South Pocffic Reglonol Envlronmenl Progromme

SPREP Reports ond Siudies Series no. 84

Palolo Deep National Marine Resenre: A Survey, Inventory and Information Report

bv Edward R. Lovell and Fouu Toloa Copright@ South Pacifio Reglsaal Euvlromrent Prograrnmq 1994

The South Facific Regioual Snvilonneat Frog,ranue autborisee the reproducti,on of this material" whole or io part, in aay for,n provided appropriatp aelinow.ledgpnreut is, given.

Origical Te.xt': E:aglish

Published in Novern'beJ 1994 by:

South Psclllc Reglonol Envlfonmenl Progrwnme P,O. Box 240 ApJo, Western Somoo

Printed by: Comrnerciql Prfniers Aplo, WesternSomo-o

P #.t 94-rc

Printed with finonciolossistsnce hom the Govemment of Aurtrollo

Loyout by Wecby lford, SPREP.

Edited by Joy Heenon.

Fronl cover photogroph: oeriofl phofo of Pololo Degp, w€.lt€rn:Sqinoo Oeooil4hs;rl 6f F.nvlrdnmEht,dnd iSors*\rotitn.

SPRDP Oataloguiagdn4ublieacion Date

Io-r,ell, Edlrard B. Palols Deep Natiimal M*iae Reserve: a Eun/ey, iuveutory aad in&rpction report / by' Edward B. IovBIl irnd Foua Toloa. 'Apla, Westexo $anoa: SPEiEF, 1994

iv, 88 p. ; gg om. (SFREP report and etudy ; no. 84)

Inclnrdes refereaces. ISBN 982.04-011r.9

l, Marine reserraeg . YYestera Sanoa 2" Erlr.iionneotel Fotlsy - Tllest€ta Sanoa I' Oonceil'adon af na'Lunl lesourtes. I. fitle

368.70723 South Pocific Regionol Environment Progromme

SPREP Report ond Study Series no. 84

Palolo Deep National Marine Reserve:

A Surv€y, Inventory and Information Report

by Edward R. Lovell Bialogical Consultonts, Fiji Lami, Fiji. and Foua Toloa Field Consultq,nt Apia, Western Samoa

Published in September 1994 in Apio, Western Sqmoo Foreword

The Palolo Deep National Marine Reserve was one of the first Marine Reserves to be established in the South Pacific. fts iocation adjacent to Apia Harbour, with easy accesg for Apia residents and tourists staying at the main hotels, makes it a valuable resource for Western Samoa. This document provides the results of the first detailed survey and inventory of the Palolo Deep. In the process of collecting the information, staff members of the Western Samoa Division of Environment and Conservation (DEC) were trained in basic marine survey, data collection and analysis techniques. Long term monitoring sites were established during the project to enable the DEC staff to record changes in the Palolo Deep Marine Reserve area over time. With the expected increased use of the Reserve by Apia residents and tourists alike, it is essential that this monitoring programme is in place. The information contained in this report was used by the DEC staff to prepare a detailed management plan for the Palolo Deep Marine Reserve. I would like to acknowledge the financial support provided by the Australian International Development Assistance Bureau through SPREP's Coastal Management and Planning Programme. In addition, I would like to thank the many people who have contributed to this project. SPREP encourages the Western Samoa Government to continue the process of developing and implementing a management plan for the Palolo Deep Marine Reserve.

I - tl/-fr^.C.,-. \) ,-- Vili A. Fuauao Director ,Gontents

1.1 Introduction .------..-- 1 1.2Genc*alAsseesnent'..'.',..'...... t....t...r..'!:.tir; 1.3, Er,rviionmental Impacts -..,.--'.-.-..1 L,4 Reeommendations..,-..,,:, !..... r...... !r.

2.L Fielcl Meth 2.2 Data Analyeis,....-...... -..q..'.,t..n....-.riii-..!...i,.'.., -.'..".-.^--..4

3. The eompositioo of Palo-lp Deep: Eabitat Deseriptio:r ...!...--.'-!.....---.-.r..i'r.,.-,-...... -..-- 6 3,1 Inehore Reef Flat... 8.2 Eastern Reef Ftat (f,raiuests !'la,b:;18a,b; 19a,b; Z&; ZA)...!i.r...--.-.r....,'...'.-,.-'.. 16 3..3 Western Reef Flat (Transecus 20a,b; 21a'd;2%,b)".,.,..- -.....-...--...'-.-21 9.4 Soirthern Palol,o Deep (traneects 4-6; ?-f t D-fG) .--...".24 3.5 Transverse Ridee and tbe Ssutheaet Margiu of the Main?alolo Deetrr (Transects 26;26ah;27a,b128a,b)."..,,.'-i.i'...,r'.,r.,,i --.--',29

3,6 Seaward Beef F-lat... ';-i,-!jii-i..r...i. g.? Seaward Reef Slope...... r...... '..,. --..--.-.-..---....33 3.8 Fish Strtrrney ...... rr.r-r.r...rj--:.-;;!r,q..'...,ra.. -."-".-...... 37

4.1 Reef Origin ...r'.i.r.'.i.;.1-r.,.r.'-rri....r.-.-.. -...... --. gB 4.2BlueHpIeorErubaynrent?...... '.."...t.-...;l.....a.... 4.3 EnvironentalCtange,. 4.4 History of the Palolo Deep for the last 39 years '5. Monitoring Progrramme, -;--.;.r.-!..i'i.-..'..".-.-..-...'----'...-48 5.1 Site I 5.s

5.4 55 Manitsmg Fequency r!--....!.':ii;.i!ii """'"'""""" 43 7. Speries Inventory."..",.....".... 1"1 Scleraiotinia. Glard Ooral) .....,...... ;....;riN.;..,i.r..,..{...... ".65 1.2 Ms,llwce...... -....69 7.8 F'ishes of Palol,o Deep...... i.....i....;..!i!b....;ia.!. ..,....69 7,4 Algae ...... -!r-.qr ...... -...... 21 I Euviroumental Impac'te. 8.1 Cy,ctronee... ..,...... j..!;...... r.!r..!,q!,,...... -,----Tz 8,2 $edimentation...... ^.,"...74 8.3 Crown of Thorns Starfieh or al4nea (Acsnihaster plana,.1 ...... -.V6 8.4 Sewage Outfall"..... ,...,.i.r...... -. 9 lVlanagement Recomneudations..i...... :...... i...i..i....!....!,!r ...... 77 gf Featuree and Inprovenente...... r....,.....,....,...... ?z 9.2 Reeommendatioru 10. 1O.1 Suggegtefl Beadin g...."..."...... -.

t0-Z Fietrd Referenee Texte ....!..,;r....,.....-.,,!..ttr_i.:i!.!!', -...... -- 94

Uee of, the lotus 123 ver. 2.4 Spleadsheet Progrqnme for Transect Aualyeic ...... ,.....86 InFoduction ...... ,86 Inputing and Frocessing the Data..,,,...... ;...... -.;r.,! .....98 Keyshoke and @ function macros used in the processeseing of sp.readeheet infornation

tv 1. Summary

1.1 lntroduction 1.2 General Assessment

The Palolo Deep Marine Reserve is a fringing Palolo Deep is a thriving and diverse coral reef. reef encompassing a lagoon comprising a total As a Marine Reserve it is a good example of a area of 137.5 hectares. The area was initially variety of habitats. Its public appeal Lies in its folmalised as a Marine Reserve in 1974 under access and as an example of a protected coral the National Parks and Reserves Act though no reef area where marine Me exists in natural management plan had been prepared. This abundance. It serves the surrounding reefs as report is to provide a greater understanding of an important source of recruitment for fish the marine environment within the Reserve. A stocks and in the re-establishment of marine basic inventor-v- and classification of the marine life after cyclones. Despite the lack of public environment was undertaken together with funding, the proprietors of the Reserve are provision for continued monitoring. responsible for developing facilities for the public and providing protection for the Reserve. The project was principally one of coral reef description and secondarily a trairung programme techniques of biological in the Environmental lmpacts survey. Transects were used to detail the 1.3 occurrence of coral, algae, fish, sea urchins and substrate. Species inventories were developed 1.3.1 Gyclones tbr hard coral, fish, molluscs and algae. General features of the habitat such as current, depth, Cyclones are part of the climatic environment profile, community type and dynamics were within which the Reserve develops. All habitats described. Monitoring sites were established as are conditioned by their occurrence. The major a means of assessing change in the reserve. cyclone Ofa has had a pronotrnced effect on the Interpretive material, through specimen reef with the total removal of coral fauna from collection and photography, was compiled. The the seaward reef slope, Tons of rubble were bathymetry v'as detailed. deposited on the reef and the fauna elsewhere reef flat was severely affected. Good Aerial reconnaissance was employed to assist in on the on the seaward reef the development of the present survey and to recruitment is occurring reef flat has developed good complement the historical aerial photographic slope. The eastern result of recolonisation by record. Impacts of cyclones, sedimentation and luxuriance as the and new settlement. the proposed Apia sewage outfall are discussed fragmentation and management recommendations made. The Deep has proved to be a haven during this Video footage, for archival and interpretive period with little damage sustained during this purposes, provided a habitat record of the severe storm. Coral spawning the following Palolo Deep with particular reference to the year, as indicated by prolific recruitment on the monitoring sites. seaward slope, was most probably due to such The Palolo Deep Marine Reserve Project will protected areas. also take part, through the collection and dispatch of reference material, in the revision of 1.3.2 Sedimentation the genus Acropora through t};.e Biogeography sedimentation represents the most of the Coral Genns Acropora Worldwide being Chronic to the nature of the Palolo Deep. conducted by Dr. Carden Wallace of the serious threat which coral can Museum of Tropical Townsville, A reduction in the depth at Queensland, from many large dead coral Australia. Additionally, samples of the crown- survive is evident coral presently of-thorns starfish, Acantltaster plarrci, were heads that occur deeper than grows. water clarity is chronically affected sent to Dr. Leon Zannof the Great Barrier Reef The Marine Park Authorrty to assist the by suspended silt. continuation of work conducted in Western Samoa. The fine bottom sediments are evidence of the process of silt capture as the result of terrestrial 1.4 Recommendations run off. A plume of silt from the Vaisigano river was observed along the western reef. Unless this situation stabilises, the composition Visitor inforntation qbout the Palolo and Reserve of the fauna flora of the will Deep should be prouided ort erttry (coral change. A community dominated by algae with reef ecosystem descriptiotl, nlap, a reduction in general would be diversity likely. r egulatio rt s, hazar ds).

+ Deuelop a 1.3.3 Sewage commercial shop to better cater for the uisitor and to generate Of considerable concern is the proposed sewage reuerLue. outfall. As with the capture of silt, effluent entrapment is likely to occur. This effluent may + homote the Deep as ant ecotourisrn affect the Reserve in terms of elevated nutrient uertue througlt promotiort by th,e Visitors' levels which both retard the growth of coral and Bureau and tourist ir*erests. Deuelop a promote the proliferation of algae. This results fornt al ecotouristn progratn me with h*erpretiue in a change in the biota where the algae display rnaterial. overgrow and/or displace the coral. There is a Deuelop an educatiortal programme consequent change in the fish fauna with larger iltuoluhr,g excursiorts by local schools to numbers of herbivorous fishes. The environ- the Reserue to foster a better ment is aesthetically degraded. understanding of coral reefs and their Of potentially greater concern is the possibility irnportance. of health threatening organisms polluting the + Prouide access to the Deep the Reserve making it unsafe for swimming. Toxic for con uenien ce of uisitors. elements may concentrate in the Deep. The main problem with the proposed outfall is its Prouide an ittshore attractiott for thase proximity to the Deep. This is of particular unable or not inclined to uisit the Deep concern as during the summer season the wind by deueloping ant. inshore pool, complete blows directly from the outfall toward the with coral and a variety of biota from Reserve. Further investigation is required as other parts of the Reserue. the value of this important natural resource and recreational site may be greatly + Regulate usa,ge to protect the ntore diminished. fragile areas of the Reserue. Upgrade conueniettces and the reef 1.3.4 Crown-of-thorns Starfish or alamea platform. (Acanthaster plancil + Prouide o,ssistance to the management irt Though several episodes of Acantthaster their task of policirtg tlrc Reserue. infestation and consequent coral reef destruction have occurred in Western and Conthtue ant on-goirtg monitorbtg American Samoa, there is no threat at present programm,e lo ossess changes in, the in the Palolo Deep. Though higher than normal Deep and to prouide a greater numbers do exist, they are not adversely understanding of the dynamics of the affecting the reef. As a tourist interest, they Reserue. should be considered an asset. There are few coral reef animals as impressive as these starfish with their large spines and size, and bright colouration. 2. Methods

2.1.2 Belt transects 2.1 Field Methods Due to the aggregated distribution of sea Initial reconnaissance of the Palolo Deep urchins, the small but numerous newly settled Marine Reserve allowed inturtive decisions to coral colonies, and the transient nature of fish, be made as to the definitions of the general belt transects were used to assess their habitats and areas of interest. A survey abundances. As with the line transect, a tape strategy was then developed to best describe was laid over the substrate. This was usually these environments. 20 metres in Iength though the abundance of coral settlement in some areas limited the Surveying was by means of manta tow survey to a much shorter length due to the observation (Done et al. 1981; Done 1989), and difficult conditions of survey. A width was Iine and belt transect methods (l\4arsh, decided upon to best provide a meaningful Bradbury and Reichelt 1984; De Vantier et al. measure of the organisms to be assessed. In 1985). The methods were adapted to the survey the case of coral settlement and sea urchins a of marine benthos following Done (1989, 1991). one metre width was used. In the case of fish it Qualitative information was gathered by was two metres. swimming and walking. 2.1.3 Specimen collection and identification 2.1.1 Line Transects Coral was collected both for reference and The 20 metre transect iengths were linked, as display. Specimens were collected and'locality, tagged required, to provide a reasonable sample for the with information describing their description of the habitat. The transect line depth, date of collection and other relevant was laid with regard to habitat features to be details. They were bleached until clean in a 5% described. Before the line transects could be solution of household bleach (swimming fool run, familiarisation of the benthic elements was chlorine: sodium hypochloride), washed, fuied required. This was accomplished by compiling and catalogued. a reference collection and by constant verif- ication in the field concerning identifications. Coral specimens to be used in the Reserve for Abbreviations of the names of benthic items or educational display were photographed in' situ attributes were used to provide consistency and so that comparisons could be made between the faci,litate data entry. living organism and its display skeleton' Reference specimens to be identified by the Survey was by SCUBA and snorkel. Data was Museum of Tropical Queensland were collected recorded on plastic sheets or underwater paper. in duplicate, allowing identification of the Where possible, the origin or end of the Reserve material to be made retrospectively. measuring tape was placed at a point relative to descriptions were a buoy base or monitoring transect stake. This Temporary collection or field would allow a more exact repositioning in made for other taxa to provide consistency in replicating the transect. Readings were taken identification during the transect survey and books (see 10'2) along the tape where there was a change in the general inventory. Reference underlying benthic attribute. Attributes comp- provided identification. Photographs were rised species or generic presence, dead coral taken for interpretive display and subsequent and uncolonised substrate. These were reference. measured to the nearest centimetre. The Specimen information and arm samples were categories generally considered were coral, taken from the Acanr'thaster population. The algae and substrq,te. samples were preserved in alcohol and sent to Townsville, Australia for assessment. 2.1.4 Monitoring programme 2.1.6 Undenuater photography Monitoring sites, designated as Slres 7-4, were Habitat and specimen shots were taken with established to provide a detailed description of Nikonos underwater cameras. This involved small reef areas. They consisted of one metre the use of extension tubes and close-up lenses square quadrats for the macro photography, a lSmm lens for habitat and assemblage shots, which were placed at 10m intervals along the and an SB 102 flash. line or belt transects. At sites l-3, the quadrat detail was described in the field. The sites were A video record of the Palolo Deep environment photographed using a Nikonos 15mm lens and was made using a Sony Hi-8 Camcorder and a square metre framer to allow subsequent underwater housing. The unit had autofocus monitoring replication. A set of the photo- and exposure, as well as wide angle and macro graphs was laminated for use as a visual lenses. The raw tape was kept for archival framing reference allowing accurate re- purposes. An edited version was developed for photography of the monitoring sites. This is a more manageable and informed description. essential for a useful comparison.

2.1.3 Bathymetry and currents 2.2 Data Analysis

The bathymetry was detailed from transect Information from the line and belt transects records, soundings, and comparison of aerial was entered into a Lotus-123 spreadsheet photographic features with known depths. The programme with respect to rtumber of records, bathymetry of the seaward slope was treated in relatiue couerage and compositiotl, tneatl, a general manner as time did not allow standard deuiatiort, rnaxinturn anrd, ntitr,imurn description of the channelled relief. occu.rretlce, percent couerage and/or density for The profiles have been constructed to provide a each habitat. Macros, a spreadsheet feature, semi-quantitative appreciation of the bathy- were written to facilitate cursor movement and metry of the reef zone. They represent a minimise keystrokes. Details of these and the summary of information derived from the general use of the programme are found in transects which are numerically listed after the Annex 12.1.2. habitat headings in Section 3. The habitat The pie graph representation of the percentage zones are defined in Figure 3. The position of composition is taken from the table summaries. the profile, within a particular reef zone, is These can be enhanced either through the designated as a broken line in Figure 5. Its WYSIWYG add-in programme or ported orientation is further explained by the through the programme Slide (see Annex 12.1). locational information below the x-axis. The The algal categories have been clumped into profile section is an adaption of the information red, green and brown. The coral names have contained rn Figures 4 and 5. It reflects the been arranged by genus. The substrate is bathymetry described in Figure 4 and requires clumped to reflect the range of types without an understanding of the zones in Figure 3. confusing definition. The assessment of the curent structure in the Consistency in recording can be difficult in a Marine Reserve is of a qualitative nature. survey conducted by several indirriduals. This Information was derived, generally, from field problem was minimised by instruction, inspection as part of the transect information. verification of the attributes, and comparison of Reference to seasonality was derived from results. As data entry requires an ordered inecdotal information. format, the transect data was checked for A helicopter from a local operator was used for consistency on entry and at the frrst data-sort. aerial photography, to provide a more complete und.erstanding of biological and geographical features. 3. The Composition of Palolo Deep: Habitat description

The nature of the reef system is consistent with Fig. t shows the location of the Palolo Deep current reef fonnation theory, involving the Marine Reserve. An aerial photograph shows balance of present physical and biological the Palolo Deep Marine Reserve (Fie. 2). The processes together with changes in sea level. Its location and extent of the general zonation is zonation conforms to general reef development shown in Fig. 3. The bathymetry is shown in within the context of environmental change. Fig. a. The position of line and qualitative This includes such short term considerations as transects is shown in Fig. 5. The habitat photos cyclones, mid-term changes such as show representative views of the area in Fig. sedimentation, and the longer term processes 6a-p. as eustasy and tectonism (Connell 1973; such The following sections describe the eeven major 1980, 1981; Purdy 1974). Chappell habitat zones: Inshore Reef Flat; Eastern Reef This area is unique in that it is subject to FlaU Western Reef Flat; Southern Palolo Deep; tectonism with substantial land movements; its Transverse Ridge and the Main Palolo Deep; volcenic history involves lava overlaying reefs. the Seaward Reef Flat; and the Seaward Reef ln addition, the occurrence of the largest Slope. The descriptions comprise details of cyclone in a hundred and fifty years (Ofa, Feb. location, exposure, depth, profile, substrate, 1990), followed by a major one (Val) the algae, coral, community t5lpe, and biological and following year, have had a pronounced effect on physical dynamics. Line and qualitative the present condition of the reef. transect information is incorporated in the text as well as in a summary table. A pie chart diagram details the percentage composition.

l6ooE lEoo l6oow

Pocific chuuk N Oceon +

^nu*-7

Fig. I: Location map for PaIoIo Deep Marine Reserve, Upolu,Western Samoq- Fig, 2: Aerial photograph of the Palol,o Deep Marine Reserve. Reservc Boundary Reef Slope

Spur and Groove

Reef Crest

Seaward Reef Flat

Main Palolo Deep

Western Reef Flat 2

o- o) o0) o s (g

Pr"troF "k Inshore Reef Flat Scale l:46fi) '/----\.--"' lfi)m x I 4a

Fig. 3: General habitat zonation in the Palolo Deep Marine Reserue.

Boundary marker buoys are numbered. e |.e -/

Scale l:46i00 H 1fi)m D€pths in mehes

8^^

Etg, 4: futhtnetrl of tlta Pshola Deep, r+ El: I

3rl a.lc"

---:- \F\- ), il' l'-4 ) -JL,/ {/{r \ I

.k Seele t:4d0Q

100rn --=s -* ** - -i

Fte. 5: Positiloa of tlnli*/rlanrr;cts (nwnibad;);qwlttqdlwt s,otydbotfun*tryptofi,Ies (*slerr*d. bt brplccn hiw"s). Fig. 6: Habitats of the Palolo Deep:

Ind.ex:

a) An overturned' Porites lutea bommie on the Westenr. Reef FIat.

b) A luru,riance of coral growth on the Eostern Reef FIat north of the platforn.

c) An irnpouerished, North-eastern Reef FIat, with less coral couer and more Dictyota sp.

d,) Clumps o/Dictyota sp. growing on the Southern Deep margin extending fron the shallow algal/coral crest.

e) The brown algaes Turbinaria ornatus and Dictyota sp. with the coral Porites sp. comprise portiotr's of the perimeter of the Southern Deep, bord'ering the Eastern. Reef Flat'

f) The coralline algal rim/reef flat of tlrc seaward, portiort. of the main Deep.

g) With their origins on the ou,ter reef slope, the rubble crest is littered with thc tentnanr'ts of coral colonies. These are uictins of ertreme waue action. Eristing as prominent ban'hs after cyclotte Ofa (Feb. 1990), their relief is now reduced. At present, they are represented, as ntbble couering the seaward reef flat.

h) The outer reef slope is d,euoid. of the nornal extensiue coral couer. Closer in'spectiort reueals large quantities of hard corol and, algal settlement, most probably originating from plaees such as th'e Deep'

i) Porites cylindrica on the southeastern wall of the Sou,thent' Deep'

j) A large stand of Actopora nobilis monopolising the Southern' Deep'

k) A colony o/Pocillopora damicornis defend.ing its spaee uithin the sprawling expanse o/A- nobilis colonies.

switntning t) A feed.iny scor o/Acanthaster planci within the A. nobilis thicket. ,Sorne Chromis viridis aboue-

m) The hard. coral Plerogyra sinuosa in. the Main, Deep. Relic P. rus colonies in th'e nurhy bachgrou'n'd-

n) The rubble slope near the seaward, entrance of the Main Deep. This talus slope is tr'nstable with little colonisation.

o) The uertical reef wall near the seaward opening, characterising the east northeast side of the Deep.

p) Columns of laryely dead,P. rus rise o*t of the silty bottom qt l?n-

l0 Fis. 6. Habitats of the Palolo Deep (o-h).

