches, hovever ~ extend vit hoot interruptioa for many eeters into the reef.Colony cross sections can be inspected in areas of tbe reef that have beendredged or vhere slumping occurs on reef margins.

The branched framevork is secondarily infilled vitn sedieents as the coralgrovs evav from the Substratum Hears the Skeletal net vOrk of t he Coral formS anatural sediaent trap or "sink". Tn deeper environments, the coral can survivesmothering only if xts grovth rate eguals or exceeds rate of sediment depositionbetve n the branches. fieasure men 1. of the rate of linear gr'ovth in this specresprovides an estimate of the theoretical maxi ~ um rate of reef fraaevork grovth.Such data on grovth rate of this series vas lacking. l lao, there vas littleinforaatron on the rate of recovery of reefs that vere damaged by sevage drs-charge that occurred in the bay betveen circa 1960 and 1978. Therefore, tarsstudv vas unfer taken to measure arovth rate in various environments on Coconutys land roef f ol lovrng the term matron of sevage discharoe into Va neohe Bay rn1979.

materials and methods

Grovth Frneriment

gl l mr ge[2mdranet er!o1'onyo f nor r tesc os or os-as a scare f u1 1 ydivrde d rut o smaller colonies of 19 cm diameter. These vere all oved to heal orsevera 1 veeks all a~in and then car ef ullr moved into lar oe cont i neo us floe labo-«tory aguaria for starning vith Alizarin Bed-9 tl.asserts, 1974! . On 20 ray1979 ' he flov of eater vas stopped aud Alrzar n stain vas added an suffrcrertguantr t y to bring t he concentration to 20 ppm. Ske letal sta ining vas allotted tocontinue from 0800 h to 1700 h under full sunlight in heavily aerated vater.ylOV Of «ater vaS then res~med. On tbe folloving day, the coraJS vere care fullvtransported in pails of eater onto the reef. Eight colonies vere attached vi! hvrnyl-covered vire to vinvl-coated steel mesh platfo res at each of our sta-tions.

~ b sta tion" vere located along a horizontal tra nsect -' unnrng directly nortr.�froa the Havaii institute of marine Hrology through several reef zones on

tho vindvard Coconut island reef tyig. 1!. station 1 vas located on a coralthe sheltered charnel adjacert to tbe laboratory. Depth of this sr eb 1 n V. St t n 2 vas located 100 m sravard an the outer viniv<rq

'- lat margin at a point that nearly uncover ~ at exv, at extreme lov trde. stat>onat a deoth of 3 m rn an area of hroh coral cover.

located on the reef f ace a

101

June 1gn3 the frames vere removed and brought oa k into the Laboratory,~ >! a ls vere dried and cut longitudinally with a diamond sav. The original

t d " air band as well as the nev vlite carbonate material that had been depos-itr ', t n t he sucoeeding u yeacs. has evident.

'.i. i ar ht'anch extenrion, branch thickness and branch s pacina distance be-tv~r n cr nters of ad!acr nt branches! was measured. These data were used to psti.-mati the maximus potential mass accretion and amount o open space created be-tver n t.he branches. The open space bet veen branches is a site for possiblese]t m nt accumulation. also, several branches vere imbedded in polyester resinand cut i.nto 1Ongitudioal thin sectiOno. Theee seCtions vere analyZed using x-raaiouraphic technique knutson et al, 19 y2!, in order to determine growth in-crements and annual changes in branch diameter.

Hiring the summer of 1983 visual surveys vere sade around "ocoout Island andthroughout the bay. Daaage caused by Hurricane I va shi ted and split large col-onies, This alloved eraaination of old skeleton laid dovo durisg the period ofertersive pollution in the bay circa early 1960's to 1978!.Result s

I'hvsi eel and 3 iological conditions at the various grovth sites during theper ~ ol of grovth vera quite dissimilar. Station 1 vas characterized by re-stricted circulation, high nutrient loading from land ronof f, large amounts offine sediment, high t cape ra t ore f 1 uct nations aad occasiona 1 lov sa linity stre ssfree runoff during star ~ conditions. Station 1 vas vithin 20 a of a stand ofaanuroves and vas also influenced by high nutrient effluent froat an experiaentalaquaculture opera!ion. Corals vere continually subjected to partial overgrovthby t ho algae Dictloslhafrl'd cavyrnosa and experiencer} ad verse gr ovth condi tious.stat ton 2 vas located on the vindvard margin af the coconut Island reef flat andnearly uncovered at lov tide. This shallov site vas sub jected to hea~y vave ac-tion luring high vinds and lov salinity during heavy rains. Solar radiation vnsvery i nt ense. the station experienced rapid teaperat ore fluctuations due to theinfluence of vater f loving off the reef flat at falling tide. Station 3 vas lo-cated at a depth nf 3 m and therefore vas not iofluenced by surface fluctuati«sin vater quality. Solar radiation vas not as severe. Station u vas at a depthof 7 m near the l over limit of coral grovth. Little vater motion occurs at. thisdc t th .n vaneohe I'av so the environs nt is characterized by extreaely heavy dep-osit ion of f'ine "ed iments. irradiance vas rela t i ve ly lov. Growth rates andrelet t ve trtadiance are reported in Table l.

~ ' v mount of open SpaCe vi hin tho Ske let al fraeevnrk hetv can the bra uCheScolony va~ cu ! cu la t ed. an equila'terat triangle v it h each leg being equalvr mr, t t anch '. pacir 9 vns drawn. Circles having a radius of half the mean0: u sot or ver< drawn with centets at each vertex of the t r inn gle. lt wasra mr Le neomet r ical crt reise to calculate the percentage of t..e triangleivr t ed by -I.clat al framework and the nercentage that is open space betweeniiirhes Table 2I . skeletal carbonate mass accretion rat e vas calculatedsean density of 1,5 o ce ' Jokiel and covdiu, 1976I .

1in 1 nlhr oth 'i Iali I«' I tIsint

I'i a nr h t t t cyme '. ts inversely relate d to depth, but branch spacing is direct.-ly 1 I itr I to drptl Tello 2I Tr other vo ds, branches become thinner aod morevt 1 1 v ra ei vitb increasing depth. Linear growth vas lowest at the t'eef fla't.t ii . ir, vi th maxi x,ua growth rate at 3 a, but decreasing at 7 m. as a result ~;Xi l ~ il .arne rate accretiOn rate shOvs a maximus �u kg m-a yr � t! at 3 m butb if ti.at value at 7 a. ttovever, rate of creation of space between thent a ..r I,<; ~;. a t out the saae at both dept hs Table 2I .

