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REPORT DOCUMENTATION FORMWATER RESOURCES RESEARCH CENTER
University of Hawai i at Manoa2 FCST Category1Report
Number Technical Report No. 122~ Report Date
February 1979Ecological Observations off theMOkapu~ O'ahu~ Ocean Outfall:A Post-Installation Study
6Author (s)
Anthony R. RussoSteven J. DollarE. Alison Kay
SNo. of Pagesix + 49
b No . of 17NO. ofTables 13 Figures 17
9 Grant or Contract Agency
Department of Public WorksCity and County of Honolulu
lOGrant or Contract No.F-637-78
11 d D •Keywor s escPbpto~s: *Benthic fauna, *coral, *marine animals, aquaticlife, marine plants, Hawaii.
Identifie~s: *Micromollusks. biomonitorin~, marine outfall, Mokapu outfall.12Abstract (Purpose, method, results, conclusions)
An ecological study of the benthic and fish communities at Mokapu,O'ahu, was completed in the summer of 1978 approximately 1 yr subsequent tothe installation of a submarine outfall by the City and County of Honolulu.Data were obtained from five transects between Mokapu Point on the northeastern tip of O'ahu to Alala Point, approximately 6 034 m (3.75 miles)south, at depths of 6 to 24 m (20-80 ft). This study is subsequent to aninitial survey completed in 1975 prior to outfall construction.
Results show little or no effect from the operation of the outfall onthe benthic and fish communities. There are not significant differences inthe abundance, diversity, or composition of fishes from 1975 to 1978 exceptat the outfall site where new substrate was formed by construction. Betweenthe 1975 and 1978 studies there are some differences in coral species coverindices, which are attributed to patchy substrate distribution rather thanstress from the sewer outfall. Differences in species composition and distribution of micromolluscan assemblages may also be explained, at least inpart, by patchy distribution of the substrate.
2540 Dole Street, Holmes Hall 283 • Honolulu, Hawai i • U.S.A.
ECOLOGICAL OBSERVATIONS OFF THE MCKAPU, O'AHU, OCEAN OUTFALLA POST-INSTALLATION STUDY
by
Anthony R. RussoSteven S. Dollar
E. Alison Kay
Technical Report No. 122
February 1979
Interim Progress Reportfor
MOKAPU BENTHIC ECOSYSTEM AND FISH POPULATION STUDY
Principal Investigators: Anthony R. Russo and L. Stephen Lau
Project Period: 1 June 1978 to 31 May 1979
The research reported herein was funded by the Department of Public Works,City and County of Honolulu, and the Water Resources Research Center,University of Hawaii.
v
ABSTRACT
An ecological study of the benthic and fish communities at MBkapu,
o 'ahu, was completed in the summer of 1978 approximately 1 yr subsequent to
the installation of a submarine outfall by the City and County of Honolulu.
Data were obtained from five transects between MBkapu Point on the north
eastern tip of O'ahu to Alala Point, approximately 6 034 m (3.75 miles)
south, at depths of 6 to 24 m (20-80 ft). This study is subsequent to an
initial survey completed in 1975 prior to outfall construction.
Results show little or no effect from the operation of the outfall on
the benthic and fish corrmunities. There are not significant differences in
the abundance, diversity, or composition of fishes from 1975 to 1978 except
at the outfall site where new substrate was formed by construction. Between
the 1975 and 1978 studies there are some differences in coral species cover
indices, which are attributed to patchy substrate distribution rather than
stress from the sewer outfall. Differences in species composition and dis
tribution of micromoUuscan assemblages may also be explained, at least in
part, by patchy distribution of the substrate.
CONTENTS
ABSTRACT....
INTRODUCTION.Description of Study Sites
FISH AND ALGAE .Methods and Materials.Results. . . ...Discussion.
CORALS. . . .Methods.Results.Discussion
MICROMOLLUSKSMethods.Results.Discussion
CONCLUS ION.
REFERENCES.
APPENDICES.