Photos: E. lovell 1991].

tl Fig. 6: Habitats of the PaLolo Deep: (i-p).

Photos: E. L,ovell 199:1.

l2 Depth 3.1 lnshore Reef Flat The shoreward margin of the beach slopes through the tidal range to a shallow inshore (Transects 1a-h; 2a-h; 3a,b) gutter which is characterised seaward by a shallow bathymetry (0.6m depth mean low water OIL!\D). It deepens at the base of the General description platform into a channel of 1m. The zone is characterised as generally flat with coral rocks This zone is the inshore area of the Marine occurring sporadically. Shallowing through Reserve. It comprises two habitats: one of a mounding occurs as sand and rubble aggregate largely sandy substrate with reduced biota to around the rocks (see Fig. 4). the southwest of the southern Deep and near shore; the other a more diverse area resulting Substrate from its proximity to the southern Deep and the presence of a more stable rubble substrate. The substrate comprises rubble and sand (74o/o). This latter area is transitional in character, The area in proximity to the southern end of grading into the Eastern Reef Flat. the Deep has the higher proportion of rubble (26%) and extends west. Inshore of this area is Location a tongue of sand which extends from the Western Reef FIat. A largely sandy substrate The area is located inshore of a line drawn dominates shoreward. Coral rocks are between the two inshore buoys (no. 1; east and scattered, and are prominent as sites of west) marking the Reserve boundary (see Fig. macroalgae and associated organisms. 3). More generally, it is the area inshore of the southern end of Palolo Deep extending to the Algae beach. Dictyota dichotoma (see Table 1) is the Exposure and Current dominant algae with its presence mainly confined to the area peripheral to the Deep. It Wave action is confined to a wind chop or much is associated with rubble, and its occurrence reduced ocean swell due to the broad reef flat reflects this substrate elenent. Coverage expanse. It is subject to the effects of tidal ranged from 48% near the Deep, to a sparse 3% exposure with shallowing depths and an in the sandy areas. Its occurrence in this zone unstable substrate. There is a pronounced represents the inshore extent ofits larger range current originating from the Deep, where it which encompasses the Eastern Reef Flat. wells over the southern crest, flowing to the Elsewhere it is co-dominant with an east. Near-coastal effects occur through uniilenti{ied red algae and ScrgosEunx sp.. freshwater run off and resuspension of sediments due to wave action.

M

.3 Depth m.6

o 75 150m metres Inshore Reef Flat Plaform

General profiIe of transects in the inshore reef area (see Fig. 5).

l3 Sa,rgassum sp. occur as a band of foliaceous Near the Deep the diversity is higher with the algae in raised areas of rubble in the central community elements being quite different. This and western area. Other algae are Halimeda is the result of three factors: a more stable sp. (Fie. 34f), Tfu,rbinaria ornata, green rubble substrate, proximity to parent stock fi.lamentous and turfed algae (Table 1). The found in abundance within the Deep, and total algal cover in this area is 9%o, where the shelter from wave action by the Deep. The sample near the Deep was entirely Dictyota, current that flows from the Deep promotes the covering 48%. algaUcoral rim to the south of the Deep and the predominance IJncommon in the Reserve, the sea grass of Dictyota sp.. The area of Sargassum sp. ocurring Halophila oualis occurs in this sandy area. on islands of rubble, provides good habitat and refuge for schools of juvenile fishes. These find shelter in the Coral shallower water and among the foliaceous algae. Hard coral cover near the Deep is 16% with five genera represented. Acropora nobilis (Fig. Bbc) is the most abundant (9%), although patchy. Dynamics The near shore area is sparsely colonised by Tidal exposure, unstable hard coral; the total coverage is 3%. Species substrate, current, occurrence is all less than one percent with wave chop and effects from the shore all play a Porites cylindrica part in making this area one of extremes. The @ig. 35b) the most prevalent. current Some medium size (60 cm) massive colonies flowing eastward from the Deep, ameliorates the land exist, but generally the colonies are isolated. effects. Current and wave action are As with the algae, hard coral is more prevalent responsible for the movement of the sand from the Western Reef in the area near the Deep. The assemblage is a Flat. For most of the year, the mixture of the genera Acropora, Montipora and current flows into the south-east trade-wind chop which Porites, with greater diversity closer to the creates a more vigorous Deep. shallow water environment maintaining a cleansing mechanism for the fine sediment. This area is shallow and subject to tidal Community type fluctuation, which gives rise to exposure and temperature The inshore zone represents two community extremes. These are moderated by types grading into each other, differentiated by the current and wave action. A principal Iimiting feature diversity and degree of development. The area is the sandy substrate. With limited benthic inshore is characterised by an impoverished attachment coupled with the vagaries shallow benthos where an unsuitable substrate and of water, a diverse and luxuriant inshore community harsh conditions moderate development. is not possible. Organisms are adapted to an unstable substrate or are found associated with isolated coral colonies or rock.

Transect I a-h Transect 3 a,b

Lrorr djrr,ll33 nlltr l.rrd tgjt tclrtopor 035 l*lllepon 5,13 H$tf,il.'fsc Porltr 2J3 cor.l ?*t l.ag Lrlt .l.tl Prvclr l.ll rrd dgr !.lt Acnpn Lt3 ErlopUle u7 nbblr ud uril 0.5 bnvr rlgrLil ccrd rrcl 7.3t

conl nbbh lf.ltr onl nbblo 26.15

Transect 2 a-h

Fig. 7: Graphs of the percentage composition of transects la-h; 2a-h; Ja,b

t4 Table 1: Conbined inshore transects in the sand,y zone perpendicular to the beach (260m): Tlansects Ia-h:2a-h.

Tara N T'oH lrean' . Sd, l{ax. ,tMh;,,.,., ::.,':,. :i ::.1.:: 'P€tt.l*::..,f . Interce$ O-e-y.

(7n Coral Acmpon hyacinthus J 32.3 7.6 30.0 10.0 0.23% Acropora nobilis 1 10.0 10.0 0.00 10.0 10.0 0.04% Montiporasp. 100.0 80.0 8,37 30.0 5.0 0,407o Porites cylindica 2 210.0 70.0 84.85 190.0 '10.0 0.84% Poriteslutea 6 150.0 75.0 22.11 80.0 5.0 0.60% Ponfes sp. 70.0 43.3 15.00 50.0 10.0 0.28o/o Porites rus 50,0 7.2 3.94 15.0 3,0 0.20%

Total 2.39%

Algae A/gae sp. 1 5.0 5.0 0.00 5.0 5.0 0.02% Coralline algae ? 34.0 17.0 7.00 24.0 10.0 0.1404 Dictyota sp. 16 721.0 45.0 12't.95 385.0 5.0 2.87% Green filamentous 5 25.0 5.0 2.50 5.0 5.0 0.10% Halimeda sp. 21 1.0 23.4 24.08 72.0 5.0 0.84% Red algae 11 485.0 48.0 119.02 405.0 5.0 1.93% Sargassum 10 370.0 37.0 62.32 20s.0 5.0 1.47% Tubinaia omata 20 150.0 7.5 3.28 15.0 0.0 0.60% Turf algae 2 155.0 n5 17.50 95.0 60.0 0.620/o

Totrl 8.5S%

Seagnss Halophila ovalis 1 570.0 570.0 0.00 570.0 570.0 2.27%

Substrate Coral rock I 416.0 52.0 94.21 300.0 7.0 1.65% Dead standing coral ( 74.0 14.8 s.00 24.0 5.0 0.29% Rubble 27 2s50.0 118.3 150.27 615.0 0.0 10j20h Rubble and sand 89 18560.0 208.s 230.24 1430.0 0.0 73.81% Sand 8 448.0 56.0 73.59 210.0 5.0 1.78%

Total 260m Totrl 87.65%

Table 2: Inshore reef flat parallel to shore near th,e southern end of the Deep (40m): Tlanseet ?o,b

Taxa Total Ititean Std, Max. Mn. PeiEqit lntercept Dev. , .,00vei (qr.r)

Coral Acropon nobilis 4 u2.0 85.5 141.21 330.0 1.0 8.55% Pavona decussata I 5.0 5.0 0.00 5.0 5.0 0.13% Porites cylindria z 12.0 6.0 4.00 10.0 2.0 0,30% Pocillopora eydouxi o 201.0 33.5 28.95 80.0 5.0 5.03% Polifes rus 7 81.0 1 1.6 7 .78 30.0 5.0 2.03% Seiatopora hystix 1 10.0 10.0 0.00 10.0 10.0 0.25% Tolal 16r9n

Algae Dbtplasp. 5 1934.0 386.8 366.34 9690 40.0 48.35%

Substrate Coral rocl 3 303.0 101.0 108.18 253.0 10.0 7.8% Rubble 15 1046.0 69.7 68.24 290.0 1.0 26j5% Rubble and sand I 16.0 16.0 0.00 15.0 16.0 0.40% Sand 1 4.0 4.0 0.00 4.0 4.0 0.10%

Total 39.5fm Total 3,,23%

l5 Table 3: Eastern reef flat ertending from inshorc buoy (1) westward, (40n): Transect ITa,b

i;, . . ':t .,i ,., ii.i,' ffi :i i'.l:. :,, i...'i'.,Uar. .' ,., , i:,,,,,,i,,,i,,i :, .,:.....,..QliJ;,,,,,,;,,,1,..,.,.,.:. ,,. ,:.,. ii,i:,,:::'t ii,i,,,t,i:ii,ri: ,,,i;,,. ,ii:iii:i.i'i'j.:l:,:'ii't:, i t i.i.r.riiii!.,:ririj.,i.:j.

Acmpnhyrcinthus 2 35.0 17.5 2.50 20.0 15.0 0.88% Acnpon nobilis ,| 2.4 2.0 0.00 2.0 2.0 0.05% Montipora sp. 1 15.0 15.0 0.00 15.0 15.0 0.38% Puillopon damicomis c 155.0 31.0 39.79 1'10.0 5.0 3,88% Pocillopora eydouxi 2 30.0 15.0 10.00 25.0 5.0 0.75o/o Poritx cylindin 14 216.0 15.4 9.31 38.0 5.0 5.40o/o Poites tutea 2 12.0 6.0 4.00 10.0 2.0 0.3070 Poriles rus 5 90.0 18.0 18.86 55.0 5.0 2.25% Pontes sp. 4 35.0 8.8 4.15 15.0 5.0 0.88% Seriatopora hystrix 1 10.0 10.0 0.00 10.0 10.0 0,25Yo Tohl 15.020h Algae Diclyota sp. 23 2480.0 107.8 97.85 410.0 20.0 62.00% Halimedasp. 4 39.0 9.8 4.54 17.0 5.0 0.98% Sargrassum sp. 2 30.0 15.0 5.00 20.0 10.0 0.75% Twbinaiaomatus I 10.0 10.0 0.00 10.0 10.0 0.250h

Total 63.98% Substrate Conal Rock 3 85.0 28.3 22.48 60,0 10.0 2,130/0 Dead standing coral 3 230.0 76.6 71.79 177.0 13.0 5,750/o Rubble 16 526.0 32.8 30.09 133.0 5.0 13.'t50h

Totrl 1000 Total 21.03.h

Table 4: Transect for eastern reef flat, extend.ing from the Reserve bound.any between buoys 1 and, Z (20m): Tlansect 2J.

Tua Tobl [4ean Std. Max. Min. Percent lntercept Dev, cover (cm)

Coral Acropora nobilis I 10.0 10.0 0.00 10.0 10.0 0.50% Montiporasp. 1 5.0 5.0 0.00 5.0 5.0 0.25% Pwites cflindica 11 128.0 1 1.6 3.08 20.0 10.0 6.40%\ Psammocora contigua 2 50.0 25.0 20.00 45.0 5.0 2.50% Pocillopora daminmis 8 85.0 10.6 4.64 20.0 5.0 4.25o/o Porites futea 1 5.0 5.0 0.00 5.0 5.0 0.250A Pontes rus 1 5.0 5.0 0,00 5.0 5.0 0.25o/o Seriatopora hystrix 1 10.0 10.0 0.00 j0.0 10.0 0.50%

Total 11.901t Algao Didyotasg. 4 195.0 48.8 37.14 110.0 10.0 9.75% Subttnte Coral rock 8 445.0 55.6 62.72 190.0 5.0 22.250/o Rubble 17 1062.0 62.5 46.44 170.0 10.0 53.10%

Totrl 20m Total 75.35%

l8 Table 5: Transectsof eastern reef flat, north of the platform in an area of high coral cover (10m): Tlansect 24.

l:t:l:il:!: r:i.:::i:

Coraf Acroporahyacinthus 1 15.0 15.0 0.00 15.0 15.0 1.50% Acropora nobilis I 20.0 10.0 5,00 15.0 5.0 2.0010 Porites cylindrica 21 590.0 28.1 23.07 90.0 5.0 59.0070 Pocillopora damicomis I 110.0 13.8 4.U 25.0 10.0 11.00% Porites lutea 2 15.0 7.5 2.50 10.0 5.0 1.5070 Porles rus 5 65.0 13.0 9.27 30.0 5.0 6.50% Zooanthids 1 15.0 15.0 0.00 15.0 15.0 1,50%

Total 83.00%

Substrate Coral rock I 105.0 11.7 7.07 30.0 5,0 10.50% Rubble 3 40.0 13.3 2.36 15.0 10.0 4.00% Sand 2 25.0 12.5 2.50 15.0 10.0 2.50To Total 10m Totaf 17.00'/t Table 6: Transectsof central eastern reef flat ertend.ing from tlrc middle buot G|m): Tlansect 18a,b.

Conl Auoporc hyacinlhus 2 30.0 15.0 5.00 20.0 10,0 0.75% Acroporc nobilis 90.0 12.9 11.29 40.0 5.0 2.250h Pocillopora damicomb 13 2s4.0 19.5 13.93 50.0 10.0 6,35% Porites cylindrica I 95.0 1'r.9 7.04 30.0 5.0 2.350/0 Porites lutea 1 15.0 15.0 0.00 15.0 15.0 0.38% ,| Pofites rus 205.0 205.0 0.00 205.0 205.0 s.13% Pontes sp. 27.0 5.4 2.58 10.0 2.0 0.68% Total 't7.8$A

Algae Dictyotasg. 24 3089.0 128.7 104.66 370.0 18.0 77.23% lurbinaia omatus 2 15.0 7.5 2.50 10.0 5.0 0.38% Total n.61.4

Substrate Dead standing coral 1 1s.0 15.0 0.00 15.0 15.0 0.380i Coral rock 3 50.0 16.7 9.43 30.0 10.0 1.25o/o Cool rubble 2 1 15.0 57.5 17.50 75.0 40.0 2.88o/o

Total tOnl Total 1.511

Table 7: North eastern reef flat extend,ing from' the seaward, buoy (40m): Tlansect 79a,b.

Acropon nobills 5 35.0 7.0 2.45 10.0 5.0 0.88% Montiponsp. 4 30.0 7.s 2.50 10.0 5.0 0.75% Pavona decussata 4 20.0 5.0 0.00 5.0 5.0 0.50% Pocillopwa daniconis 1 5.0 5.0 0.00 5.0 5.0 0.'l3olo Porites cyllndica 14 178.0 12.7 4.32 20.0 5.0 4.45To Poiteslutea 1 25.0 2s.0 0.00 25.0 25.0 0.63% g Poflles rus 90.0 18.0 '16.61 50.0 5.0 2.25% Pon{es sp. 1 20.0 20.0 0.00 20.0 20.0 0.50o,6 Total 10.09/o

Algae Drblptasp, 7 902.0 128.9 65.01 200.0 45,0 22.5s% Twbinaria omatus I 70.0 23.3 '18.86 50.0 10.0 1.75% fobl ?1.nh

Substnte Coral rocl< I 20.0 20.0 0.00 20.0 20.0 0.50% Conl rubble 24 2477.0 't03.2 91.74 335.0 20.0 61.93% Rubble and sand 1 128.0 125.0 0.00 125.0 125.0 3.230h

Total {0m Tolal 55.66%

l9 Table 8: Abund'ance infonnatinn. on Echinometra mathaei from, the eo,stent. reef flot. Five 20nt2 belt transects were ru,n along lhe line transects as indicated.

Trangect l7 a,b Traneect 24

Acnrur 0.1! t.rltil lJ! lri lt.:rd ll.lt hcUbgur 4.f! crrrl nltL t.?5 ?rcllblrrr ll ItoralD.n fJl lcrlt.3, tgi'11.1'11'r to...all& lJ

crnl rrct lf.t

ord rrlbh I read l.3 Acregcrr !J lron rl;r lt

Fig. 8: Graphs of tln prcentage compaitbn ol tran*ctc t7a,b; d 24.

Transect 18 a,b Traneect 19 a,b

lcllbpnIJJ crrrlrct fJ t rli.r t5l )rrr rl3rr ll3 Acrlnn !

lerl rL c*rl fJt llorllgcn l.?5 crrd rrlDh lJl onlml lJ5 lcclllogcn f.l! lcrltrr TJil lr.rr dl.. 7ttrl lcmgon l.ll nllb I rild t.lil

crnl nbLb 6l.tl

Fig.9: Graphs of the percentage conposition of transects 18a,b; and lga,b

20 Depth 3.3 Western Reef Flat The Western Reef Flat is generally deeper than the Eastern. The excavated nature of this area (Tra nsects 20a,b; 21 a-d; 22a,bl results in a depth of lm MLW. There has been a movement of material towards and into the Deep. The bathymetry deepens gradually with General description its approaches to it. Exceptions to this are areas near the perimeter of the Deep where the This area is a homogeneous reef flat comprising presence of large coral colonies (Porites sp.) a relatively impoverished fauna. Similar to the cause material to aggregate in mounds. There inshore area in nature, it appears as a is also a shallow area of principally sand and disturbed unconsolidated environment, where rubble on the north-west margin of the Deep. the flat has experienced deepening through erosion. The benthic community is depauperate The remainder of the reef flat is shallower with with sparse coral occurrence (Fig. 6a). the south-west margin of the Reserve characterised by a depth of 0.3m. At the Deep, shallows to Location margin of the southern it 0.6m, then grades into the inshore area to a The area is to the west of the Paiolo Deep. It is depth of 0.2m, as shown below. bordered to seaward by the rubble banks of the reef landward or southern outer flat. Its Substrate margin grades into the inshore zone, defined generally as a line between buoy no. 1 and the The bare substrate percentage varies between southern end of the Deep (Fig. 3). 44o/o and 650/o, with a greater substrate component seaward. As with the eastern flat, greater percentage of rubble, varying Exposure and current there is a from 27-460/o. The sand component is much Relative to the Eastern Reef Flat, this area is Iarger inshore and around the rim of the Deep, more exposed. Though protected by the reef where it grades into the talus slope inside. crest and the rubble banks, it is subject to greater wave and current action, particularly Algae during the summer months. This is a period characterised by the presence of strong north: The algal percentage cover ranges between 28% west winds, creating a strong current through and 44o/o. The principal algae is Dictyota sp., the Reserve. occurring between 28%o and \Lo/o. Other algae present are Halined,a sp., Sargassum sP,, T\trbinaria ornata, and a green frlamentous algae. In the inshore area (transect 2O), Sargassum sp. (7%) and Tfu.rbinarin ornata (60/o) occur.

General profiIe of transects in the area of the weste t. reef flat (see Fig. 5)

MLW

o 1OOm

Reserve Palolo Boundary Deep

2l Community type Coral A predominance of Dictyota sp. covers a rubble The coral composition is impoverished with substrate. Apart from the prevalence of Porites total cover ranging between I.5o/o and l7%. bommies, coral colonies are patchy in Unlike any of the other areas, it is largely occurrence. Porites with. Porites lutea as the predominating species. Its occurrence ranges between 1.8% Dynamics and IlYo. There are large Porites bommies which in many cases are overturned or partially Comparison of the aerial photos (FiS. l9) shows buried. These bommies, often heavily damaged, the deepening of the Western Reef Flat to be a provide prominent relief from what is otherwise relatively recent event (post 1990). Being less a flat expanse. This is a stable surface for other protected than the eastern flat, the occurrence benthos, which form islands of diversity in what of the major cyclones may have altered the is otherwise an impoverished area. nature of this reef flat through the erosional effects of extreme All other species have less than B% coverage. wave or current action. The presence of the rubble Other species present, in decreasing coverage, banks may have altered the pre-cyclone are Acropora nobilis, A. hyacinthus, Porites rus, Ofa influence of ctrrrent and wave action. These Montipora sp., Acropora humilis, Porites now form a seaward barrier to the excavated cylind.rica, and Pocillopora damicarrr,ls. Unli-ke area of the Western Reef Flat. The high energy season the protected Eastern Reef Flat, this area has for this area is during few small colonies. The lack of benthic the summer when the north-west wind and current are strong. development and the loose rubbly nature of the substrate give the impression that the area is The western margin of the Deep is a talus slope periodically disturbed. of rubble which must have traversed the reef flat, grading it in the process. The rubble continues to be transported towards the Deep during periods of rough weather as the rubble banks migrate shoreward (Fig. lg: photos 1g90, 1ee3).

Ttansect 21a,b

Dr.rt rlt.. tua lro da.. a.ta .hl Et a.ta

l|lEdtmt.' ,-lliJTf

.l6t6t t rfidlt.rl

wlhlurttlt nl$r r iol l(tl A6t drJt orlnllhl''l ,.raLrt

arodtstjl Ino d3a lt5

t 11L. t,al Ea l.a

rhlHt4'Jt r..rl nllL tlJ

Ttanrect 22a,b

Fig. I0: Grapla of tle prent4e *,mp',itbn for weetcrn reef trw*o,b.

2 Table 9: Western reef flat, ertending eastward from the inshare buoy (40n): Transect 20a,b.

Acropora nobilis 120.0 120.0 0.00 120.0 120.0 3.000/6 Porites lutea 350.0 350.0 0.00 350.0 350.0 8.75o/o Total 11.75% Algae Dictyota sp. A 1200.0 200.0 200.83 490.0 10.0 30.80% Halimeda sp. 1 1.0 1.0 0.00 0.0 0.0 0.00% Sargassurn sp. 1 290.0 290.0 0.00 290.0 290.0 7.25o/o Turbinaia omata 2 250.0 125.0 115.00 240.0 10.0 6.25o/o

Total 41.304/o

Substrate Rubble 1 1090.0 1090.0 0.00 1090.0 1090.0 27.25o/o Rubble and sand 670.0 134.0 164.75 430.0 10.0 16.75% Total 40m Total 4o/o

Table 10: Central westenr reef flat, extending eastward fron th,e middle buoy (78nt): Tlansect 27a..