' ~ x-r au ' nor apus r vvealed a annular cycles of branch diameter betweent..I 1'' 1 ! 'Q 3! . lt appea.s that brancheS becOae very thin in 'lec.-Jan. wheni ~ ape rat uro are at the annual ainisiia. Sub-annual Imonthlyylv, b rt. F run~hxr l occurs at the onset of spring grovth.Obs~rvstrotts

nf r el lot i oh

1U2

st. t ~;. h vas located a< 7 m vhich appeared to be the 1 over li mit of coralsr nut h c n the reef face. The stations vere visually sonitored over a period of4 y

Fig. 1. pap o the toconnt Island Beef shoving Sta tions 1-4, direction o «andsgenerated bI Hnrr icane Iva and areas «here porites co~ares-a colon>en «ere di -..�lodaed and transported dovnslope as a result of the .,tora.

103

reef co. d 1 ! 1. Lrr». f. 9 rO vtn O1 la! ref f J n var rouS envir«i! sf; ve r rradrariCe va s calPd Y Ir 'r r f,f're! n,.! foll Of ieV

r oFcuiaqf

} ehn Annual Relatr VeGrovth Irradiance

cm yr- f! t! Surface!

ff V-an T OtalGrovth

fch! >S. 0- r

jai.on loratron Depth m!

'I 50.5 5.9t 2.4 16 6087

protectedchannel

500 5 9 7t � 1867

1' r n d v a r d& eef Cr est

3.53.0 13.6tt.e 3242

!'indvardmeef Slope

vhrdSlope

i!rod 7,0 11.212.1 52 2.8 13

R � v ss � t i

Their '. Grovth and marreue accretron rates of skeleta1 frasevork arid acCueu-lated se lrmd nt vrthzn fraeevork Of reef coral !I~orit coml!r~Sa on the «indvardCoconuf Is.and Rfeef. Value for skeletal densrty of cora1 = 1.5 g ce-a, densityof sei jment = 1.3 q ca-s.

De pth s!3 00.5 7.0

Prhf«'h f!jhmf''ter CIII! ZS,D,era!I-I I.pacrnq cv! i: . O. '.1 .'I .I!li I rii f vef n cr'anch cel tf r -!

1. 2i0 - 41. 7t0. 3

1.! t0.21 .7t 0.4

'1. 1t0. 41.9+0. 6

I i nv I f,f I ' f' 1 rI '1 r a lf Ii V o 1 }I r!

..i'd f: in fi u '.1 qrqvf h Cm!55 72

3 5}i ! f -ii ~d!bonatv arf rctron rate

ii; m-v vr-I! 1912

] f 1 i il addvd }»f vi f n hrhfIChe« II I V ' Vr I! 11,000 19,300 20,200

tron pntvn trhiI!!

e yr-} 1.1

141.925

2.026

cr f' f. r o li r h t cort p! us sedimf nt

38

104

.14 ' f a :.o'I! jcf Ii ' S {C

!',1 m

1 I' 'II" if f I' ll]i ~ d !

! f-V

r Idir avt.v} Z-I!

hesr nate

dr

ts overfrom 1

scharge

ral soritesthe 4 yearight extinct Ssi h et al

TotalSortalrt y

q!

con!!r essa onqr out h per rodron data tak

198 1!

t he Coconut�979-1963!

en rr! Yaneohe

LEGENO

solid skeleton

1983 b or ed s ke I e ton

Dict yospItaer ia

VnCOn Solidated

mote rial

]963

Fig. 2. sketches of colonies of sorites c~om ressa living along the reef sar Sinsof COconut island. 11iagraas represent condition of Cora ls in pre-po 'lutior.period �963!, during tiae of pollution �9'73! and during recovery �'3831

-12

1983�<~h

-10

1982 � '

1981

4

1980

1979

stain li-0c ttt

!. 06

t-radtogra! ! o. a coral branch fror a colony used zn the gcovth study� a 'e~ t!i! . Location oy tna original alitarin stain line '!979'! and subsequentai r ui:: chanoes !9RO- 'I983! f or oeely deposited skeleton are shovo.

Fig. n. photoqraph of couals oveztur ned by Hurt icane 1va. Original centralstalk of the coral is badly weakened by boring organisms a nd covered wrt b en-cznsting barnacles and bivalves. such massive enc:ustation and borang of ske3.e-tal material ceased upon terminatiou of sewage dischauge in 1979. ante thatcoral branches have begun to grow upward 8}ong the edges in the 6 mont! s betweenthe time. when the storm passed and the time thai the photogtaph was taken. Thecoral head is appzoximatelv 1 a in diameter.

107

ln f l u ~ b~f ref

Un9 'I var Iof terla r.dg,a!urrrpaOSIiir nr rver

'ovtaber 23, 1982< Burricane Iwa soved across the verr. art-hipelago along a northeasterly path and passed

Th ~ island of oahu caught the eastern edge of the. a 1 u, vinds case froa ti e southwest as the hurri

l 1 rrhi f ted tthe e t he soutane,t as the storm incre s da e inpreva iraq diriiction Of the Trade rrrnds.-'' ,ui a t 1 r, nrtrma l ly ] rote c ted ale~ward" side of de ho.

v I i iy avarreil pfof far arid Tribhle, in pres ! .

stern portion of theclose to the isla nd

storm. On th is-a ne approached, but

int ei sity. Tham isves generated by the

Reefs in this area

' I< i I .I r h urine«rr< na ocCirr: r 1 in KaneOhe Baeo e Bay. The bay is prot ec! ed f rorrr large. vi ny a arun oerrier reef. Reefs within thacr tv c ~ vi ri vavi st ir e bay normally do not re-ve vi st re:;, the r,or th~ est marry t ~ pr r vsilinq VE Trade gine c f reefs r ecei ve moderate wave

ar i ii9, v irrotr ctr rl arid cal ~ . Tl er de rrinds, but the baCk-Side Of tbes e~ .. e norma 1 l v pro tee t ed s out h ea st. ed ge ofpoef reCeiVed t he Orurrt of"irr."ir.i ',r a, with Vind .;- edS r I o the storm vave energy induCed ayvi n speeds gustinq up to over 80 ailes per hour F . 1ig. !.' or. irir ' ! 1 inr! I r~' va "t rut It at lOv '. ide

ov . ide The t ee. crest, doai na ted bo I it a trrrl~25 COmn~ Ssa vas i yo I,, f s OVereu with leSS thari 0.5 S Of~ r I V,t ruck, Behind t,he reef1r!ri a to the island shore.ee crest is a broad sand- r eef fl ty aore. Seaward Of t.he reef crest the

y 60 1' to the flat lagoon floor de thangle approxima'el 60o ep! 'fe I - "t F' - 1 eer y a of the la ge coloniesiga. and V! . Beerl all

e eton located 10 c0 a roa the branch tips. Theer. -. ov tne period of ol lotp u ion as described above.a �. Colonies up to