ILLUSTRATIONSFIGURES
1. Map of Sampling Stations, 1975 and 1978, M6kapu Outfall,Kailua Bay, O'ahu. . . . . . . . . . .. . ....
2. Comparative Standing Stocks of Fish at All Stations,1975 and 1978, M6kapu Outfall, O'ahu .
3. Variation of Total Coral Cover and Species Cover Diversitywith Depth, 1975 and 1978, Mokapu Outfall, O'ahu ....
4. Percent Cover of Four Dominant Coral Species. Station A.Mokapu Outfall. O'ahu, 1975 and 1978 .
5. Percent Cover of Four Dominant Coral Species. Station B,Mokapu Outfall, O'ahu, 1975 and 1978 .
6. Percent Cover of Four Dominant Coral Species, Station C,M6kapu Outfall, O'ahu, 1975 and 1978 .
vii
v
1
3
3
3
4
6
11
11
11
19
27
27
28
35
36
37
39
2
9
13
14
14
15
viii
7. Percent Cover of Four Dominant Coral Species, Station 0,M6kapu Outfall, O'ahu, 1975 and 1978 16
8. Percent Cover of Four Dominant Coral Species, Station E,M6kapu Outfall, O'ahu, 1975 and 1978 . . . . . . . . . . 17
9. Total Coral Species Richness (Number of Species) forEach Transect, 1975 and 1978, Mokapu Outfall, O'ahu. . . . 18
10. Abundance and Species Diversity of Micromo11usks on TransectsB through E, 1975 and 1978, Mokapu Outfall, O'ahu. . . . . . .. 30
11. Summary of Micromo11uscan Species Distribution and Abundanceat Depths of 60 to 100 Feet, 1975 and 1978, Mokapu Outfall, O'ahu. 31
12. Micromo11uscan Species Composition at Depths of 20 to 100 Feet,MOkapu Outfa11, 0' ahu, 1975 and 1978 . . . . . . . . . . . • . 32
13. Summary of Micromo11uscan Species Composition and Abundance atDepths of 20 to 40 Feet, Mokapu Outfall, O'ahu, 1975 and 1978. 33
PLATES
1. Newly Formed Boulder Substrate Surrounding Mokapu PointSewage Outfall Pipe, O'ahu, 1978 . . . . . . . . . . . . . . . .. 7
2. Diffuser Vent in Mokapu Outfall Pipe, O'ahu, 1978. 223. Exposed Mokapu Outfall Diffuser Pipe, O'ahu, 1978. . . . . 224. Bryozoan, Triphyllozoon hirsutum Busk, Growing on
Outfall Boulders, Mokapu Outfall, O'ahu, 1978. . . . . . . . 24
TABLES
1. Comparative Statistics for Fish Populations from 1975 to1978, Mokapu Outfall, O'ahu. . . . . . . . . . . . . . . . . . 4
2. Algal Biomass, Mokapu Outfall, O'ahu, 1975 and 1978. . . . . . 63. Percent of Benthic Cover by Coral Species for Each
Transect, 1978 and 1975, Mokapu Outfall, O'ahu • . . . 124. Abundance, Species Diversity, and Species Composition of
Micromo11usks at Transect Stations, Mokapu Outfall, O'ahu, 1978. . 295. Mean Percent Composition and Ranks of Six Dominant Species
of Micromo11usks at Depths of 20-40 Feet, 1975 and 1978,Mokapu Outfall, O'ahu. . . . . . . . . • . . . . . . . . . . 34
6. Mean Percent Composition and Ranks of Five Dominant Speciesof Micromo11usks at Depths of 60-100 Feet, 1975 and 1978,Mokapu Outfall, O'ahu.......•............... 34
INTRODUCTION
During the month of December 1975 a study of the benthic marine commu
nity was made at Mokapu, O'ahu (Russo et al. 1977). This study was made
prior to the completion of an outfall with a discharge of secondary sewage
effluent; the outfall, constructed for the City and County of Honolulu,
began operation in December of 1977. During February 1978 the Kane'ohe Bay
outfall was diverted to Mokapu, and in May 1978 the Kailua Bay outfall was
connected to the Mokapu line. The total volume rate of sewage discharge is
approximately 0.4 m3/s (9 mgd). The total outfall length is 1 524 m
(5 000 ft) including the diffuser which has a length of 293 m (960 ft) and
lies in 25.9 to 30.5 m (85-100 ft) of water. The outfall pipe was
laid on the bottom, and its entire length was covered with large boulders.
In compliance with the request of the City and County of Honolulu, this
study is a follOW-Up to the initial survey made in 1975. The main purpose
of this study is to report on changes, if any, in the benthic community
made by the construction and discharge of the outfall. The following have
been monitored and changes reported:
1. Fish population abundance and diversity
2. Biomass of attached algae
3. Diversity and cover of hermatypic corals
4. Distribution, abundance, and diversity of micromollusks.
Ecological surveys of the marine environment prior to installation of
sewer outfalls are now a part of most local water quality studies (Turner
et al. 1964). Follow-up surveys are important in order to assess any major
changes to the coastal marine environment caused by the construction of
outfalls (Turner et al. 1965). These surveys also provide outfall engineers
with feedback on the design of outfall systems and their feasibility for use
under different environmental conditions.
A study of the Kailua Bay area was conducted in 1973 (Kay et al. 1973)
to examine the benthic biota in a baseline study from M6kapu Point to
Mokulua Island off Lanikai. The 1975 study (Russo et al. 1977) gathered
data on the benthic biota at five stations from Mokapu Point to Alala Point
north of Lanikai (Fig. 1). These same stations were reexamined in June
1978; comparative findings are presented in this report.