.-'',Tffi, lilar;

,'t ,:,,,,,:,r,(g!l):

Coral Acropora hyacinthus 1 205.0 205.0 0.00 205.0 205.0 2.56% Acropora humilis 2 25.0 12.5 7.50 20.0 5.0 0.31o/o Montipora sp. 1 40.0 40.0 0.00 40.0 40.0 0.50% Porites cylindica 3 20.0 6.7 2.36 10.0 5.0 o.25% Pocillopora damicomis 1 5.0 5.0 0.00 5.0 5.0 0.06% Porites lutea 4 893.0 223.3 321.48 775.0 3.0 11.16Vo Pontes rus ? 170.0 85.0 80.00 165.0 5.0 2.13% Total 16.97to

Algae Dictyota sp. 10 2053.0 205.3 133.65 470.0 10.0 27.96% Green filamentous 1 5.0 5.0 0.0 5.0 5.0 0.06% Total 28.020h

Substrate Rock 14 491 .0 35.1 33.44 1 30.0 0.0 6.14% Rubble 25 3688.0 147.5 112.60 4?5.0 2.0 46.lQo/o Sand 2 220-0 110.0 30.00 140,0 80.0 2.75o/o Total 78m Total 54.99%

Table 1I: North western reef flqt, ertending from the seaward buot GOm): Tlansect 22a,b.

:1 1:1:::::li ii:r:::r:: i:i:::: ::1: : ilil ii!i11: : ::rr.. i: l4(:.:l:,tiii:;:,:.1,,':lfifi.ii,i..iil: ,',,i,1,:,::lii.,P$!!4:1,:,: : ::l: r:.i:l - r :. l:l rl ;:l i :r:i:r i : :::r1i:l t: t,:::::.::: : : :r, :6(**t:ll i :''t ''..'.. ,, , ...,,,.PFr

Goral Porites lutea 10 54.0 5.4 3.04 11.0 1.0 1.35o/o Poites rus I 5.0 5.0 0.00 5.0 5.0 0.13o/o Total 1.48Yo

Algae Dictyota sp. 10 1248.0 ',t24.8 118.21 410.0 8.0 31.20% Halimedasp. I 95.0 10.6 7.62 30.0 3.0 2.38o/o Total 33.58%

Substrate Coral rubble 12 1738.0 144.8 173.90 612.0 15.0 43.45o/o Rubble and sand 4 836.0 209-0 161.58 400.0 13.0 20.90%

Sand 1 24.0 24.0 0.00 24.0 24.O 0.60% Total 40m Total 64.9s%

23 Depth

3,4 Southern Palolo Deep The algal margin of the Deep, to the east and south, has a depth of 0.1 metre below mean low (Transects 7 -11 4-6; ; 12-161 water. The margin to the west is deeper with a depth of 0.3m. The eastern border descends to lm to 2m where a terrace extends along the General Description eastern side, sloping to 5m. From there it The southern end of the Palolo Deep is descends to a sandy bottom at 7-10m. The slope relatively shallow by comparison with the main below the terrace is gradual along the length of body of the Deep. This site contains the most the Deep from the southern end but is progressively luxuriant coral development in the Reserve. It more abrupt toward the seaward occurs almost exclusively along the eastern end. The rubble slope on the western side rises side. The remainder of the area is deeper, with at the angle of repose from the sandy bottom patchy coral cover on a sand bottom. The (see Fis. 4). western slope comprises a talus slope of rubble. Substrate Location The uncolonised substrate component increases This area extends from the main body of the with depth. It varies between 29%, in the Deep shoreward of the Transverse Ridge to the shallow area, to 46% at the terrace margin. In seaward margin of the inshore zone (Fig. 3). the upper mid-terrace zone, coral rock is the most common substrate (18%). Standing dead coral is the next most common component (6%). Exposure and current Other elements are rubble and sand. Mid- This area is the most protected environment in terrace, standing dead coral forms the largest (L7o/o). the Reserve, being surrounded by reef flats and component Rubble and sand are the the main body of the Deep to seaward. Current next most common at 7o/o and 8% respectively. is constant except at extreme low water. It Rubble is the most common substrate at the flows along the length of the Deep, spilling terrace margtn (21%). Coral rock, standing Iaterally over the rim to the south-east. As the dead coral and silt and sand are the next most (6Vo, tide falls, the current is confined to and can be common components 5% and 6.bo/o seen to flow over the southern end, rising respectively). noticeably. It then forms a strong inshore At depth, the substrate is characterised by silty current which flows past the platform. This sand with the presence of large coral boulders results in the presence along the south-east rim and varying degrees of rubble. The substrate of a consolidated coraValgal margin with component is high here (>80%). Dictyota sp. (Fig. 6d) and higher coraL cover immediately shoreward.

3

Dspth 6 m 9

0 25 50m metres West Southern Palolo Deep East Rim Rim

General profile of transects in the area of Southent, Palolo Deep (see Fig. S)

24 The we-qtern margin is a rubble/boulder slope Comparing the mid and shallow terrace comprising coral fragments similar to those transects 7-11 and 12-16 respectively, it can be found on the seaward reef flat, grading to sand seen that the Acropora rrcbilis and Porites rus and rubble at the upper margin. Some live are relatively equai in occurrence in the shallow coral exists at its base but it is generally area with Porites cylhtdrica half as common. uncolonised. Within the mid-terrace, Acropora nobilis becomes twice as common (3lo/o) as the two Porites species which are of equal incidence Algae (15% and 17%). With depth (transects 4a-d; 5; The percent of algal cover varies from less than 6), Porites cylind.rica becomes dominant (LI%) IVn to 2-l%. depending on depth. The area of wit}a Acropora rtobilis and Poriles ru.s more least algal pl'esence is in the mid-terrace region equal (6% and 8% respechively). coral covel is highest and the algal where ihe The other corals in the assemblage occur presence (Clodophoropsr.s?) is confined to the individually at less than ZYo, with the exception base of the Acropora rnbi,lis beds (Fig. 34d). of Seriatopora hystrix with a 3%-4% coverage. This rvas not measured in the survey (Figs. 6k and 6l). It is ahvays present at the base of the coral so that a neasurement of A. rr.obills is a Gommunity type of the algae. measurernent Four distinct habitats exist, two of which are co- The proliferation of the algae is the result of the dominated by three species, the other two protection afforded by the pugnacious and having very little living cover. The first habitat territorial damselfish, Stegastes liuidus (see is the vertical south-east margin comprising an section 6; Fig. 34c, 34d). The algae also occurs algaUcoral ridge at the crest or margin of the on standing dead coral but is generally grazed Deep. This grades down the slope becoming a by herbivorous fishes. As the area is largely relatively diverse assemblage (10 spp. coral :5 monopolised by this coral, there is little area for spp. algae) not subject to the monopolisation as other algae to develop. By contrast, the areas seen on the terrace. peripheral less monopolised and to this area are The nore vertical portion of the slope has less algal diversity is greater. coral cover and is characterised by smaller In the shallow area, there is 13% algal presence colonies, generally dominated by P. rus and P. (Fig. 6e). This figure would be higher if the cylbtdrica with small colonies of Seriatopora slope below the algal ridge or the ridge itself hystrix. A. nobilis is a minor component in this were assessed. The greatest percentage is area. The algal component is higher than the unidentified (8%). Halhneda opurt'tia and adjacent terrace (13% vs. 1%) though this fails coralline algae were the next most prevalent. to take into account the filamentous green By contrast with the reef flat, Dictyofo sp' was algae that exists within the Acropora n'obilis poorly represented (0. 1 5%). stands. The deeper area was represented by algal The terraced area is characterised by a greater species. Jaria sp. were the next most common prevalence of Acropora n'obilis (3Io/o), component, characteristic of this deeper area monopolising large areas. An important and occurring extensively. Halimed,a opurvtia component of this area is the association of the was present as a minor component. damsel Stegastes liuidus and the frlamentous green algae Gotts 1977). Though not measwed in the transects, the algae co-occur with all of Goral the A. nobilis and other branching coral. Its Here the hard coral is the most luxuriant in the presence is at the base of the colonies and is a Reserve. The shallow and mid-terraces have a dominating feature of the assemblage (see living cover of 6to/o and 64% respectively. The Section 6: 34c, 34d). assemblage has a 30% deeper margin of the The deeper portion of this area is characterised assemblage is co- coverage. The hard coral by a greater abundance of algae, such as Jant'ia nobilis, dominated by three species: Acropora uncolonised substrate becomes (Tigs.6i,6j). sp.. The Porites rus, and Porites cylindrica greater. The deepest portion of this area is is 2oo/o, Their relative cover in the 2-3m depth impoverished with little Iiving cover. It is ZOYo mid-terace and 12% respectively. In the largely a sandy substrate with scattered coral cover is 31%, 15% depth of 3m, their relative rock and rubble. and 17% respectiveiy. At the 4-5m depth, percentage cover is 60/o, 8o/o, and II% respectively.

25 Dynamics The relic existence of large Porites rus or P. cylhr.drica colonies in the silty, murky bottom A process common to the Palolo Deep, this is a of depositional environment characterised by its the Deep is evidence of a more suitable environment turbid water and silty depths. Chronic, current in former times. They presently occur only Deep, born siltation is a continuing problem for the about the margin of the suggesting environmental change. benthos. Cleaning mechanisms require energy. Successful growth strategies such as the open Substrate is the principal factor limiting much branching form of Acropora nobilis, minimise of the living cover. Reef development is this requirement. The colony maintains its consolidated on the eastern side but is living presence above the substrate with its represented by an unstable boulder and rubbie ramose branches cleaned by the current action. substrate on the western margin. Rubble Another factor is turbidity, limiting the generated by successive cyclones continues to penetration of light. Consequently coral create an unstable slope. The bottom is occurrence is generally at less than 5m depth. composed of fine silt, old bommies and imported material from the shallower areas.

Transect 4-6 Transect ?- I I

?.rll.. ll.7a ccnl nlllc ll.2l d..d r(,.orrl tJ.at AcdD.r.3l.t

irod l.Jl

Morllgcn lJ,l nf,Lto I tr-r.ltr. l.at rrrd t.i! na rlt.. a.l2 G.hr.. l.la r.rd 2.lt conl nbltr ?.0J g.rbaoDorr t.2a Acr.lEr. a.0t a..d rt- ccr.l a.?a .oml ro(L 0.27 lod(l !i.J!

d.rd rt..o.rl d,t, conl rrbbh 1.0{ l.r.Doh 2t.tl

.odl r*t lt.Ot P.to.r C.rs

gnt rl3rr l.l7 rod rl3rr l.,| 6.rLtoDoh l.Jt Poclllogorr 0.?l Pr?11.. t7.r7

Transect l2-16

Fig- 11: Graphs of the percentage composition of transects .

26 Tabb 12: Southern Palolo Deep, 4-5m dcpth profiIe (118m): Tlansects 4-6

,,, :,;:$fl, it.r.j T.s8 'l:,Tob[ 169:: !Sg!, :.: : :.:,t,t',,,.:.:Lff n, F . . ::::r::i:r: :tnlCr pt ::oByi .',,00y0f,r,, ',:,:: .{qn)

Gonl Aaopora hyadntrus 3 35.0 1',t.7 6.2 20.0 5.0 0.4996 Acmponnoblis 10 648.0 64.6 53.4 150.0 10.0 5.60% Galaxea d*eda 3 175.0 58.3 33.5 100.0 18.0 1.66% Monfiporasg. I 17.0 't7.0 0.0 17.0 17.0 0.34% Puitescyli'ldrica I 1310.0 163.8 174.6 54't.0 10.0 11.12% Porfies rus I 890.0 98.9 118.1 420.0 10.0 7.62% SedatWratlrydrix 5 365.0 73.0 90.2 250.0 10.0 3.24%

Tohl 30.07%

Algrc ?Algalspp, 12 1845.0 153.8 1 15.1 460.0 15.0 15.58% Hdlnedaquntia I 50.0 50.0 0.0 50.0 50,0 0.62% Jaria sp. 5 950.0 190.0 1f 1.3 435.0 35,0 8.12% Totrl aSrh

Subfffi Conl mlr 6 650 108.3 143.1 1't0 10 5.62% Rubble 12 2530 210.8 317.0 1190 20 21.28% Dead sbndirU conl 11 548 49.8 44.6 170.0 4.0 1.76% Rutrble ad silt 5 670.0 131.0 63.1 230.0 40.0 3.87% Sand 2 2$.0 't15.0 35.0 150.0 80,0 2j2% sitt 2 150.0 75.0 25.0 1m.0 50.0 1.45% Siltand sand 4 755.0 188.E 52.5 265.0 110.0 6.19%

To0d llSn Tobl $.5Sr

Table 13: Southern fulolo Deep, mid-tcVw, 3m fupth profiIe (109m): Tlanseets 7-11.

tlcln ,, SS. l!14il, .,,,r,,litn; P6rwrt DeY. oriver

Conl 17 325{.0 191.1 199.18 n6.0 10.0 30.80% ^crqorrno0tbFoclbpondadcome 2 32.0 160 6.m 21.0 8.0 0.2996 Podascyftdtbr- 11 1920.0 171.5 208.57 640.0 30.0 17.48!I Podasrus t3 1656.0 127.1 112.86 440.0 2.0 15.07% Sordopor!r16th 5 110.0 n.0 12.08 45.0 10.0 1.m%

Tobl 6a.61%

Algil ?Ahslspod€s 70.0 70.0 0.00 70,0 70.0 0.Bf% lldlrlpdacp-. 5.0 5.0 0.m 5.0 5.0 0.05% To'trl 0.6$t

Subttntr Coral toct 1 30,0 30.0 0.00 30.0 30.0 0.27% DBd lbndhg conl 19 19t0.0 102.1 92.07 320.0 5.0 16.18t6 RubUe 12 n2.0 6t.3 W7 n2.0 5.0 7.03% Rubble and sand 4 280.0 70.0 32.60 115.0 25.0 2.55% sand 11 917.0 83.f 118.58 450.0 r0.0 8.34% Tobl t0.9nt Totrl 34.6rr

27 Table t4 fuurllzm fulolo rbp, upper miS-terrry, B-ilm fupth profiIa (9gn): rlanea.,ifs 12-16.

AcloForaraqpona 3 190.0 63.3 40.28 120.0 30.0 1.90% AcraBonnobllis 2A 1891.0 94,6 114,75 490.0 6.0 19.91% Galarea as-lieata t 38,0 38.0 0.00 38,0 38.0 0.38% tulontipnsp. 1 60.0 60.0 0;00 60.0 60-0 0.60% Pawnavaians 1 35.0 35.0 0.00 35.0 35.0 0.3516 Forllcsc?/idnca '2.0 33 1171.0 35.5 29,2'l 125.0 11.71% Pcxlllqoradanicomts I 73.0 73.0 0.00 73.0 73.0 0.73% Fuiteshtea 'l 't00.0 100.0 0,00 100.0 100.0 1.00% Poribsrus 52 2074.0 39.9 53.99 310.0 2.A 20,26V0 Seilatryrchystlx 11 439.0 39;9 18.36 65.0 12.0 4.39%

Tolal 6tlfrltlc Algae ?A$al speohs t0 625,0 62.5 45,8S 180.0 10.0 8.25% 0oralllneapae 4 144.0 36.0 24.60 750 13.0 1,44% Dlctyob sp. I 15.0 15.0 0.00, 15.0 160 o1s% Hdindaopunta 7 217.4 31.0 20.08 60"0 7.0 2.17s6 Tuibinaiaomala 2 65.0 32.5 17.50 50.0 15.0 0.650/6

Tatal 12.6670 Subdrats Coral ru* 29 1611.0 55.6 60,23 280,0 3.0 18,0170 Ddstrndingconl u 618.0 28"1 31.10 110.0 2.0 6.18% Rubble I 206.0 25.8 19.95 60.0 4.0 2,06% Rubble and sand 5 216.0 43.2 18.30 65.0 10.0 2.16% sand I 50.0 s0.0 0.00 50.0 50,0 0.50% Toial 983nl Tohl n91%

28 Exposure and Current 3.5 Transverse Ridge and the The area is very protected by its depth and Southeast Margin of the Main being bound on both sides by the margins of the Deep. The current is a pronounced featwe of Palolo Deep this area. The boundary affects the current which (Transects 25i 26a,b; 27 a,b; structure by creating a constriction causes the current to accelerate with velocity 28a,b) second only to the Deep margin near the platibrm. This flow provides improved conditions for the coral development which Description General floruishes on the ridge. The ridge is a natural boundary between the Main Deep and the Southern Palolo Deep. This Depth area is similar to, and extends from, the coral zones of the Southern Deep. The bathymetry of The depth in this area varies, being generally the area is different, sloping to a greater depth- defined by the 6m depth contour on either side Laterally, the Transverse Ridge mounds from of the ridge. From here it rises to 3-4m at mid- 10rn on the southern side with a gentle slope ridge, shallowing to 2m at the base of the into the Marn Deep (Figs. 6m, 6p). The area eastern rim. The profile across the ridge is extending along the perimeter of the Main Deep gentle with the steeper slope on the southern progressively steepens to a vertical relief (Fig. side. The slope becomes more abrupt along the 6o). The coral cover is high and principally south-east side of the Main Deep' Here it conrprises Acropora spp. descends to depth with a 45% slope. The profile consists of a 2m level at the base of the rim with the coral cover being iimited to 5m. The base of Location the slope is at 10-12m where the sandy bottom The area is midway along the north-south axis grades slowly to >20m (Fig. )' of the Palolo Deep. It begins from the southern Dead standing coral and the combined side as an extension of the terraced area of the substrate of rubbie and sand occur in equal Southern Deep, extending to the western quantities on the transverse riclge (11% and an area perimeter (Fig. 3). It also comprises 12% respectively). In the coral covered area extending to the north-east, along the south- along the south-east rim, the dead standing of east margin of the Main Deep for a distance coral component becomes dominant with 29% 120m. coverage. Rubble extends to depth from 5m- Lateral to the ridge to the north and at a depth greater than 12m, fine sand. predominates-

MLW

2.5 D€Pr! m5

7.5m 0 50 l00m metre East West Traosverse Ridge Rim Rim

General profile of transects irt the area of Tlansverse Ridge (see Fig' 5)'

29 Algae Dynamics In all erreas where there are branching The ridge area is consolidated by sprawling Acropora spp. is also the filamentous algae branching Acropora spp. This consolidation (Clad,oph,oropsr.s sp.?) growing from the base of extends along the Deep margin. Below this, in the coral thickets (Fig. 6I). As with the rest of the steeper areas, is a talus slope. Here the the southern margin, the rim is predominantly source of material is from the growing rim, coralline algae, and other genera such as which is greater than 5m. The bottom of the Turbinaria and Halimeda present (Fig. Ge). Its Deep comprises the fine silt found at the bottom seaward development becomes pavement-like , of this zone. capping the area of vertical relief @ig. 6f). The water is generally turbid and the rock surfaces are covered with silt. The current, at Coral depth, is minimal with the transport largely taking place the shallower The Transverse Ridge is characterised by in layers (< Bm). Deposition branching Acropora nobilis with clumps of occurs throughout the Deep as material is washed Acropora. grandis. Porites cylindrica is in and settles as the current slows with depth. Coral growth represented as a minor component (4o/o) is limited by substrate, the constant becoming dominant along the eteeper Deep rain of silt and the availability margin (29%). Porites rus is the next most of light in the deeper areas. Zooanthids, as principal dominate with 10% coverage. The Porites monopolisers, flourish in the absence colonies are often large along the steep rim, of other benthos such as coral or forming buttress or cornice-like features. In alsae Gig.34j). some cases these have toppled. The Remainder of the Palolo Deep Community type The south-east margin of the main body of the Palolo Deep continues The community composition varies with depth to the north-east with the relief steepening and habitat. This includes the tidal and wave to vertical surfaces often inJluenced undercut (FiS. 6o). Rubble slopes occur rim, consolidated by coralline algae intermittently. (Fig. 60. At greater than 2m, the area is The algal crest is most mainly covered by branching coral. Branching developed in this area with the surface of the reef heavily cemented coral with the periodic occurrences of Porites flat Gig. G0. Coral growth limited rus and P. cyclind,rico is present on the steeper is by the relief with larger colonies appearing to be portions and in uncolonised areas of rubble and nearly toppling. silt. At depths greater than 5m a rubble slope The north margin comprises rubble dumped occurs with little coral, colonised by expanses of over the edge of the Deep, forming a talus slope Jania sp. and concentrations ofzooanthids. (Fig. 6n). This area extends to and along the western At the base of the slope is a sandy, silty margin. The profrle becomes more vertical substrate with large coral rock or relic bommies as it approaches the transverse ridge. These surfaces appear which have been recolonised by zooanthids and to be in a state of more limited hard coral. Some Latge Porites rapid erosion, as there is little coral growth. rus profile colonies occur with subetantial dead, silt The depth is through the section of maxinum width and depth. covered areas. The sand is devoid of obvious benthos with the exception of several species of holothurians (Holothuria sp., Bohad,schia sp., and Stichopus sp.) (Fig. 6p).

MLW

Dcptb.15

25r 0 metre 480m West Rim Main Palolo Deep F-qt Rim

General profile of tronsects in the area of Main palolo Deep (see Fie. S)

30 Table 15: TYansuetae ridge uross thc fulala Dep (40m): Tlaneect 25.

Coral Auopora grandis 4 110.0 27.5 8.29 40.0 20.0 2.750/o Acropora nobilis 6 930.0 155.0 21250 610.0 10.0 26.35% Porites cylindrica 1 170.0 170.0 0.00 170.0 170.0 4.250h Sotl coral 1 20.0 20.0 0.00 20.0 20.0 0.50%

Total 33.85Yr

Algae Filamentous algae 1470.0 735.0 405.00 1 140.0 330.0 36.75% Coralline algae 160.0 160,0 0.00 160.0 160.0 4.00%

Total 10.15'/o

Subskate Dead standing coral 3 440.0 146.7 157.92 370.0 35.0 1'1.00% Rubble and sand 1 480.0 480.0 0.00 480.0 480,0 12.Moh Sand 1 100.0 100.0 0.00 100.0 100.0 2.500/0 Total 38.8m Total 25.50%

Table 16: Sou,theast ,ndrgin of the Main Palolo Deep (120m): TYansects 26a,b; 27a,b; 28a,b.