I; iru il est anat t s th the reef slope intoa. am avera e of abfr >t tr.< ..hallo 9 .. out 2 cubic aeters of'a II ~organ~ and reef flat shif ta'd dov aslope alon 9.- ~ ., c .'a vs rc au ved arid +ranece o. 1 ka. kn estimated t t lo a ofir .ransportcd iota deeper eater b ~h

y

lDB

War ~ .. it anSparenCy vaS ver y hiqh pr iar tO tlie onset Of See sac di =Charge.vrri 1 lit 7 was oeneraily 1 0 to 15 s. The steeply-slopirg reef faces t ha t fringeCacorrr island vere Covered vith healthy standS of the reef Coral Zo~r"Brfpi~.".;0 to depths of 10 m s. d. Tovnsley, . Rukerarr, 'e van lreukel em, per-sonal nvaunrcetionl . skeletal material laid dovn in this period can still beoaservrd to be relatively free of boring and encrrstinq organisms Fig. 2!, asshow r rv retrosoective analysis of freshly spli t colonies exposed by Hurricane

rr iting the survey of 'I943, the Old Skeletal aeter ial im the nra ncheS aOre' I ar at ce frow tl e grOving tips vaa found to be free from boring and encruSta-t tonn- aiiterial between 10 ce and 5fr ce period of poilu tion! was severelybor«. srverly bored material vas too crumbly to section with a rock eaw, evenvhorr the branch vas i a bedded in polyeeter resin. Comparative Saaplea taken rO ~unool; Ited reefs in the northr rn portiori of Kaneohe Bay did not show the boredarrd enCrusted band between 10 and 50 Cm f'roa the groving tipS. Zf feCtS of pol-lutior in the bay have been previously docuaented e.g. Banner and Bailey, 1970;varaq I, 1972; Banner, 197m; saith et el., 1981! . Eutrophication of K'aneohe Baybetveon 1960 and 1979 had a severe impact on the I'~r' t~s ~o~s~s community.rhytoplankton and benthic plants vere sti ~ ulated by the higher availability ofnutrient n. Increased parti.culate concentration in tbe water decreased lightpenetr atiori. papid growth by the green "babble algae" lrict~os Qae~ ~~vrnoslchoked out corals ]Fig. 2!, although a fev coral "aubbins" survived in aost are-as. 'r genic material accumulated and sediments became anoxic Guinther andteart!vt t, 1 985} . The increaSe in Suspended particulate orgaaiC mat ter led to aqr at increase in standing crop of filter feeders. Barnacles, oysters,bryo.oanr, vermetids, serpulid vorss and other encrusting organisms guickly col-or ized a 1 1 a veils hie hard substratum. Even the bent hic alga pi~et sfrha~eri~cfggraosd vaa extensively colonized by filter-feeding ePifaunal coaaunities-necruite~nt of nev coral colonies vas seldom observed during the period of pol-lutior, hut bas becoae extresely common since pollution abatesent. Boring aol-lusco..ronges and polycheetes riddled and veakened coral skeletons. The colo-nies b»c i ~ r. so sof t that ve could not walk or stand on coral beads withoutha vr na . hem Shatter ender our veight. hf ter polluti.On abatement, the SarViviagbrar ches rapidly grev into nev colonies with hard slreletons that cao support thev.igbt o'. a ean <Fig- 21-

months af ter the storm, it vas apparent that abundant coral fragaen'tsbroken hy the storm vere regene ating and groving into oev colonies [e. gFsguivel, 1985!. many of the large overturned beads had reoriented the growthaxis of the branches in an upvard direction Fig. u! . Different "morphs" ofthis s pecies can he identif ied by color and branc! si=e f!unter and Kehoe,>985!. This allovs one to recognize material broken froa parent colonies.ragaentS Of readily identifiable morphs vere Strewn as muCh as 20m ro 30a dovn-

stream of tbe parent colony during the store. Storm waves vashed the finesediments from the outer reef flat, thereby creating a suitable hard substratethat promoted fragment survival and cementation. Subseguent regrowth and fusionof cloned colonies derived from fragments led to a fairly rapid recoverv of are-as damaged during the storm. storm daaage vas no longer obvious hy 1985.

Discussion

Branched corals such as norjtys ~mressa adapt to life in dee per vat er bydecreasing their branch diameter and increasing the space betveen branches.such morphological changes can be vieved as as an adaptation to reduction inlight, reductioa in vater motion and higher sediment accusulatioo rate- coralsgroving in deeper vater receive less light. Increased branch spacing allovsmore of the limited available light to penetrate between the branches of thecolony and reach the zooxanthellae located in tissues on the sides of thebranches. Rater motion is generally reduced in deeper vater. I'ider spacing ofbranches allovs greater circulation of water through the colony. Thickening ofthe branches in shallov water is advantageous because greater structuralstrength is required in turbulent, shallo» conditions. Thus< morphologicalchanges that are beneficial as hydromechanical and pbotobiological adaptationsia the environment can also be viewed as an adaptation to accoamoda te the highersediment accumulation rates experienced in areas vhere vater motion and lightare lov.

The estimated aaximua calcification rate of 2u kg m-x yr-~ for solid standsof Porites is in agreement with mariana accretion rate reported for others e g.Chave et el., 1972! . Infilling of the spaces betveen the br ancbes vitb sedimentderived from the reef flat could increase local maximom potential accretion rateto 50 kg m-x yr->. This value is an order of aagnitude larger than the meanreef flat calcification values previously reported, and cer tainly vill only ap-ply to a narrov zone at the edge of the reef. The edge zone is, hovever, thearea t hat is growing o utvsrd, so it is o f great interest. in tares o f unde> stand-ing maximum horizontal extension of patch reefs in Raneohe Bay. Patch reefs andfringinr, reef aargins in Kaneohe Bay often shov 80% to 100% coverage ty thisspeci s 1e. g. Holth us, 1985!

