HAWAU-T-79-004 C. 2

MARINEOPTION PROGRAMDATA ACQUISITION PROJECT

PAPOHAKU BEACH AND MOLOKINI ISLAND,

Guy A. Anzai

Geoffrey Akita Lisa Boucher Richard Fantine Tad Kobayashi Gordon Muraoka Holly Price Steven Takenaka Leonard Torricer

1

August 1979

SEA GRANT COLLEGE PROGRAM

University of Haggai' , ffawaii MARINEOPTION PROGRAM DATA AC UISITION PROJECT

PAPCHAKUBEACH, MOLOKAIAND MOIOKINI ISLAND, MAUI

Guy A. Anzai

Geoffrey Akita Lisa Boucher Richard Fantine Tad Kobayashi Gordon Muraoka Holly Price Leonard Torricer

WORKING PAPER NO. 39

August, 19?9

SEA GRANT COLLEGE PROGRAM

University of Honolulu, Hawaii

This morningpaper is pub2ishedvith funds provided in part by the University of Hara ii SeaGr'ant C'o22ege Wogxam under 2nstitutiona2 Grant No. 04-8-258-44028 from NOAA,Office of Sea Grant, Departmento f Commerce,tuith additiona2 funding support fr'om t he State of Hawaii Office of' t h e MarineAffairs Goordi- nator under Task Order No. 220. The USGovernment is authorised to pr'oduceand distribute reprints for governmenta2pur poses not- nrithstandingany copyright notations that mayappear hereon.

TABLE OF CONTENTS

INTRODUCTION . ~ ~

MATERIALS AND METHODS.

Transect Site Selection . ~ ~ ~ ~ Fish. ~ ~ ~ ~ 4 ~ ~ 13 2 Substrate . Algae . ~ ~ ~ ~ ~ ~

TRANSECT AREA DESCRIPTION. ~ 5 PapohakuBeach, Molokai 5 5 Molokini Island, Maui

PAPOHAKUBEACH, MOLOKAI.

Results 6 Discussion. ~ 12 Conclusions . ~ 14

MOLOKIN! ISLAND, MAUI. 15

Res ul ts 15 Discussion. ~ ~ ~ ~ . 22 Conclusions . . 25

SUMMARY s ~ ~ ~ a ~ ~ t ~ ~ ~ ~ ~ ~ ~ ~ s ~ ~ ~ ~ ~ ~ ~ ~ a ~ . 25

ACKNOWLEDGMENTS+~ ~ ~ ~ ~ a ~ ~ ~ ~ ~ ~ . ~ ~ ~ ~ e ~ ~ ~ ~ 26

REFERENCES CITED . 26

APPENDICES . 29

Appendix A. Scientific, Hawaiian and/or English Names of Fish Seenat PapohakuBeach and Molokini I sl and e ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 31 Appendix B. Presence/Absence and Frequency of Occur- rence of Fish Species by Transect Stations at PapohakuBeach. 33 Appendix C. PapohakuBeach Fish Data with Bottom Description by Transect. . 35 Appendix D. Comparisonof 1974 Study with 1976 Study by Biomassand Numberof Fish Per Substrate at Papohaku Beach. 36 Appendix E. Similarities Between Coral Species and Substrate Types Between Tr ansects at PapohakuBeach . 37 Appendix F. Frequency of Occurrence and Percentage of Cover 3.75 m ! for Algal Species and Substrate Types at PapohakuBeach. 38 Appendix G. Fish Statistics and Substrate Types by Transect off Molokini Island . 39

Appendix H. Presence/Absence and Frequency of Occur- rence of Fish Species by Transect off Molokini Island. . 40

Appendix I. Similarities Between Coral Species and Substrate Types Between Transects off Molokini Island. . 42

Appendix J. Molokini Island, Algal Species and Sub- strate Types with Frequency and Percentage of Cover for 11.25 m~. 42

LIST OF FIGURES

Figure

Transect locations at Papohaku Beach.

2 ' Transect locations off Molokini Island. 16 Fish biomassby feeding type off Molokini Island.. 17

LIST OF TABI ES

Table

1 Transect locations at Papohaku Beach. ~ i a I 8 2 Fish biomassby substrate type at PapohakuBeach. ~ ~ ~ ~ 9 3 Comparison of 1974study with 1976study at Papohaku Beachby biomassand numberof species of fish per 9 transect. ~ ~ ~

4 Percentage of substrate types by transect at PapohakuBeach. ~... 10 5 Frequency of occurrenceand percentageof total live coral cover by coral species at Papohaku B each ~ . -.. ~...... ~....,... 10 6 Frequency of occurrenceand percentageof total substrate cover by substrate type at Papohaku Beach ~ a ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 7 Frequency of occurrenceand percentageof cover by dominant a'lgae at PapohakuBeach . 12 8 Transect locations off Molokini Island . 16 9 Fish biomassby substrate type off Molokini Island...... 17

10 Fish biomass by feeding type off Molokini Island . 17

11 Biomass of desirable food fishes off Molokini I sl and o ~ ~ o ~ ~ ~ a ~ i ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 18 12 Percentageof substrate type by transect off Nolokini Island. ~ ~ ~ . 20

13 Percentage of coral cover and number of coral species per transect off Nolokini Island . 20

14 Frequency of occurrence and percentage of cover of live coral off Molokini Island. 21

15 Frequency of occurrence and percentage of cover by substrate type off Molokini Island. ~ . 21 16 Frequencyof occurrence and percentage of cover by dominant algae off Molokini Island. . 22 INTRODUCTION

Someopen shoreline areas in the state of Hawaii are being lost to commercial, urban, and recreational development. Such development may subject the adjacent marineecosystems to changesdue to the drastic alter- ation of the land. For example, prevailing winds may sweepexposed topsoil from defoliated land and deposit it in nearby waters, or surface nmoffs may increase along with great amountsof effluent from outfalls. Also, increased recreational use of a once secluded and pristine area without regulatory measuresfor its protection maycause the eventual depletion of the resources in that given area. This concern for and the lack of infor- mation dealing with such developmentprompted students from the Data Acqui- sition Project of the MarineOption Program MOP! at the University of HawaiiManoa campus to organize and participate in a study to observeand record physical andbiological data from PapohakuBeach, Molokai andMolo- kini Island, Maui. The first part of the study consisted of a six-day survey off Papo- haku Beachin early June of 1976. The initial phaseof land development by KaluaKoi Corporationfor an urbanresort complexprovided .a unique opportunityto observeand record the effects of landdevelopment on the marineecosystem. A similar study conductedby other MOPstudents in 1974, prior to the constructionof the urbanresort complex, served as a basis for comparison. Thesecond part of the study consistedof a four-daysurvey around Molokini Island. Uninhabitedand situated in the Alalakeiki Channeioff Maui's west coast, this area wasrecognized by the state of Hawaii as a uniquemarine environment of inestimableeducational and scientific value. At the time of this study, the State Departmentof Landand Natural Resour- ceswas considering the establishmentof the watersof Molokini Island as either a natural reserve area or a marine life conservationdistrict 976 SenateResolution ¹416 and SenateConcurrent Resolution ¹110!. Thewaters surroundingMolokini Island have since been established as a MarineLife Conservation District July 1977!, Molokini Island is a 19-acre, crescent-shapedtuft crater which exhibits unique features including a sharpbottom transition from shallow,to deep; water clarity permitting greaterthan 50-m visibility; and abundant, colorful marine life including relativelyunique coral species J.E. Maragos,1977: personalcommunica- tion!. However,little recordedinformation exists as to the physicaland biologicalmakeup of the area. It wastherefore realized that a baseline studyaround Molokini Island would contribute useful information to those workingto preserveand to protect the island's natural resources.

MATERIAS AND METHODS

Transect Site Selection At eachtransect site, three suitable landmarkswere sighted from the waterwith a hand-heldcompass the landmarks'bearings were recorded in degrees!.The landmarks were then located on a chartand a singleline drawnthrough each at the appropriatedegree. The approximate start of each transect was placed at the intersection of the three lines and recorded in degrees latitude and longitude to allow accurate repetition of the transect without the use of permanent markers. All transects ran parallel to shore at an approximately constant depth and were selected, rather than randomly chosen, to adequately cover the Papohaku Beach and Molokini Island areas in the 10-day survey period.

A modification of the visual census methods utilized by Brock {1954!, Odumand Odum955!, Bardach 959!, and McVey 970! was employed off both PapohakuBeach, Molokai and Molokini Island, Maui. Divers equipped with either SCUBAor worked in teams of four, consisting of one line roller, two counters, and one safety diver. The counters followed behind the line roller as the 100-m plastic transect line was being deployed to record any fish being frightened out of the immediate survey area. Each counter was responsible for ! identifying species, ! esti- mating the number of individuals of each species, and ! estimating the standard length tip of snout to middle of caudal peduncle! of each indi- vidual in an area 5 m wide and within 2 m from the bottom, adjacent to the transect line. The combined area covered by both divers from the beginning of the transect line to the 100-m mark equaled 10 m x 100 m or 1,000 m and was designated as station A. Upon completion of station A, counters exchanged lanes and proceeded to record from the 100-m mark to the begin- ning of the sametransect line station B! to thoroughly cover the survey area. In addition, points of interest and species of fishes outside the transect area were noted on underwater vinyl slates for later transcrip- tion to data sheets. Collected data were processed via several computer . programsdevised by the Hawaii Coastal ZoneData Sank HCZDB!at the University of Hawaii.' The HCZDBprintout included data on ! numberof species, ! biomass standard length x constant value for particular species!, ! standardlength, ! frequencyof occurrence numberof stations!, and ! presence/absence of species. The species of fishes were categorized by feeding habits based on studies by Hobson974! at Kona,Hawaii. The three categories included ! herbivores--strictly algal and detritus feeders; ! omnivores-- opportunistic feeders on both plant and animal matter; and ! carnivores-- feeders on animal matter, including coral, plankton, and other fishes Appendix A!. Dominantgroups of fishes were determined by high frequency of occurrence > 0.541 at PapohakuBeach and > 0.833 around Molokini Island! rather than bligh biomass and large numbersof individuals since itinerant species occurring in large schools at a few stations wouldhave large bio- massand numbers, yet would not truly indicate the commonnessof that species. Fishes were grouped by occurrence over similar substrates in an effort to determine preferred habitat. Habitats included steep slopes with abrupt dropoffs; areas of less steep, sloping bottoms with rich coral cover; areas of boulders and ledges; and areas with sparse coral cover on a silt or sand bottom. It must be noted that information collected by the visual census methodyields only a rough estimate. Discrepancies betweenthe actual and recorded length of individuals, total biomass, and population size can be attributed to ! the several families of nocturnal fishes, such as apogo- nids 'upapalus!, holocentrids squirrelfishes!, and priacanthids 'aweo- weos!, which retreat into cracks and crevices during the day Gosline and Brock, 1960; Hobson, 1968!, making sampling difficult; ! the behavior and relative abundanceof certain cryptic species which appear primarily during dawn and dusk periods Hobson, 1965, 1968! and which would be over- looked by a visual census at other times; ! observers "spooking" fishes out of the area under study; and ! observers over- or underestimating the size of individuals.