2
oIo
KILONETER
IOQ
8 - SAMPLING STATION
SUBSTRATE DESCRIPTION
1 - SAND, SILT
2 - SOLID BASALT PAVEMENT
3 - SOLID LIMESTONE VENEER
4 - BASALTIC BOULDERS
WATER DEPTHCONTOUR
(METERS)
I~
FIGURE 1. SAMPLING STATtONS. 1975 AND 1978, MOKAPU OUTFALL, KAILUA BAY,O'AHU
3
Description of Study Sites
Station B coincides with the Mokapu outfall; three stations, C, 0, E,
are located 1 609, 4 023, and 5 632 m (1, 2.5, and 3.5 miles) south of the
outfall; and station A is located 1 609 m (1 mile) north of the outfall off
Mokapu Point (Fig. 1). Station C is located north of Kailua Bay, station 0
in Kailua Bay, and station E off Alala Point. At each station, depths of
6, 12, and 18 m (20, 40, and 60 ft) were studied as in the 1975 study. In
1975 the 30-m (IOO-ft) depth was examined, and results showed that all
stations except A had a sand substrate at the 30-m depth. In this study
the 24-m (80-ft) depth, which seemed to be the limit to hard, coral-bearing
substrate, was surveyed at stations A, B, and o. The results from this
depth will be used as an additional area for comparison in the follow-up
study proposed for the summer of 1979.
FISH AND ALGAEMethods and Materials
Fish surveys were made at stations A through E at depths of 6, 12, and
18 m. A 30-m (lOO-ft) line marked off at every meter was laid on the bot
tom. A diver swimming along the line counted and identified species of
fish 3 m (10 ft) on either side of the transect line so that a 186 m7
(2000 ft 2) area was swept. At station B, a fish count was made at each
depth on the outfall and 10 m (33 ft) to either side of the outfall. Fish
diversity was calculated at each depth using the Shannon-Wiener information
index, H' (Russo et al. 1977). To compare fish communities before and
after outfall operation, Sorenson's Similarity Index was used (Mueller
Oombois et al. 1974). The index in effect measures the redundancy of fish
species from one spatial community to another or, as in this case, from one
point in time to another. The equation for the index is SI = 2CjA + B where
C = number of species common to both transects
A = number of species in 1975 transects
B = number of species in present (1978) transects,
Mueller-Oombois and Ellenberg (1974) suggest that an index of 50% (.50)
represents a threshold value; that is, if the index value exceeds 50%, the
similarity is great enough to indicate that the species are a part of the
same association or community. In general, common species may be emphasized
4
in defining communities since they are most involved in the bioenergetics
of the community.
Algal biomass at each station was determined by cropping all the algae
in a 1/4-m2 quadrat placed randomly at five points along the transect line
at each depth. Each quadrat sample of algae was sorted into genera, weighed
wet on a Mettler balance, dried at 38°C (100°F) for 24 hours, and then
weighed again. Presence or absence of major invertebrates other than corals
was noted on each transect.
Results
Calculation of Sorenson's Similiarity Index for all stations and depths
shows very little change in characteristic fish communities between Decem
ber 1975 and June 1978 except at station B, the outfall site (Table 1).
TABLE 1. COMPARATIVE STATISTICS FOR FISH POPULATIONSFROM 1975 TO 1978, MOKAPU OUTFALL, O'AHU
Station Similarity Index % Change % Change in % Change inFish Species in HI Total Fi sht No. Species
A20 .61 -12 -8 -20
40 .60 -19 -3 -17
60 .67 -1.7 -22 +4
B20;~ .25 +123 +82 +77
40 no fish counted +100 +100on first transect
60 +100 +100
C20 .80 +30 +5 +100
40 .55 +25 +11 +16
60 .55 +11 -28 +8
020 .55 +0.5 -14 +20
40 .61 -12.5 -28 -5
60 .51 +1.1 +25 +20
E20 .45 -6.5 -7 -16
40 .57 -5.0 -16 -10
60 .58 -10.8 -20 -17
*Transect excluding outfall fish aggregations.tLarge schools excluded.
5
There was little similiarity between the 1975 and 1978 transects at station
B. Results also show little change in total fish, diversity index, and
species richness at each station and depth except for station B. At sta
tion B20 there was an 82% increase in total fish counted, a 77% increase in
numbers of species, and a 123% increase in diversity.
Fish of the families Acanthuridae and Chaetodontidae were abundant,
along with the family Pomacentridae, as they were in the 1975 survey.
Thalassomaduperreyi (Labridae) was abundant at most stations as it was in
1975. A list of species, diversity indices, and equitability components
(H'/H'max) for all stations and depths is shown in Appendix A.
At station B results showed a large increase in all major fish fami
lies, especially over the large rocks covering the outfall. Transects laid
at a distance 10 m from the outfall line showed less fish than directly
over the outfall but still showed more total fish and greater species rich
ness than in the 1975 survey. At station B depths where no fish were seen
in December 1975, fish now abound. Large aggregations (>30) of Kyphosus
cinerasoens (nenue), Chaetodon miliaris (lemon butterfly), Mulloidichthys
aurif7amma (weke), Parupeneus porphyre~s (kumu) and Abudefduf abominalis
(mamo) were seen at depths of 24 to 30 m (80-100 ft) along the diffuser.
Results of the algae collection showed that most of the biomass of
algae recorded was found at stations B, C, and D. In the 1975 study most of
the algal biomass occurred at these same stations (Russo et ale 1977; Table
1). There was no large change in biomass of algae from 1975 to 1978.