Some of the patch reefs at the north end of the tay do not eaeroe at ex'renelov tide and ar e completely covered by this species Dunter and Vehoe. 1985! .such reefs probably are calcifying near the upper end of the range �0 to 15 kg

yr '! and probably can continue an averaoe upvard grovtn of at least > cmyr-'. Again, this value applies to localized areas and disregards the fact tnatsome of the material is eroded and transported into deep water by stormsFurthermore, this value applies orly to reefs that are hiohly protected fromocean swell and large storm vaves. Regeneration of coral populations in KaneoheBay since the termination of sevage discharge in 1978-79 has been dramatic e. g.Evans et al., 1985: Holthus et al., 1985!, out a. record of the period exists inthe reef fraaevork. This strata of weakened and encrusted skeletal material israoidly being irfilled and overgrovn by stroog nev skeletal framevork. 1ateriallaid dovn durirg the period of pollution vill be increasingly difficult to de-tect i n f ut ure yea rs.

literature Cited

Banner, A. H. 197a. Raneohe Bay, Hawaii: urban pollution and a Coral reefecosystea. proc. Second lnt. coral Beef Symp. 2:685-792.

Banner, a ~ H. and .1. B. Bailey. 1970. The effects of urban pollution upnr. acora 1 reef system< a preliminary report. Hawaii Inst. sar. Biol, Tech.Rept. 25:1-66.

109

Bsluiv I, l. p. 1985. Direct retrospective analysis of the reef coral +g~tesColnre=bd: evrdenci Of Sexual vs. asexual origrns of reef coral populations.I 8: Coral Beef population pi o logy. ed. by p. L. Jokiel, p.. H. Richmond and1'. x. Rogers. Hawaii inst. Rar. Biol. Tech. Rept ~ Ho. 37.

. van s, C. K., J. F. Haragos, and ". F. IIOlthnS. 1985. Reef Cora lS in I aneObeRnv. 81X years befOre and af ter ternination Of sewage diSChargeS Oebu,Ilnvnir hrchipelaqa! . In: Coral Beef Papulatran BiolOgy. ed by P. I,.Joki 1, R. II. Fichsond and F. k. Rogers. 'Havair inst. I1ar. Biol. Tech. Rept.no. 3 7.

r.ninther, R. B. and H. l. Bartlett. 1985, h nOte on COaparatiwe guadratSad plinq fOr the in faunal and epi faunal invertebrates, 1968-71 and 1983, onthe Coconut Island reef flat, Kaneohe Bay, Dahn, flavaii. IR: Coral Reefpopulation Biology. ed. by P. L. Jokiel, R. R. prch aond and R- l Rogers.II avai t inst. Har ~ Brol. Tech. Rept. RO. 37.

Holt hvs ~ P F. 1985. Coral reef coaennities of Ka neohe Bay, Hawaiir anoverview. jul coral Reef Population Biology. ed hy P. L. Jokiel, P,. H.Richnonr! and R. k. Rogers. Havaii Inst. War. Biol. Tech. Rept. Ro. 37.

Holthan, P. 'F. C. v..vI ."�... F. anS and J. E, RaragOS. 1985. Coral reef reCOverySubsv gus nt to the f real. va ter kx 1 l of 1965. Qnl Coral Reef pa pulatiunBiology. ed. by P. L- JOkiel, R. H. FichaOnd aad R. R. ROgerS. Hawaii Inet.impar . I'xol. Tech. Rept. Ho. 37.

Hunter, c. L. and c. C. Fehoe. 1985. patchwork patchreefs: the clonaldiversit y of the coral iiogllRR ~~s~ssa in Kaneohe Bay, Hawaii. ~nr CoralBeef population Biology ed. by p. I,. Jokiel, R. H. Richaoad trad R. L.Ronors. Havaii inst. Har, Prol. Tech. Rept. llo. 37.

1 O I'. 1 e 1 <ao} 1 t.iI y and H. p. rowdin. 1976. IlydrosechanXCal adaptation in tbe

tree-livirq coral t'Hndj,d sputa. nature 262r212-213.

Xri ut «in, n. V., P. V Fiuddese exer and S. V. Sarth. 1972. Car al c brOnolaeterS:si 1- 1Ii Il qrovt h bends in roef c era is. Rcxence '1771270-272.

' ash< r I.r, k F.. 191u. sea sure aen oft of alizarin depositi on by coral. Proc..i-c~n int. Coral Reef Sysp. Bust. 2 2k 1-24q.

J, F. le72. h nt ud of ty o t he ecology of Hawaiian coral, reef s. Ph. D... ~: I dt xan, Iin 1 v. Havaii. 2qV PF.

ar . r. S Tr1tble. in pr as ~i I <'' .I la the Hdv*xldfi I landS I roc Hurricane effects on an aonariua

F1fth Ir!t. Coral Reef COnqress,

'. I.. ~ v. J. R: a ver er, F. It. Law<. R. F. Brock and T. ff Ralsh. 1981 ~I i o Ii I dy r.P1dq dlvi r ior eaperiaent: erS peer �= �y pr ur dtxOr. PaCifXC ScienCe 35:279-396.

Cnawe, i . F., S. v. Dsit.l, and K. J. Boy 1972 Carbonate production by coralr~:- f=. Ra rane neology 12: 123-1iIO.

I note on coaparative quadrat Saapling fOr infaunal and epifaunal inVertebrates,1968-71 and 1983, on the Coconut Island Beef Blat raneohe Bay, oahu, havaii

2. Guin therABC'S, !nc.970 B. Balaheo lve., Suite A300Kailos. BI 96734

11. L. Bartlettsichigan state universityhast Lansing, nl 48923

abstract

Bra antic changes occurred in the Coconut Island reef flat infauna and epifau-aa fOl laving the terninatiun Of Sevage diepOSal in the bay in 1978. Cosparisonof sanples taken in 1968-71 vith saaples taken in 1983 rerealed a sharp reduc-tion in auaber of crabs bivalves and various filter-feeding invertebr ates.Savage abateaeat apparently reduced the asount of food available to these spec-ies. On t he ot her hand. sea cucuebers increa sed in su aber. The se orgari an s a redeposit feeders. The substrate vas highly aaoxic im 1968-11 due to eutrophica-tion. "erhaps the anoxic sedieents vere an unsuitable food source for the seacucua 1 ere.