Other methods of sampling such as fish poisoning Randall, 1963! and the use of explosives Talbot, 1965! were not considered. Although they would probably yield a more accurate count, these methods would be detri- mental to the population sampled.

Substrate

The substrate and algal surveys were done in conjunction with the fish survey at each transect site to allow for a correlation of the three. These surveys were conducted along the same transect line imme- dately fo11owing the fish transect.

Ten numbers from 0 through 100! were picked using a random numbers table and subsequently matched to the meter number of the 100-m transect line. For example, if the number 17 were pi.eked, one sample would be taken on the 17-m mark.

At each of the 10 randommeter marks along the line, a modified photo- graph method devised by S.A. Reed 976: personal communication! and a visual field identification methodwere employed. The photographic method was conducted with a camera equipped with a 35-mm lens mounted on a 1.3-m high aluminumstand. A light reading was taken at the beginning of each transect line and the camerawas adjusted to the specified setting. A plastic 5-cm card, markedwith the transect and samplenumbers, was placed on the substrate to insure proper identification of the photo- graph. Bach photograph covered an area of 0.5 m x 0.8 m for a total area of 4 m at each point along the entire transect line. Visual identifica- tions and rough estimates of percentage of cover were obtained at the same time to insure somedata in the event of film failure. Coral species and other types of substrate were recorded on plastic underwater slates. Samples of unknown species were retrieved for later identification. In areas of unfavorable conditions e.g., strong currents and vertical drops! where the mountedcamera could not be used, only the visual survey method was employed.

The substrate was divided into six types as follows: ! hard basalt bottom; ! rubble--loose, dead coral; ! sand; ! silt and mud; ! algal mat; and ! live coral. The live coral could be further divi- ded into percentage of cover by species using the sameprocedure, To determinethe percentageof coverof eachsubstrate type the s1' ides wereprojected life-sized onto a screenwith 10-cmsquare grids Frequencyof occurrencesimply meansthe numberof traiisects a par-" ticular type of substrate or coral species appears.in divided by the total numberof transects surveyed. The similarity index indicates how similar one transect is to another by the value obtained from the Soren- sen quotient:

2 x E min. aq, b~!

A + B where

Total percentage of cover of coral species or other substrate types at transect A

Total percentage of cover of coral species or other substrate types at transect B

ag Percentage of cover of coral species 1 or substrate type 1 at transect A

bi = Percentage of cover of coral species 1 or substrate type .1 at transect B

The higher the value between transects, the pore alike they are, in terms of physical and biological substrate types,

Al gae

The algal survey was conducted using SCUBAand a modified point- quadrat method. A 50-cm2quadrat divided into 10-cmgrids was placed at five points every other point of the substrate team's 10 random points! along the 100-m transect line. At each point, the dominant alga occupy-. ing each of the 25 10-cm squares of the quadrat was identified and the percentage of cover was estimated and recorded on a plastic slate bearing a scaled-down grid.

Unidentified dominant algal genera occurring on the transect were col- lected and recorded. A pressed sample of each unidentified alga provided a basis for identification upon return from the field. In the absence of algal cover, the type of substrate beneath each 10 cm square was recorded. To standardize terms, the substrate team's definitions of specific substrates were followed except for "rubble," which includes live coral in this portion of the study.

Information was also recorded on genera of algae observed on or near the transect line but not appearing under the algal quadrat, interesting features of the substrate, physical factors, and unusual events occurring during the transect. Transcription of the raw data from the plastic slate took place immediately after each transect to avoid error. Valuescorresponding to the proportionof 10-cm~squares that each genusoccupied per transectwere tabulated using the HawaiiCoastal Zone DataBank computer. The area surveyed amounted to 1.2Sm~ per transect. Eachsurvey area was analyzed for frequencyof occurrence the numberof transectsa genusappeared in dividedby the total numberof transects!and percentageof cover the numberof 10-cmsquares occupied by a genus dividedby the total numberof squares!.Algae with a frequencyof > 0.5, or appearingon at least one-halfof the transects,were considered domi- nant forms. A Sorensensimilarity quotient test wasrun to determinesimilarities between transects of the two survey areas.

TRANSECTAREA DESCRIPTION

PapohakuSeach, Nolokai

Extensiveland gradingwas observed directly behindthe white sand beachat Papohaku.Frequently throughout the four-day survey, strong gustsof windwould blow great amounts of red soil out overthe ocean. Naterclarity wasoften poor, probably as a result of this surfaceerosion. Theinshore areas had large boulders and sand-silt bottomswith lim- ited coral growth. Offshoreareas consisted of occasionalboulders, sand patches,channels, deep crevices, ledges, and some live coral cover.

Nolokini Island, Maui

Surfaceconditions off Molokini Island were extremelyvariable through- out the day. Usually,calm, flat conditionsprevailed throughout the morn- ing, wi:thwindy conditions and seas of upto 1.3m pickingup by mid afternoon. Sometimesabrupt changes in surfaceconditions from calm to roughand choppy, and back to calm, or 180' wind changeswithin 15 minutes, occurred. Thesubmerged crater off Holokini wascharacterized by a flat floor sur- roundedby a rapidly slopingbottom and abrupt drop-offs. Coral coverwas extensive, with somesand and coral rubble patches. Theexceptional clarity greaterthan 30 m visibility! of the water aided observation of fish, substrate, and algae. PAPOHAKUBEACH, NOLOKAI

Results

Fish A total of 19 transect lines Figure 1 and Table 1! were laid at Papo- haku Beach in June of 1976 in the same area surveyed by MOPstudents in June of 1974. Since the precise locations of the previous transect lines were not known, only a general area comparison between the two surveys was possible. Transectdepths ranged from 3.8 to 12.2 m. Visibility waspoor at times, particularly in the inshore areas, but averaged 13.6 m. Groupingtransects accordingto similarities in substrate Table 2! revealed that those areas which afforded sometype of vertical relief had the highest fish biomass. Thesetransects were generally located over a substrate of sand with basalt ledges or boulders. The average biomass on transects with this type of substrate was2.23 kg/1,000m~. Speciesdiver- sity was greatest in this category--an average of 11.2 species per transect. The lowest average biomasswas found on transects in areas which appearedto havethe greatest siltation Table 2!. The averagebiomass for these transects was 0.56 kg/1,000 m~, with an average of 8.9 species per transect. The remaining transects were grouped together as flat, sandy sub- strates. The average biomass for these was 1.14 kg/1,000 m , with an aver- age of 10.3 species per transect. Transects 1 through 6 were located in the area of the 1974 study. Com- parison of the offshore area transects 1 through 4! with the data collected in 1974 showeddecreases in biomass, number of fish, and species diversity . Table 3!. The average fish biomassof transects 1 through 4 in 1974 was 6.79 kg/1,000 m in comparisonwith 1.80 kg/1,000 m in 1976. The average numberof fish per transect decreasedslightly from 76.3 in 1974 to 70.6 in 1976. The average numberof species per transect declined from 15.1 in 1974 to 13.6 in 1976. Greater decline in biomass, number of fish, and species diversity was found in the inshore areas tr'ansects 5 and 6'!. Here, average biomassper transects droppedfrom 5.83 kg/1,000m in 1974to 0.64 kg/ 1,000 m~ in 1976, with the average numberof fish per transect declining from 13.0 to 5.3. The most abundant fish in the Papohaku Beach area was Tha7assoma 2upezrepi see Appendix8!. This fish was found at 16 of the 19 transects for a frequencyof 0.757. Other specieswith high frequencyof occurrence included Pomacentvusjenkinsi, G'hromis oval is, Stethoj u his haLteata, Pampeneusp/eur ostigma P~peneus porphyr eus, and.Chaetoaon mi7i~s. The numberof species, biomass, total numberof fish, depth range, and bottom description are given for each transect station in Appendix C. A comparisonof the averagebiomass and averagenumber of fish from the 1974 study with this study is shown in Appendix D. I.ESENOI I d7 %>'';.;,.!+I',~',""' ,-"'. ""<"-. tooIIO4 sort Coho rocks trohooCI

d00 meters

horth

214 I I 'd0"

/'7 /Il/ / /I /'.' / I /'/ '/ J

210 I I '00 N

LANAI NAUI

Figure 1. Transect locations at Papohaku Beach. See Table 1 for specific locations.! TABLE 1. TRANSECT LOCATIONS AT PAPOHAKUBEACH

Transect Latitude Longitude Direction of Transect

21' l l ' 39" N 157'15 ' 23" 131' T 21' l l ' 44" N 157'15'25" W 180' T 2'I ' l l ' 39" N 157'15'29.5" W 180' T

21' l l ' 37" N 157'15 I 25I ' W 180' T

21 11'52" N 157'15'05.5" W 169o T 21' l l ' 59" N 157'15'04.5" W 169.5' T 21' l l '23" N 157'15 ' 12" W 168' T

21 11'37 5" N 157'15' 12" W 192 T

9 1'Il '35 5" N 157'15' 12" W 179' T 180' 10 21' l l '22" N 157'15'25" W 21' l l ' 14" N 157'15'14" W 24'

12 21' l l ' 31" N '157'15'30.5" W 180.5' T

13 21'l l '27 5" N 157' 15' 18" W 180 14 21' l l ' 08. 5" N 157'15 ' 22" W 180' T

15 21' l l ' 07" N 157 15'27" W 180' T i6 21'10'43'' N 157'15'28.5" W 219.5' T

17 21'10'55. 5" N 157'I5'33" 219.5' T 18 21'10' 50. 5" N 157'15'2$" W 220 T

19 21'10' 57" N 157'15'21" W 119.5' T TABLE 2. FISH BIOMASSBY SUBSTRATETYPE AT PAPOHAKUBEACH

Average No. Average Biomass Substrate Type Transect No. of Species Per Transect Per Transect kg/1 000 m !

Ledges, boulders, s igni f- 1, 2, 3, 5, 11.2 2-23 icant vertical relief 6, 8, 9, 10, 13, 16, 17

Inshore areas, very silty 7, 11, 12, 8.9 0.56 14, 15

Flat sand and algae 3, 18, 19 10. 3 1.14

TABLE3. COMPARISONOF 1974 STUDY WITH 1976 STUDY AT PAPOHAKUPEACH BY BIOMASSAND NUMBERAND SPECIES OF FISH PER TRANSECT'.