Halimeda remained the most abundant algae seen. Dictyopteris, Amansia,
Asparagopsis, and Desmia were also abundant. At station B40 beds of
Dictyopteris australis were found covering large areas of the bottom. Al
though many other algal species were seen, none was abundant enough to mea
sure and record for biomass. Results are shown in Table 2.
As in the 1975 survey, the most frequently encountered macroinverte
brates, other than corals, were sea urchins of the genera Echinothrix,
Echinometra, and Tripneustes. At station C the burrowing sea urchin Echi
nostrephus aoiculatus was observed. This species was not reported in the
1975 survey. The distribution of sea urchins was spotty, and their abun
dance very low.
6
TABLE 2. ALGAL BIOMASS, MOKAPU OUTFALL, O'AHU, 1975 AND 1978
1975 1978
Station Depth Wet Wt. Dry Wt. Wet Wt. Dry Wt. Genus
(m) ------------------ (9) ------------------
A20 6 None None None None None
40 12 None None None None None
60 18 None None None None None
B20 6 None None None None None
40 12 None None None None None
60 18 None None None None None
C20 6 None None None None None
40 12 None None None None None
60 18 125 44 91 28 Dictyopteris
162 75 100 45 Halimeda
020 6 None None None None None
40 12 79 15 65 15 Halimeda
60 18 None None None None None
E20 6 229 89 None None Dictyopteris
40 12 None None None None None
60 18 None None None None None
Discussion
The fish communities at each of the stations and depths remained
essentially the same from 1975 to 1978 except for station B, the outfall
site. Similarity indices show that the communities did not change their
composition by species (Table 1). Communities would only be expected to
change their species composition under extreme stress, e.g., depletion of
food supply, destruction of substrate habitat, change of physico-chemical
environment. Changes of this type were not observed in this study. All
values of Sorenson's index, except for station B, were above 50%. Expe
rience has shown that there is rarely a similarity index based on species
presence which exceeds 60 to 70% (Mueller-Dombois 1974). Even the species
lists of two similar, neighboring communities rarely have more than 2/3 of
their species in common. At station B the similarity index indicates a
significant change in fish abundance and diversity, probably due to the new
substrate afforded by construction of the outfall. The new substrate pro
vided by the large rocks covering the outfall attracted large numbers of
fish, providing not only substrate for the growth of attached algae but
also habitat space (Plate 1). The rudderfish (Kyphosus cinerascens) was
abundant over the outfall substrate. This fish, a strict herbivore, was
seen grazing on attached algae. Acanthurids, which are also herbivorous in
general, were abundant along the outfall. Large numbers of weke (MUlloi
dichthys aurif1amma) were observed feeding on the sediment alongside the
outfall. Weke normally feed on benthic infauna and may have been feeding
on organic detritus in the sediment. Whatever their food source, the out
fall site attracted them in large numbers. These observations are consis
tent with many studies done on the increased nucleus of marine organisms,
especially fish, associated with the construction of artificial reefs.
PLATE 1. NEWLY FORMED BOULDER SUBSTRATE SURROUNDING MOKAPUPOINT SEWAGE OUTFALL PIPE, O'AHU, 1978. BOULDERSSIGNIFICANTLY INCREASED THE HABITAT COMPLEXITYLEADING TO MORE NUMEROUS SETTLING SITES FOR BENTHIC INVERTEBRATES AND TO INCREASED SHELTER ANDGRAZING SURFACES FOR MOTILE INVERTEBRATES ANDFISHES.
7
8
The new substrate also attracted large numbers of water column forag
ers. Many fish are adapted to foraging for plankton and organic particles
in a water column above a reef substrate (Davis et al. 1973). There are
several species of coral reef fish which forage in a water column. For
example, Strasberg (1966) cited the pomacentrid Abudefduf saxatiZis as being
a "particulate plankton picker." Large aggregations of the pomacentrid
Abudefduf abominaZis were seen in the water column above the outfall at all
depths. This species was described by Hobson (1974) as being mainly a
diurnal planktonivore which preys primarily on copepods in the water column.
Large aggregations of the butterfly fish Chaetodon miZiaris were observed
over the outfall at most depths. C. miZiaris is also a p1anktonivorous
water column feeder (Hobson 1974).
A fish count was made on the diffuser in 27 m (90 ft) of water. Large
aggregations of Lutjanus kasmipa (taape), Acanthupus oZivaceus, and MUZZoi
dichthys fZavoZineatus (weke) were seen (Appendix A). The bryozoan Tri
phytZozoon hipsutum, sometimes called lace coral, was very abundant along
the diffuser (4-8 colonies/m2 ). The abundance of bryozoans may be due to
the increase in particulates and microorganisms in the water because this
organism is a strict filter feeder.
In the December 1975 study, fish standing stock abundance was reported
to be high at stations A, D, and E and lower at stations C and B. The same
trend has held in this study (Fig. 2). When large, passing schools of fish
are excluded from the total counts, the percent change from 1975 to 1978
ranged from 3 to 28% at all stations except for station B which showed an
82% increase in fish seen. These counts were made off the outfall. A much
larger increase of fish was realized if those directly over the outfall
were included. Along with this increase in total fish, a significant in
crease in diversity index and numbers of fish over data reported in 1975
was seen at station B (Table 1). At all other depths except C20 the per
cent change in species richness was small. The anomalous result at C20 is
explained by the fact that an increase from 2 to 4 species was seen, that
is, there was a small absolute increase but a large percentage increase.