In trod action

The shallov nariue biotopes on the extensive fringing and patch reefs inKaneohe Bay vindvard coast of the Island of rahu, Bavaii! hare undergone sub-stanti al changes in specres conposition during the last half century. Thesechanges have been aost obvious in the southern part of the Pay vhere vater cir-culati.on is nore restricted and vhere urbanization of the ad jacent watershed hasbeen sos. pronounced. The r.eef structures of raneohe 8ay vere built hy, and ontheir outer f r'inge and fronts I slopes at least! vere a t on.e tice cost l.y pop ala t-ed by, corals and coralline algae � fores vhich have be" one increasrngly rare oraltogether absent in the south part of the Say.

Degradation of the coral/coral liae algae ree f conaunity has been at tributedto a conbination of factors related to urbanization including dredging! ~ ostlySinCe the 1940's see Banner and Bailey, 79701 Capernn. et al., 1971: Panner,1974]. prrsary anong these factors appears to have been the discharge o nunic-ipal sevage froa the !raneohe vastevater Treatnent olaat ioto the southern basin Saith, 1977! . althoogh iepaCte Of urbanization generally beCase a Sub jert Ofintense inquiry during the 1960's and 1970's see Zieaann, 1970: Barn gos, 1972;Krasnick, 1973; Grovhoug, 1976: Pastetter and cooke, 1979!, rery ltttle quanti-tative data exists pe taining to the biological coasunitres extant prior to ur-banization of the vatershed and eutrophacation of the south Bay vaters Thus,"degradative" chanoes in the biota of the Bay can be docuaented only be refer-ence to anecdotal saterial e.g., the "coral gardens" of t' he south 3ay! or bystud yi ng spatial and d'or tenporal ch anges as expression s of a eutrophication gra-dient.

In 1978 the discharge of sevaoe vas diverted frox Kaneohv say t.o a deen oceanoutfall outside the Bay. I subsequent change rn the quality of Kaueohe Bay va-ter vas carefully sonitored and do" unsated by Saith, et al �981! . a corre-spondingng decl. iae in the abundance of readily obser ved bent h i c f i lt er- feederssuch as sponges. soft corals, bryozoans. and tunicates vas noted Dr. paulJokiel, Dt. Bichard srock, personal coneunications!. Infaunal populatrons ofclans also appeared to be declining see Guinther, 1984! . changes in the bioe-ass of the cryptof anna. [specifically, the endolithic fauna! have been reportedby Brock and Saith �983! .

Baterials aad hethods

Set veen Oeceeber 1968 and day 19 73, years representative of the period of e o-trophication in south Baaeohe Bay, one of the authors I aint her! undertook a se-ries of beathiC Sanples On the fringi.ng reef of Coconut Island [5ok u o Loe!, lo-cated in the southern part of Kaneohe Bay betveen 2 aad 2 5 kn fron the sevage

Initially, ony molluscs vere quantified in the quadrate: later, the scope vasexpanded to aociude crustaceans, and still later, all macroinvertebrates gener-ally excluding spreading, color.ial forms such as sponges, bryozoans, and coralsvhich vere recorded as prese~t but not quantified in these samples! . The pco-ject for vhich the samples vere collected vas eventually abandoned b fcient nuebers oof saaples vere collected from any given sector or eefua a an one e ore suffi-enai le a ri oro s9 r us statist real analysis of the data. nonetheless in vi f tho ree zone todocumented chan e s, ill Viem O t ethe ev d c anges in vater quality vhich occurred betveen 1978 d 19 3i dence t hat cet'tain invertebrate populations in the south Bay vere declin-vid ning, a resampling of the shallov reef flat envir sent h ld th

i g interesting comparative data. 8. L. Bartlett undertook this additionalsaeplrng as a student project in the summer of 1963.

The original saaples vere collected by forcing a 1/a square aeter act 1 dmansions vere 50 x 50 x 17 ce ta' e er ac ua i-and renovin all sand dx ce!, stainless stem 1 box f rase into the substratura uater>al, removed by hand ves pld . g an coral rock to a depth of approziaately 1S Thcm. e aa-Ravair Institute of Narine Biolo k o eSot till

ves p aced in containers and transported back to tho ogy laboratory located on coconut Island! forsor nq and vashiag over brass sieves 11, 2 and u ma aesh o e s~ lecroinvertebrates vere removed and counted. an u ma aesh openings! .

The 1983 sam les rip g. ! vere collected in an ideuti=al mannsame bor. fr mme {being constr ucted of meldedexcellont condition e o ve ed, stainless steel, it had survived inon! . Because tl.ere vas insufficient time to usampling program valid for th 1decided that samples mould be collected fr thee re atively com les reef f lat environment< it vas

e co mete rom same 1ocati one as thos~ seeppling than had been don ' ' t ' lle e ore. in addational program � ex endin a rp ing a qrea ter effort in resaa-the comparrsons. kith he n one initially mould not h vea contributed to the val.idity ofbie, t h» result f houg a saaple by sa e le cp caparison is not strictly possi-

e draaatically different froa thes o t e resamolrng proved to be

1983, the lov organise counts prom ted a hu .". n ee, after the first several saaoless vere coll ected inprompted a thorough reviev of the collection andocess �y durnther to confrrm that

d hthrre vas a dramatic change.ee e same ss those used betveen 1968 and 1973. Indeed,Sess 1 t s

The r sul tra rat le 21,lgPl r nants Iled 1 snap! esahern r', statthat no saa; lthr ol 6~r ser

s of the t vo samplrng programs are re'i t1 th 1968-73 " insaeole couuts arran erraoged in the left column and the

g

e t l oa t.he particular Ser ' ld p e', it is meantp

series could oe aae same immediate area

h1t houqh laroer oolychaetes found rn the sa ~ essr golf iCanr rlumher of th fese ores caa be ex ectede samples vere removed and counted &am ar sh opening! si eve. Th p e to pass through the finest �'lrvs. ,us, counts for this t.e. , ' axon vould not be representa-Dier us ion

yhl rrnultn Of thrS st udv are adaittedly preliminarsit h, tg83! have demon" trit d h",ma, . ' c rom o er studres aoy- " . o p

e t at a decline in bent i. st pa rticula rly Broc'k andSion Of Sevage inputS ftnm th 8blar ocr r overt ebra ten of th e a y, a n d our results t hi.c bio mass f olloved diver-

e s.allo v reef f lat enviroextend these findings to theroman an be attributed to do a ecrease in organic loaironaent. The reduction in9 loading: prior to diversion,

l 12

outf al 1 saaple locations vere initially established accord rng to a "stratxfiedrandom" pattern � that. is, placement of the sampling device mas determined ran-domly from qrids established for desrgnated reef sectors representing differentexposures to vind, vs ves, and probably vater gualit y. samples vere taken inshallov, reef flat envrroneents, doeinated in most cases by loose sediaentarymaterial fsand, gravel, cobble! Depths varied froa sites representative of theplover littoral or upper Sublittcral zone tO perhapS -50 cm relative t BLLRrimarv differences in physical attributes vithin this depth range are in therelatrve proportions of hard and soft bottom types to be found in n

quad ra t. n in any gives

Pip. 1. &pprosiaate quadrat saaple loca taons for 5. L. Bartlett's 1983 sasples.