Offshore transects Inshore transects through 4 5 and 6

1976 1974 1976

Average biomass 6.79 1. 80 5.83 0.64 per transect kg/1,000 m2!

Average No. of 76.3 70.6 126.8 21.3 fish per transect

Ave rage No. 13.6 13.0 5 ' 3 of species per transect Substrate

The estimated mean live coral cover at Papohaku Beach was 0.3 percent of the total substrate Table 4!, The four species of coral encountered were Pozites Zobata 9.8 percent of the total live coral cover!, PociZZopora meandrina 2.6 percent!, Montipoza vez~oosa .7 percent!, and PooiZZopora damicomis .9 percent! Table 5!. The remainder of the substrate con- sisted of 54.2 percent algal mat, 31.3 percent silt, and 14.2 percent sand Table 6!. Of the substrate categories recorded, the algal mat had the high- est frequencyof occurrence.000!, followed by silt .947!, coral .670!, and sand .474! Table 6!.

TABLE 4. PERCENTAGEOF SUBSTRATETYPES BY TRANSECTAT PAPOHAKUBEACH

Transect Coral ! Algal Mat ! Sand ! Silt Z!

0.1 63. o 0.0 36.9 1.3 55. 4 28.1 15.2 1.2 45. o 29.8 24.o 24 35 0.1 56.9 0.0 43.o 0.0 50.0 50.0 0.0 0.0 44.3 2.8 52 9 68 7 0.2 87.5 0.0 12 ~ 3 0.1 93.9 0.0 6.o 9 o.4 82. 1 0.0 17-5 'IO 0.1 73.4 25.5 1.0 11 0.0 4.7 0.0 95.3 12 0.2 3o. 8 0.0 69.0 13 o.4 59.4 0.0 4o.2 14 o.4 41. 7 0.0 57.9 15 o.4 57.4 0.0 42.2 16 0.0 41. o 56.5 2.5 17 o.4 42. 8 5.0 51. 8 18 0.0 46. 2 32 3 21.5 19 0.0 54.3 39 2 6.5

Mean 0.3 54.2 14.2 31 ' 3

TABLE 5. FREQUENCYOF OCCURRENCEAND PERCENTAGEOF TOTAL LIVE CORAL COVER BY CORAL SPECIES AT PAPOHAKU BEACH

Frequency of 4 of' Total Species Occurrence Live Coral

Pooi ZZopoza dami oops 0.453 1.9 Poli ZZopora meambina o.158 22.6 Montipora vexmoosa 0.105 5.7 Polities Zobata o.684 69.8

10 TABLE 6. FREQUENCYOF OCCURRENCEAND PERCENTAGEOF TOTAL SUBSTRATE COVER BY SUBSTRATE TYPE AT PAPOHAKU BEACH

Frequency of 0 of Total Substrate Occurrence Subs tra te

Cora l 0.670 0,3

Sand 0.474 >4.2 0.947 . 3>.3 Algal mat l.000 54.2

Hard basal t 0.000 00.0

Rubble 0.000 00.0

The most obvious invertebrates encountered were thousands of mol- lusks known as the sea hare, Aplyaia juLuma. These creatures were approximately 5 cm in length and were found in large aggregations at every transect line. Other mollusks found were the commonspecies of Cyprea, Conus,Mirza, and Perch~. Manynudibranchs and their egg masses were also observed. In the rocky surf 2;oneof the point near Papohaku Beach, manylarge opihi were seen between7 cm and 10 cm long. Seve'ral large octopi were observed in the rocky inshore areas, two of which were estimated to exceed 2,200 grams. Also noted on several transect lines were many large spiny lobsters of the genus PanuLima. Silt coverageranged from 0 to 95.3 percent, with the highest percent- age occurring on transect 11 and the lowest on transect 5 Table 4!. Sandcoverage ranged from 0 to 56.5 percent, with transect 16 having the highest value and 10 of the 19 transects with no sand Table 4!. Transects 4 and 15 exhibited the greatest similarity to each other .9920! by coral species and substrate type, while transect 1 wasthe least similar to the other 18 transects Appendix E!. Transect 2 had the greatest coral cover at 1.3 percent of the total substrate Table 4!. The greatest number of coral species occurred at transect 17 where Porites Lobata, Poli L'Lopora meae&ina, and P. dami- oo~is were present.

PapohakuBeach has diverse benthic flora with 28 species of algae recorded--26 "known," and two unidentified " unidentified I and II"!-- and two types of recordedsubstrates--sand and silt. SeeAppendix F for a complete listing.! Frequencyof occurrence of dominant algal genera and their corres- ponding percentage of cover are given in Table 7. TABLE 7. FREQUENCY OF OCCURRENCE AND PERCENTAGE OF COVER BY DOHINANT ALGAE AT PAPOHAKU BEACH

Frequency of Genus R Cover Occurrence

Padina 1. 000 Dicty optezi s 0.947

Ni cz'odi cty on 0.895 33 O'Ladophoropsis 0.789 6

8a2imeda 0 579 1 Scu gassum 0.526 8 Di ctyota 0.526

Large, brown frondose algae, Padina sp., Dictyoptewis sp., Sax'gassum sp., and Dictyota sp., dominate the bottom along with the greens, Miczodic- tyon sp., C7adophoropsis sp., and Ha7uneda sp., a calcified algae. Large, extensive stands of Dictyoptems sp. were often observed both on and near the transects near shore. Padina sp. also occurred near shore. Both were dominant on all the transects closest to shore.

Several varieties of red algae were observed, but only t:entice>as sp., which occurred on six transects and accounted for 8 percent of the total algal cover, deserves mention. "Unidentified I" appeared only on transect 2, although occasional stands were observed near a few of the offshore transects

Many of the Dictyopteris sp. and Scugassumsp. thalli had only midribs present and were seemingly at various stages of losing their fleshy blades. Also, many of the Nicrodictyon sp. and C'Ladop'horopsissp. thalli had a cropped or trimmed appearance.

Discussion

Fish

Transects were grouped by similarities in substrate in an effort to determine what types of areas were preferred by fish. High biomass was associated with areas of boulders and ledges. This information parallels McVey's 970! findings on fish populations in artificial areas at Pokai Bay, . His study showed that fish populations increased with increasing vertical relief. Areas lacking any type of relief were less attractive to fishes.

Comparisons with data collected by MOP students in 1974 showed a marked decline in numbers, species, and biomass of fish, especially in inshore areas. A combination of factors may be responsible for this. Prior to and during the 1974 survey, no construction on the land directly

12 behind the beach was evident. Construction, which began iri December 1975, simultaneously affected the land and water ecosystems. Extensive grading removedall vegetative cover and left the topsoil exposed. Prevailing winds quickly picked up the loose soil and deposited it into the surround- ing waters. The greatest siltation occurred in the inshore areas. Lower fish numbers in these areas are probably due to the adverse effect of silt- ation, with inshore areas showing the greatest decline. Similar effects of siltation to surrounding water were found by Grigg 972! at Hamakua, Hawaii. He stated, "Changes in both species and abundance of near shore fishes have been observed off sugar mills, with catches there about one- third those of control stations." An interesting point which perhaps supports another reason for this decline is the notable absence of desirable food fishes in the 1976 survey. The 1974survey noted such large table fish as C'cmmzrneLampygua omilu!, Mpzipzistic ~an 'u'u!, Apzionuizeeeene uku!, andPcmcpenaue poxphy- z cue kumu!. Of these, only Pcmrpeneusporphyry eve waspresent in 1976, with all individuals 18 cm or less in length. Increased fishing activity mayhave been the reasonfor this because,with construction, accessroads to the construction area mayalso have led to increased use of the shoreline.

Substrate Perhapsthe most striking observationmade at PapohakuBeach is the large amountof land-derived aedimentsand the notable lack of live coral cover in the surroundingwaters. This suggeststhat the presenceof sand or silt mayregulate the distribution of coral. Studies doneby Woodand Jones 907! at Cocos-KeelingAtoll andMaragos 972! in KaneoheBay show that heavysiltation can inhibit growthand evenbury the coral. Thepre- sence of silt also contributes to an unstable environment which prevents the coral 1arvae from finding suitable attachments Maragos, 1972!. Light inhibition due to suspendedparticulate matter may also inhibit coral growth since it is light-dependent Maragos, 1972!. Studies by Finckh 904! at Funafuti Atoll and Woodand Jones 907! indicate that competition with algae mayinhibit coral growth, retard its recovery, or even kill it. The large population of Aplpeia juLnznaat PapohakuBeach probably had no effect uponthe coral population; its presencewas probably due to the algaeon whichit fed M. Dunlop: personalcommunication!. At transects 16, 18, and 19, the large amount of sand and small amount of coral cover suggest that sand is an inhibiting factor to coral presence. Silt was not a factor here becauseof its limited extent. Lowsiltation wasprobably due to thesetransects being the farthest awayfrom the grading site. Certain species of coral vary in their degreeof tolerance to the presenceof sandand silt. Poziteslobata had the greatestpercentage of cover of the four coral species found at PapohakuBeach. This is primarily dueto its polypswhich may be moreeffective at removingaccumulating silt particles thanpolyps of othercorals such as Focal,lopo~sp. as reported by Mantonand Stephenson 935! in their studyon the GreatBarrier Reef. The amount of light penetration due to suspended particles in the water can also influence the presence of certain species of coral. In Hawaii, Porites sp. may be less dependent upon light Maragos, 1972! than Pocillopora sp. and may becomedominant in such areas. PociL,'Eoporasp. was found in the offshore areas where the full effects of the silt were not felt.

The functions of algae in marine ecosystems are diverse and important. As primary producers, algae serve as a basis of food production and many calcareous forms contribute directly to reef building as well as provide sedimentation in reef environments Smith et al., 1973!. They also provide major sources of organic matter in chiefly inorganic atoll soils Doty, 1973!. Any comprehensive study of a marine environment should include a census of the algal population because of its multifaceted role in marine ecosystems.

The Papohaku Beach area is characterized by abundant, diverse algal growth dominated by fleshy, frondose Patina sp. and Dictyopteris sp. near shore and a mat of C'Ladophoropsis sp. and Nicrodictpon sp. throughout, Dictyopteris sp. apparently favors high energy environments Smith et al., 1973! which probably accounted for its predominance at the transects along the shore where strong bottom surge was present. The heavy siltation may decrease coral populations and reduce the competition for space between algae and coral Maragos, 1972!, allowing more room for algal growth. On many transects, the observer had to brush the silt cover off the bottom to note the flora. This indicated a possible recent accumulation of silt which may bury and kill the algae, Cs was found in a study by Neal 930! off Waikiki, Oahu.