Grigg (1972) reported changes in both species and abundance of near
shore fishes off sugar mill outfalls on Hawai'i. He reported the increase
of some species which could utilize the waste as food and a decrease in
others which were affected by the pollution load. Turner et al. (1965)
-(IJ,
Eoco'-o 400c-
_ 1978 June
F:::::::::::~ 1975 December
9
~
:r:CJ)
l.L.l.L.oen~
uo.....en<!)
zoz~en A B C
STATIONo E
*Excluding large schools and those fish counted over theoutfall.
FIGURE 2. COMPARATIVE STANDING STOCKS OF FISH AT ALLSTATIONS, 1975 AND 1978, MOKAPU OUTFALL, O'AHU
reported large aggregations of two species of fish feeding off effluent in
the vicinity of the Orange County, California Sanitation District Ocean
Outfall. They measured kelp bass and sand bass close to the maximum size
(0.4 m or 17 in.) for these species. However, Turner et al. did not record
whether or not there was a change in the diversity at the outfall. The
central idea is that a community has structure which is somehow defined, at
least in part, by its diversity. In polluted environments there normally
is less diversity. Pollution intolerant species give way to more tolerant
forms (Stein and Denison 1963). The tolerant forms tend to become more
abundant because there is less competition for existing resources. Patrick
and Strawbridge (1963), describing the changes in a phytoplankton community
structure, carefully point out that the use of single indicator species may
not be a valid measure of pollution load. In reporting large aggregations
of organisms, especially fish, the investigator may not correctly interpret
an increase in abundance and biomass as a positive result since pollution
intolerant species may have been replaced by tolerant ones. There is also
the remote possibility that a large change in the community might occur
with no change in diversity. This may occur where niches occupied prior to
an environmental change are replaced with new species tolerant to the new
10
environmental milieu. In the case of the Mokapu outfall station, there was
a definite increase in abundance of a few species of fish; but, more impor
tant, there was also a concomitant increase in both diversity components-
the species richness and equitability. There was also an increase in fami
lies represented. This is an indication of little or no pollution stress
on the fish community; in fact, as mentioned above, the outfall construction
probably enhanced all aspects of the fish community by increasing habitat
(shelter) space and settling sites for algae and other sessile organisms
upon which fish feed.
The biomass of algae did not change significantly. The same dominant
species were seen in this study as were observed in 1975. During dives at
26 to 27 m (85 to 90 ft) over the diffuser, the effluent discharge was no
ticeable directly over the diffuser but disappeared 15 to 20 m from the out
fall. The outfall plume discolored the water slightly as it rose and formed
a 2 to 3-m turbidity layer over the outfall. The mixing is vigorous in
these waters, and it is speculated that little nutrient enrichment takes
place except at the diffuser. This may explain why there was no apparent
increase in algal biomass at the stations. Extensive Dictyopteris beds were
observed, but these beds were also observed by Kay et al. (1973) and by
Russo et al. (1977) before the outfall became operational. At station E20
no algae were recorded in this study although Dictyopteris was recorded in
1975. This is probably due to our inaccuracy in finding the area, the
spotty distribution of algal beds, and the turbidity at this station.
CORALSMethods
The coral community and associated macroinvertebrates were surveyed
using a continuous photographic transect technique, a method which appears
to be more efficient with respect to time spent under water and area sur
veyed than either a chain transect or conventional quadrat method. In this
study, each transect was 30 m long at depths of 6, 12, and 18 m at each of
the five stations.
The photographic transect technique involved mounting a Nikonos II
camera on a supporting frame 1.25 m above a 1 x 0.7-m quadrat. The support
ing frame and quadrat were constructed of O.Ol-m (1/2-in.) aluminum tubing,
11
and the camera was mounted on a Plexiglas plate attached to the four sup
porting arms of the frame. At each transect location, a 30-m polypropylene
line was laid across the bottom at a random orientation. The camera quadrat
frame was placed on the bottom so that the l-m side of the quadrat was
parallel to the first meter of transect line. A color slide was then taken
of the 1 x O.7-m area within the quadrat. This process was repeated at 12
random marks along the 30-m transect line. Transect locations and depths
were written in large letters on an underwater slate and photographed with
the remaining film for later identification. Because small colonies may
not show up in the transect photographs, a diver with a species checklist
on a clipboard recorded the presence of all coral species in each frame of
each transect.
The developed slides were projected onto a grid with the same dimen
sions as the quadrat, and the abundance of corals and noncoral substrate
was estimated by counting the number of square centimeters occupied by each
coral colony. From these counts, estimates of percent cover and colony
size were determined.