,: : $en ,$il1 ililAii, ttt,:.i.,.:,i.: i'itffl;. 0or,

Acropora grandis 4 260.0 65.0 I 1.18 80.0 50.0 2.17% Acropora nobilis 8 2020.0 252.5 164.45 530.0 40.0 16.83% Pocillopora danicomis 1 10.0 10.0 0.00 10.0 10.0 0.08% Montipora sp. 1 15.0 15.0 0.00 15.0 15.0 0.13% Porites cylindica 14 3465.0 217.5 216.80 760.0 25.0 28.88% Poriles rus 14 1205.0 86.1 4s.83 160.0 30.0 10.04% Seriatopora hystix I 100.0 50.0 40.00 90.0 10.0 0.83%

Total 58.96%

Algae/ Filamentous algae 1 100.0 100.0 0.00 100.0 100.0 0.83% rooanlhids Zooanhids 1 430.0 430.0 0.00 430.0 430.0 3.58% Total llllr

Substrate Conl rock 3 380.0 126.7 '124.99 300.0 10.0 3.170h Dead standing coral 14 3420.0 244.3 165.99 580.0 50.0 28.50% Rubble 2 A(n 32.5 7.50 40.0 25.0 0.54% Sand 4 530.0 132.5 65.34 240.0 80.0 4.42% Tobl l2llm Total 36.dt%

Transect 25 Transect 26 a,b;27 a,b; 28 a,b

drrd rL corel ltj lncr rtgl 15,71

lcroport ll rul l.lt rrbbl. & tud 11 nd rl3r I cctd trbblr lJ{ roll conl 0,5 ccrd rocl 3.17 Porltg r1.25 tr.r. tllr. l.al rooullkb t.5t 6.rbtolron 0.tt

dcrd rL tord lt Porllcr 18.12 Acrogon 2f.1

Fig. 12: Graphs of the percentage composition of transects 25; 26a,b; 27a,b; 28a,b.

3l Augmenting this flow is the shape of the reef Seaward crest whereby waves are refracted into this 3.6 Reef Flat opening. This area, represented by a rubble strewn reef The small Deep is located at the edge of the reef flat with rubble banks (Fig. 6g), is exposed at, crest 100m to the east of the Main Deep. Its low tide and wave-swept at high. The substrate nature is simiiar to the seaward portion of the is unstable and characterised as a boulder zone larger Deep. It is mantled by coralline algae with the material being continually transported and has rubble slopes that have been cascaded shoreward by wave action. into it from the seaward side forming a talus slope. At depth the reef face is Despfte steep and often its superficial rubble natule, the undercut on the eastern and southern margins. underlying reef structure is perhaps moet the Significant coral development occurs in areas of consolidated of all the reef zones dense, with its stable substrate. Distinct from the Palolo Deep, pavement-like nature resulting from growth in this area is more conducive to coral growth, as a high energ"y environment. crest, The reef it has no suspended silt problem and possesses foom which the spur and groove system an environment where wave action aerates and. originates, forms a buttress against wave continually changes the water. action. Rubble is constantly being generated by this action. Cyclone Ofa gave rise to prominent banks by the removal and transport of fore-reef 3.6.1 The Spur and Groove System materia-l to (Zam this area and Sua 1991). This area forms the buttress structure which The zone is impoverished by the effects of buffers the force of the oceanic swells (Fig. exposure (dehydration, temperature extremes 13a,b). Reef development here is the most and wave action) limiting the biota. This is consolidated of all the reef structures. The largely colonised by coralline algae, which has bathymetric relief is irregular. The channels cementing powers to bind the rubble and create and extensions from the reef flat form the spur a wave resistant structure. The rubble exists in and groove system. These extend as ridges on varying sizes and, as a result, has a variety of the reef slope with depth, and consequently habitats. The larger, more stable pieces, reflect form channels often scarping to 2m and a rocky shoreline habitat whereby exposure descending in a branching pattern. Sand and mediates faunal occurrence. Most of the fauna rubble exist in the bottom of the scoured is cryptic, particularly at low water, or channels. nocturnal. Functionally, the spur and groove system Oecurrilg in this zone are two interesting breaks the wave force, moderating the effects of structures, the si:award entrance to the Deep, this high energy zone. The grooves or channels and a small Deep outside the Reserve (Fig. allow the transport of reef material down the 13a). The entrance comprises shallow channels front of the reef. The surging action of the resulting from a reef structure similar to the swells scour the channele with this rubble and spur a1rd groove system. Oceanic water sand. This zone is universal to exposed areas constantly enters the lagoon through this and is also described at Aleipata (Andrews and feature, beiug driven by wave action. Holthus 1989). Fig. l3: a) Photograph showing the spur and groove systerm, the seaward entrance to the Palolo Deep and the adjacent smaller Deep.

b) Photograph showing the branching chawtels of the spur and groove systen extending d.own the reef slope.

32 3.7 Seaward Reef Slope The reef slope relief is characterised by a eeries of branching channels similar in configuration to the spur and groove system. Though the General description slope profile is gradual with respect to the bathymetry at the level of the ridges, the The seaward reef slope is an area characterised general profrle is one of undulating relief with by exposure to oceanic waves, rugged relief and channels of varying depth (2-7m). The channel a bathymetry ranging from 0-30m. It is unique sides are steep with a sandy floor. in that it has been totally denuded by cyclone Ofa and is now regenerating. It is covered by The reef substrate is largely consolidated coral patches of encrusting algae and small coral rubble and rock with loose rubble and sand in colonies (Figs. 6h, 30-32). It is the zone which the channels. Unlike the fine silt of the Palolo extends from the exposed spur and gxoove Deep, the margin of reef base is coarse, white system at low water, to the sandy margin carbonate sand. Loose sand and rubble are offshore. found in the channels and are subject to constant movement by wave action. Location Algae This area is the northern most seaward margin of the Palolo Deep Marine Reserve. It is the The most prolfic benthos at site 3 are patches central part of the reef front of East Reef (Figs. of an unidentified brown algae that encrust the 1,3), which extends from the eastern margin of reef surfaces at >3m. Halimed'a sp. is common the Apia harbour entrance to the west, to the among the rubble hollows. Coralline algae Viala river reef pass to the east. occurs in light coloured patches, bordering each other as they grow together on the reef surface. They are the principal colonisers, collectively Exposure and current making up most of the living cover. This habitat is the most exposed in the Reserve. It forms a buttress against oceanic swells with Coral the dense reef structure and benthos conforming to a high energy system. Depth is a The occurrence of coral is represented solely by substantial buffer, with breaking wave action recolonisation after cyclone Ofa. All previous conlined to the upper margin. Strong surge coral was removed by the cyelone. At present, occurs at depth. The current is longshore and numerous small coral colonies are evid.ent on seasonally variable. the reef slope. These comprise five genera. The only recruit identifiable to species level is Pocillopora eydauxi. Acropora spp. form the Depth overwhelming majority of coral settlement, From the intertidal spur and groove system comprising 640/o at site 3 and 87Yo at site 4 inshore, the reef slope descends to a depth of (Tables 17, 18). By assessing the modes of the 30m. Its initial descent from the spur and colony diameters, age classes may be inferred: groove system is 2m, followed by a 15o-30o Densities of 7.8 colonies /mZ were recorded at slope. At its most seaward extent, it descends site 3 and 33 coloniesimZ at site 4. 8-10m vertically, to a coarse sandy bottom.

MLW

D"pth a

30m 0 75 l50m metres Reef Spur & Reef Slope Sand Crest Groove Bottom

General profile of transects in the areo of the seaward reef slope (see Fig. 5).

33 At site 3, L7L colonies were measured, The lack of living habitat has simplified the reef representing four genera (Table 17). They had community with the absence of commensal a mean density of 7.7 colonies/m2. The fauna. S$ corals were established, and with Acropora spp. were the most numerous with 5 their reladively faster growth, colonies were coloniesr/mz. The relative numbers illustrate observed up to 30 cm diameter. Large the preponderance of recruitment from the encrusting colonies of Diploastrea heliopora and Acroporidae: Acropora and Montipora spp. Porites sp. have survived the cyclones, though (Fig.15). Recruitment by a six month (4cm dead patches indicate some damage. diameter), eighteen month (6.5cm) and two and a half year (8.5cm) old colony was discernible in Dynamics the Acroporo (Fig. 14). The colony size range was 1-12cm. For the Montipora, three age The reef slope, typically the most luxuriant of classes were discernible for similar time periods all the reef environments, has suffered the with 2cm, 5cm, and 7cm. removal of virtually all of its living material. That, together with other reef debns, has At site 4, five genera are represented with a cascaded on to the reef been be semple size of 598 (Table 18). The composition flat, or left to consolidated on the slope. This process of the sample is overwhelmingly Acropora (Fig. shows a of reef development where the reworking of the 17). There are two size classes discernible at substrate yields a new varied 3cm and a 6.5cm size class. This presumably surface of relief. Recolonisation has been underway for at least represents recruitment at srx months and two years, with good settlement. 'eighteen months old (Fig. 16). The lack of a third size class indicates no recruitment the The reef structure has developed in response to frrst year after cyclone Ofa, which may relate to a high energ-y environment with respect to its the occurrence of cyclone Val. The minimum topography and consolidation. Through the and maximum colony size was lcm to 13cm. presence of channels and buttresses, the spur and groove system breaks the force of the wave Fish action. This also conveys sand from the reef crest or flat system to fore-reef areas. This Large numbers of herbivorous fish frequent the sand and rubble transport is vital to the reef front. The fish are principally scarids, formation of the system. The depth of the labrids and acanthurids grazing on the channels is the result of sand-abrading wave encrusting fleshy and coralline algae. Habitat action. dependent species like the damsel fishes were The understanding of eustasy is important to seen colonising the many spaces amongst the the explanation of reef morphology. The rubble. Top level consumers were rare general reef topography was formed as the although a coral trout and grouper were result of aerial erosion during the interglacial sighted. Pelagics were represented by a small periods of the Pleistocene (Fig. school of unidentified scombrids. Schools of 18). This process is discussed geomorphology antherinids and caesios were seen about the in the section (see Section reef surface. 4).

Gommunity type This area is characterised by the recolonisation of the fore-reef, after the total removal of coral fauna two years previously by cyclone Ofa. Small colonies which had settled afber cyclone Ofa were widespread and appeared unaffected by cyclone Val. Both coral and algal size categories indicate that the small colorues are the result of mass spawning @allace and Bull 1981; Wallace 1985 a,b; Babcock et al. 1986). This would have occurred in the months of October and November, for the two or three seasons after cyclones. Coral growth appears good, discernible at generic level.

34 Table 17: Combircd ruf slope belt trensects at site g.

tue , r6e| trfi, MeL ' qqBkl Inbfpet Dlv' nci# ,:.:'.'{qin}

Acroprasg. 110 621.5 5.65 2.10 11 1 5 MffiWraW. 18 239 1.98 2.02 12 2 2.18 Poribs sp. 1 13 3.25 1.30 5 2 0.18 Podlopraeydwrt I 51.5 6.06 3.12 12 2 0.41

Gouprffrdcr: Totrl no. tum tcrn s'8. Ier. Iln. Drnrfi

171 928 1.98 0.76 3.4 1 7.77

Table 18: Combincd reef slope. belt tran*cts at site 4.

1,1lB,.. Mh, Dertsry .l' t{ Ioa r 86, iril. lhbttrpt De{. nosJrrtt' :' ffi

Asqponsp. 521 2630.1 5.0 3.05 55.0 1.0 15.88 Gorns0oa sp. 2 9.0 4.5 1.50 6.0 3,0 0.16 tulcrlrlipadq. 21 107.0 5.1 1.62 8.0 2.5 1.87 Pcr//opraeydouxi 15 79.5 5.3 2.11 8.0 2.0 1.03 Ponfles sp. 39 1E8.4 {.8 2.07 12.9 2.0 0.4r

Goupdrtr0c: Idlm. Sun Icrn $d.dfi. tu. Iln. Ihnrfi

598 3014 1.95 0.07 55.m r.00 19.35

J rn/ltopoo

El,,' , porlrcs H tt WI./iif/Ii Mooripon g o A"mpor" rlI W

120

100

80 1 2 3 1 5 0 7 I I 10 11 12 Slze (cm) ou

Fig. 14: Frequency/size of coral colony dianeters of four genera at site 3.

Numbers of colonies

Fig. 15: Colony numbers per genera at sitc 3.

35 120 J peltlopon 100 l:ffi ?otlrot 80 Itffflffffr Mootlpon 60 W ,l.ropot &

20

0 | 2 3 .a 6 6 7 I 010t.t t2rs She (cm)

Fig. 16: Frequenqt and size of coral colony diameters of four gencra at site 4.

Fig. 17: Colony numbers per generd at site 4.

36 3.8 Fish Survey balistids, serranids and holocentrids found. only in the Deep. The Seaward Reef Slope (transect The survey of fish at the Palolo Deep involved 3l) is characterised by an abundance of frsh. compiling a species list (Sec. 7.3) and an initial Eighty-eight individuals in 5 families were description of differences between habitats. recorded. Occurrence here did not correlate One hundred and eight species in 32 families with Iiving coral cover but rather with rubble were identified within the Palolo Deep Marine reliefand an abundance ofalgae. Reserve. The swimming transects detail the The Palolo Deep offers protection to the fish presence of families and relative numbers fauna in a variety of ways. Fishing is (Table 2l) at monitoring site 2. Though ' :lcing prohibited, allowing a reproductive population replication, it does provide a ;risrdl for recruitment. It would have been description. Presence/absence information and particularly important as a refuge during relative abundances describe the differences cyclone Ofa when other areas were inundated between the habitats (Table f9). by the storm. During the diurnal tidal rhythm, The most impoverished area was the Western fish retreat to the Deep at low tide, returning to Reef Flat (transect 21) with a low complement the r',ef flat to graze only at higher water. of fami,ly presence and abundance. By contrast, Species associations, exclusive to habitat, are the two transects (I, II) run in the southern evident with the high abundance of coral eating Palolo Deep had a relatively high abundance of chaetodons among the Acropora nobilis 62 and 101 individuals respectively, in nine thickets. The territorial Bluntsnout Gregory, families. The Transverse Ridge had 213 Stegastes liuidus, is also associated here in individuals, largely due to a school of large numbers (4 individuals/mz). acanthurids that entered the survey area. The Eastern Reef Fiat had intermediate numbers, At monitoring site 2, fish transects (I, II) were with a decrease seaward. conducted for comparison with other habitate and to provide some basis for resurvey as part The degree of coral cover in the habitat of the monitoring programme. Table 19 appeared to be controlling abundance. Habitat provides a record of this information. preference is shown by the presence of larger numbers of pomacentrids and chaetodons, with

Table l9: Conparatiue fish abundance at nonitoring site 2, the eastern reef flat, the transuerse ridge and th,e seaward reef slope. (The sample transect area is 40m X 2m.)

Transect no. I tl l7 t8 t9 21 25 3f

Family:

Holocentridae 43 0 0 00 0 0

Senanidae 1? 0 0 00 0 5

Mullidae 41 3 o 56 o 0

Chaetodontidae 14 17 0 40 I 0

Pomacentridae 32 49 1 612 33 20

Labridae 35 CI 14 JO 10 18

Scaridae 00 0 12 60 0 15

Acanthuridae 46 1 0 00 130 30

Siganidae 116 8 ? 30 30 0

Monacanthidae 00 1 0 00 0 0

Balistidae 31 0 0 00 0 0

Others 00 I 4 00 3 0

T6ht;

Transaci locations:

l, ll : Monitoring site 2 transecls 17-19 : Easbm Reef Flat Westem Reef Flat 25 : Tnansvene R'xlge 31 : Seaward Reef Slope

37 4. Geomorphology

It is here that the decreased bathymetry has 4.1 Reef Origin allowed the formation of barrier-like reefs. Reef development to the east is much reduced where the oldest volcanics (Fagaloa) of much older The living reef types found in Western Samoa Pliocene age have lost their barrier reefs during are wholly fringing reefs, although Moreton et the Pleistocene submergence or sea level rise. al. (1989, MS) in following the terminology of The bathymetry here is much steeper inhibiting Stearns (7944), classif.es the seaward crests of reef development as it was unable to keep pace some of these as barrier reefs. with the submergence. The formative elements of accretion and erosion Relic or drowned barrier reefs (at b0-80m near combine with the processes of conditioning by shore and 80-100m depth, occurring l-l.b km sea level fluctuation and periodic aerial erosion, offshore) have been speculated (Moreton et al. vertical movement through tectonic activity, 1989 MS.). These would be much older than and the overlaying and establishment of new the present reefs and would have been exposed surfaces by lava flows, to create the present aerial reefs to erosion several times during the Pleistocene when sea levels were at their Assessment of coral reef history is complicated. lowest. Rather than their demise due to rapid Early development was in the post-pliocene, submergence, it is likely that volcanic activity where it grew on an erosion built platform was responsible. reeulting from the degradation by pliocene The area overlaid by Pleistocene lava flows was volcanic activity. On Upolu, the situation was extensive and those areas peripheral to them unique as many of the old reefs were buried by volcanic activity (Kear must have been affected and probably and Wood lgbg), extinguished. providing new surfaces for development when the sea level rose. The Pleistocene is the epoch of the Ice Age. It was characterised by several major periods of The Palolo Deep reef developed on the eastern glaciation separated by warm intervals. During end of a gentle sloping she1f, was created each glaciation the polar icecaps increase their during the mid to late Pleistocene by the volume, which consequently causes a drop in Mulifanua volcanics. The . nost extensive sea level. Eustatic fluctuation during the Plei- Western Samoan reefs also occur on this stocene (see Fig. 18 below) alternatively caused substrate extending along the north coast. these reefs to emerge and drown. Sea level They are much more expansive to the west with fluctuated between present mean sea level and their seaward margins two miles off the coasr. minus 150m (Chappell 1981).

f;,rss 4,tss-t$rr$(- \Nr:cr.t, -4s.rqqsrl-- 4txrru- ir'rra.5r.rg]s

30 o 50

I

:.fr- -50 ryc 60 40 nt t.o 50 Years beJore Preselt (rl0r)

Fig. 18: Sea leuel fluctuatinns in the late pleistocene era.

38 During the last glaciation (20,000 years ago), 4.2 Blue Hole or Embayment? the sea dropped to over 100 metres below its present level. Worldwide, coral reefs were The most interesting feature of the Palolo Deep exposed, often left high above sea level. This Marine Reserve is the Deep itself, a steep sided period was so dominated by sea level changes lagoon embedded into the reef flat. Moreton et that for about three-quarters of its history, most al. (1989) attributes this feature to the of the reef areas that we see today have been progressive closure of one of the convolutions or out of the water. Acid rain can etch through embayments "by which the barrier reefs outer limestone with comparative ease, so that during margin is inturned into a grotto, or closely the centuries when rain fell on the carbonate sheltered'deep hole"'. rock it was dissolved. At those times the reefs physical theory. would have been eroded, producing caves, Two features support this gutters and valleys in the limestone matrix. Firstly, the maximum depth in the Deep is After they were flooded by the sea, marine life equivalent to that at the base of the seaward would gradualiy re-establish and corals would reef slope. Also communication between the Deep and the ocean exists through a start to grow fVeron 1986). still shallow channel in which the spur and groove The reef is built by layers of coral and coralline system is shared. The embayment prior to algae, growing upon the eroded gutters and closure may have been the old river course of valleys. To a large extent, these eroded the Vaiala River, now responsible for the substrata govern the shapes of modern reefs. adjacent embayment. The caves formed through this dissolving process subsequently collapsed, forming Deeps Alternative hypotheses concern freshwater pipes which or Blue holes. upwelling via volcanic tunnels or concentrate the emerging groundwater into During the period of submergence from the late wells (Kear and Wood 1959) inhibiting coral Pleistocene to a Recent or Holocene high sea growth. Solution is a likely mechanism for the level, drowning of reefs occurred on the steeper formation of the Deep, whereby a cavern has coasts. As well as submergence, emphasis must been created by freshwater dissolving the be given to the erosional aspect of the steep reef carbonate, leading to a cave which has slope, where early coral colonisation and subsequently collapsed (Palmer 1984). consolidation is prevented through lack of recent protection in a high energy environment. The This hypothesis mitigates against the because the prgcess present reefs owe their occurrence to the age of the reefs. This is periods, when the reefs reduced slope of the lava flows, which permit occurs during subaerial exposed a lowering sea level. successful settlement and development have been by of formation of the reefs (< 10,000 (i\4oreton et al. 1989 MS). The recent years ago) would not have allowed this to occur Due to volcanism, development of most of as older reefs had been covered by volcanic Upolu's reefs is very recent. It occurred on a material in the Pleistocene. If this volcanic surface that was laid down in the late activity occurred prior to 150,000 years ago, Pleistocene when the sea level was lower than then the mechanism is valid. This is because the base of the existing reefs. This provides us sea level reached present levels twice during with a minimum reef age of about 8000 years. the Riss-Wurm interglacial before receding for B.P. for the Palolo Deep. This presumption is 100 thousand years (Fig. 18). The development based on known sea level fluctuation without of the blue holes on the Great Barrier Reef in the complication of tectonic activity. The Australia is thought to be the resuft of several seaward reef base is 30m deep and would not low sea level periods @achshall et al. 1979). have been subject to colonisation until that time known as holes', submerged (Fie. 18). Variously 'blue sinkholes or drowned dolines are thought to be Though tectonics and the likelihood of the reef a karst feature developed during the structure being greater in extent than is Pleistocene lowering of sea level (Agassiz 1894). evident may modify this presumption, it is These occur throughout the world with most of unlikely that it is older than the Holocene (< the examples described in the Caribbean 20,000 years B.P.). This means that unlike (Agassiz 1894; Jordan 1954; Dill 1977; Stoddart most of the coral reef systems of the world, the 1962). Blue holes exist in Australia @achshall present reef relief is not the result of a living et al. 1979) and at Bikini and Eniwetok atolls veneer, overlaying the karst or weathered @mery, Tracey and Ladd 1954). topography of the pre-Pleistocene reefs.