Siltation may partly explain the cropped appearance of a few of the dominant algal genera. Harger 972! reported the same state of thalli lacking fleshy blades and with only midribs present for Dictpoptez'is aus- tralis at Waikiki. He postulated that the condition was due to abrasive sand scour. At Papohaku, the heavy silt layer stirred up by bottom surge may have provided abrasive scouring partly responsible for the condition of Dictyopteris sp., Sazgassum sp., and the "cropped" Ni.croCictpon sp. and C'7adophoropsis sp.

Conclusions

The survey at Papohaku Beach represented a unique opportunity to study and record an ecosystem under transition. Initial comparison of the 1974 study with the 1976 study revealed that drastic changes on the land can alter the physical and biological structure of the adjacent marine commu- nity. The most notable changes were the great amounts of silt present in the surrounding areas and its subsequent effect on the marine life. Marked decline of fish and coral populations illustrated the adverse effects of the presence of silt. On the other hand, certain species thrived in the new environment as evidenced by the great numbers of sea hares and certain species of algae.

14 Nineteen transect lines were laid and marked in this survey, provid- ing future surveysa basis for moreaccurate comparisonsof the area. The next phaseof study, the period after construction, should provide valua- ble information on the effects of ! fertilizer and freshwater runoffs from the planned golf course and urban dwellings, ! runoff from sewer outfalls, and ! increased recreational uses.

MOLOKINI ISLAND, MAUI

Res uk ts

Fish Nine transects were completedoff Molokini Island betweenJune 16 and June 19, 1976--seveninside the bay area enclosedby the crescent-shaped crater and two outside, off the ends of the island at Lolilali and Pahee o Lono Figure 2 and Table 8!. Transect depthsranged from 7.6 m to 18.3 m. A quickly sloping crater bottomwhich gets progressivelydeeper away from the island waspresent. Outsidetransects 1 and 3 off Lolilali and tran- sect 5 off Paheeo Lono, suddensteep dropoffs to more than 30 m occur. A strong northwesterly encounteredoff Lolilali madetransecting difficult. A total of 94 fish species were recoxded off Molokini Island. Of these, 87 were "known"or identified species and sevenwexe "unknowns" from the families Balistidae, Carangidae, Holocentridae, Labridae, Poma- centridae, and Scaxidae. Total biomassfor the 18 stations was 234.216kg calculated from 7,680 individuals see AppendixG! for a meanof 13.012 kg/1,000 m~and 426.7 individuals per station. Thetotal axeasurveyed included 18,000 m Results were similar to that of PapohakuBeach in that the numberof individuals, numberof species, and biomasswere greater in ax'easnear steepdropoffs and areas of rich coral growththan in areaslacking substan- tial vertical relief Table 9!. Carnivores outnumbered both herbivores and omnivores almost two to one by numberof species present and three to one by numberof individuals Table 10 and Figure 3!. Thebiomass of carnivores wasalmost twice that of herbivores and omnivores combined Figure 3!. Specieswith a high frequencyof occurrenceincluded Suj'@@men hu sa, Co~sgaunoxdi,, ThaZassoma dupemeyx', Pancpaneua muZtx faaeiatue, So~a sp., NasoZx,tuxatus, Paraoimhites aroatus, CLxetodon kleinx',, Ctenoohaetus stxigosus,ZancZus eanesoens, and Zebx'asoma Qm!esoens see Appendix H!. Thirty species of desirable food fishes with a total bxomassof 73.472kg were recorded Table 11!. This correspondsto roughly 30 percent of the total fish biomass off Molokini island. Figure 2. Transect locations off Molokini Island. See Table 8 for specific locations.!

TABLE 8. TRANSECT LOCATIONS OFF MOLOKINI ISLAND

Transect Latitude Longitude Direction of Transect

1 20 38'16» N 156 30'00» W 03.5 T 2 20 38'09» N 156'30~02» W 3455, T 3 20 38'16» N 156 30'03.5» W 208.5 T 20 38'06» N 156 30'00» W 122' T 20 38' 14" N 156 29'49» 83' 5 6 20 38'08.5" N 156 29'50.5» W 07 T 7 20 38'05.5» N 156 20'53" 42 8 20'38~10» 156 20~58» W I 31.5 T 9 20'38~10.5» N 'I56 20'59.5» W 157 5 T

16 TABLE9. FISHBIOMASS BYSUBSTRATE TYPEOFF MOLOKINI ISLAND AverageNo. AverageBiomass Substrate and Transect No. of Species Per Transect Bottom Type PerTransect kg/1,000m~!

Abrupt dropoffs with 28.2 13,82 I i ve cora 1 g row th 3, 5 L i ve co ra 'i g rowth, 30. 4 ]4,77 gradual slope 2, 4, 6, 7, 8 Sparse coral growth, coral rubble, 18.O 6. 36 silt, and algae

TABLE10. FISHBIOMASS BYFEEDING TYPEOFF MOLOKINI ISLAND

R of No. of No. of Biomass Feeding kg/1,000 m2! Total Biomass Species Type Individuals 12 77.13 32. 9 Herbivore 1,655 11 6.05 2.6 Omnivore 308 62 151.03 64. 5 Carnivore 5,717 85 234.21 100.0 Tota 1 7,68O

EOO

g IDD

X 100

Hororsoros Orooivoros cororvoros T0TII. PEEDIIOOTYPE Figure3. Fishbiomass byfeeding type off Molokini Island TABLE 11. BIOMASS OF DESIRABLE FOOD FISHES OFF HOLOKINI ISLAND

Species BiofAsss kg!

Myrip~stis murdjan 3.002 Caranx sp. Scomberoides lysan 0.149 Decaptems pinnuLatus o.o74 Apron virescens 0.917 Mulloidichthys vanicolensis 3.374 Mu2loidichthys f Lavo2ineata 0.024 Pavupeneuschryserydr os 1.18o Pctvupeneusp2eur ostigma 1.035 Pavupeneusporphyr eus o.283 Pampeneus mu2ti fasciatus 8.187 P~peneus bi fasciatus o.o28 Abudefduf sor'didus 0.161 Abudefduf abdominalis 1.091 Cir'r hitus pinnulatus 0.009 Chei Lio inemis 0.0»

Bodianus bi2unuLatus 0. 207 C'hei 2i nus r hodochr ous 1.886 C'oris f2avovittata 0.100 Seams sp. Seams per spici l latus o.428 Acanthuses triostegus o.»4 Acanthu~s dussumieri 3.o62 Ctenochaetus strigosus 14. 661 Naso 2ituratus 26.647 Naso hexacanthus S.432

Naso brevirostris o.683

Naso unicormis 0. 192 Me'Lichthys niger 0.075 Eyphosus ocy~s o. 46o

Tata I 73.472

18 Six other species of fishes were seenat Molokini Island by DAPper- sonnelduring the four-daysurvey. Theseincluded three individuals from the familyMobulidae manta rays! with an estimatedwing span of 3 m,one Aetobatus~nazi spottedeagle ray! with an estimatedwing spanof 1.5 m, 150to 200Decapte~ pinna'Laths opelu! at 18 to 20 cm,one Abno- taxisgz'andocu'Lis mu! at 38cm, two Atutera saripta 'o'ili lepa!at 50 cm,and three sharks 1.5 to 2.5 m fromthe family Carcharhinidae.

Substrate Thenine transect surveysconducted off Molokini Island revealeda total of six substratetypes hardbottom, rubble, sand,silt/mud, algal mat,and live coral!. Of the total areasurveyed, 18.4 percent was made upof live corals1 species!with other substrate types making up 81.6 percent Table 12!. Coral coverreached 35.4 percentwith diversities of up to eight spe- cies in someareas Table13! . PoritesLobata, Poci llopora mearu&ina, andNontipora ve~cosa were common throughout the areas surveyed and occurredwith a frequencyof 1.000 Table 14!. Abntipo~yermLLi and Pontescompressa were also fairly common,both occurring with a fre- quencyof 0.889. The other live coraisoccurred with frequencies of 0.111 or 0.222. Of the live corals recorded, P. lobata accountedfor 54,7 percent, Pocilloporameandrina 15.8 percent, N. uezmccosa10.5percent, and the total of the less prominentcorals 3.9.0percent Table14!, Ontransects 4 and8, Pontes lobata representeda large proportion of live corals, 20.8 percent and 19.5 percent, respectively. Thesubstrate around Molokini island consistedlargely of rubble deadcoral!, hard bottom basalt!, live coral,and sand Tables 12 and 15!. Transects3, 5, and6 showedlarge percentages of coral rubble at 70.8percent, 72.5 percent, and 50.7 percent, respectively gable 12!, Largepercentages of hard bottom in theareas near the sea cliff tran- sects2, 4, 6, and7! hadpercentages ranging from 28.6 percent to 68,2 percent.Sand pockets occurred frequently and accounted for thelarge percentageof sandcover in transects1, 2, and8, Of the outerareas surveyed, transects 3 and5 hadthe highestsimi- larity indexof 0.8210.Transects 4 and 7, whichwere next to the sea cliff, showeda similarity index of 0.7810.Transects 1 and 8, both occurringat depthsof over17 m, showed a similarity index of 0.7240, See Appendix I.!

Twelvespecies of algae--10"known" one "unidentified" blue-green algaepossibly Schizot&ix sp., andone "unknown"--were recorded off Molo- kini Island. SeeAppendix J for a completelisting of algal speciesand substratetypes.! Thefigures for frequencyof occurrenceand the cor- respondingpercentage of cover for an11.25 m~ total transectedarea show that encrustingand filamentous red algaeare dominant Table 16!.

19 .TABLE12. PERCENTAGEOF SUBSTRATETYPE BY TRANSECTOFF MOLOKINI ISLAND

Hard Bottom Rubble Sand Silt Algal Mat Coral z! a!