Results
Tabulation of the coral coverage at each transect in terms of percent
coral and noncoral bottom cover is listed in Table 3 for each species and
substrate type for both the 1975 and 1978 surveys. Also in Table 3 are the
differences in total coral coverage between the two surveys. The total
coral cover and the species cover diversity (Shannon-Wiener index H' =s
~L Pi/In/Pi, where Pi = the proportion of the ith species) at each transect'l-=l
for both the 1975 and 1978 surveys are shown in Figure 3. For the thirteen
species encountered. five can be considered dominant species as they
occurred on most of the coral-containing transects. These five corals-
Poai Uopora meand:l'ina, Porites lobata, P. corrtpressa. Montipora verI'Ucosa,
and M. patula -- are the dominant corals on most Hawaiian reefs and have been
shown to comprise very high proportions of the coral cover off the Kona
Coast of Hawai'i (98%) (Dollar 1975a. b) and in Kane'ohe Bay, O'ahu (95%)
(Maragos 1972). Figures 4 to 8 show plots of abundance of these corals
at each transect for the 1975 and 1978 surveys. The various species of
12
TABLE 3. PERCENT OF BENTHIC COVER BY CORAL SPECIES FOR EACHTRANSECT, 1978 AND 1975, MOKAPU OUTFALL, O'AHU
Spec I..A20 A~O A60 A80 820 840 860 C20
Stat IonC40 C60 020 040 060 080 £20 [40 £60
).95 0.69 0.09
4.59 )4.)0 )0.)0
000
0.22 0.36 0.51
2.28
0.52
0.82
5.02
1.62 16.87 10.76
12.62 '.)4 4.'2
o 7.11 0.)0
0.0' 5.15 14.78
7.)6
1.)
6.)7
27.7
0.39
1.46
0.86
15.5
1.% 24.28 12."
16.16 5.20
2.50 2.40
0.87 11.7)
4.20
1.91
2.'7 26.37 10.53 12.00
4.1) 8.19 '.7) 1.57
0.14 O.~ 0 0
0.30 5.28 '5.2 0
O.,~ .1.90
8.6) 3.21
0.14
9.50
0.83
17. )4
2.76•• t
4.60
).54
7.H
10.60
19.70
1.15
0.20
7.05 7.70
8.SO 16.7)
ot 0.09
0.02 0.05
1978 7.62 10.78
1975 2).00 1).8)
1,,8 O.~ 0
'''5 0 0.02
0.0)
3.14
o62.42
o 0
o 0
o 0
o 0
o 0
0.08 0.09
).)9 0
3.13 0
o 0
o 0
O.I~ 0.03
0.16 0
0.85
0."7.)0
".00.22
o0.03
o 0
o
o 0
4.40 0
o 0
o 0
o.h 7.74
o 0
o 0
o
15.9) 0
o
o 0
o 0
o 0
o 0
3.62 15.60 '.1'
o 3.82
8.44 40.11 11.4)
49.86 53.12
o 0
o 0
18." 0
o 0
o 0
o 0
0.23 0
o 0
14.57
3.19
0.80 8.52
3.68 0
000
000
0.47 0 0
0.02 0.01 0
000
0.28 0 0
000
000
000
000
0.06 0 0.51
0.09 0 5.0
1.55 0
0.63 60.45 67.71
0.06
2.62
o 0 0 0
o 0.05 0 0
o 0 0 0
o 0 0 0
000 0
o 0 0.11 0
o 0 0 0
o 0.46 0 0
o 0 0 0
o 0 0 0
0.58 0 0 0
4.70 0 0.4) 0.3)
0.15- 0.14 0.05 0 0 0.35 0.41
0.16 0 0.'9 0.68 0.47 6.45
o0.)1
o 0
o
o 0
o
o 0
o
o 0
o
o 0
0.02
O.O~ 0
o
o0.)0'
0.19
o
.,78 0 0
1"5 0 0
,,,8 0 0
'''5 0 0
1,,8 0 0
1"5 0 0
.,,8 0 0
1"5 0 0
1,,8 0 0
1"5 0 0
1,,8 0." 0
1"5 0 0.05
1,,8 80.67 69.70
'''5 51.82 40.09
1,,8 4.26 11.71
'''5 16.71 29.60
7).35 86.16 94.60
51.20 Z).49 40.33
).80 1.60 0.01 ~1.2) ~.6
28.30 14.66 45.34 89.6
".17 58.24 0.53
8.33 10.34 I.~
oo
oo
2.52
I.~I
o'0.37
oo
1~.03 1.50
0.84
oo
o
o
oo
79.32 13.6~ 66.40 25.61 55.93 86.60
18.39 9.10 fl.77 38.70 14.16
o28.09
oo
oo
65.35 85.50 61.69
17.55 7.76 51.28
0.59 0.09 0.19
0.06 1.62
oo'.0
1.14 °o
o
°
12.31
".90
o
°1,,8 0
1"5 0
L....atona
Sond Rubble
lIoncorel
Ia.elt
Totol CorelCover
1978 15.07 18.57
1975 31.69 30.98
14.33 12.45
27.64
5.46 0.83 0.53
29.47 44.26 44.13
4.86 34.04 14.41 38.00
9.07 82.67 92.20 21.49
33.94 86.36 53.13 12.59 41.58 13.39
82.~ 91.14 17.71 60.06 85.37
Difference In coralcoyer bot""'.n 1975end 1978
16.62 12.41 13.32 24.01 43.43 43.60 '.81 48.63 77.79 16.51* 49.04 4.78 5.12 18.48 71.98
H'-SpeclesCover DI ver'Sl ty
1978
1976
0.47
0.62
0.94
0.76
0.63
0.71
0.59 0.61
1.1~
0.45
1.21
0.36
0.95
1.00
1.27
0.82
0.95
0.55
0.85
1.29
1. 14
1.21
1.06
1.61 0.94
0.70
1.61
0.83
0.58
0.83
"Higher coy.r In 1978.t_ .... No dat8 collected.to • Spec Ies not found. no l.