113

TAPLE I. Quadrat counts Number of individuals per I/4 m2!, forthe coconut Island I!oku o Loe! reer flat, comparingsamples taken from 1969 to l973 and samples taken in1983.

1969-73 1983'C C tC 4RC tF t CC 5C 0 RSRR ac i c -x s esca

SAIIPLE: 6 HN41-85-04! **DATE: I/05/69

SAIIPLE: IDATE: 0 8/07/ 83

mi sc. po1 y chae tes 18

I:birgdpia rigida 17

GAIIPLE; 9 %64-95-D9! ~DATE: 05/02/69

SA!IPLE: 2DATE: 07/07/83

mi sc. poly ch set esCrggi bl!lcm sgiaIIsgaICcllgJ,b!rlIIIR Sglsgs!!al

f!i%I,lgs! Zcbrl�!f'I Da ba1laother molluscs

gl!la radii

243

123I I ir!g!!la raa3 ii

SAI:PLE: 11 NE2-24-9! *DATE: 07/16/69

SANPL E.' 3oATE: 06/ 23/83

mi sc. po 1ycha ate sve rmet ldSEf esa bullaXe3,li Da mb!!SLa

P41I

UglgLI! gria a!9999aria 3

SAÃPLE: 8*DATE: 03/ 28/6 9

SAMPLE: 4DATE: 08/07/83

29aoib!IS gati.fi99Smssc. polychaetes

l425

Al ph ex de e89lghb!!cia 0!9099 ari.a

I13

1 l4

Ul!rlc9dr Vga fl! llplrllUiIs9P D90!9I! gacaaf:leos ba11aXelilsa 6!alafas!SI,>sl s aMMraElis!IS sp.Lepigdi!IS Saaguioegscabala~ ada!are

I;I 99 Qia 9GI.a peaI;ir.isg!t aebrg sIallga ggal Jkclaoa

bg "iggdrgaa f.gclggkgs>eris eroia SDl9r9SZ9aalsgb999bl199 Esl.algidaS

s?go959D per�aClaoa baal'aI�996.a 9UJ.ggilioeaI,aZellil!a 9Mala!9ot!,er moll uses

I I2235

2 2

1020

I

I 5 3 2106

I 8 5

19831969-73

21

Edwardaia spPalEibQa sp.

115

SA lPLE: 3 NE41-09-08! *DATE: 12/07/68

Pla!'Qhraar hue Qrellaiea 1Creea belle 3Tellina r Qheaka 4I!aro Da Qhlignilineaka 1

SAPIPLE: 7 SE41-16-03! ~DATE: 03/11/69

@ella 5Xellina x Qhnafa 3iiaQQIoa Qbliguilinea& 2

SAIIPLE= 1D SE64-77-07!*DATE. '07/10' 69

crena belleXel1ina rQhuaEaHarQma Qhligrilinea2.a

ShFIPL E: 3 8DATE: 06/23/71

EnDQdeQQere me!i!!aQ1!lee

SAMPLE: 35DATE; 04/2 9/71

i!in' aeia nulrhella 11hnrhnplaura sp. 4bn~laura nigrearer!e l 95enngdeQQRJ.e medlkeQir!eotner anemones 4

Z32eWQleine !LraQeeii

m aglnQeem 1Biifigm xehrQm 1 !afire gealrerianaleQi!QQEQD garne 1herren hanlezanaBrarbidQniee rrehrisrri ~Tellina ualaram 3

e elerrra 13

fbirnrkata rigid'!!el LEh!!ria mnnQraria

csee w w c c � � C maw

SAMPLE: 5DATE: 08/1 8/ 63

misc. polychaetesuniden.

:hired Qla rig' ia

SAMPLE: 5 above! !

<no comparable sample>

shePLE: 6DATE: 06/18/63

Ed!razdsra sp.hl!ibQQlerxa sp.misc. poly cha e teaCrena hellaTellina palaraD!!IQlnihQXie mrnnrarla

SAHP1,E; 7DATE: 06/11/63

misc. polychaetes

GQQadarr&~s gueriniChirnr!rra rigida!iQlrberia mgnQraria

231

6287

19831 96 9-73s e s pcs msc=s a =c = = =ca casatr.ac c e.ca ca =s ace-sacr e esece3=aeas

SAHPL E r 8DATE: 06/Xls 83

<no comparable sample!

mk sc. polychaetesb Dblbbbxb

!72

SAMPLE: 34DATE: 04/ 2 7/ 71 SAMPLE r 9

DATE r 06/09/83

972699

P12

anemone

misc. polychaetes 36IbcbDX?cbiix2b l2MblSi~b IUISXxx20& sP

IbtlrQQsbiib0 Sef,blx?Idea 2

Eiiblxb elbMcb

SAHPLE: 33DATE: 04/26/ 11 SAPIPLE r 10

DATE: 06/x3/83hxpibbib Swlabellbotner anemones 3

82

HxxDiil2ox b Yb~Dilbb

other polycheates

EuxYibxxb r0brX2lbnbibother polychaetesD4Qd DLb xxCMQDQbZerxbicxcib cblbxbsiamb

avblibrxbnb

71

ISChnxxchiiDD 12biblx2Mbb 4

16Portuni daeXanthidae

ib XlroxxDCariXXSbYi4ia Yi

116

AxSibbib Dxl1Gt! RJ.lbanemoneZxXL 1 Mb CQSr121bkkbZbzb~s KtklxbbiiZWXgJmb SamPlbDbib

lbxlgbirzDXl mrnaCienb bbllbZblimb iabxrbiborner raolluscsFiiibxrS lbfYJJrbQxxbXiibxrb SlbcirbCblribxxb Jaibz"or.net c r usta ceaZbirudx2ib ii gidbbXXIrx&xxrxb brbaxtcbrib