10. 5 38.0 42.2 0.0 0.3 9.0

10.0 55.5 0.0 31. 1 0.0 3.4

0.0 70.8 0.0 .0.0 0.0 29.2

48.2 14.4 7.5 0.0 29.9

0.0 72 5 9.8 0.0 0.8 16.9

28.6 50- 7 3.0 0.0 0.5 17.2

68.2 13. 8 8.2 0.0 0 ' 5 9 3

5 7 38.i 20.8 0.0 0.0 35.4

8.8 0 ' 0 0.0 19.2 53.6

6.6 18.4 Mean 24. 1 35.2 13.6 2.1

TABLE13. PERCENTAGEOF CORAL COVER AND NUMBER OF CORAL SPECIES PER TRANSECTOFF MOLOKINI ISLAND

4 of No. of Transect Coral Cover Coral Species

9.0

10.0

29. 2

29.9

16.9

17.2

9.3

35.4

8.8

20 TABLE 14. FREQUENCYOF OCCURRENCEAND PERCENTAGEOF COVEROF LIVE CORAL OFF HOLOKINI ISLAND

Frequency 4 of Total Substrate Species of Occurrence

Porites 2obata 'I . 000 54. S

Poci 22opora meanch'ina 1.000 15. 8

Monti pora verrucosa 1.000 10.5

Montipora verri22i 0.889 9.3

Pontes compressa 0.889

Poci L7opora epdouzi 0.111 0.8

Leptasbrea bottae 0.222 0.4

Montipora patu2a 0. 111 0.1

Montipora fLabe22ata 0.222 0.1

Psammocora Stephanaria! 0.111 trace ste22ata

Pavona varians 0.111 trace

TABLE15. FREQUENCYOFOCCURRENCE ANDPERCENTAGE OFCOVER BY SUBSTRATETYPE OFF HOLOKINI ISLAND

Substrate Frequency C of Type of Occurrence Total Substrate

Coral 1.000 18.4

Rubble 0.889 35 3

Hard bottom 0.667 24.1

Sand 0.778 13.6 Algal mat 0.667 6.5 2.1 Silt 0 ~ 111 TABLE 16. FREQUENCYOF OCCURRENCEAND PERCENTAGEOF COVER BY DOMINANT ALGAE OFF MOLOKINI ISLAND

Frequency of Type of Alga 4 Cover Occurrence

Encrusting Rhoctophyta 1.000 17

Filamentous Rhodophyta 1. 000 32 Diotyota sp. 0.888 6 Fi 1 amentous Chlorophyta 0.889 5 Cora l l inaceae 0. 718 "Unknown" 0.667 62 7 Rim'odictyon sp. 0.556 Filamentous Phaeophyta 0.556

Delicate filamentous reds, and other filamentous algae in general, growing on and between fingers of live coral and on rock and hard sub- strate as encrusting forms, were the dominant algae. Many of the fila- mentous algae were short, usually not greater than 1 cm long, with a cropped appearance. Members of the family Corallinaceae, mainly crustose, encrusting forms, and Cora'LHna sp., were observed both on and near transects and com- prised 8 percent cover.

Fleshy frondose algae were either small in number or not present . Even the Dictyota sp. observed grew with a somewhat reduced thalli, con- forming and adhering to the substrate usually rock!. Very few Palea sp. Diatyoep'IM&a oavemoaa, Neomemesp., and HaHmeda sp. were observed. A large stand of "toilet paper algae" was noted on the first 40 m of transect 9, but was not recorded as it did not occur within the quadrats.

Discussion

Fish The number of fish species present in an area is influenced by several factors, e.g., currents, water clarity, and the nature and complexity of the substrate Brock, 1954; Bardach, 1959; Randall, 1963; McVey, 1970! . The major contributing factor is probably the latter, because it incxeases the number of habitats providing food, shelter, and other essential requirements. This relationship of number of species to substrate was seen off Molo- kini Island where the areas of extensive coral growth and steep dropoff had higher biomass than the areas of sparse coral cover and little verti- cal relief. Transect 9, an area of sparse coral growth and little vertical

22 relief, wasunusual in beingthe onlyarea where this typeof condition existed. Thispocket of coral rubble,sand, and dense growth of algaewas surroundedby rich coral coverin nearbyareas, indicating that somesort of ecological disturbancehad occurred. Diverswho frequent the islandreported that therewere several intact bombslocated within the crater,probably as a resuLtof military training exercisesat thenearby island of .Nhile transecting, DAP divers cameacross two of theseintact bombswhich present potential dangersto unwarydivers and should be avoided Figure 2!. Fromhis studiesat PokaiBay, Hawaii, McVey 970! foundthat there wasa relationshipbetween the numbers of fish countedand water clarity. Unusuallyclear waters off MolokiniIsland perhaps enhanced observation of fishesand a moreaccurate or highercount may have been possible than in locations of poorer visibility. Currentspresent off MolokiniIsland may provide adequate amounts of oceanicplankton. The feeding by reef-dwelling fishes on this plankton asit is beingswept by canrepresent a substantial import of energyinto the coralreef communityaswas found by Randall's963'! study in the VirginIslands and Emery's 968! studyin the FloridaKeys. Thecarnivorous fish populationwas significantly greater than the herbivorousand omnivorous fish populationscombined. Randall 963!, in studyingthe fish population of artificial andnatural reefs in the VirginIslands, and Talbot 965!, in his studyof coralreefs in Tangan- yika,also noted this disproportionate occurrence. Onereason for this is that, althoughthe herbivore biomass is probablyas greator greater thanthe carnivore biomass onreefs, the largest proportion of thereef herbivorebiomass is composed of mollusks, crustaceans, and annelids, with fishmaking upa smallerportion Bardach, 1959!. Another reason is that ediblealgal forms could become less conspicuous andhighly calcified due to selectivepressures such as strong surge and grazing invertebrates see"discussion" section under "Algae" !, limiting the herbivorous fish populationbythe amount of available food. Still anotherreason is that carnivoresare more diverse in their feedinghabits Fhrlich,1975!, thus allowing moie species to occupythe samearea. Fishingpressure has a considerableamount of influence upon the abun- danceof fish withina givenarea. Hooking,trapping, netting, and spear- ingcan be devastating to the resident population of reef fishes Randall, 1963!.Off Molokini Island, a highproportion 1s4 percent! of thetotal biomassencountered was composed of desirable food fishes, indicating little fishingactivity there. Fisheswere very tame and showed little signsof "spooking,"a trait usually associated withpopulations sub- ject to intensefishing activities.

Substrate Therelative distribution and abundance of hermatypic corals off Molo- kini Islandwere influenced by local circulationand substrate types. Theisland's shape is suchthat thenorth side is opento theocean and I is subjected to various currents and surge Palmer, 1930!. The high degree of species diversity and bottom topography is characterized by the shape and location of Molokini Island. Of the 11 coral species represented, Poritee lohzta, PooiHopora meand2ina, and Montipora ver'mooea occurred at frequencies of 1.000, suggesting that these species of coral thrive in areas where surge and currents are present Maragos, 1972!. P. Lobar showedthe highest per- centage of cover on transects 4 and 8. These transects occurred in areas subjected to surge and abrasive , conditions for which Pomtee spe- cies are particularly adapted, as found by Cary 931! in his study on Tutuila reef, American Samoa. Transect 3, which was subject to currents and high surf action, had a high percentage of cover 0.7 percent! of the coral species Poecil.Lopora meandzina. This occurrence can be explained by the greater preference of the genusPoli Llopoza for areas of greater surf action than other genera of coral Maragos, 1972!. Montipora vermLLi and M. ve~cosa were usually found in low encrus- tations around Molokini Island. These species take low forms in areas where wave energy is high Maragos, 1972!. Around Molokini Island, the regional distribution and abundance of corals were effected by other substrate types, mainly rubble coral!, hard bottom basalt!, and isolated pockets of sand. The outer, steep-sloped reef areas transects 3 and 6! had large amounts of loose coral rubble, suggesting exposure to great mechanical stress.

Erosion of the inner sea cliffs, as noted by Palmer 930!, was caused mainly by wave action, accounting for the large percentage of hard bottom basalt! in these areas. Transects 2, 4, 6, and 7 had large amountsof loose basalt rocks that were unsuitable for coral growth. Isolated pockets of sand occurred at transects 1, 2, and 8 and were present in less abundance at other high-energy areas. The reasons for the large percentage of. algal growth and siltation at transect 9 remain unknown. This area seemed denuded of live corals and other typical substrate. It has been noted by Wood-Jones 907!, in a study at Cocos-Keeling Atoll, that algae may cause sediments to accumu- late in protected pockets, eventually leading to restricted coral growth.

It seems that the dominant algal forms off Molokini Island prefer low- energy environments and hard substrates since no bottom surge was noted in the crescent area and the algae grew on rocks and live or dead coral. The light growth of filamentous algae betweenfingers and on the heads of live corals provided evidence for probable competition for space Maragos, 1972!.

24 Perhapsthe thalli observedat depthsof 14 to 20 m weredeepwater forms. Harger972! notedthat Phaeophytaat Waikiki tended,with increasingdepth, to be slightly morecalcified, smallerin size, andmore often of prostateform, closely adhering to the substrate,rather thanthe erect, free-fLoatingform. Healso notedthat an apparentincrease in grazingpressure with depth might contribute to a changein formsince small, inconspicuousalgal formsare moredifficult to removefrom the substrate. Thismay explain the xeducedsize and adherance of the Diatyotasp. observed, and also the dominantnumber of filamentousand encrusting algae over large, frondose types. Althougha smallpercentage of the fish populationoff MolokiniIsland is herbivoxousor omnivox'ous,a large population of seaurchins is pxesent. Seaurchins graze on algae,preferring crusty algae, filamentous browns and reds, and larger, leafy forms such as Padinasp. The low diversity and high occurrenceof "unknown"algal generaaxe probablydue to theobserver's insufficient knowledge of algal forms found off MolokiniIsland, as there weremany small and/orfilamentous forms, making identification very difficult.

Conclusions

Thewaters off Molokini Island truly representa uniqueand beautiful marineecosystem. Areas within the cratexexhibit a greatvariety of marine life in its natural environment,and the pristine quality of the waterpro- videsfor safeand favorable diving conditions. Thegreat abundance and overall friendlinessof the fish populationare unmatchedeven by suchareas as the HanaumaBay Marine Life ConservationDistrict on Oahu. Yet, cautionmust be exexcisedwhile diving off MolokiniIsland because of thepotential dangex of live ordnanceand strong cuxrents outside of the crescentarea. Surfaceconditions, too, at timesbecome rough and warrant attention.