39 The origin of the blue hole has been considered The strongest cyclone ir 150 years (Ofa) in the context of the antecedent karst followed a year later by a strong cyclone (Val), hypothesis whereby recent reef growth is but a altered the physical nature of the Palolo Deep thin veneer over the aerially eroded Pleistocene and its adjacent reef area. The outer reef slope reef growth @urdy 1974). The term 'blue hole' was assaulted by huge waves for a week. at comes from the colour of the deeper water of the which time the existing coral reef was removed. feature in contrast to the lighter green/brown This material progressively scoured the reef colours ofthe reefflat. front down to bare rock, with a large portion of it finding its way on to the reef rubble From the description, the Lighthouse Reef Blue crest as banks. During this period and with rhe Hole in British Honduras (Ditl 1977) is a subsequent cyclone Val, similar feature with respect to its superficial much of this material was deposited appearance from the air and with respect to its into the Deep forming rubble slopes on the northern and western ecological zonation, debris concentrations and sides. fine sediment bottom. Elements of profi-le such Secondly, though cyclonic activity adds material as the undercutting of the vertical wall areas to the Deep, its turbulence may serve to remove indicate that it may indeed be a collapsed cave. substantial quantities of the suspendible Porosity in the walls, where rubble material has material. Though the large quantity of rubble also been consolidated, is another similarity. created by cyclone Ofa may represent a once in a hundred and year The blue holes on the Great Barrier Reef are fifty event, this is but a moment in the history of the Deep where such similar to the Palolo Deep in that they have a an event must have occurred many times. The coral rim Slving way to a steep slope with a fine rubble sandy bottom. There is high coral cover near slopes had been well established prior to cyclone Ofa (Zann and Sua 1991; 1978). the surface, diminishing with depth. Below the Dahl shallow area of coral development, rubble predominates, often occurring in chutes whereby it slides down depressions in the slope. 4.4 History of the Palolo Deep for the The relatively flat bottom is characterised by last 39 years creamy, sticky mud. The most recent changes in the Palolo Deep The description of collapsed dolines (Bachshall concern terrestrial changes around it. et al. 19?9) provides a good description of a Particularly over the last 20 years, urban feature similar to the Palolo Deep. Futher development, including land fill reclamation, evidence for this theory is provided by the and the clearing of the watershed have presence of a small Deep to the east. This increased the silt load in the rivers and streams structure appears as a collapsed cavern. It is discharging into the Currents, most improbable that this small but relatively ocean. particularly with respect to circulation through deep structure could be created by a closing off Apia harbour, convey this silt across the East of a convolution of the reef margin. reef. Here, the Palolo Deep acts as a sink where accumulation occurs. This silt is continuously resuspended, affecting the living 4.3 EnviromentalChange environment within.

From the coral growth in the bottom of the Inspection of aerial photography taken Deep it appears that change has occurred. Now periodically since 1954 provides an insight into appearing as relic material, coral growth must some of the changes that have taken place on have beeu luxuriant at greater depth in a the reef flat (see Fig. 19). Though irter- former time. pretation of the aerial photos can be difficult due to differences in perspective and The following processes appear to maintain the photographicaily induced contrast, valuable Deep. Firstly, coral growth about the edges is information can be gained by a general Iimited by the angle of the slope. Consolidation comparison. is confined to the rim with the area below less so and the coral colonies subject to toppling as The presence of rubbie banks is a consistent they increase in size. Depth and turbidity limit feature of outer reef flat topography. Though the vertical extent amenable to growth. The now more extensive after cyclone Ofa (Zann and east/south-east portion of the Deep is the best Sua 1991), these banks are always present as example of this, with its steep slope and shear the result of the constant accumulation of fore- faces @ig. 6o). reef material over time from varying degrees of storm and large oceanic swell conditions.

40 The reef flat benthos is seen to change in At the rim is an area of surface consolidation. location and concentration. The protection of Underwater inspection reveals erosional the eastern reef flat by the Deep, allowing features such as undercutting at the large greater benthic development, is evident in the Porites bommie (Fig. 35d) and in the vertical 1954, 19?0 and 1993 photos. The western reef reef face near the entrance (Fig. 6o). A flat shows a marked change in the 1954 and prominent talus slope occurs along the south- 1970 photos. This may relate to such events as east margin of the Main Deep. It is likely that the extensive landfill and coastline alteration these features are a result of cyclone Ofa, as the dwing the harbour dredging in the 1960s and perimeter profiIe indicates growth and not the construction of the main port in the 1970s. progressive erosion. The current regime was altered, changing the Other post-cyclone Ofa features, evident from western reef flat environment. Ttris change is the 1990 photo, are the large rubble banks with coincidental with the rural and urban the reef flat benthos of 1970 removed. These development, where river borne silt loads have rubble banks represent a change in the two progressively increased. The early 1980s saw habitats. With their origins from the reef slope, the development of the Royal Samoan Hotel this habitat is now open to colonisation. The project with its xtensive landfiIl and dredging area of deposition is now one of constantly adding to the suspended sediments. changing substrate, which, due to its relief, is more like a rocky shore habitat with varying margin of the Palolo Growth in the south-east degrees of intertidal exposure. Deep is evident since the 1954 photo with a change in the perimeter profrle. The intertidal In the 1993 photo the continued excavation of Porites flat extending from the algal rim into the Western Reef Flat is obvious. This had its the east reef flat shows greater development beginning as early as 1987, but became more when compared with the 1954 and 1970 photos. developed in 1990 and is now a major erosional An exception to this is the area buried by rubble feature. This is an area of substrate instability at the seaward end. and must contribute to the coarser component of the Deep's sediment.

4l Fig. 19: An. aeriol view of the Palolo Deep from the years I9S4 to 199J

1954 1970

1987

42 5. Monitoring Programme

Four monitoring sites were set up to allow continued assessment of the condition of the benthic assemblage over time (Fig. 20). For 5.3 Site 3 three of the sites (1, 2 and 3), each station comprises a series of quadrats located 10 Site 3 is located on the Reef Slope. It is metres apal't. In sites I and 3 there are three represented by a combination of three quadrats quadrats per station while in station 2 there are with two line transects extending along the six (Fiss. 21-32). edge of the quadrats and two belt transects adjacent (FiS. 20). Information from the line The composition of each was noted and transects is compiled in Tables 22 and 23. The photographed. The quadrats are one metre belt transect information is used in the square and are permanently positioned by steel description of the seaward reef slope (section stakes. These quadrats are positioned along 3.8; Table 17; Figs. 14, f5). The site location is transect lines of known position. The in line with the western boundary of the configuration of the lines and quadrats vary Reserve. The compass bearings are: western with the site. The line transects were analysed headland 2800 compass (most seaward), eastern as spreadsheet data and tabulated in the headland 99o compass (most seaward), eastern nanner of the general survey data. Site 4 is end of the tiled dome on the government represented by belt transects only. building 2100 compass, and east side of the Palolo Deep platform 1930 compass. The site depth is 10m. 5.1 Site 1

Site I is posit.ioned along Transect 18a,b 5.4 Site 4 originating from the middle buoy (no. 2) on the eastern Reserve boundary. The first quadrat is Site 4 comprises two belt transects. The located 10m from the base of the buoy, while information is compiled in Table 18 and Figures the second and third are located at 10m 16 and 17. The location of the site is marked on intervals along the line at 20m and 30m Figure 20. Its position is marked by steel respectively. stakes. The site depth is 12m.

5.2 Site 2 5.5 Monitoring Fequency

Site 2, near the transect ridge, is located in the The monitoring frequency should be at six Southern Palolo Deep in the general area of the monthly intervals or after an event of probable survey transects numbered 6, 11 and 16. This inJluence. At times when a particular effect or area represents the seaward end of the area process is occurring, the frequency of surveyed in the Southern Palolo Deep near the monitoring should be adjusted so that it is Transverse Ridge. Six quadrats have been adequate for description. The procedure should established with line transects connecting include photography of the quadrat areas and quadrat pairs, located on the same depth profrIe reassessment of the line transects. The (i.e. quadrat nos. 1-2, 3-4, 5-6). The line'transect monitoring effort must be commensurate with information is compiled in Table 20. the change to be assessed. A short comparative A fish survey for the same area is summarised description may suffice if little change is in Table 21. The sample is from two belt evident from the original photographs. Coral transects which run from quadrat one to five bleaching or death, storm damage, crown-of- and from quadrat two to six (see Sec. 2. thorns presence, changes in the relative Methods). composition of the substrate and biota are elements that should be recorded.

43 Ftg. 2O: trlncation af the nwni.tarittg sitcr.ctfi, fult, tnan;*et

Site 4 Stte 3 t2 I ,l-Tz g$ ff iT$ +Y

$cale 1i4600 Tm EFo 'd.,4.; ;

&4 Species identification: 1 Dictyot,a sp. 2 Montiporo sp. 3 Pocillopora datnicortis 4 Porites cylhtdrica 5 Porites ru.s

Fig. 92: Eastent, Reef Fla.t: si.te 1; quadrat 2

Species identification: I Acropora tr,obilis 2 Pocillopora da.micornis 3 Porites cylindrica 4 Porites rus 5 Psanrnocora cotr,tigua

45 Fig. 23: Eastern Reef FIat: site 1; quadrat 3 .

Species identifications: o*" L- Montipora sp. lY@/\-'l 2. Pocilloporo da.micornis K, j?= 3. Porites uerrucosa \:/ {7r:_4 3. Porites cylind,rica -f) \14:-t 4. Psammocoro corttigua eefi g;

Fig. 24: Palolo Deep: site 2; quad,rat 7.

Species identifr cation: L. Porites cylindrica 2. Porites rus

46 Fie. 25: Palolo Deep: site 2; quadrat 2

Species identifrcations: l. Acropora rrcbilis "vEfv 4 2. Janda sp. (- f^ n.-- \\ 3. Porites cylhtdrica r/ 7'"N\ 4. Porites rus &;trxi '4ffi$v% Fig. 26: Palolo Deep: site 2; quadrat 3.

Species identifrcations: 1.. Acropora rnbilis 2. Cladophoropsis sp. ffi_ffi

47 Fig. 27: Palolo Deep: site 2; quad,rat 4.

Species identifrcations: )j,#)s/ 7. Acropora,nobilis 2. Clad,ophoropsis sp. 3. Porites rus

Fig, 28: Palolo Deep: site 2; quadrat 5.

Species identifications: 7. Jani,a sp.

2. Pauona d,ecussata, 3. Halimeda sp, I A; f l^il J^v--- Jrr\S= D Fig. 29: Palolo Deep: site 2; quadrat 6.

Species identifi eations: L. Acropora rwbilis 2. Pocillopora damicornis 3. Seriatopora hystrix 4. Cladophoropsis sp.

Table 20: Moninring eip 2:Tlannfi A, B and C run between quad.rats 1-2, 3-4 and 5-6.

TAli; ... ;:,,:,,,;,

Tnneost A

Gonl Acropora nobilb 3 58.0 19.3 12.28 35.0 5.0 3.87% Podtes cylindrica 't 100.0 100.0 0.00 '100.0 100.0 6.67% Poriles rus 3 s20.0 173.3 78.78 250.0 65.0 34.67% Seriatopona hystix 1 685.0 685.0 0.00 685.0 685.0 15.670h

Tohl 90.88%

Subelrals Rubbb n.0 25.7 11.73 42.0 15.0 5.13% Sand 60.0 30.0 10.00 40.0 20.0 4.00% Totrl 9.13?l

Tnnsect B

Conl Acropona nobilb 2 n5.0 387.5 232.fi 620.0 155.0 n.50% Sedabpom hystix I 195.0 195.0 0.m 195.0 195.0 19.50% Totrl 97.00%

Subslrats Rubble 30.0 0.00 30.0 30.0 3.m%

Tobl 3.111196

Tr|nioct C

Conl Acropora nobilis 3 45.0 15.0 1030 30.0 5.0 4.50% Podtes cylindrix 2 110.0 ss.o 5.00 60.0 50.0 11.00% Porites rus 1 150.0 150,0 0.00 150.0 150.0 15.00% Sedatopora hystix 1 5.0 s.0 0.00 5.0 5,0 0.50% Tot|l 3r.004

Subsffie Rubble 150.0 0.00 1s0.0 150.0 15.00%

To0d t5.lxl!a

49 Table 21: Comparative fish abund,ance between the monitoring qtrcd,rats at site Z. (The segnents are 10m.)

0 0 4 0 0 3

0 0 1 2 0 I

0 0 4 0 0 1

t0 4 0 4 11 2

32 0 0 14 I 27

0 3 0 0 0 5

0 4 0 2 0 4

1 0 0 3 3 t0

2 1 0 0 1 0

Fig. 30: Reef Slopa site 3; qu.d,rat I.

Species identifications: I. Acropora sp. 2. brown algae 3. Montipora sp. 4. Pocilloporaeydouri

50 Species ide ntifrcatiou l. Acropora sp. 2. brown algae 3. Morttipora sp, o 4. Pocilloporaeydouxi Bo OV Fig. 32: Reef Slope: sitc 3; quadrat 3,

Species identification: l. Acropora,sp. 2. brown algae 3. Dictyosphaeria cauernosa 4. Montipora sp. 5. Porites sp.

5l Coml Acmpora sp. I 58.0 6.4 2.75 12.0 3.0 2.90% illontipora sp. 4 20.0 5.0 2.71 9.0 2.0 1.00% Polbssp, 1 3.0 3.0 0,m 3.0 3.0 0.15%

Totrl: 4.05?t Algae

Conllinealgae 17 19't5.0 112.6 80.s8 315.0 9.0 95.75% Halimeda 1 1.0 1.0 0.00 4.0 1.0 0,20%

Totrl: 2000 Totrl: 9535%

Tablc 23: Monitoring eitc 3: Raf alap transect ad.jrcent qudrat on westsidc.

:,",, Iira, , Tohl :llbao ,, ;,,,,$il. iltu; Min. Perwrt :i. lnbroed Ddlr. : ..... ri:. r*!cr (cm)

Coral Aqopora sp. 11 60.0 5.5 0.89 8.0 5.0 3.00% I{ontpom sp. 2 10.0 5.0 0.00 5.0 5.0 0.50% PoliHsp. 2 13.0 6.5 1.s0 E,O 5.0 0.65%

Tobl: 4.15% Algae Coralline apae 11 1903.0 135,9 't15.17 397,0 6.0 95.1s% Dkfyuphaah sp. I 11.0 11.0 0.m r4.0 14.0 0.7.% Totrl: m00 Tobl: gtt.Eli?t

cotdllno rljrc 95.? Hettaodr rP. 0.2

Acroporr 2'9 MoBtlPosr I

But Sldc

corrlllm djrc 95.1

Wert Sldc

Fig. 33: Graphs of the com.position of the site 3 transects: east and. west side.

52 6. Natural History

The following is information on the natural history of selected species found in the Palolo Deep.

Fig. 34a: Ech,inometra mathaei: Burrowing Fig. 34 b: Echinothrix calamaris: The sea urchin on coral rock. nocturnal, banded sea urchin.

?'.: .-l

Echirrcnetra rnathaet Echinothrix calanaris The common name is Mathae's Sea Urchin or Commonly known as the banded or striped the Burrowing Sea Urchin. sea urchin because of its white and black banded primary spines. Adult size is about In adults, the upper spines vary from a unit'orm 25cm in diameter (spine length included). grey to red, black or a dark green, occasionally Juveniles often have pure white primary pink, with white tips. Juveniles can be brown, spines. It has a spotted bubble-like anal cone. green or purple but the spines usually have a white ring at the base. Adult size is about It has been found from the Indian Ocean to the 8.5cm in diameter including the spines. Red sea, and from Hawaii to Tahiti. This striking sea urchin is confrned to shadowy One of the most common sea urchins on the recesses within the reef during the day, and reef, they range from the Red Sea to the South- feeds on algae and detrital deposits at night. A west Pacific. Usually found under slabs of dead night dive in the Palolo Deep will reveal a coral in shallow water areas (0-5m) of the reef surprising number of urchins grazing the flat often in exposed situations such as the reef surface ofthe rocks. crest. They are found to reside in cup-like excavations or burrowed out grooves which Extreme caution should be taken when serve as protection. They make these encountering this species as its secondary excavations by abrading away limestone spines are very sharp, poisonous and cause material with their stout spines. Often the painful wounds. Along with the long spined sea excavations represent many generations of urchin (Echinothrix diadema) they may be effort. considered the most dangerous creatures on the reef, as the careless reefobserver will attest. The urchin is a herbivore, scraping algae from boulders and rubble close to its hollow at night, and catching drifting algae. At the Palolo Deep 'burrowing sea urchins' are common on the reef flat where they iahabit coral boulders. Numbers up to 51/m2 were recorded, though the average is between 5-10 urchinsimz. Their abundance is related to the presence of coral boulders, which provide shelter and on which their food source is found.

53 Fig. 34 c: Stegastes liuidus: The territorial Fig. 34 d: Aeropora nobilis with the algae Bluntsnout Gregory grving rise to ?Cladophoropsis sp. coraValgal relationship in next photo.

Photo: Fishes of the GBR: Randall, Alleu, and Steene.

Coral with algal growth The coral Acropora is commensal with the dense algal growth which is a result of the territorial behaviour of the damselfish, Stegastes liuidus Stegastes liuidus. Many damselfish on coral A member of the Pomacentridae @amsel reef are territorial and some defend the area for fishes), this frsh is known as the Bluntsnout months or years. Strongly territorial fish are Gregory. It can be identified by its overall often rasping or browsing herbivores whose brown colour, frequently with an ill-defined territoriality seems based primarily on the blackish area below the soft part of the dorsal defence of food resources. fins. The aggressive behaviour reduces total grazing Adult eize reaches 17cm, total length. A very pressure to such an extent that a denee mat of aggressive and pugnacious fi.sh, it occurs in algae develops within their territories. In some colonies that can emit staccato sounds. An instances, this could shade the coral thereby average density of 4 individuals/square metre reducing coral growth itself. In the Palolo Deep was observed in tbie Acropora nobilis beds in there is a balance. The coral growth occurs in the Southern Palolo Deep. an open manner above the algae. This allows enough light to penetrate allowing algal The species occurs from South Africa the and development and the colony surface to be Red Sea to the Society Islands and Line Islands. cleaned by the current of the constant rain of They usually reeide among staghorn corale in ggdiYnsalg. depths of 1-5m. They defend territories against all intruders" including nipping the legs of divers. As a result ofkeeping the herbivores at bay, algae tends to proliferate at the base ofthe coral colonies.

54 Fig. 34 e: Chaetodon, auriga: Threadfin Fig. 34 f: Halimeda sp.: This algae is a Butterfly fish. principal contributor in the sand cycle.

Halim.eda discoid,ea Chaetodon. auriga Belonging to the calcified green algae Also known as the threadfin butterflyfish. It (Chlorophyta), this genus can be easily can be identified by its converging black stripes identifred by its distinctive chains of [ttle hard (chevron-like) yellow on its sides. It has dorsal, segments made of green kidney-shaped disks. posteriorly, caudal and anal fins and a black It occurs profusely in both the Caribbean and part a spot on the softer of its dorsal fin with the Indo-Paci-fi.c region and is usually found on black band through the eye. Adults usually the leeward side of the reef. It is found around pennant have a long type {ilament extension of margin of the Palolo Deep from the rim to just the the dorsal fin, after the black spot, which is <10m. It is also found on the reef flat but is less used in territorial defence. Juveniles Iack the abundant due to unsuitable substrate and wave grow filamentous trailing. Adults up to 20cm in action. It flourishes on the outer reef slope, total length. particularly in the shelter of the cracks and Occurring from East Africa and the Red Sea to shallow crevices. Polynesia, can be the Indo-Pacific and they T}rre Halimedo plays an important part in the found singularly or in pairs on shallow sand cycle. The plant deposits limestone within protected reefs, and also on outer reefs to 30m. its disks and each chain may add an entire disk The Threadfin butterflyfish does not seem to be everyday. Although the disks are hard, their territorial during the day but at night expects connections are fragile so that water movement to sleep in its own spot, and beware any fish or disturbance from browsing or grazing that gets in the way. It feeds on algae, animals breaks them off quite easily. The disks polychaetes, prawns and coral polyps. In the soon become part of the sand and eventually Palolo Deep, it occurs in the proximity of the become minute grains. large stands of Acropora. Sand not only forms cays, beaches, and lagoon floors, but also fills in the spaces between the coral framework of the main reef structure.

55 Fig. 34 g'. Acanthaster planci: The crown-of- Fig. 34 h: Bohadschia graeffei: A grazing sea thorns starfish enjoys a meal. cucumber.

Acanthaster planci Bohadschia graeffei

(see Sectiotr, 7. 3: Crown-of-thonts Starfish.) Commonly known as Graeffe's sea cucumber. The species varies in colour but generally has Commonly known as the crown-of-thorns short white tipped papillae, covered in dark This can grow up to 60cm in starfrsh. starfish spots with larger mottled brown areas. Adults There are many diameter with 13-16 arms. grow up to 30cm in length. The holothurian's surface that can sharp spines on the dorsal body is relatively firm to touch but it must be be avoided or cause severe injuries and should treated carefully as it throws out its guts and colour range can be handled with care. The associated organs (which will be regenerated) quite variable from bright red to deep purple or as a vigorous defensive action. shades of green. In the Palolo DeeP, Acanthaster is particularly colourful with It occurs throughout the Indian Ocean and bright orange spines. West Pacific. It can be found on the sand or rubble areas of the coral reef, usually in tropical Pacific region, It occurs throughout the sheltered back reef or lagoon areas. At the has separate sexes usually among the corals. It Palolo Deep, its occurrence is limited to the summer months and spawns during the sandy, silty bottom of the lagoon. (Birkeland and Lucas 1990). A single female may spawn many millions of eggs. The larval It is a detrital feeder. Feeding occurs through stage may last up to 25 days and travel 500km the intake of sediments and small animals (Moran 1986). caught up in the soft, sticky 'catching pads' of its feeding tentacles. In some areas, this This spiny starfish feeds on living coral polyps species has been found to 'clean' detritus that stomach outwards, pressing it by turning its settled on fronds of living algae. against the coral and digesting it. Upon settlement, the cryptic juvenile feeds on coralline algae. Within 6-12 months, it begins to feed on coral. It reproduces after its third year and lives to a maximum of 9 years (Zann et al. 1987; 1990). Known predators include the puffer fish, the giant triton, and recently it has been suggested that xanthid crabs may help to protect their host corals by nipping the crown-of-thorns' tube feet thereby causing it to withdraw.

56 Fig 3l i: Pterois loiilons: The Firefish or Fig. 34 j: Palythoe heideri.'. At depth in the Turkevfish. Palolo Deep, the very common zooanthid.