13
80 STATION A 1.6 STATION A
60 1.2
40 0.8
20 0.4
0 0.0
80 STATION B 1.6
60 1.2
-J:40 ~ 0.8
>-20 I- 0.4
(/)
a:: a::ww 0 > 0.0
>0
STATION C0
STATION Cv 80 w 1.6.J Cl<X <Xa:: 60 a:: 1.20 wv >.J 40 0 0.8<X vl- I0 20 (/) 0.4I- w~ 0
u0.00 w
a. /j
STATION D(/)
-,,-"STATION D80 .J 1.6<X
60 I- 1.20I-
40 0.8
20 0.4
0 0.0
80 STATION E 1.6 STATION E
60 1.2 A--- 1975 Ii --- 1975
40 -- - 1978 0.8 - --1978I
20 0.4 I-
0.020 40 60 80 100 120 0 20 40 60 80 100 120
TRANSECT DEPTH, It TRANSECT DEPTH, It
NOTE: Ft x 0.304 8 = m.
FIGURE 3. VARIATION OF TOTAL CORAL COVER AND SPECIES COVER DIVER-SITY WITH DEPTH AT EACH TRANSECT, 1975 AND 1978, MOKAPUOUTFALL, O'AHU
14
20
1978STATION A Montipora spp. 0
Poct/lopOra meandrina 1//1Porites lobata f:;/rJPorites compressa •
10080
1975
////
~~ ;;I ;; ~;:.~;:.[;.·.•i;;11;;;;:..:••......:..;:;.•.::;;
/ /;;/;; :~::.•..:~;::: :~ ::::::;: ;~~~i·II:III\!oL....c:::::I.<::..L.~a. __---'=~~;;;:;:;:l__...c:=:L~=~
-1..--1LL..=
20 40 60
10
a: 10lJJ
6<..>~ol-I-o(I)
cfl 20
DEPTH, It
NOTE: Ft X 0.304 8 = m.
FIGURE 4. PERCENT COVER OF FOUR DOMINANT CORAL SPECIES, STA
TION A, MOKAPU OUTFALL, O'AHU, 1975 AND 1978
1978
10
0::::::;:;
a:lJJ
1975>0<..>~ 300l-I-0(I)
~ 200
10
020 40
STATION 8
Ill'
;;160 80
Montipora spp. 0Pacillopora meandrtna[22]
Porites lobata [J]
Porites compressa •
100
DEPTH, It
NOTE: Ft X 0.304 8 = m.
FIGURE 5. PERCENT COVER OF FOUR DOMINANT CORAL SPECIES, STA
TION B, MOKAPU OUTFALL, O'AHU, 1975 AND 1978
15
30
1978
20
a: 10w>0u~0 0~ 1975~0CD
~ 200
10
STATION C
IMontiporo spp. 0Pocilloporo meondrino k /1Porites lobata (::::::::::1
Porites campresso •
20 40 60
DEPTH, ft
NOTE: Ft X 0.304 8 = m.
FIGURE 6. PERCENT COVER OF FOUR DOMINANT CORAL SPECIES, STATION C, MOKAPU OUTFAll, O'AHU, 1975 AND 1978
Montipora are grouped together because many of these species are so morpho
logically similar that they are indistinguishable in the transect photo
graphs. Figure 9 shows the number of coral species at each transect during
both the 1975 and 1978 surveys.
By conducting repetitive surveys at the same locations in successive
years, one can distinguish changes and effects in both space and time.
When the benthic-cover data obtained in the 1978 survey is compared to the
1975 survey data, no clear and consistent patterns associated with the out
fall location are evident. The only pattern that does occur is that coral
cover was lower on all but one transect in 1978. This trend does not ap
pear to be correlated with outfall stress since some of the greatest dif
ferences occurred at stations farthest from the source of discharge. In
stead, the major differences that did occur appear to be the result of
patchy distribution of benthos and substrate types rather than biotic re
sponses to outfall stress. ($ee discussion below.)
The greatest differences occur at the outfall site (station B) where
coral cover of all species, number of species, and species cover diversity
16
60 ....--------------------------------.....,
50
1978 STATION D
Montipora spp. 0Poci//oporo meandrino 1221
40
30
20
a: 10lJJ>0u::E0 0t- 1975t-Oco~ 500
40
30
Porites /obato
Porites compresso
Iiill
•
20 40
DEPTH, ft
60
NOTE: Ft X 0.304 8 = m.