SUMMARY

Often too little concernis paid to the sometimesfragile marineeco- system.What is neededis to facilitatea betterawareness andappreciation byproviding information dealing directly with the marine resources within givenareas. With this in mindthe DAP members gathered baseline data and establishedtransects for future surveysat PapohakuBeach and off Molokini Island. ACKNOWLEDGMENTS

The Data Acquisition Project teamthanks Dennis T.O. Kamof the Hawaii Coastal Zone Data Bank for his help and advice with statistical computer analysis; Dr. S.A. Reedof the ZoologyDepartment at the University of Hawaii for providing technical advice in the area of substrate photography; Jeff Hunt, an instructor at WindwardCommunity College, and Marilyn Dunlop for their assistance in algal identification; John McMahon,Director of the Marine OptionProgram, University of Hawaii, for timely advice and commentspertaining to the manuscript;Dr. JamesShaklee of the Zoology Departmentat the University of Hawaii for reviewingthe manuscript;and CaptainWillard Austin and the crew of the schoonerMachias for their excellent surface support. A very special thanks to Donna K. Noborikawa for donating so much of her time in organizing the student team and pro- viding the draft figures in the manuscript. Funds for the PapohakuBeach and Molokini Island surveys were provided by the State of Hawaii Office of the Marine Affairs Coordinator and the University of Hawaii Sea Grant College Program.

REFERENCES CITED

Bardach, J.E. 1959. "The summerstanding crop of fish on a shallow Ber- muda reef." L mnology and Oceanography1;77-85. Brock, V.E. 1954. "A preliminary report on a methodof estimating reef fish populations." Jceumtl of wildlife Management.18:297-308. Cary, L.R. 1931. "Studies on the of Tutuila, AmericanSamoa, with special reference to Alcyonaria." Publications of the C'cumegie I'nstitution 413:53-98. Doty, M.S. 1973. "Tropical algal ecology and conservation." Nature Con- semation in the Paci fic. Australian National University. pp. 183-196. Emery, A.R. 1968. "Preliminary observations on coral reef plankton." Limnology and Oceano' aphy 13:293-303. Erlich, P.R. 1975. "The population biology of coral reef fishes." Annual Review of Ecology and Systematics 6:211-247. Finckh., A.E. 1904. "The atoll of Funafuti: biology of the reef-forming organisms at Funafuti atoll." Royal Society of London, Co~2 Reef Community 5:125-150. Gosline, W.A., and V.E. Brock. 1960. Handbookof' Hawaiian Fishes. Honolulu: University of Hawaii. Press. Grigg, R.W. 1972. "Someecological effects of discharged sugar mill wastes on marine life along the Hamakua coast, Hawaii." Vates Resou2ces Seminar Series 1llo. 2. Water Resources Research Center, University of Hawaii, Honolulu. pp. 25-46.

26 Harger,B.'W.W. 1972. "Studies on benthic algal flora seawardfrom the reef flat at Waikiki, Oahu,Hawaii." Thesis for Masterof Science Degree,Department of Botany,University of Hawaii,Honolulu. No. 1058, 185 pp. Hobson,E.W. 1965. "Diurnal-nocturnalactivity of someinshore fishes in the Gulf of California." Copeia 3:291-302 ' Manton,S.M., and J AT.A.Stephenson. 1935. "Ecologicalsurveys of coral reefs: Great Barriex' Reef expedition 1928-29." Scientific Reports 30!:273-3].2. Maragos,J.E. 1972."A studyof the ecologyof Hawaiianxeef corals." Ph.D.dissertation, Department of ,University of Hawaii, Honolulu. No. 501, 290 pp. McVey,J.P. 1970."Fishery ecology of the Pokaiax'tificial reef." Ph.D.dissertation, Departmentof Zoology,University of Hawaii, Honolulu. 267 pp. Neal,M.C. 1930. "Hawaiianmarine algae." BerniceP. BishopMuseum Bul'Letin No. 67. 84 pp. Odum,H.T. and E.D. Odum. 1955. "Trophic structure and productivity of a windwardcoral reef communityonEniwetok atoll." EcologicalMono- gmphs 25: 291-320. Oishi,F. 1974."Papohaku Beach survey." UNIHI-SEAGRANT-WP-00-14. Universityof HawaiiSea Grant College Program, Honolulu. 13 pp. Palmer,H.S. 1930, Geo'Logyof Molokini. BerniceP. BishopMuseum, Honolulu. Vol. 9, No. 1 Randall,J.E. 1963."An analysis of the fish populationsof artificial andnatural reefs in the Virgin Islands." C~bbeanJo~l of Science 3:31-47. Smith,S.V., K.E. Chave, and D.T.O. Kam. 1973. Atlas of KmeoheBay: a z'eefecosystem undex stress. UNlHI-SEAGRANT-TR-74-04.University of Hawaii Sea Grant College Program, Honolulu. Sorenson,T. 1948."A method of establishinggxoups of equalamplitude in plantsociology based on similarity of speciescontent: and its applicationto analysisof thevegetation of Danishcommons." Biol ogiske Skxi f fex' 4!:1-34. Talbot,F.H. 1965."A descxiption of the coral structure of TutiaReef TanganyikaTerritory, East Africa! and its fish fauna."Proceedings of the ZoologicalSociety of London13:431-470. Wood-Jones,F. 1907. "On the growthforms and supposed species in corals." Proceedingsof the ZoologicalSociety of Lonely:518-556.

27 avvmorccs Appendix A. Scientific, Hawaiian and/or English Namesof Fish Seenat PapohakuBeach and Molokini Island

Feeding Habit a Family Scientific Name Hawaiian and CorrmonNames s!

ACANTHURI DAE Acrmthurus achi l.les paku'iku'I, achilles tang Acanthurus dussunneri paiani Acanthurus nigro fuscus mal'i Acanthurus nigr oris maiko Acanthurus olivaceous na'ena'e Acrrnthurus trioetegus manini, convict tang Acanthurus xanthopterus pualu Ctenochaetus stz igosus ko I e Paso bz eviros tris kala lola Naso he@acanthus opelu kala HCH 0HH iyaso lituratus kala, c'Iown tang Haso unicornis kala uni cornfish Zanclus carl scene kihikihi, moorish idol Zebzasoma flave scene lau'i-pala, yellow tang Zebrasoma veli ferum sai Ifin tang

APOGONIDAE Apogon kallopterus 'upapalu

AULOSTOHIDAE Aulostornus chinensis nunu, trumpetfish 'o' i 1 i BALISTIDAE Cantherines dumer ili HeLi oh thy s vi dua humuhumu-hi'u-kole HeLichthys niger humuhumu-'ele'eie Pervagor spi losorna 'o'ili uwiuwi, orange-tailed f1 ledfish C0 00 Rhinecanthus r ectangulus humuhumu-nukunuku-apu'a Sufflamen bursa humuhumu-umauma-lei Sufflamen frenatus humuhumu-mimi pao'o kauila, o'opu pao'o BLENNIDAE Etallis br'evis

CARANGIDAE Cararnn mel ampygus omi I u Decapterus pinnulatus opelu, mackerel scad Scomberoides lyean lac, leatherback runner 'Prachuropscrumenophthalmus akule, haialu, aji, bigeye scad

Potter's angel CHAETODONTIDAE Centropygepotteri Chaetodon auriga threadfin butterfly Chaetodon fremb li i blue-stripe butterfly Chaetodon kleini Chae Codon l.unula k ikapu, masked/raccoonbut ter f I y Chaetodon mi liaris lemon butterfly Chaetodon multicinctus pebble butterfly Chaetodon ornatissimus kikakapu, ornate butterfly Chaetodon quadrimaculatus four-spot butterfly Chaetodon trifasciatus Chaetodon unimaculatus kikakapu,one-spot/teardrop butterfly Forcipiger fLavissimus lau-wi liwi Ii-nukunuku-ol'oi Heni ochus acuminatus false ki hi ki hi, poorman' s idol

pi liko'a CIRRHITIDAE Ci rrhi tops fasci at us Cirrhi tus pinnulatus po'o-pa'a, o'opu kai Paracir r hi tee arnratus piiiko'a Paracirrhi tes for steri hi lu pili ko'a, pi I i ko' a

white eel CONGRIDAE Conger sp cornetfish FISTULARIIDAE Fistularia petimba ala'ihi HOLOCEHTRIDAE Adiorym sp. F Lcrrrrneosarnmar a lrfyripri ates rnurdjan u'u, menpachi nenue KYPHOSIDAE Kyphosusbigibbus

Appendix A. Scientific, Hawaiian and/or English Namesof Fish Seenat PapohakuBeach and Molokini Island continued!

Feeding Habit* Fami I y Scientific Hame Hawaiian and Co>mr>onHame s! i.t>>SRI DAE Anampseschrysocephalus An>r»>pensr uvieri snowflake wrasse, Uncle Sam wrasse rrr>ji.rr>us 7>i7 »>rrr 7 ot>re a'awa, table boss >ei 7.'».'>." rh>r>inchr>>ns po 0 v 'he jr j»er»>is kupou'pou, ciqsr wrasse 'nri s hi > 7,7 i arri Coris f Lavovi t tata hiiu ;nr'is gaimnrdi loio .orr s ver>t>sr.a Ci >mpho»»sv>r»',us 'aki- lolo, hinalea i ' iwi, bird wrasse Halichoe>ws orr>at tissumrs Ia'o, 'ohua, pa'awela Hemipteronotus taeniourus dragon wrasse CC CC lwbr'>ritacrolharyn>ladon geoffreyi Potter's wrasse Stethojulie balteata 'omaka Th>zjlasomaifrr> LLi eui hinaiea luahine, mongoose wrasse Thalasoma duperreyi hinalea Iauwi I i ~ 'a'ala' ihi '7'halasoma purpureum 'oiani, 'olal i, palea'a, 'awela, hou

LUTJAHIOAE Aprion virescens uku twt,jan>>s 7 asmira taape, blue-line snapper wekw' ul a HULI.IDAE Hulloidichthys flavolineata hfuLLoidichthys vanicolensis weke'a'a, spot weke Parupeneus br'.fasciatus munu Parupe»euschryserydzos moano kea mona kali Parupeneus multifasciatus moano, moana Parupeneusplerrrost-igma malu Parupeneus por7hyr>..us 'kumu

HURAEHIDAE :ymnothborax meleagris puhi moa,mamoa, waa, oopakaku,boxfish OSTRACIOT>DAE Ostraci.on rneleagris lllaoalao,memo, sel'geant maJor POHACEHTRIDAE Abudefdu f abdominalis C black memo Abudefduf sindonis 0 Abudefdu f sordidus kuplpl, grey memo Chromic leucurue Chromic ovalis C C Chromia vanderbi lti Chr m>is verator black damsel Ltasryllus albiseLla 'alo'llo'I, three-spot Eupomacentrusfasciolatus PLectrogLyphidodon ir»paripennis PLectroglyphidodon johnetonianue uhu SCARIOA'E Seams perspici Llatus Scarus sp. uhus leaf fish SCORPAEHIDAE Taenianotue triacanthue 9 rouper' SERRAHIDAE Perudanthias thompsoni 'o'opu-hue, maklmakl,keke TETRAODOHTIDAE Art>thron hispidus Aroth>on r>releagris C0 0C Canthigaster janthinopterue sharp-back puffer Cant higadter coronatue