Pterois LrolitztLS Palythoa heideri Commonly knorvn as firefish or turkeyfish. Looking similar to the butterfly scorpionfish, Knoq'n commonly as Heidi's Zoanthid. These this fish can be distinguished by its separated animals appear as stalked anemones and are pectoral lin rays. Colour ranges from pink to intermediate in structure between anemones black, the body being crossed with pairs of and hard coral. They appear as a dense colonial white bancls. Adult size is approximately 38cm veneer of brown polyps, each up to a centimetre in length. in diameter. The colour ranges fronr green to brown. It ranges from Western Australia and Malaysia to south-eastern Polynesia and north to Japan. They are found in rocky habitats below the It can be found about lagoon reefs and around turbulence ofthe surf zone to 10m in the warm coral heads. This firefish usually occurs in a temperate regions. They occur throughout the pair or group and inhabits the same cave, ledge tropics from the Indian Ocean to the Pacific. In or overhang for a number of years. It is mainly the Palolo Deep, they monopolise surfaces found in the southern portion of the Palolo below the proliferation of hard coral. They are Deep or along the southeast margin of the main abundant as carpet-like colonies on dead coral Deep. bommies in deeper areas. Here there are few benthic organisms. Their proliferation in such Although it is known to hunt during day or an environment indicates less dependence on occurs night, higher activity around twilight light and/or a tolerance ofpersistent siltation. and darkness, where it feeds on shrimps. A painful and poisonous sting, that usually They feed on plankton and suspended subsides in 6 to 12 hours, can result from the sediments. The polyps are not very sensitive to penetration ofthe dorsal spines. touch.

57 Fig. 34 k: Fungia fungites: A free living or Fig. 34 I: Anphiprion ocellaris'. Anemonefish unattached mushroom coral with commensal with Stoichoctis sp. tentacles expanded.

Amphiprion ocellaris Fungta fungites Common name is Clown Anernonefish. They One of the many solitary, free living corals are only associated with large sea anemone which are generally known as mushroom such as Radianthus ritteri and Stoichactis coral. A solitary polyp can be up to 28cm in giganteum. They can be easily identified by diameter. The corallite, which is usually their bright orange colour with 3 white stripes somewhat oval in outline with a convex upper running dorsal-ventrally. Adults grow up to surface, has a large number of vertical ?cm in total length. radiating septal ridges. The dent in its centre is the position of the polyp mouth. Unlike most Clown Anemonefish occur from the Tropical Ocean the Western are corals, the polyps do not divide to form colonies. Indian to Pacific. They found up to The mushroom coral also do not cement on inshore and offshore reefs, in themselves to the reef. At the time of larval 20m depth. Commensal with the anemone, settlenent, they attach and grow to a size of they are immune to the stinging cell of their host. three to five centimetres before breaking free to assume the adult mode. They are omnivorous feedere. At night, many Clown In the Palolo Deep many of these larval stages anemones close and enfold the Anemonefish. The summer, can be seen to be attached to the rubble, species breeds in protecting They clustered in depressions along the rim. The with both adults the eggs. pair, female occurrence of the adults indicates a preference normally occur as an adult the juveniles for this area. The specimen pictured is unusual being larger with several occupying in its bright colouration. the same anemone. The species ranges from Eastern Africa and the Rdd Sea to the Paci.fic region. This coral is very common on some reefs, and has been confused with F. danai which is more circular. Mushroom coral because of its unattached nature generally prefers the quiet waters of the pools, lagoons or the deeper water. It seems very tolerant to silty environments often seen in abundance. In the Palolo Deep it is not very common.

58 Fig. 34 m: Millepora platyphylla: "Fire coral or Fig. 34 n: TYidacna squaftLosqi Giant Fluted stinging coral" is actually a Clam. Elydrozoan.

Tridacna squa,tnosa Millepora platyphylla Commonly known as the giant lluted clam. It More commonly klown as the fire coral or the can be easily identified in field by its relatively stinging coral because of the burning large size and the characteristic sculpture ofits sensation it causes when touched. Millepora leaf-like flutes. The giant fluted clam has are colonial and hermatypic but are not a true beautiful mottled mantle lobes with variation in or stony coral belonging to the class Hydrozoa. colour including blue, green, brown and black. Adult clams can g"row up to 40cm in length. Colonies are arborescent, platelike, columnat or encrusting with a smooth surface covered by It occurs from low tide mark to 20m. This near-microscopic pores. Tentacles project from species ranges from the Red Sea to South Africa the colony surface. Visible as fine straight and from the Tropical Indian Ocean to the hairs. the tentacles connect with the mouth or Western Paeific. It occurs in the Palolo Deep as gastropore. Sexual reproduction ls well as on the reef flat. accomplished through a phase involving minute During low tide when the mantles are exposed, medusae. These sturdy colonies are capable of produces mucous cover that prevents withstanding strongest wave action. it a the dehydration. The giant clam has a unique Although several different growth forms occur, symbiotic relationship with single celled it is now believed that only one species exists. microscopic algae, zooxanthellae. The minute This genus ranges from east Africa to the algae in the mantle produce carbohydrates Tuamotus and Marquesas. This species is using the sun's energy and provide some of the normally found on projecting parts of the reef clam's food. This hermatypic clam is also a where tidal currents are strong. It is also filter feeder. abundant on upper reef slopes and in lagoons and may be a dominant component of some coral communities. In the Palolo Deep, its occurrence is limited to the upper rim area.

59 Fig. 34 o: Carpilius rnaculata'. Three Spot Fig. 34 p'. Octoprts c),oneus'. Blue octopus: Crab. generally noctural, this one was seen late afternoon.

Carpiliu,s trlo,culata Octopu,s cyarueus Also known as the three spot crab. Its body is oval in shape and more broad than long. It is The common name is Blue Octopus. Can be generally a light brown or fawn colour with identified by its large head with a pair of eyes three conspicuous deep red spots on the that bear a remarkable resemblance to carapace and two behind each eye. The pincers vertebrate eyes. It has 8 arms covered rvith two are large in proportion to the body and the rows of suckers. Usually weighing up to 4kg. fingers can be tipped with black. This species The octopus with its arms extended has a can often be found in crevices on the reef where diameter ranging up to 50cm with a length up it wedges itself in backwards with its strong, to lm. T,ike other members of the family it has spine-tipped walking legs to face the world with the ability to change its colour to blend with its its pincers. background. The colour ranges from dark brown to pink. The species can be seen at dusk They range throughout the western Pacific. In or dawn, and is generally nocturnal. the Palolo Deep, they are seen mainly at night when they come out to forage. They are Its distribution is throughout the tropical Indo- herbivorous and scavengers. west Pacific ocean. Octopus Iive in a liar comprising coral rubble, a crevice or overhang surrounded by dead shells of molluscs. They are carnivorous, feeding on other molluscs. The mouth has a beak resembling that of a parrot which is strong enough to fracture thick shells such as that of the robust Turbo sp. Their saliva is toxic inactivating their prey.

60 Fig. 34 q: Dald.orfia horrida: Arare and Fig. 34 t: Padina comnessoni: Fan-like brown unusual crab. algae.

Daldorfia horrida Pod,ina cornrnessorti This is one of the largest crabs on the coral reef. The fan-like brown algae @haeophyta) is A specimen collected in the Palolo Deep had a very common on the reef flat and inshore' It carapace diameter of 15cm with the distance usually grows in a cluster. The genus Padin'ais between the end of the pincers 75cm. of particular interest as it is the only brown algae that deposits limestone. It has been described as gtotesque with its mass of lumps and knobs but it is these features The base or thallus branches out into one plane that allow it to blend so well with its rubble with an iaroll of growth towards the lower side. environment. It has a stippled, cream to ruddy Concentric lines of hairs mark the reproductive red colouration. Slender antennae, eyes that organs ofthis algae. withdraw into sockets and a claw in which the moveable portion is angled, are other features. Its distribution is tropical, extending to Australia. In the Palolo Deep, it is rare. It inhabits burrows in the coral rubble. This nocturnal species is an omnivore, feeding as a scavenger as well as a grazer.

6l Fie 34 s: Diodon liturosus: Black-blotched Fig 34 t: Diod,on litrlrosrr,s: Black-blotched porcupine fish. porcupine fish.

Dindon liturosus Diod.on liturosus Commonly known as the black-blotched The porcupine frsh presents a formidable porcupine fish. It is characterised by its large sight to any predator. In stress, when head and spines. Its colouration is light brown camou{lage and evasive swimming fails, it has a to cream on the body with black blotches specialised defence system. Embedded in the extending from the eyes and forming a saddle skin are dozens of erectile spines, which in the across the back. This colouration disappears relaxed position lie backwards along the body. upon inllation. The pectoral fins are yellow. By pumping water (or air when at the surface) The fish can reach a total length of 50cm. into the abdomen the skin cavity is in-{lated and They are found in the Indo-Pacific, mainly on the spines stand rigidly up. These spines are the reef on sandy or muddy bottom. They are needle sharp and can cause painful, slow- carnivorous and feed on crustacean, molluscs healing wounds. The flesh of this fish may and echinoderms usually in the early hours of contain tetraodontoxin making it very the morning. Their hard dental plates and poisonous. It can also inflict a serious bite. strong jaws are well suited to crushing the tests and shells of their prey. They are poor swimmers and can only move il short bursts over small distances.

62 Fig. 34 u: Choetodon ephippitLrtt: Saddled Fig. 34 v: Corxthoichtys sch.ultzi: Schultz's butterfly fish. pipefish.

ChaetodorL ephipphrnt Corythoichthys sclt ultzi Commonly known as the saddled Commonly known as Schultz's pipefrsh, they butterflyfish. They have a yellow snout and are relatives of the sea-horses. They have a pectoral fins. A large black area exists very small gill opening, no spines in the fins, no posteriorly and on the adjacent dorsal and pelvic, pectoral or anal fins and only a single caudal fins. Adult.s have a long filament dorsal fin. Their maximum size is about 16cm extending back from the dorsal fin. They grow in length. up to 23cm in length. They can be found from the Red Sea to the This species is widespread in the Pacific Ocean Society Islands. Though most often observed as far north as Japan and Hawaii to the among the corals of the shallow reef, they occur Polynesian groups in the east and Papua New in depths up to 30m. Guinea and Cocos-Iieeling in the west. They Peculiar to this group, the female deposits her are common in depths of less than 10m. Their eggs on the ventral surface of the male. The diet is composed of coral polyps, algae, shrimp males incubate the eggs in a pouch or and polychaetes. In the Palolo Deep, they occur carrying them until hatching. throughouf the extensive beds of Acropora specialised area, nobilis. They usually occur in patrs.

63 Fig. 34 w: Parribacus antarcticus: Slipper or Fig. 34 x: Heniochus chrysostomus: Pennant spanish lobster. bannerfish feeding.

Par r ib acu s arltar ctieus H eniochus c hr y so s to mus Commonly known as the Antarctic Slipper or Known as the Pennant Bannerfrsh because of Spanish lobster. The slipper lobster has a their elongated dorsal spine with an expanded knobbly carapace with short, flat plate-like pennant-like margin. They are characterised antennae. Its camouflage colouration is a by two prominent alternating brown and white mottling of cream, black and brown. It is large, vertical stripes with a yellow snout. Adults can up to 25cm in length, and is slow moving. reach 25cm in length. They are usually found in Females can sometimes be determined by the solitary or small groups. presence egg mass held swimmerets of an by They oecur from Cocos-Keeling Islands to under the tail. Pitcairn Island. This species is one of the most They occur from the Indo-Pacific to the commonly seen fish on the reefs in depths of Caribbean inhabiting rock and coral reefs in the Iess than 20m. In the Palolo Deep, they occur tropics and sub-tropics. They secret themselves on the eastern reef flat and in the coral areas of in crevices amongst the coral and rubble by day the Deep. They feed mainly on coral polyps but but can be found foraging at night. also on zooplankton and other benthic invertebrates. They are carnivorous, feeding on molluscs and small invertebrates. At times, they prey on giant clams.

64 7. Species Inventory

A reference collection was compiled. Identifications of the predominant genus 7.1 Scleractinia (Hard Coral) Acropora were carried out by Dr. Wallace of the Museum of Tropical Queensland, Townsville' The Scleractinian or hard coral fauna of the Other identifications were made using the Palolo Deep is trnique by virtue of its varied Scleractiria of Eastent' Australia (Veron et al' environrnent and its recent history involving 19?6-1934). A taxonomic listing was compiled two major cyclones. Though a u'ide variety of as well as a reference list of the specimens with species is present, pledominance in the habitat such details as reference numbers, collection zones is the result of a history of recolonisation site, museum numbers and irl silru photographs. after majot' cyclones and environmental of the collection is the considerations suclt rrs tolerance to exposure, An interesting feature presence possible endemic species, or those unstable sub-strates and sedimentation. of whose occurrence is confined to a restricted The Reef Slope is in the Process of part of the Pacific. The identification of some recolonisation with a predominately Acropora specimens remains to be finalised as they are assemblage. Affected by wave action and unusual in comparison with species occurring current. the \\'estern Reef Flat is relatively more centrally in their range' The collection inrpoverished. dorninated by large Porites httea contains 11 families, 32 genera and 48 species. colonies. The protected Eastern Reef Flat is co- A maximum generic number for the Samoas, donrinated b1' trvo species of Pori'tes (P' rus and based on world-wide distributional data, is P. cylinclrico) aud Pocillopora danicontis. between 50 and 60 fferon 1993). Forty-ni-ne American The Palolo Deep is monopolised by areas of genera have been recorded from (Birkeland This compares sprawling Acropora nobilis as rvell as Iarge Samoa et al. 1992). genera from Fiji' colonies of Poriles rrus and Porites cylind'rica. with 58 and 148 species Chronic sedimentation affects this area. The Main Deep is characterised by relic coral bommies (>5m depth) in the silty environment with the principal coral growth along the south- east perimeter. The inshore area is depauperate due to tidal exposure and substrate limitations. Fig. 35: Hord Corals of Palolo DeeP.

e) Poriles nts b) Porites cylindrica

65 Fig. 35: Hard Corals of Palolo Deep (cont'd).

c) Acropora nobilis Galaxeo fasciatlaris

d) Porites lutea d Fungta fungites

e) Acropora nasuta h) Plerogyra sinuosa

66 7.1.1 Scleractinia: taxonomic listing OCULINIDAE

POCILLOPORIOAE Galaxea fascicularis

Pocillopora damicomis PECTINIIDAE Pocillopora eydouxi Echinophytlia aspera Pocillopora verucosa Orypon sp. Seriatopora hystrix MUSSIDAE ACROPORIDAE Lobophyllia sp. Acropora aspera Acropora caduus MERULINIOAE Acropora cercalis Merulina ampliata Auopora ct. grandis Acropora cf. aculeus FAVIIDAE Acropota sp. Acropora cuneata Diploa*rea helbpora Acrapora cytherea Favia matthaii Acropora cf. cytherea Favites cf. flexuosa Acropora danai Favileshalicr,n Aaopora cf. danai Leptasfrea pupurca Acroporc divaricata Leptoia phrygia Acropota sp. Leptoda phrygia Acropora formosa Acropora genmilera Acropora humilis Acropora hyacinthus Acropora sp. Acroporc sp. 7.1.2 Scleractinia: reference collection Acropora nasuta Acropora nobilis Ref. Splcies Collection [uirnin or Acropora cf. nobilis Ilo, , . sito pho{ognphed:,, Acropora paniculata sp|clm'|r Awporarobusta Acropora ct. samoensis 48) Acroporasp.(myilerf) Palolo Deep eA3q2 A$opora secale 12) Acmpon aspra Palolo Deep G43464 Pholo Acropora valida Acropora sp. 13) Acmporaanduus Palolo Deep G43461 Astreopora listeri Astreopora nyiophth alna 43) Acmwncr,nalis Mulinu'u ft G43478 Montipora cf. auslralbnsh 54) Acro@neredis Palolo Deep G43471 Photo Montipora turgescens 60) Acropon ceredis Palolo Deep G43468 PORITIDAE 44) Acropon cf. gnndis Palolo Deep G43495 Alveopora cf. allingi Gonbpora lohata 66al Acropnct, awleus Palolo Deep G4U82 Porites cylindica Acroponsp. Mulinu'u G43473 Porites lobata 28) Pt ('frne Poiles lutea acaleus 3) Porfies nlgresoens 3l) Acroporawneda Palolo Deep G43491 Porfles rus 33) AcrocE,rauneata Mulinu'u Pt G43492 SIDERASTREIDAE 53al Auoponqfierea Palolo Deep G13470 ND' hscinaraea columna Psamnocora contigua 571 AcrcNraqtherea Palolo Deep G43487 Psammocon haimeana 38) Aaoporact.cytherea Palolo Deep G43484 AGARICIIDAE 59) Acropondanai Aleipab G43489 Leploseris rnyceloseroides Acmporacf.danai Mulinu'u Pt G43490 leplosens purpurea 35) Pactrysens speciosa 40) Aaopond.danai Abipab G43493 Pavona decussata Pawnavarians 30) Actoo{,ndivaticata Palolo Deep

FUNGIIDAE

Fungia tungites Fungia repanda Fungia scruposa Herpdihalimax

67 Rcf. :Spcctss Girflltllofr fluteuin''or llo: 3!b , plrotuoirph.d rBeclmrm

65) Acroporasp. Palolo Deeo G43477 87) Gonioporalobata Palolo Deep ('echinata -like') 56) Herpolitha linax Palolo Deeo 63) Acropora formosa Palolo Deep G43497 11) Leptastreapurpurea Palolo Dee0 55) Acropora gemmibra Palolo Deep G43469 83) Leptoriaphrygia Palolo Deep 39) Acropora hunilis Aleipata G43462 25) Leptoriaphrygia Palolo Deep 34) Aaoporahyacinthus Mulinu'u Pt. G43481 Photo' 85) Leplosensrnycefoserordes Palolo Deep 32) Acroponhyacinthus Mulinu'u Pt. G43467 711 LeptosedEpurpurea Palolo Deep 15) Auopora sp. Palolo Deep G43475 84) Lobophylliasp. Palolo Deep flNen4ike) 19) Merulina anpliata Mulinu'u Pt. 58) Acropora nasuta Palolo Deep G43466 9) Montiporacf.australiensrs Palolo Deep 26) Acropora nasuta Mulinu'u Pt, G43457 46) Montiporaturgescens Palolo Deep 67) Auoporanasuta Palolo Deep G43458 17) Montiporc turyescens Palolo Deep 37) Actopota nobilis Palolo Deep G43465 20) Monliporaturgescens Mulinu'u Pt. 62) Acropora nobilis Palolo Deep G43460 Photo 72\ Oxyporasp. Palolo Deep 61) Actopota nobilis Palolo Deep G43463 2l Pachyserisspeciosa Palolo Deep 36) Auopora nobilis (thick) Aleipata G43498 74) Pavonadecussata Palolo Deep 64) Actoporanobilis(thick) Aleipah G43496 5) Pavona decussala Palolo Deep Photo 52) Acroporapaniculata Palolo Deep G434s9 ND 7) Pavonavaians Palolo Deep Photo 45) Acroporapaniculata Palolo Deep 79i PocilloporaCamimmis Palolo Deep Photo 3) Acroporapaniculata Palolo Deep 4) Pocilloporaeydouxi Palolo Deep 8) Acroporcrcbusta Palolo Deep G43494 231 Pocilloporaeydouxi Mulinu'u Pt. 69) Acroporacf.samoens,s Palolo Deep G43476 Photo 771 Pocilloporaeydouxi Palolo Deep Pholo 471 Acropora secale Aleipata G43486 75) Pocilloporavenucosa Palolo Deep 50) Acroporasecale Mulinu'u Pt. G43479 16) Poitescylindrica Palolo Deep 66b) Acropora sp. Palolo Deep Phoh 80) Poiteslobata Palolo Deep Photo 41) Actoporasp. Palolo Deep 21) Ponleslulea Mulinu'u Pt. Photo 49) Acroporavalida Mulinu'u Pt. G43485 76) Ponles nrgrescens Palolo Deep Photo 68) Acroporavalida Palolo Deep G43488 78) Pordesrus Palolo Deep Photo 51) Actopora sp. Aleipah G43480 ('Sanoan valida') 6) Poriles rus Palolo Deep

53b) Acropora sp. Aleipata G4U74 18) Poriles rus Palolo Deep (Sanoan valida') 70) Psamnocoracontigua Palolo Deep 10) Alveopora cf. allingi Palolo Deep 73) Psammocora ct. haimeana Palolo Deep 271 Astreoporalisteri Palolo Deep 82) SeriatoporahYstrix Palolo Deep Photo 241 Astreoporanyiophthalna Palolo Deep 81) SetiatoporafiYsfnix Palolo Deep 1) Coscrharaea columna Palolo Deep

14) Diploastreaheliopora Palolo Deep

42) Echinophy'liaaspera Mulinu'u Pt. Index:

88) Favia nafthaii Palolo Deep Photo: Refers to a file photo of the species.

in reference material. 90) Favlles cll /?exuosa Palolo Deep ND: No duplicate specimen existing lhe 89) Favites halicora Palolo Deep 29) Fungiatungites Palolo Deep 22) Fungiarepanda Mulinu'u Pt. The Palolo Deep material was collected dwing 20) Fungia scruposa Mulinu'u PL the period of 20 April to 31 May 1993. 86) Galaxeafascicularis Palolo Deep

6B 7.2 Molluscs 7.3 Fishes of Palolo Deep

Locallty Scisnlific ilensB Englirh Samoan

GASTROPODA AUNTIIURIDAE Suryeonfirh Pone, palagi

CONIDAE Acanlhurus Orange-socket surgeonfish auranticavus Conus|r-rtile Palolo Deep Conus vexillum Reef flat A.gultatus Whitespotted surgeonfi sh Maogo Conus cf. frigidus Reef flat A. lineatus Stipped suEeonfish Alogo Conustulipa Reef fiat

VASIDAE A. nigicans A. nigicauda Blackstreak Pone-lusina Vasa ceramicum Reef slope suqeonfish Bluelined VERMETIDAE A. nigrois surgeonfish Ponepone A. olivaceus Onngeband Ponepone Dendropoma maximum Porites colonies surgeonfish

LITTERINIDAE A. hosfegus Convic{ surgeonfish Manini Pone Lifterina coccinea lntertidal shore Ctenochaetus Brisile-toohed suryeonfish Ceithium nodulosum Reef flat slflalus Dlslorsio sp. Naso annulalus Whitemarg in unicomlish

MURICIDAE N.lituratus Smooh-head un'rcomflsh lli'ilia Morula granulata N. unicomis Bmwn unicomfish Umeisu

CYMATIIDAE Ze0rasoma scopas Brorm lang Pitopito Cabestana cl. spenglei Z,veliferum Sailfish surgeonfish lliu

CYPRAEIDAE APOGONIDAE Cardinalfshes Fo Cypraea tigis Apogon angustatus Stipped cardinalfish Fofusiloloa Cyqaea moneta Cheilodipterus Tiger cardinalfish Fotaoto, luganini STROMBIDAE macrodon luhuanus Reef flat Strombus ATHERINIDAE Siherfishes Sali Lambrs scorpius Palolo Deep Lambis (Lanbis) truncata Alheinonorus Robust hardyhead lacunosus TURBINIDAE AULOSTOMIDAE Trumpetfishes Taoto, taotito Turbo petholatus Reef flat Tutbo sp. Aulostomuschinensis Trumpeffsh Taoto+na