"'ix b bbnlembbksixeb Sbll~bbsisI.xrS Debtco pbibbCiczb bblibc the r mo11uscs

lb b Yxxr b busZx 'xxs Elccwb

ZCLDxxs lbkffbttner crustacea

12

1 12

1811

6 2

l192l18

22 4C

19831969-73== = = =so== == = ===co== ===|=a= seas == - c= = =- i = x=- - c=--e= =a r==

SAMPLE; llDATE: 0 8/24/83

SAIIPL E: 3 9DATE: 06/24/71

PPI~IIa 1 01r!ala sp. 4

ot acr anemones 14sp.

misc, po'ychaetesXIIXgrrIPe RPmglanara 15mrsc. polychaetee 2 36

1415

Crena rLellaZallina Palaram

SAMPLE: 12DATE: 06/Vm/83

SAMPLE: 31 �42I71H-0022!DATE: 04/23/71

hiPraaia PIIlghelIa 6BIIr!IPPIei!xa sp. 6otner anenione s 184 other anemones

ZalxthPa PaammmPIIiliaPPmarPIeiua kra!iaaliPmmaf.PIeima kxameii 379

XiIIWhPe PPmmlai!ara 1laShnPEI!irPD PeralPirie' 2aipdipXa pcragp!!a 6

label!IIPSbi r PII Per alP&ea

pais sP.

tlapmma Dhlig!uliaeara

Eriaiia elearxa

!!Plmrni!I Ia monagaxia

SAMPLE. 13DATE; 08/08/83

<no comparable sample>

531

C!IixPOPra XigidaHPImr!IIIIia moaPPaxxa

SAMPLE: 16 �01270-0392!DATE: 10/12' 70

SAMPLE: 14DATE D6/24/83

mr sc. Po lychaetes

Al pha erriseChiIP!!Pre Iigi&IIPImr1IPIia mPIIPIaxiaHPIPr 1!IIIfa mPIIQPaxia

117

RirriPm zehX1!mISPDGPmPR PermaCreaa hellaTallil!a Palaram

other molluscsXrialia e1eprXaI!erPPPSIaPSPa rhIIkII!!axI ePrmdiIIS aangPiaeIIIPaEyXIxaga!la sp.otner crustacea

hiPraaia PIIIEI!ella>1!rhmglemxa sp.mr ac. poly chaetesCreD a tell 8XaJJ.ina xmIIaHaPP!IIa PIII2IIIIII!eara

3l344

3

12 4 1 7 230

J!IPPPDII sp.Creme hella

un re!en�. shr imp

19831969-73a'cRESCcc ~ azL'h92hggCCCCRkhgRNCRI%RLsscsk CCCECR ~ CCRCRRCCRMC

SAMPLE: 15DATE: 08/24/85

~no comparable sample>

misc. polychaetesiirvpvvix sp-

174

EA PLE: DL! S7.S.3DATE: 8 5/ 2 8/7 3 <no comparable sample>

4 7ll 36

anemoneCrsva hei&Chirr>dv& rividaS,i Rgvla rebziiZJ' YJJLAdkrs ii U b

SAMPLE: 32 �4 217148-82!DATE: D4/23/71 <no comparab I e sample� >

bigiasis ~slit 2Mrbvuibvrs Rii raa~s 13otner anemones 30Zv mrv1eivs krzvvsii 53 DSigbvbsria RD~ 8Bit.&v s xbhrv s 18EvriSr,srviS Rbkvrvlrtv VL 3UvricildrVR4 Wntvv&Z 21SVVRrv V>R kernr 7Gsrrsa baalswns 3Osrrsa sbbdyirr:axis 2Zrdrbidvaker Vrehribrrisfvs 2Creva hslls 19other molluscs 20K izvs slsvrrs 2RsrbX�rsVsvs sp- 4l ertsvdiva SARVvibeVS 5Cslrrbvs J4 ir.RV 1xanthldae 6other crustaceans 66Zbir vdr r4 ri rrdbbvivibvria zvvvrvri4

only molluscs and brachiopods> enumerated in sample*+ Dnly mcl l us s, brachiopods, and crustaceans enumerated ihsample

1 96 9-73 1953

Ci! I DA 8 I A Co EL ERTER AT A !ACT IMPAR IA

3lir!ha ia DviEhellaBarhDglelrra lreaceQshaLhgDlelira sp.I!DDdeazyiS ~aideSI,'rhrarrisr'a sp.

--- all anemones, inc 1. s!iso. IECL ERACTI> IA

Zuxires cr!maressaHaahiD0ra gerraaasa

KOANTHIHIARIAzZalXtha DSammqghilia

s

99925

ll21550

110

POLYCHAETAZu~hae aaiaplaDMazomarekeios Krausii

--- all polychaetes, inc. misc.

10406

7!183

I-!OLLUS CAGASTROruDA

Birnam zehrira!Cruaihalua! sDi Daa!lmi3iitdDra Dckagai!aHlgg52Dia zilDs&Qzri<DdraI2a ~El!liLE5aWDa Qllal2criasaPerisrerDia MDraslmmaZlajLDhraoEII!ls DEellakvsDiahDDari a a0rzallaVermetidae

BIVALVIARL40h&DDXes GrehriaLriaLas<ReDa hellalar�Dr!mDD meri!aPacer!!a DbligwliDearaDsrrea haalegaaaQa r I ea sail 4 Yi re0 si 5Teliiaa galaramTelliaa rmhllata

POLYPLACOPHORAM~hifan Deralaides

all molluscs, incl. misc.

1164

1 13

<l2

17714

221

164

10164

119

Table 2. Taxonomic summary. Counts normal rzed to numberym2 basedon all samples sor ted f or the phylum.

hRTHROPODA, CPVSTACEhr'E cAPODA

Al p tier ua "3CalclsiiS lsXROsMisgs slexxxsZbiggs JSRylzSDirSXf.ksgs sp.1SXttgdigS SADggiDRus5RT.Dpggxspsgs XbgirgbaxRscbygx4psLls sp.Zaxtrtgsgs psgific vsPort uni dae, uniden.shrimp, uniden.Xbalsmita SdmsteXanthidae, un iden.

STD flMGPODAGDDDdapiylrfS ggSXlDJ

all crustacea, incl. misc.