*H ~ Herbivore; C ~ Carr>ivore; 0 w Omnivore AppendixB. Presence/AbsenceandFrequency of Occurrenceof FishSpecies By Transect Stations at PapohakuBeach

Transect Station prequencyof Species I 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Occurrence AS AS AB AS AS AB AB AS AB AB AB AB AB AS AS AB AS AB AB

ACAHTNURIDAE Acanthuruaap. X 0. 027 A, duaateeieri. X X X x x x r 0.216 A leuntUeu'erua x 0.027 X X x r x 0.243 A. rtiani'fuaoue x x r x 0.081 A. ntgirilca X X X x x X r 0.2 70 nl i vnoeus x r r r x 0. 35'I A Lriiiatafgue X X X X x x X X X X rx A. ecnthoptarus X 0,054 Ctenoohoatua e trig osue O.'108 le braetera oali famer 0.027 APOGOHIDAE Apiyurt ep. X X X X X 0. 162 A. hal l opterwe 0.027 BALISTIDAE Balietid Sp. X X X 0.081 Neliohthye niger 0.027 Petnruyoaepiloeorea Xx xr X X X X X X 0.405 Bhinsoan thus reotangulus X 0.027 Buff lamenbursa X X X X 0. 162 S. franatus x r 0.054 CHAETODOHTI DAE Chastodon pot teri X X 0.054 C. frambbii X X X X X X xr 0. 216 C. kleini X X 0.054 C. miliarie xx r X X X r x xx XX XX X X X X xr x 0.595 C. multicinotue x r X X 0. 162 C. ifuadrr'auroulatue X X X 0.081 poroipigar flauissvnue X 0.027 Hanioohueacuminatue X 0.027 C RRHITIDAEI Cirrhitops faeoiatus X X 0. 108 Cirrhitua pinnulatue 0.027 Paraoirrhi t st aroatue X X X Xr Xx 0. 189 COHGRIDAE Congertp. 0.027 -HOLOCENTRIDAE Adiorye sp. x x X X 0. 189 LABRIDAE Labrid sp. XX'X 0.216 Anampees chryeocephalue X r x X X X 0. 189 Bodianuapilunulatua X X X X X X X X 0.243 Chailinus rhodochroue X X X 0. 135 Corie ballieui X 0.027 C. gaimardi X X X X 0, 108 C. can@etc XX XX 0.162 Hali oboeres oreuztissbmia X X 0.135 H taili p ca roric t ue taeniouru e XX XX X 0.135 Labroidea phthirophagua x x 0.054 tertr opharyingodon geof f royi x 0.027 gtathojulie buLteitt.it XX XX X X XX X X X XX XX Xr XX X 0. 541 Thnlaaacmritbitllraui X X X X X 0. 135 T. duperrayi X X X XX XX XX x xx x xx X X 0 757 T purpur'eum X X X 0. 081 LUTJAHIDAE Lu ,iorruskuamiro X X 0.054 Appendix B. Presence/Absenceand Frequencyof Occurrenceof Fish Species Hy Transect Stations at PapohakuBeach continued!

Transect Stat.ion Frequencyoi Species I 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 IS 19 Occurrence A 8 A 8 A 8 A 8 A 8 A 8 A 8 A 8 A B A S AS A 8 A B A 8 A 8 A 8 AS A 8 A 8 NVLLIDAE pule p .nttua i i!!uk i ttue 0.027 I . 'll I'ilsl?1'lt iros X X X X 0.108 k tI I i gtte .i t1 ue X X X X X x x x x 0.297 P. p I unvetigmu XXXXX X X XXXXXAXKKXXXXXXXXXX 0.616 I'. p?t'p!lj rene X X X XX XX X XX A!t XK XX XX XX XX X 0. 595 OSTRACIONTIDAE tot,t??.tivn ttluI au0 t 'i t 0.021 POIIACEIITRIDAE Abuds fdu f ah b?m?nal i e X X 0.135 Pleotmg I Aphidodon ?mparspent? so X 0.027 A. sindonis 0,021 A. eondith s 0,054 t.iuomis ottulis K X XX XX X K XX XX XX XX X XX x x x 0.622 ttanderbilti X X 0,054 Itaeosllus albieella X X X X X X 0,162 P. joiptetonlanue 0.021 Stegaetesfasoiolatue X XX XX X X X XX XX XX XX X X X 0.676 SCORPAEIIIDAE Tueniunotus tniuoanthue 0.027 SERRAIIEDAE P. thompeoni 0.027 TETRAODOHTIDAE At' ?litt'vtt!tt epidus X 0. 108 A, meleugtis X X X A 0. 108 C ttI hi Sastttn o tt'vtk it e X X X 0. 108 C. J tt thittvptente X X X XX XX X A 0. 297

34 AppendixC. PapohakuBeach Fish Data with Bottom Description by Transect

Fish Data Transect Ho. of Biomass Total llo. Depth Bottom Descript.ion Station Species kg/1000 m?! of fish m!

IA 0.898 26 Flat bottom, mostly sand, algae and si I t, 9 9.1 to 10.7 18 10 1.299 28 someboulders

2A 21 104 Coral and algae cover boulders. One cliff 4,80g 7.6 to 12.1 28 area with great depth change--sand patches.

2.457 121 First 50 m of transect over flat, sandy 3A l5 9,1 to 10.7 38 17 2.011 102 bottom. Somecarel. Bottom flat last 50 m

4A 8 44 Flat bottom~ some rocks or small boulders, .4g8 9.1 to 10.7 4e 15 0.647 6g sand-silt and algae.

6 Flat bottom, sand and algal eever. Area 5A 3 0 075 4,6 to 7.6 58 11 2.135 53 bare except near rocks.

0. 011 I Flat sand-silt with a few rocks 6A 6.1 68 0. 278 22

10 Flat, very si lty bottom. Hoch algal cover. 7A 0. 099 6,I to 7.6 78 0.043 '12

10 Bottom flat with occasional large rocks, BA 3. 619 78 4.6 to 7,0 Be 10 6.075 covered with mud and a Iqae Hard, flat bottom covered with sand and 9A 12 1. 074 30 10,7 98 16 1. 452 52 rocks inshore surf/surge zone, large wave-worn IOA 14 6.355 132 3.0 to 4.6 IOB i6 4.333 139 boulders with sand inbetween. Sand, hard bottom with pockets in rock, I IA 0. 958 97 4.6 I IB 0. 482 75 silt. Fiat sand-silt and coral rubble. I2A 13 0.235 31 9.1 128 12 0.348 28

122 Flat sand-si lt algal bottom with large 13A l9 1.193 9.8 138 17 3- 332 97 ledges. Mud and si it--no vertical re! ief or large 14A 0.001 10.0 148 O.OOI rocks Flat bottom very silty . 15A 16 2.622 68 12.2 158 17 0.84i Bi Hostly flat with sand-silt and algae, 16A 2,240 19 4.6 16e 3.738 20 some rocks. 62 Sand, algae, coral ledges 17A 18 1. 072 10.7 178 1.718 79 0 7'l3 Flat sand bottom, I BA 8 43 7,6 iBB 10 1.153 59 Flat sand and algae 19A 2 0. 003 2 4.6 198 10 0. 518 38

35 Appendix D. Comparison of 1974 Study With 1976 Study by Biomass and Number of Fish Per Substrate at Papohaku Beach

Papohaku 1974+ Papohaku 1976i.

Transects 1 through 4 Transects 1 through 4

Average Average Average Average biomass No. of biomass No. of fish per pel f ish per per transect transect transect transect

o.41 18.4 Omnivores o.48 10.9 o.26 8 ' 4 Herbivores 1.29 14.0 44.3 Carnivores 5.02 50. 5 1.13 7o.6 Tota 't 6. 79 76.3 1.80 Average No. Average No. of species' 15. 1 of species: 13.6

Transects 5 and 6 Transects 5 and 6

Average Average Average Average No. of biomass No. of biomass fish per per fish per per transect transect transect transect

0.02 1.3 Omnivores 1.25 3o.8 0.05 2.0 Herbivores 3 50 4o.5 0 57 17 3 Carnivores 1.08 55-5 o.64 21. 5 Total 5.83 126. 8 Average No. Average No. of species: 13.0 of species: 5.3

+Calculationsbased on 1974Papohaku fish data--trophic levels. The sum of transectsup and back was divided by numberof transects i.e., sum1A through4B wasdivided by 8!. i Calculationsbased on 1976 Papohaku fish data--trophiclevels. Thesum of transectsup and back was divided by numberof transects.There is no2B for 1976,so the sumof transects'IA through 4B was div'ided by 7.

36 «ICt 0 0

0 0 0 a «A

«3Ct «tt ~ A IA

0 tA0 I0 Oo co

«ICt «A0 0 oCt «A '0 0 o O 0 0 0 0O 000 th«A «A C« O 0 «I« 0 0 0 0 0 N IA n 00 CO 0 a a

0 0 0 0 «A0 0 Ct CACt g N OtOl NIA «A N I O CO I «0 0 0 0 0 0 0 0 0 0 0 0 «A 0 N Ih 0N N Ct N «O I O IA 0 0 0 0 c 0 Ct g 0 I «O 0 0 R m 4 0 0 0 0 0 0

0 0 R lh0 0 N 0 0 N CO t IAOl lAt t 0 0 CI 0 0 0 0 0 0 o 0 «I 8 NO IA I th ltl CO «IO CO CO I NCt 0N 0 0 0 0 0 0 0 0 0 0

o O 0 0 0 0 000 IA Ol N «A lh N «t 0 0 Olth 4 I 0 0 0 0 0 o o 0 0 0 0 0 0 0 0 0 R I 0 IA 8 3 00 IAI M CO ~0 IA 0 0 0 0 0 0 0 0 0 a 0

0 0 g O 0 0 g O 0 0 a g ill Ct 0 0 I 00 I IA0 I/l 0 0 0 4 lh 00 0 ' 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 R Ol CO N IA IA COn I Ol I«A I COO IF CO 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 CI «I '0 o 0 ~ g 0 0 o O P O «3 0 «A COI t««N I IA ~ A N 0 0 o 0 0 o 0 o o 0 0 0 0 CI 0 0 0 0 0 o 0 O g 0 0 0 o r- I «A 0 I Itl O0 COCO «I CO I I 4I 0 0 0 0 0 0 0 0 0 0 0 0 0 Cl 0 0 0 0 0 CI 0 0 0 0 0 N0 0 0 0 0 0 g 0 00 0O OI Ch «A 0 I/l Ch O IXI Ol 0 IFl «A «Ct I 00 I 0 0 0 a «I o 0 0 0 0 a a 0 0 0 0 «I 0