TROCHIDAE BALISTIDAE Tdggerfishes Sumu

Trochus pyranis Palolo Deep Mistapus undulatus Orarpelined tdggerfish Sumuaimaunu

MITRIDAE Rhinecanlhus aculeafus Whilebanded figgerfsh Sumu+o uo

Stngate//a sp. Palolo 0eep Suffianen fieanatus Bridle f'ggemsh Sumugase'ele'ele Mitra cucunerina Palolo Deep Atule-loto, ulirega Vexillum sp. Palolo Deep C4ES/O|,/,DAE Fusilierc Blue and gold fusiliers Atulstoto TEREBRIDAE Caesio cawlaureus CARANGIDAE Trcvally Terebra dinidiata Palolo Deep

NASSARIDAE Cannxnelanpygus Bluefintrevolly Malauli-apamoana

Nassanus albescens Shoreline CHAETODANDAE Buterffyfth Tililin

NERITIDAE C. awiga Threadfin butlerflyfsh Siu Nerita undata Shoreline C. ephlppium Saddled butterfy{sh Tffi tuauli Nerita dicata Shoreline C. nelannotus Blackback butterflyfnh Tiffi pa'ipa'i OPISTHOBRANCH - Phyllidae C. refbulatus Retiq.llated butterflyfish Tifitifi maona Phyilidia wellata Palolo Deep C. tnfascialus Redfin buteffih Tiftifi rnnif

C. ul,bfensis Double-saddlebutlerflyfsh Tffigutu'uli

C. uninrculatus Teardop butterffish Tifitifi pulesama

HeniocDus Pennantbannerfish Laulaubulaumea ch4aoslomus

69 Scientific l'lamer Englirh Samoan $lntHri'lllnre E@h Srmorn

CIRRHITIDAE Hawtflsh La'o, laugiva LUTJANIDAE Smppcn Iu

Pancfuhites arcalus Arc+ye hawkfish Lausiva Lullanus fu/vus Yellow.margined Tamala, taiva seaperc{l P. forstei Blactside hawkfish Lausiva t. gibbus Paddletail Malai DIODONTIDAE Porcupinefieher Taulo l. kas Diodon hystrix Porcupinefish Tauh, tautu Manlor niger Black and white seaperch Matala'oa D.lilurosus Black-blochedPorcrpinefish Tauta,tautu MICRODESMIDAE Dartfishes Mano'o-ui FISTUt/'RIIDAE Flutemouth Taotao Ptereleotrb evides Twotone lartlish Mano'oui Fidulaia Smootr flutemoutr Taotoama. taotao commeF;onii MONACANTHIDAE Leatherja :ket Pa'umalo

GOBIIDAE Gobies Mano'o Oxwonaanthus Beaked leatherjacket Pa'umaloguluumi ,ongiroslns Valenciennaesligata Bluebandgoby Mano o-sina Monacanthus Fan-bellied leatherjacket HAEMULIDAE Sweeilips ciinensr's Pleclorhynchus Many-$potted I'amai-moana sweeUips Cantherhines Yelloweyed leatherjacket Pa'umalo chaetodonoides dumerilii P. pidus Dotted sM,eetlips MUUIDAE Gostfishes HOLACENTRIDAE Squinelfshes and llalau Soldierfsh Mulloides Yellowsfipe goatfish l'asina (<8cm), Myrprfsts aduslus Shadorfin soldierfish Malau tuavela ilavolineatus vet6 goailish Neoniphon Blackfin squinelfish Malau loa Parupeneus Dash-and{ot Tusia operculais bad,einus goat{ish N. sammarc Spoffin squinelfish [,ilalau fui P. cyclaslonus Goldsaddle Moana

|"/'BRIDAE Wrasees Sugale P. indicus Indian goatfish Ta'uleia

Cheilinus chlorourus Flonal maori-wrasse Lalafi-mahpua'a NEMIPIERIDAE Bream

C. tilobatus Tripletailmaoriwrasse Lalafi-malamumu Scolopsis fiilinealus Threelined monocle bream Tivao

C. unifascrblus Redbreasted wrasse Lalaf, tagafa, OSTRACIIDAE Borfishes llloamoa malakea Ostrrcion cubicus Yellow boxfish Moamoa-lega Epibulus insidiator Slingiaw wrasse Si'umutu, lalafi-fua'au 0. mefeagns Spotted boxfish llloamoa-uli, Gomphosusvaius Bird wnasse Gutuslo, sugal+lupe moamoa-sama

Halichoercs Pink belly wrasse Sugale ulwela POMAC/.NTHIDAE Angelfishes Tu'u'u naeaitacF"us Centroppe Lemonpeel angelfish Tu'u'u-sama H. timaculatus Threespot wrasse Lape, sugalepagota llavrssr:mus

HemBynnus Blackeye fiicklip Sugale-laugutu Pomacanlhus Emperor angelfish Tu'u'u-moana melapterus impentor

Labichthys Tubelip wrasse Sugal+htuli Woplites REal angelfish Tu'u'u-moana unilineatus diacanthus

Labroides bicolor Bicolor cleaner wrasse Sugale i'usina POMACENTRIDAE Damselfishes Tu'u'u

L. dinidiatus Bluesteak chaner Sugale moobi Abudetduf Banded sergeant Mutu wftlsse seplemfascialus

LoD,ppsis Yellowback tubelip A. sexfasc,alus Scissor-hil sergeant Mamo xanthonota l. vargrbnsis Indopacific Mamo Macropharyngodon Black wrasse Amphiprion Red and black Tu'u'u-lumane aegrosensrb melanopus anemonefish Ihalassorna Sixbarwnsse Sugal+a'au Chronis Black+xil chromb Tu'u'u-segasega hartutickii afiiopedroalis LETHRINIDAE Emperors Mah'ele'ele, filoa C. viridis Eluegrcen chrcmb I'alanumoana Lethrinus harak Thumbprint emperor Filoa-vai Dascy'lus aruanus Humbug dascyllrc Mamo L. cf. nebulosus Spangled emperor Ulusa'0, mulogo Ponacentrus paw Blue damsel Tu'u'u-segasega, Monotaxis Big+ye bream Mu-malavaivai, teatea gtand@ulus malamu

70 Sdmflbftrn r E{glui $r,r., 7.4 Algae

SC/qR/DAE Parrotlishes Fuga, laea, galo CHLOROPHWA

Hipposnrus Pacific longnose Ulapokea Codialeg panotfish longiceps Hdindla opuntia Scarus dintdialus Yellowbaned Fugaelosama H. macroloba panotfish Chlorodesmusfasllgiala Caulena ncenosa 5. ovrlseps Egghead panotlish Fugaelosina hdiun sp.

S. sordrdus Bullethead panotfish Fugausi-tuavela Siphonocladalm

SEFR/4NI0IE Groupers and Gatala, 'ata'ata Vabnia venbiqsa Sea Baeses Oadoplroropsissp. adyosphaerla cavemwa cephatophotis argus Blue-spotted grouper Galala-uli, loi PHAE0PIIYTA E. hexagonatus Hexagon rockcod Gatalas'au Ec{ocarpahs E. mena Dwarf spotted Gahla-aloalo rockcod Hydroctattws sp. C'alrynenia sP' E. tauvina Greasy rockcod Gatala-tane Padinacf/nnersoni Anthiasdispar Redlin anthias Segasega-moana Dictyotales A. pasca/us Amethyst anthias Segasega-moana DWta dilrtoo,na Pseudanthias Stockyanthias fiypsetosoma Fucales stGANlDAE Rabbitfishes Lo sargassumaidaeloliun furbinatiaomata Siganus argenleus Forktiail nbbiilish Loloa RHODO,HWA S. fuscescens Dusky rabbitlish Crytonemiales S. sp,hus Scribbled rabbitfish Anefe, pa'ulu peyssone,ra sp. SyNGwAIH,DAE Pipefishes - Anphina sp. corythoichttus Banded pipensh 'iflSit*r*, *. tinlesfinai's Lilhophfilun sp. syiloooNr/DAE Lizardfishes Ta'oto Potdithonq' Sauida gracllis Slander lizardfish Ta'oh TERAPONTIDAE Terapon Perches Avaava

Tenpon jatbaa Ctescent grunter Avaava TETRAODONTIDAE Pufien Sue Arothron Blackspotted puffer Sue-uli, nigropunclafus su*lega

C. am0or'l'rensrs Ambon loby Suelape

C. bennefti Bennetfs toby Su+afa C. sdadi Solanders loby Sue-mimi C. valenlini Bhck-saddled toby Sue-mu ZANCLIDAE ttloodsh ldols Tanduscomutus Moorish idol Peape'a, lauhufau

7l 8. Environmental lmpacts

From a coral reef perspective, the Palolo Deep Marine Reserve is an area of contrast. History 8.1.1 Cyclones Ofa and Val and circumstance have developed a variety of Between I and February 1990, tropical habitats which have been differentially storm 5 cyclone Ofa, the strongest cyclone years, affected and are now changing as the result of in 150 devastated Western Si moa. year later in sedimgntall.r. The reef fauna has developed A January, another stro cyclone, also within this framework to give rise to a distinct lg Val, These greatly reef zonation. occurred. two cyclones have affected the present nature of the Palolo Deep The most well developed and luxuriant parts of Marine Reserve. Cyclone Ofa stripped the the reef, in terms of coral development, occur in seaward reef slope of its coral which, together the embedded lagoon and on the Eastern Reef with older rubble, was deposited shoreward. Flat. The reef flat to the east of this area has The subsequent cyclone, VaI, degraded the greater diversity but is constrained by depth storm banks, spreading them across the reef and the period of development. Due to cyclone flat and moving more material into the Deep. damage there are virtually no large coral Details of the storms' path and general effect on colonies on the reef slope. Recolonisation of this Western Samoa is found in Zann and Sua area is prolific, with high densities of small (1ee1). colonies developing. Rubble banks and slopes The effects of cyclones depend on their intensity reflect the recent history ofcyclone effect. and direction, and the nature of the reef Following is a discussion of some of the events community. The northern coast is generally and processes which have been inlluential in protected from the prevailing winds and is the conditioning of the biological and physical subject to greater damage as the coral growth is aspects of the reef as it is today. less robust. The storm banks were largely made of rubble formed prior to the cyclone and dislodged and canied on to the reef flat. Cyclones 8.1 Storms have different effects on different parts of the reef system. It is this feature which The most significant natural occurrence capable allows the coral reef to respond successfully to of causing immediate catastrophic coral catastrophic events. The Palolo Deep is a good mortality reefs cyclone on is a tropical or example of the ability of the system bo respond hurricane. The effects of extreme wave action in terms of recolonisation and regrowth. The as well as the reduction in satnity, increased presence ot' the Deep provides a protective turbidity, the change in the reef morphology environment in which the reef fauna and flora and current structure, result in the alteration of can survive the devastating nature of storm the reef communities and, at times, wholesale waves. It is from this reservoir of living destruction. material that the reef is subsequently restored. These community changes involve the reduction of monopolising species, the dispersal of coral 8.1.2 Devastation and Recovery fragments which may develop into new colonies, and new substrate becoming available The type of catastrophic coral mortality as for settlement. As part of the system within witnessed by cyclone Ofa is impressive as it which coral reefs have evolved, cyclones have represents nearly the complete removal of many bene{icial effects on their growth and benthos from the outer reef slope, the development. Literature concerning the effects deposition of extensive rubble banks on the of cyclones on coral reefs is found in the seaward reef flat and major alteration of the suggested readilg portion of the references. reef flat fauna. On the geological time scale, this is not an uncommon event and has occurred many times during the reefs existence. The tropical phenomenon of coral reefs and cyclones represents proof enough that they can survive and even thrive as the result of their interaction.

72 (1972) found that coral recovery Records of 'such devastation provide an Shinn as to the nature of the following hurricanes Donna and Betsy in understanding quick' Despite apparent destruction and subsequent redevelopment of Florida was relatively one year the damage coral reef assemblage. In the Caribbean, extensive damage, within the In four years repair was British Honduras reefs were stripped of living was barely noticeable. corals over a zone 8km wide during a hurricane total groove in 1961. The barrier reef spur and A more remarkable example of rapid recovery is system was stripped in an area of over 40km in in Indonesia. Here a situation occurred where width (Stoddart 1963). d,estruction was total as the result of a reef bare substrate affected by the center of being overlaid by lava' The At Rendezvous Reef. and coral to survive was the e*pe'rienc"d remarkable recruitment the storm, the only years (Tomascik, in Mon!ostrea orvrtu'Ioris' It was the general recovery in five nrassive with the Palolo Deep, it is an coral Diploastrea which was one of two fress). As nrassive assemblage which has species rvhich survived the reef slope Acropora dominated prolifically on a totally denuded reef' devastation at Palolo Deep. A simrlar effect settled occurrecl with the total disappearance of the Stephenson et al. (1958) suggested that Palolo more fragile branching Acroporas at both settlement of planulae larvae is inhibited by the Deep arrd Rendezvous Reef' Devastation by mobility of debris formed by the storms, and C-vcione Ofa. however, represented destruction their estimate of recovery following the Low oi a greater magnitude. Unlike the Honduras Isles, Great Barrier Reef storm is minimal' In situalion where corals survived with depth' the Palolo Deep situation the rubble on the reef there rvere almost no corals left alive' Rather slope was largely removed due to the steep than the reef surface and slope being mantled slope, and wave action' lBall, Shinn & Stockman 1967)' at wfth rubble (1971) discussed the theoretical Palolo Deep it was deposited in large ridges on Endean which affect the rate of recovery the reef crest. considerations of devastated reefs. These include the The clegree of exposure to the storm force is availability of pianulae larvae and their importJnt in reef sut'vival' At Jaluit Atoll survival rates, competition with other tMarshall Islands), massive quantities of rubble organisms and corals, substrate conditions' were an indication of the coral destruction on raies of growth and the succession of successful the seaward reef, while the more delicate colonisers. Certainly the frequency of ryclones branching and foliose corals survived unharmed affects the nature of a coral reefs development' in the lagoon (Blumenstock 1958, 1961)' It may account for major regional differences cyclonic and non'cyclonic areas A range of recovery rates has been proposeil by between 4)- *,ariou]s authors (Shinn L972 {1-4 yrs}; (Stoddart 1969e, L97 Stephenson et al. 1958 {10-20yrs}; Stoddart The prolific settlement on the reef slope at 196'9a l2O'25 yrs)) which reflects the variability Palolo Deep is indicative of a reef well on the in a reefs response to storrn damage' Grigg way to Fish occur in abundance' The and Maragos (1974), in examining lava flows of hard coral"*"otury. assemblage is represented by three known age, found the recovery time for the urntt"t periods of settlement since cyclone Ofa' types of rlef communities in exposed conditions The origin of this settlement is probably from tl'Ue Zo years, while 50 years may be required the Deelp itself. The benthos is dominated by for the more complex reefs found in sheltered encrusti'ng algal colonies' The substrate has areas, been consilidaled by coralline algae' With the ever increasing relief it is resurvey of the British Honduras coral developing an The 1965 that in another four to sxK years reefs showed little reef recovery in the area of estimated coral cover will occur, creating a greatest damage. The only corals occurring in substantial lny ,rumbers were those which survived the flourishing reef. storm itself, with large areas colonised by the The eastern reef flat is a good example of the algaes Pad'hua and Halitneda' Substantial re-establishment of hard coral with L0-l7o/o ,u"orr".y had occurred il areas where damage occurring overall, and an astonishing 83% had been slight. The survey af 1972 revealed one localised area' The size of the little or no recovery in areas of extreme damage "orru"ugJincolonies indicates that they are the result of ten years later but complete recovery in areas fragmentation. of moderate damage.

73 The substrate is coral rubble which has only Loya (L972) concluded that heav1, moderate stability. As a result, many of the sedimentation may be a very significant factor colonies are adapted to a semi-mobile existence, in determining Scleractinian community unattached, with their growth form conditioned structure. He attributed a reduction in species periodic by over turning. In the context of a abundance and percentage cover in areas on shallow reef flat situation, coral growth is Eilat Reefs, Red Sea, to this agent. He poinrs flourishing. By contrast the paucity of the out that the few massive species found in this western reef flat provides evidence that the zone have probably evolved cleaning Palolo Deep offers both shelter and, most mechanisms. Marshall and Orr (l93t), in probably, living fragments assisting in the re- studying the effects of sedimentation at Low establishment of the hard coral assemblage. Isles, Queensland, found the predominant bay genera to be those most tolerant to siltation. 8.2 Sedimentation Deposited sediment may limit the establishment of sessile organisms such as coral. Nlotoda (1940) explained the paucity of The origins of sedimentation in the Palolo Deep reef corals in certain areas in Palau being due are from two principal sources: run-off carrying to unfavourable substrates. This is certainly topsoil and landfill, and the degradation of reef the case in the bottom of the Deep where burial rubble following two major cyclones. The most is evident and there are expanses of sand important long-term source is soil carried by between relic coral bommies. The rubble slopes run-off, caused by the clearing the of also present an area devoid of coral growth. watershed, the general development of Apia Kissling (1965) found substrate to be the pr-ime and the port. The deposition of many tons of factor in regulating coral distributions in the reefal material resulting from cyclone Ofa, and shallow its subsequent water environment at Spanish degradation have given rise to Harbour. an abundance of silt. The finer component is responsible for the persistent turbidity present Lovell (1989), in cornparing the underlying in the Deep. It is speculated that the principal subfossil reefcorals with the recent asseurblage, sedimentary influence on the marine found a marked change in composition had environment is of terrestrial origin. An occurred. The hypothesis for this change increase in suspended sediments and general included, as a major factor, the chronic siltation occurs during times of flood and is presence of suspended sediment and silty persistent as general run-off. The nature of the substrates in the present environment. Palolo Deep as an enclosed lagoon in the reef Given this persistent silt load, the reef system flat facilitates the deposition of silt. The has become pre-conditioned to some extent. current structure in and about the Deep creates The presence of persistent silt is characteristic a sink situation, into which transported silt is of a coastal environment. Lovell (1g8g) continuously deposited. has shown that a reef subject to periodic It is well known that siltation directly affects catastrophic floodrng can survive. If the species diversity and the degree of living cover. environment returns to its pre-flood statc, Bull (1982) in comparing two bays on Magnetic species number and to a lesser extent, diversity Island, North Queensland, found a marked are maintained when re-colonised. It may be reduction in the number of species in the site presumed that if the period of heavy siltation is most affected by siltation. a short-term event, then any alteration to the reef may be temporary. If the suspended The Palolo Deep is characterised by limited sediment becomes chronic, and heavy silting a spebies with a co-dominance of three coral permanent featur.e, then the coral reef species in the Deep, where the siltation is assemblages will be dramatically reduced. This greatest. Roy and Smith (1971) found a EOo/o seems to be the case at the Palolo Deep lower coverage in turbid areas where of the Fanning turbidity is significant. Island lagoon. Porter (1922a,b) attributed diversity reduction in back shelf regions of Caribbean reefs to sedimentation.

74 8.3 Crown'of-Thorns Starfish or 8.4 Sewage Outfall alamea (Acanthaster Plancr-l A proposal is being investigated for the offshore disposal of effluent from the Apia sewerage The location of the outfall is 1300m (See also Section 6: Natural History' ry.t"-. from Mulinu'u Pt. at a depth of 40m' It is 2'5km Fig. 34 g) to the northwest of the Palolo Deep (GKW The period^ic i.ncrease in the population of this Consultants et al. 1984)- has made it an in{luential coral-eating starfish of this scheme, from the standpoint During periods Advisability feature of ihe reef community' of pollution of the Marine Reserve, warrants when there are large numbers of these starfish' Of concern is denuded of living furiher investigation. areas of the coral reef are the sewage effluent which change to contamination by coral. Infestations cause a dramatic of waste products including removal of comprises a host the coral reef community' With the cleaners, and all is initially fae"es, toxins from domestic the coral's tissue, the skeleton material for which the domestic associated fauna manner of colonised. by algae. The considered a common receptacle' respect to the fish sewer is changes, particularly with by faecal coliforms provides the largely herbivorous' Contamination r.*"-bltg" which becomes disease. Nutrient enrichment in reduced, making potential for Monopolising species are lead to a suppression of coral by other ih" R""urrre will substrate for recolonisation growth and a marked change in the benthic Species".'Lil"bl" diversity has been shown to species. communrtY. increase as the result of an infestation (Porter 19?2b). The occurrence of terrestrial run-off As the Deep acts as a sink accumulating has been linked to Acanr'thaster outbreaks material by virtue of its current structure' the (Birkeland r982)- nature and transportability of the effluent to be considered. Nutrient elevation' have been recorded five needs Documented outbreaks phosphorus, is known to affect the 1969-70' 1977' particularly times on Upolu (1932-33, 1950s, corals to calcifu and lead to a the infestations haven't of 80, 1980s), though progrlssive"tititv degradation of the coral community in the Palolo Deep during necessarily occurred 1988) and inshore areas (Rasmussan periods (Zann and BelI 199f)' With iXi""uy these 1988). A sewage plume is developed as the ,ro.*.1 population densities at six specimens effluent is carried to the (Endean 1982)' the t.rgety freshwater per kilometre of reef where the wind driven current crown-of-thorns in the Palolo Deep is n.rmber of "*f"."distributes it for some distance' higher than normal' Feasibility studies have indicated that current On this basis eight specimens would be the west (GWK twice that flow is consistently to expected for the entire Reserve; et al. 1984) as the result of in the Southern Palolo Consultants ,r.,tb"t were observed southerly winds which predominate in the rarely elsewhere' Little coral had Deep, though winter months. The sampling period failed to with feeding scars usually evident in been eaten take seasonality into account whereby the wind proximity bf the individuals. No threat to the and surface current directions reverse' Winds from these starfish is perceived in the the coral in the summer months blow from the northwest future. A large and colourful animal immediate and are likely to convey contaminated water which has been a focus of interest from and one into the Reserve. the scientific community over the past 30 years' at makes its presence of particular value to the When all costs are counted, dumping sewage option' Reserve from a tourist point of view' If the sea can prove the most expensive population numbers and consequent coral General categories of expense would have to pipe destruction were to become unacceptable then include the discharge facility and effluent collecting would be appropriate' This strategy construction, maintenance and the unaccounted pollution lends itself to the Deep where the habitat is but often unacceptible costs of chronic and confined and easilY collected' ieading to both potential health hazards degradation' In a region where samples for enviroimental Seven specimens were collected as are frequent, the budget for The size of observed tropical cyclones purposes. costs would be hard to assess' The "u""t""h varied between 18'5-25'5cm' The individuals ""p1u""-"rrtaliernative of a secondary or tertiary treatment a female, had a large ovary largest specimen, plant whereby the solid waste can be sold as readiness for a second annual ind-icatini fertilizer may prove more attractive' spawning.