2l8

2

1 4<1<l<1

2

63CshCH ZOPODA

1 XDg&s reemi

E Cjt 1ROD ERNATAOP H18 ROJ UEA

QpfLi Rciis szxigDyxtl OLD TH 0 RO1 D Eh

Cbixodgio xxgi daIgl gbbgxks mDDDcaxxs

all echinoderms56 27

814 36HEHlCHORDATA

EHTEROPHEUS'l'ARXyCbgdtxs fldys

Several SpeCrer. of ZOanthidS vere v rnut I sl and r eef du vere very common on the Coco-ur>ng tne l968-1973 sam linmetr d>str>but>on only poorl coithe am 1 es reported herein.poor y coincided with the l.ocations

Sampling to a depth greater than lSrurface vo 'ld h an cm below the substratum

tne aJ placid shrrmp.e required to obtain a re presents ti v e sampl e

120

lThe ma or |

form of itlpia ls1 ty o. other" anemones ar>sos p~ >Jell>. re probably the smal l

the concentration of particulate or ganic carbon varied bet veen 20-40 mmoles/main the south part of Kaneohe Bay Smith, et al. ~ 1981!; after diversion, partic-ulate organic carban dropped to 15 mmolesf ma.

Potentially, other factor s may have influenced the small change in soft bot-taa infauna recorded by Brook and Smith In Our Study, one group of organiSmS,the holothurians, appears to have actually increased in abundance between 1973and 1993 uf all the infaumal species recorded, only the holothurians partxcn-larly Chirodota rigid@! feed directly on ozganic particulates vithin the sedi-~ ent ~ all of the other soft bottom species e.g., the bivalves, Ctena bella andyell~a 21kgagymj, although found vithin t.he substratum along vlth C. Ihgrda,feed on detritus and/or phytoplankton fro ~ off the bottom or out of the vatercolmmn the bivalves extend a siphon out of the substratum! . k decline in theriohneas Of organiC partiCles within the sediment mould lag behind that in thevater column and on the bottom surface. F urther, if during the period of eutro-phication the sediment environaent vere over loaded by the a" cueulation of argan-ic matter, the resultimg tendency tovard anoxic c'onditions could become detzi-mmntal to species «ith no or minimal dir'ect contact vith the overlying vatercolumn, depressing the populations of these species.

The data provide little information on changes. in the see Table 2! The 1968-73 samples Ibearing in aind thatlooked at in all samples! are on the vhole more diverse tb

~umber of species collected during the period prior tofrom the Bay vere not collected 'in the post-diverszot, samplethe apparent densities of most species enumerated in loH3<samples would ns od be collected to reasonably expect tnspenieS reCorded earlier In the Hrock and Smith �983! stuno taxa vere eliminated follovrng sewage diversion: 'he decprimarily due to a reduction in t.he numbers of indivxduals.the same may be true for the groups of org anisms st udzed hergive no clear indication that the specxes composztxon of tvertebrate assemblaqe on the shallow reef flat at oconutcally following diversion of the sevaqe from Kaneohe Bay,may have drsappeared from the habitat.

species compositionnot all groups vere

an the 1983 samples.drve ra i on o f seve ges. However, gzven

c onside ra nly morencounter all of thedy rt vas noted thatrease in bromass vas

our da ta suggestThe 1983 resultS

he cryptxc macrorn-Island changed radr-lthough some species

Literature Cited

Banner, h. H. 1974. Kaneohe pav, Bavaiir uroan pollution and a Coral reefecosystea. Proc. 2nd Intl. coral Beef Symp., Brisbane. 2 r685-702!

Banner, A. H and J. H. Bai ley. 1970. The effeCts of u rban pOllution upon acoral reef system, a preliminary report. Onr v. of Ba vair, Bavaii Inst. Har.Biol., Tec h. Pept. No. 25 66 pg s,

Brock, P. E. and S. v. Saith. 1963. Nesponse of coral reef cryptofaunal coaau-nities to food and space. Coral Reef s 1: 'l79-183.

Caperon, J., S. A. Cattail, and G. Fzasnick. 1971, phy toplankt on kinet ics rn asubtropical estuary; Eutrophication. Lianol Qcean oqr. 1 6 uf 1 599-607.

Gro»»g, J. G. 1976. A preliminary evaluation of envzronmental indicator sys-.ems in Havaii. maval undersea Center, Tech. Note 16 E9. N.U.C.. San Dzegoand Hawaii. 97 pgs.

121

our results are of interest in viev of the conclusioas expressed by Brook andSaith �983! that the soft tottom cryptofauna as opposed to the hardhottomcryptof auna1 displayed neither a lar ge biomass response to nutrient loading oor'a great decline following tereination of nutrient inputs. Ne would suggest thatthis conclusion is somevhat method dependent. That is, by sampling a greatervolume of sediment and concentrating on the larger organisms, the mobile infau- ~nal and epifaunal organisms of the zeef flat can be shovn to have inczeased dur-ing nutrient loading and declined a f ter termination of n utri eat loading.

""uinther, B. B. 198u. BiOasaay aod biOacCuaulatiOn SPeCieS liSt fnr dredgedant erial testing in tbe Hawaiian Islands. prep. for 0. S. Arsy Engineert'ivision, pacific Ocean kKCOK, Inc., Ho. 395, 57 pgs.

KraSnack, C. J. 1973. TeupOral and Spatial Variationa in phytoplanktOn prOduC-tivity and related factors in the surface vaturs of Kaneobe Bay, Oaha,Mavaii. H.S. Thesis, I1niV. Of Ba Waii, BonOl ulu. 91 Pga.

Saragos, J. Z. 1972. k study of the ecology of Ha vaiian reef corals. Pb. B.Bivsertation. Univ of Havaii, Honolu3u. 209 pgs.

Bantet ter, E. B. and u. J. Coake. 1979. BeSpenueS Of sarine fouling coununi-tiee tO uevage abateaent iu Kaneohe Bay, Oabu, Bavaii. Bar. BiOl.53: 271-280.

saith, s. v. 1977. Kaaeohe Bay: I prelieinary report on the responses of acoral reef/estuary ecosysteu to relaaation of aevage stress. Proc. 3rd Iutl.coral Reef sysp, Hiaui. 2rn26-a31.

saith ~ s. v., u. J. Kiauerer, E. k. Lass. B. E. Brock, and T. Q. Halsb. 1981.KaneOhe Bay Sevage diuerainn erperiuent: perupectivea on eCOSyetea reepnnaeuto nutritional perturbation. oac. Sci. 35:279-395.

Sieuann, 0. a. 1970. The horizonatal diatributiOn of ZOOplanktOn in KaneOheBay, rahu, Bavaii. univ. Bavaii, Havaii Inst. Bar Biol., Tech. Rept. No. 23.56 pgs.

122