I « th -0 lh 4! P IO Ol ««CA AppendixF. Frequencyof Occurrence and Percentage of Cover 3.75 m2! for Algal Speciesand Substrate Types at PapohakuBeach

Frequency Cover Species of Occurrence

Phylum Cyanophyta Lyngbya sp. 0.158 Symp7oca sp. 0.053

Phylum Ch'iorophyta Dictyospheria cavernoea 0.053 1 1 Dictyospheria vezs7uyeii 0.421 Microdictyon sp. 0.895 33 Cladophoropeie sp. o.789 Codium sp. 0.053 61 11 HaLimeda sp. o.579 Neomerie sp. 0.158

Phylum Phaeophyta Fi 1 amentous--genus undetermined 0-053 Encrust ing--genus undetermined 0.053 1 1 Family Dictyotaceae 0.053 1hctyopterie sp. 0.947 152 Dictyota sp. 0.526 Padina sp. 1.000 128 Sazgaeeum sp. 0.526

Phylum Rhodophyta Genus undetermined from t ransect 6 0. 105 Liagora sp. 0.211 Ga7azama sp. 0. 105 Amphizoa sp. 0-053 o.158 1 1 Coral lina sp. Gracilaria sp. 0.053 Spyzidia sp. o.158 Centzocerae sp. 0.316 Amaneia sp. 0. 158 8 1 Laze encia sp. 0.211

Phylum Undetermined un i dent i f i ed 0.053 unidentified il o.158

Substrate Type 0.316 sand 5 2 silt o.474

38 Ig Vl Ig L0V Ig Ig I I 0 0V 4I Ig 0 Vl I 0 V O V 4I 0E D 0 Vr E0 8 Vl C 30 l7> L 0 ~ L cg Ig 4I 4I Vl Ih ICI C 4I 3 I Vl Ig 4I M 4I M 0 CB Ig 0 O V O ~ C Ig I/I L II- 0 0 0 0 Ig CD Ig Ig L0V 4I O L V L 4I 0 l 0 L 0CL Ig D D I 0 CL 0 II- O 0 V D D 0 C 0 0 D Ig C Ig C 0 cg 4I L Ig Ig O V I O 4I D D D I L 0 0 CIIL 0 V 0 0V 4I O 4! 4I 4I 0 Q L V Vl I o ~ V CL CII Ig L Ig D Ig 4! 4I D 4I D L C ! D ! D 0 Ig D C0I4I L D 0 ~ 3 0 O. 0 L cg 0 L O I/I L Ig «4: cg I/I O «f. V C0V

N N N O

0 0 V 0 0 0 0 V a. E 4! ~ N Cl O

D4! 4- 0 r Z Ih CO O' O CO 0 ~ W A O N 4 N% N lA O CO rh 4- & CO Ccg Ig a ~ N «- D 0 0 I- 0 3 V

V cVE I/I VI I Vl O cg O LA ~ NN O W O N N O ~ CO ICI E O O h COM CO M N ~ 0 CO O CO O O ~ ~ ~ ~ Vl O 4' Q ~ CFl N NIg CON ~ Q N ~ Ol N N N O 4I CL Vl

Vl 4I C0 O' O N N QO xaOJ 0 4I ~ N D4I X CL Ch N N N CL I/I V Ig CI. 0 VIg V Ig V C V 0 4I 0 «4: CO VI ~- C Ig Ig I NN mM WW nm ~~ r r COCO I- In

39 AppendixH. Presence/Absenceand Frequencyof Occurrenceof Fish Speciesby Transect off Molokini Island

Trenscct Station I 2 3 4 5 6 7 8 9 Frequency Species AS A B A 0 A B A 8 A B A 8 A B A B

ACAHTHURIOAE X X X 0,167 Aaanthurus aohi lies 0.111 A. dussumisri X X x x X X X X X X XXXXXX 0.778 A. nigrafuscue 0.778 A. nigrorie X X X X X X X X X X X X X X X X X X X X X X X X X 0.667 A. alivacsue 0.056 A. triostsgus x X X 0,111 A. zanthoptsrus 0.833 Ctsnochastus strigosus X X X X X X X X X X X X X X X X X X X 0.222 gaea breviraetris 0.389 y, hssracanthus X X X X X X x X X X X X X X X X X x X X X X X x 0.944 .V. lituratus 0.111 N. uni carni s x X x x X xx xx xx xx xx xx 0.833 Zanclus oanescsns 0.833 Zsbrasamaflaaescsns X X X X X X X X X X X X X AULOSTOHIDAE 0.222 Aulostamus chinsnsis X X X X SALI STI OAE X X 0.167 Balistid sp. 0,056 Cantherhinss dMnerili 0.056 ideiichthys nigor X 0.1'l1 AI. tridua X X X X X XX X XX XX 0-556 Psrvagor spi l osoma 0.056 !%i nscanthus rsocongulus X X X X X X X X X X X X X X X X X X 1.000 ~flamen bursa 0.056 S. frena cue X SLEHHIIDAE 0.056 gaal liae brevis CARAHGIDAE 0.056 Cerangid sp, 0,056 Dsaapterus pinnulatue 0.056 Scambsroides lysan CHAETODOHTIDAE X X 0.722 Centropygs pat teri X X X X X X X XX XX X X 0.111 Chastadan auriga X X 0.278 C. frsmblii X X X X X X X X X X X X X XXXXX 0,889 C. klsini 0.389 C. lunula X X X X x x X X X X X X X 0.333 C. miliarie 0.556 C. multicinotus X X X XXXX X X X X X X X X X 0.389 C. arnatissimue X X 0.556 C. rluadrimaoulatus X X X X XX XX X X X X X X X 0.38g C. trifaeciatus X 0.056 C, unimaculatus X X X XXXX X X X X 0.667 Forcipiger fiaaissimus O. 167 iisniochus a~~inatus X X C RRHI IT I DAE 0.222 Ci r r hi t ape fascia tun X X X X X 0,056 Ci rr hi tue pinnu lc run 0.944 Para>irrhi ten arcatue XXXXXXXXXXXXXXXXX X X X X 0,222 P, foster i FISTULAIII IDAE 0.222 Fistularia petbnba X X X X HOLOCEHTRIDAE 0.056 Adiaryx sp. X X 0.111 Flantnsa eawnara 0.444 Nyripristie murdJ'an X X X X

40 AppendixH. Presence/AbsenceandFrequency of Occurrenceof Fish Speciesby Transectoff HolokiniIsland continued!

Transect Station Frequency Species I 2 3 4 5 6 7 8 9 of Occurrence AS A B A B A B A B A 8 A B A B A B

KYPHOSIOAE O. 111 Ayphosus bigibbus X X LABRIDAE 0. 389 Labrid sp. X X X XX XX 0.056 Anampseschrysocephalus 0.056 A. cuvisri 0.111 Bodianue bi Ltctulatus X X X X X X X X X X X X X X X X 0.778 Cheilimue rhodoohrous 0.056 Chailio insrmie X X 0.056 Coris bal lisui X 0. 056 C. flavovi t tata 1.000 C. gaimardk. XX XX X X X X XX XX XX XX XX X 0.056 C. vsnusta 0.222 Gortphosusvar ius X X X XX XX X X X 0.444 galichoeres ornatissimus 0.278 Labroides phthirophagus X X X X X Hacrop~odon 0. 278 gsoffroyi X X X X X X XXXX X X X X O.556 gtethojuli baLtsata 0.056 Thalaesctnaballieui X XXXXXXXX XX XX XXXXXX 1.000 1'. dupsrrsyi 0,056 1'. purpure um X LUTJANIDAE 0.056 Aprion vircscens HULLIDAE fsrLLoidichthys 0.056 flavolinsata X X X X 0. 167 AL vonicolsnsis 0.056 Barupensus bi fasciatus X X X 0. 'I 11 P. ohryssredros 1.000 P. multifasciatus X X X X X X X X X X X X X X X X X X X X X X XXXXX 0 556 ?. pleuroetigma 0 167 P. porphyrsus X HURAEHI OAE 0.222 CLymnothora»ms Lsogri s X X X OSTRACI OHT I DAE 0.056 Os~ msLscgris POHACEHTRIOAE 0.111 PomacentrId sp. X X X 0. 11'I Abudsfduf abdominaLie 0.056 A. imparipennis X X X X 0.222 ~OPCfhdue 0. 6'I1 X X X X X xxrtx Chromia Lsucurus 0.056 C. ovalis X X X X X 0. 222 C. vanderbilti 0.056 C. veratsr X X XX XX XX X X 0,611 Oascyllus albiseLLa 0. 667 gtegastee fcscio Latus X X X X X X X X X X X X Plsctrogly phidodon 0.61'I johnetonianus X X XXxXrt xx X X SCARIDAE X X X X X X X X X X X X X X X X X X 'I . 000 Scarus sp. 0. 222 gcarue perspioillatus X X X X TETRAOOONTIDAE X 0.056 Arothron hispidus 0.222 i. meleagris X X X X XXXXXXX X X Canthigaster 0,611 janthinopl,srus

41 AppendixI. Similarities BetweenCoral Speciesand Substrate Types BetweenTransects off Nolokini Island

Transect 2 3

1.0000

0.4890 1.0000

o.462o 0. 1000 1.0000

0.414o 0.6440 0.2880 1.0000 o.566o 0. 1840 o. 8210 0.3870 1. 0000 o.6o4o 0.4030 0.6300 o.616o 0.6710 1. 0000 0.3890 0.7110 0.2250 0.7810 0.3050 0.5450 1.0000 o.724o 0.3620 0.5530 0.546o 0.6360 0.6180 0.3630 1.0000 o. 261o o. 1060 0.2700 0.2220 0.2650 0.2640 0.1940 0.2670 1.0000

AppendixJ. NolokiniIsland, Algal Speciesarid Substrate Types wry% Frequencyand Percentageof Coverfor 11.25 m2

Frequency of 4 Cover Spec ies Occurrence

Phylum Cyanophyta Genus undetermined from transect 20 0.222

Phylum Chlorophyta Filamentous-genus undetermined o.889 Micxodictyon sp. o.556

Phylum Pbaeophyta Encrusting-genus undetermined 0-333 Filamentous-genus undetermined o.556 Dictyota sp. o.889

Phylum Rhodophyta Encrusting-genus undetermined 1.000 17 Filamentous-genus undetermined 1.000 32 Fami'iy Corallinaceae o.778 6 Corallina sp. O. 111 2

Phylum Undetermined Unknown o.667

Substrate Type basalt rock 0.333 rubble 0.778 sand o.667

42