75 An environmental impact that can be predicted What is apparent is that Palolo Deep is a silt is that there will be a change in the benthic trap into which a continuous current feeds from community as algal cover will be reduced as a seaward. If the plume encroaches on the Deep, result of reduction in light around the outfall. as it most probably will due to the prevailing Deposit feeders proliferate with the settlement sumrner winds, there will be deposition. Faecal of large quantities of particulate material. The contamination may render the Deep unhealthy extent of the effects of the plume is dependent for public use. The continuous input of nutrient on two factors: quantity of effluent and the material may change the nature of the benthos current structure @r. Hutchings pers. comm.). in the Reserve with a flow-on affecting the fauna and fish. Nutrient loading has a most pronounced effect on reef corals. Effluent concentrations are seen to greatly enhance the growth of algae at the expense of the corals. Coral morbidity and mortality may not be related to effluent toxicity, but are the result of competition with algae for space and 1ight (Marszalek 198fb). If degradation does occur then the value of the Reserve as a reservoir for recruitment, recreation and tourist attraction will be greatly diminished.

76 L Management Recommendations

9.1 Features and lmprovements 9.2 Recommendations

The successful development and operation of the.Palolo Deep Marine Reserve is the result of 1. Provide information about the Deep. resources Mr. Siaki (Jack) the efforts and of Information about the Palolo Deep should be Their efforts Laban Toomalatai and his family. provided to visitors: are responsible for the development of the visitor facility and the policing of the Reserve. a. At the entrance, a general summary of the There has been some monetary and material varied aspects of the Deep should be support from the Government. provided on admission. Acquainting visitors with the Reserve at this point would provide proximity the capital, has a Despite its to it a more informed basis for their subsequent atmosphere about quiet, secluded it. It is experience. uncommercial and has the feeling of personal discovery. The family run business is Presented in the form of a fact sheet this unassuming and friendly. Recommendations to information should include: Reserve must necessarily be improve the 3 Map, habitat descriptions, history of the integrated into the existing formula of success, Palolo Deep where appeal has been found in modest development. 3 Cautions against habitat disturbance, coral collection and breakage Improvements to the Palolo Deep could involve two areas: 3 Detail hazards: urchin, cone shells, coral cuts, currents, general water and sun safety M Improvement of the financial base of the Reserve operation. Without increasrng the b. Interpretive information should be provided entrance fee, revenue should be generated in the form of displays within the Reserve within the facility to provide a more which should include: promotional and adequate cash flow for 3 Dynamics of the coral reef system: maintenance requirements. trophic relationships; processes of reef There is a need for rnunicipal and national building; destructive processes (i.e. Bureau) to bodies the Visitors' 3 Threats: siltation; freshwater; dilution; value Pa,Iolo Deep appreciate the of the pollution; Crown-of-Thorns starfish; National Manne Reserve and to provide cyclones; other effects of man assrstance in its promotion and the development of interpretive material. 3 Description of cyclone Ofa and its effects M Development of public information within the facility and implernentating measures 2. Develop a commercialshoP required to conserve the Deep and enhance Visitors will be better catered for and visitors' expenence. revenue will be generated by the presence of a shop. Commercial items, such as educational and souvenir material, could be sold: for example, an expanded and more elaborate version of the information on the fact sheet in a booklet. It should include a summary of material similar to that presented as interpretive display information. Memorabilia of the Palolo Deep and coral reefs to include clothing, photographs and postcards showing the Deep, its habitats and inhabitants are examples. Q6ld dlinlrs and snacks should be available.

77 3. Promote the Deep as an ecotourism high or mid-tide. Though there is no venue evidence of a problem thus far, its shallow depth presents hazard A formal programme should be developed to a to the area over time if care is not taken. cater for groups. The Visitors' Bureau should be approached for their advice and participation. 6. lmprove the Platform With regard to the Deep, visitors' experience Renovate the platform to provide a partially would be enhanced by the redevelopment of covered facility which will allow visitors to the underwater trail. As detailed in the escape the sun ifthey desire. visitors' fact sheet, an appreciation of the main features of the Reserve as identified in 7. Police facilities the field, should include dominate species, ecological zones and unusual features. The task of policing the Reserve against such activities as illegal fi-shing has been a job 4. Pond development assumed by the proprietor Mr. Laban and the DEC park ranger Kelati. To make The possibility of developing a pond near the this task more effective, a punt and shore for viewing by people unable or binoculars should be provided. qnq,'illilg to trek to the Deep has been proposed by Mr. Laban. This is an 8. Research and educational opportunities interesting idea with good potential and should be pursued. To develop a progressive understanding of the Palolo Deep, the Reserve's advantages as Fundamental to robust coral development is an area of research opportunities be adequate circulation. This is characteristic should promoted. It has easy access and shore of the inshore area. Live coral could be brought in to create a habitat which would facilities. Good baseline information rs available with a variety environments. attract its own assemblage, providing an of Emphasis should be placed on the sheltered, example of the larger ecosystem. The secure nature Deep for inshore reef flat is impoverished but would of the Palolo conducting experiments. be enhanced by creating a deeper habitat with varied relief. However there would be The site should be used for field excursions initial difficulties such as establishing corals by secondary schools to provide education on amenable to transplantation and the the importance of the coastal environment stabilisation of substrate. The development and the conservation of coral reefs. of a stable and viable display would rely on Discussions should be conducted between initiative, industry, experimentation and the park management, DEC, and the continued maintenance. Education Department to facilitate this.

5. Regulate visitor access and usage 9. Monitor the Deep Unregulated visitor usage with time could A monitoring programme is essential to result in the degrading of aspects of the provide a more complete understanding of Reserve. An example is access to the Deep, the dynamics of the Deep. Baseline which at present is generally by wading information is fundamental to the across the inshore flat. The presence of assessment changes experienced in the boulders, coral colonies, and other reef Reserve which are the result of natural inhabitants makes approaching the Deep causes or otherwise (see Section 8). hazardous for visitors. Marine life, to a limited extent, is also damaged. The 10. Review Proposed Sewage Outfall approach to the Deep should be confined to a cleared and marked path. The access route All important to maintaining the natural would be designated on the fact sheet and by state of the Reserve is the prevention of a sign at shore's edge. pollution from the proposed sewage outfali. As the feasibility study failed to monitor the Damage from reef walking in the more seasonal luxuriant areas behind the Platform (Fig.3) variation in current direction, further investigation is could easily occur as the assemblage has is required. It suggested SOPAC approached good relief and is composed largely of young that be to conduct such a studv. (< 3.5 yrs) branching corals. It should be recommended as an ideal snorkelling area at

7B 10. References

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82 Dodge, R.E. and Vaisnys, J.R. f977. Coral Bazzaz, F.A. 1983. Characteristics of populations and growth patterns: responses populations in relation to disturbance in to sedimentation and turbidity associated natural and man-modified ecosystems. In with dredgir'g. J.Mar. Res.35(4): 715-730. Mooney, H.A. and Godron, M. (eds). Disturbance and Ecosysterns. Components of Dodge, R.E., Aller, R.C. and Thomson, J. 7974. Response. Springer-Verlag, Berlin, 258-27 5. Coral growth related to resuspension of bottom sediments. Nature 247: 574-577. Birkeland, C. 1982. Terrestial runoff as a cause of outbreaks of Acanthaster planci Dollar, S.J. and Grigg, R.W. 1981. Impact of a (Echinodermata: Asteroidea). Mar. Biol. 69: Kaolin clay spill on a coral reef in Hawaii. 175-185. Mar. BioI.65:269-276. Birkeland, C.and Lucas, J.S. 1990. Acanthaster Guicher, A. 1985. Natural and human change planci: Major tnanagetnen"t problem of coral of sedimentation in lagoons behind barrier reefs.Boca Raton, I'A: CRC Press. 257 pp. reefs in the humid tropics. Proc. 1th hfi. Corol Reef Cottgress 4:207-212. Borowitzka, M.A. 1981. Algae and grazing in coral reefecosystems. Endeauour 5: 99-106. Hubbard, D.K. 1986. Sedimentation as a control of reef development: St. Croix, US Brown B.E. 1987. Worldwide death of corals: Virgin Islands. Coral Reefs 5: ll7-125. natural cyclic events or man-made pollution? Mar. Pollution. BuIl. 18: 9-13. Hubbard, J.A.E.B. L973. Sediment shifting experiments: a guide to functional behaviour Brown, B.E. and Howard, L.S. 1985. Assessing in colonial corals. In: R.S. Boardman, A.H. the effects of "stress" on reef corals. Cheetham and W.A. Oliver, Jr. (eds). Aninal Aduances in Maritte Biology 22: L-63. colonies. Dowden, Hutchinson, and Ross, Cairns, J. 1980. The recovery process in Inc., Stroudsburg, Pa.: 3l-42.. damaged ecosystems. Arttt Arbor Science, Hubbard, J.A.E.B. and Pocock, Y.P. 1972. Ann Arbor Michigan. l67pp. Sediment rejection by recent Scleractinian Choi, D.R. 1984. Ecological succession of reef corals: Paleo-environmental a key to cavity-dwellers (coelobites) in coral rubble. reconstruction. Geologische Rundschau BulL Mar. Sci. 35: 72-79. 61:598-626. Connell, J.H. 1973. Population ecology of reef Lasker, H.R. 1980. Sediment rejection by reef building corals. In: Jones, O.A. and Endean, corals: the roles of behaviour and R. (eds) Bialoey and Geology of Coral Reefs morphology in Montastrea ca,uenlosa 2 pp. 47-75. Academic Press, London. (Linnaeus). J. Exp. Mar. BioI. and EcoI. 47: 77-87. Downing, N., El-Zahr, C.R. 1987. Gut evacuation rates in the rock-borrowing sea Levin, 1970. A litera,ture reuiew of the effects J. urchin, Echinometra mathaei. Bull. Mar. of sartd rentoual on. coral reef community. a Sci.4l:579-584. University of Hawaii, Dept. of Ocean Engineering. Univ. of Hawaii Sea Grant Endean, R. 1976. Destruction and recovery of College Program Technical Report. coral reef communities. In: O.A. Jones and R. Endean (eds). Biology and geology of coral Motoda, S. 1939. Submarine illumination. silt reefs. 3:2L5-254. content and quantity of food plankton on reef corals in the lwayama Bay, Palao. Hodges, L.T. 1977. Coral size and orientation Palao Trop. Biol. Stn. Stud^ 1(a): 637-649. relationships of the Key Largo Limestone of Florida. Proc. ?rd Int. Coral Reef Syrnp- Rogers, C.S. 1983. Sublethal and lethal effects Miami, Florida 2: 341 -352. of sediments applied to common Caribbean reef corals in the field. Man Poll. Bull. Jackson, J.B. 1977. Competition on marine 14(10): 378-382. hard substrata: the adaptive significance of solitary and colonial strategies. Amer. Nat. ILI:743-767. 10.1.3 Ecology: Kinsey, D.W. 1988. Responses of coral reef Bak, R.P.M. and Criens, S.R. 1981. Survival systems to elevated nutrients. 55-65pp. In: after fragmentation of colonies of. Madracis (Baldwin, C.L. ed) Nutrients in the Great mirabilis, Acropora palmata A. and Barrier Reef Regioru. Workshop Series 10. ceruicontis (Scleractinia) and the subsequent Great Barrier Reef Marine Park Authority, impact of a coral disease. Proc. 4th htt. Townsville, Australia. Coral Reef Symp. Mardla,2:22-227.

B3 Pearson, R.G. l-973. Recovery of coral reefs on 10.2.2 Molluscs: the Great Barrier Reef following attack by Wilson, 8.R., and K. Gillett. Australiart Shells Acandhaster. Micron esia: I:223 A.H. & A.W. Reed, Sydney, Wellington, Pearson, R.G. 1981. Recovery and London. recolonization coral reefs. Mar. Ecol. of Cernohorsky, W.O. 1967,1972. Marfite Shells Prog. Ser. 4: lO5-I22. of the Pacific Vol. IJL Pacific Publications, Potts, D.C., Done T.J., Isdale, P.J. and Fisk, Sydney. D.A. 1985- Dominance of a coral reef Habe, T. 1964. Shells of Westenr. Pacific in community by the genus Porites Colour Vol. II. Hoikusha Publishing Co. (Scleractinia). Mar. Ecol. Prog. Ser. 23: 79- Osaka. 84. Kera, T. 1962. Shells ol tlre Westent Pacific irt Randall, R.H. 1981. Morphologic diversity in Colou.r Vol. I. Hoikusha Publishins Co. the scleractinian genus Acropora. Proc. 4th Osaka. hr.t. Coral Reef Symp. Manila2: 157-164. populations on Sheppard, C.R.C. 1982. Coral 10.2.3 Fishes: reef slopes and their major controls. Mar. Ecol. Prog. Ser.7 pp. 83-115. Grant, E.M. f 978. Guide to Fis/r.es. Department of Harbours and Marine, Brisbane, Queensland. Australia. 10.2 Field Reference Texts Munro, I.S.R. 1967. The Fish.es of Neut Guitr,ea. Department of Agriculture, Stock and Fisheries, Port Moresby, PNG. 10.2.1Gorals Randall, J.8., G.R. Allen, R.C. Steene. 1990. Scleractinia of Eastern Australia Aust. Irtst. Fishes of the Great Barrier Reef and Coral Ma,r. Sci. Monogr. Ser. 1, 3-6 Vol I-V 1976- Sea Crawford House Press. Bathurst. 1985. Australia. ISBN 186333 0f 2 7 Veron, J.E.N. 1986. Corals of Australia attd tlte Indo-Pacific. Angus & Robertson, Sydney, 10.2.4 General: London. ISBN O2O7 151164. Coleman, N. 199f. Etr,cyclopaedia of Marine Veron, J.E.N. and Pichon, M. 19?6. Arintals. Angus & Robertson, Australia. Sclero.ctinia of eastern Australia, Part 1: ISBN 0 207 L64290. Fanilies Thamrlq,steriidae, Astrocoenidae, Pocilloporidoe. Aust. Ins. Mar. Sci., Monogr. Endean, R. 1982. Australia's Great Barrier Ser. L,86pp. Reef . University of Queensla'rd Press, Australia ISBN 0 70221678X. Veron, J.E.N., Pichon, M. and Wijsman-Best, M. L977. Scleractinia of eastent Australia, George, D. and J. 1979. Maritre Life: Arr, Part 2: Families Fauiidae, Trachyphyllidac. Illustrated En cy clopaedia of h wer te brates in. Aust.Ins. Mar. Sci., Monogr. Ser. 3, 232pp. the Seo. Rigby Ltd., Australia, 1979. ISBN o 7270 1107 3. Veron, J.E.N. and Pichon, i\d. 1980. Scleractinia of eastent Australia, Part 3: Fa,milies Agariciidae, Siderastreidae, Fungiidae, Oculhr,id,ae, Merulhtidae, Mussidae, Pectinida,e, Caryophyllidae, Dendrophylliidae. Aust. Inst. Mar. Sci., Monogr. Ser. 4, 422pp. 86pp. Veron, J.E.N. and Wallace, C.C. l-984. Scleractinia of eastent Australia, Part 4: Farnilies Acroporidae. Aust. Ins. Mar. Sci., Monogr. Ser. 6,485pp.

B4 Annex:

Use of the Lotus-l23 ver. 2.4 Spreadsheet Programme for Transect Analysis

Combine files if several transects are to be used for a particular area: /FC. Introduction Edit problems or unusual entries: F2. The translation of data from the field notes to Backup fi,le: IFSB. the spreadsheet and its subsequent analysis is discussed in a step by step process. Macros are Save the fiIe under a new name so as to found in the macro library (section 12.1.2). archive the original transect data and continue processing the original: /FS *.new. The field survey technique is discussed more fully in the methods section. Briefly, it involves Write a formula which subtracts one recording benthic attributes under a transect measurement or cell from the next (i.e.+A5- line. The format for each entry, such as the A4) and place it adjacent to the first entry coral Porites lutea, is to abbreviate it to Pl. under the Intercept column. Proceeding along the transect line, a Copy it down the length of the column: /C. measurement of the coral is taken from the line where its presence finishes, As an example, the Edit inappropriate numbers such as those format for P.lutea finishing at 235mm would be occurring as the result of combined 235 pl. This figure is not a measure of extent transects: Delete. but one of sequence whrich, through the Convert the formula figures to absolute programme, will be converted to a lenglh values: /RV. measurement. List them in the adjacent column: Value convergion. Inputing and Processing the Data Total the value conversion, which is the true transect length and edit the percentage The method used for input and processing of denominator in the data processing macro the data in the Lotus spreadsheet is as follows: \x GbAA1,/AAI,F2,save). It is the last line 3 Create a spreadsheet lile with titles to serve of that macro. as a basis for the input of all transect (i.e. Sort the taxa and the absolute values: lDS. TFRAME.wkI). Set the column width for the taxa (column B) at 22 and that for the Hide C column:/WCH. percent (column J) at 12: /WCS. Center the In the taxa column, type the name at the "N. Right align the titles ": \. base of each grouping of similar attributes. 3 The titles appear as in the following: Backup frle:1FSB. Transed no. and descriotlon To obtain the number of taxa (N), enter the

Transec{ | Taxa Intercept I ValuelNlMeanl Std.lMax.lMin.l% taxa value ranges at the row ofthe attribute Measurement Convenion Dev. name using the function @count bY 3 Retrieve the file: /FR TFRAME.WKf. employing the macros \c to enter the first cell in the range and V to $ive the last cell in 3 Rename the file to identifu the transect: /FS. the range. The formula format is 3 Add in the macto library: lA @count($cell..$cell). Use the $ to specify an absolute address. 3' Input the data: Use \e macro.

85 With the N formula highlighted, use the Insert the rows in the table to allow the macro \x to calculate the values in the other headings Coral, Algae and Substrate: \i or columns. /wir. Continue calculating the @count function for Center the N column figures: \r. each attribute name and employing the \x Calculate totals for the transect length, the macro. percent cover per category: \u. Save and Backup: IFSR,IFSB. Convert these values to absolutes and to Use the absolute value function /RV to percentage formats: /RV, /RFP. convert the formulae and to tranfer the data Save and Backup: /I. SR, /FSB. to a part of the spreadsheet where there is no row or column continuity. 3 Print the tabie: /PPR",G. Hide the columns concerned with the 3 Printing the Graphs. transect measurement, the intercept length, and the formula conversion: /WCH. Graphics Remove the rows above the taxa names: The pie charts in the report have been macro \w or /wdr. developed using the programme Slide. This is Reorganise the table into alphabetically a graphics programme which ports to Lotus'and listed categories of corals, algae and fulfils the same function as that of the graphics substrate. Create an empty rorfi/, move in the enhancement add-in WYSIWYG. appropriate entry and remove the empty rows to consolidate the table: \i,/m,\w.

Keystroke and @ function macros used in the processessing of spreadsheet information

\e {?}-{right}{?}-{down}{left/xg\z- Data entry from field notes.

\c @count($d Used after the value conversion to calculate N and to establish absolute addresses.

The following is the pincipal macro used for calculations used rn fhe lransecl analysis. It is a combination of macros for the various functions listed below. To start, the cursor needs to be at the N value or at the @count(*) column. Calculates:

\x /c-{right}-{rightfledi${homei{right}sum{del 5}-/rff+1-- Sum or total intercept /c-{right}-{righffidit}{home}{right}AVG{del 3}-/rff+1 -- Mean /c-{right}-{righti{EDIT{homeflright}std{del 3}-/rff-- Standard deviation /c-{right}-{right}{edit}{home}{right}max{del 3}-/rff+ 1 -- Maximum value from transect value /c-{right}-trighti{edit}{home}{right}min{delete 3}-/rff+1-{left 4} Minimum value of transect entry /c-{right 5}-{right Sfrv--{edit}/1 200O-/rfp--{left 7} Percentage cover - requires editing of the transect length before use. \o /c-{right}-{right}{edit}{home}{right}sum{del sl-hff+1-- Calculate sum

\u @sum($d Assist in sum calculation. \r ..$d To be used in association with \"

B6 Calculation of individualvalues: Galculates: \a h-{right 2}-{right 2fieditflhomefidght]AVG{del 3}-/rfi+1-- mean \m /c-{right}{righffidit}{homeflright]max{del3J-/rff+1- maximum

\n /c-{right}-{righffidiffiome}[right2]in{delete2prtr+1*{|eft5} minimum

\p /c-{ri gh t}-{righ$/rv--{edit/4000-/rF* percenlage

\d /wdr Deletes rows.

\i /wir lnserts rows. v ffs-{right}- Saves file.

V {edit}{home} -{down} Centers numbers in cell and goes to he one below.

\t {edit}{home}{del}-{down} Deletes cellentry and goes to he one below. V ieditflhomefldel]"- Right aligns label. Used to center'tolal'labels.

87 Acknowledgements

The Palolo Deep Marine Reserve Project was a cooperative effort between the Western Samoan Division of Environment and Conservation (DEC) and the South Pacific Regional Environment Programme (SPREP). Funding was provided by the Australian International Development Assistance Bureau (AIDAB). The project was co-managed by Dr. Andrew Smith, Coastal Management Officer (SPREP) and Dr. Tony Robinson, Resource Management OfEcer (DEC). The co- consultants were Edward R. Lovell, Biological Consultants, Fiji, and Foua Toloa. Participants in the training and survey programme were Lavaasa r.\ralua and Cedric Schuster (DEC) and Brendan Pasisi of the Department of Fisheries, Niu:. Many thanks to Dr. Carden Wallace who promptly identified the hard coral specimens and provided helpful comments on the assemblage, and to Dr. Leon Zannfor his insights into the project. Appreciation is expressed to Mr. Siaki (Jack) Laban Toomalatai who ably manages the Palolo Deep and to Kelati, the chief ranger, for valuable information and advice concerning the priject. Many thanks to Michelle Lam, Liz Wilson and Antoine D.R. Nyeurt, University of the South Pacific, Fiji, for their valuable assistance and contribution to the report preparation. Also thanks to Nancy Daschbach of the Fagatele Bay National Marine Sanctuary for information and material on the marine reserve, to Andrew Smith, Kelvin Passfield, Joy Heenan and Juana Lovell for editing the manuscript, and to R. B. Lovell for managing the colour photography. The Authors.

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