FIGURE 7. PERCENT COVER OF FOUR DOMINANT CORAL SPECIES, STATION D, MOKAPU OUTFALL, O'AHU, 1975 AND 1978
1978
20
10
ol-__--l=t;;;;;:;:a..- ~L.£i:i:i:i<
1975
0:: 60W
~U
~o 50l-I-om;f!. 40
30
20
STATION E
Montiporo spp. 0Pocil/opora meondrina I22JPorites lobata liDPorites compressa •
17
DEPTH. ft.
NOTE: Ft x 0.304 8 = m.
FIGURE 8. PERCENT COVER OF FOUR DOMINANT CORAL SPECIES, STATION E,MOKAPU OUTFALL, O'AHU, 1975 AND 1978
19
were lower in 1978 (Figs. 3, 5, and 9). In contrast, observations showed
that an enhancement of the community had occurred at the outfall structure
itself due to settling of various benthic forms on the complex substrata
formed by the diffuser pipe and boulder cover. The other area of greatest
cover difference between 1975 and 1978 occurred at station E60, the station
farthest from the outfall diffuser. Stations C and D, between stations B
and E, had relatively similar community structures at both surveys while
station A, closest to the outfall, showed consistently less coral cover and
lower diversity in 1978 relative to 1975.
Appendix B gives a brief summary of each transect location during the
1978 survey, describing both the physiography of the substratum and the
dominant macrobenthos that inhabit the area.
Discussion
The environmental variables seeming to affect coral community struc
ture most directly are wave energy which causes skeletal breakage and tis
sue abrasion, available light energy associated with photosynthetic and
calcification processes, available solid substrata for larval settling,
sedimentation that may smother corals, and interspecific competition be
tween corals and other benthic forms. The relationships of these natural
factors to the coral communities located at the five study stations in
Kailua Bay are discussed by Russo et al. (1977).
Possible causative factors of stressed benthic communities which have
received particular attention in the past include increases in nutrients,
turbidity, sediments, and various toxins (EPA 1976). It is possible that
all exert· some influence on the distribution and abundance of the benthos,
and some effects are probably synergistic (Grigg 1978). Human activity can
also alter coral community structure by creating artificial changes in some
of the above variables. The major environmentally significant aspect of
the delivery of organic matter (sewage) is nutrient loading and consequent
cultural eutrophication of the ecosystem (Smith 1977). Accumulation of
organic-laden sediment emanating from the outfall discharge or from passage
through the food chain can modify or cover substrates causing them to be
come unsuitable for settlement of many epibenthic species (Grigg and Kiwala
1970). While sewage outfall discharge can increase turbidity, causing a
decrease in the light energy available for photosynthetic activity, an
20
effect of sewage effluent on reef corals appears to be domination of the
hard bottom substrates by epifaunal filter feeders dependent upon suspended
organic materials in the water column. These filter feeders can success
fully outcompete corals for space and light. Benthic algal blooms that are
attributable to eutrophication from sewage outfalls also have resulted in
reduced settlement and smothering of corals in nutrient-rich outfall areas.
Two sewage outfalls on O'ahu have been shown to cause these types of
effects on coral communities. Within 500 m of the old Sand Island sewage
outfall, Grigg* has observed bottom communities dominated by mounds of
deposit-feeding polychaete worms (Chaetopterus sp.); corals were totally
lacking. Between 500 and 1 000 m the Chaetopterus mounds disappeared and
substratum consisted of either sand or limestone. Echinoids, zoanthids,
and calcareous algae were the dominant macrobenthos in this zone. Beyond
1 000 m, living corals and associated species gradually increased in abun
dance up to a distance of 6 000 m. The highest species diversity of the
macrobenthos was observed at stations of intermediate distances where nor
mally dominant species were less abundant and species associated with the
outfall were present.
Likewise, Kane'ohe Bay, O'ahu, has been chronically polluted by sewage
as well as subjected to substantial terrestrial input from sediment-laden
stream runoff. Today, the southeastern sector of the bay, the area receiv
ing large amounts of both sewage and stream flow, bears little resemblance
to a coral reef. Most areas once supporting coral? now have a variety of
filter feeding organisms--barnacles, sponges, tunicates, oysters, poly
chaetes, etc. The central sector presents a picture of a coral reef commu
nity with conspicuously altered biological characteristics. In particular
the green alga Diatyosphaeria cavernosa has displaced corals, primarily
Porites compressa on the reef slopes of that sector (Smith 1977). Present
ly the sewage discharge into Kane'ohe Bay is suspended, and the resulting
responses of the communities to this relaxation of stress are being moni
tored.
However, these types of responses to sewage stress do not appear to be
taking place in Kailua Bay approximately 1 yr after commencement of outfall
operation. On the contrary, the outfall structure itself is creating addi-
*R. Grigg 1978: Personal communication.
21
tional hard substrate area for coral colonization; and numerous small colo
nies, most of which appear to be PooiUopora meandrina and Porites lobata,
were observed on the large basaltic boulders that are placed over and adja
cent to the outfall pipe. The bryozoan Triphyllozoon hirsutum Rusk was also
observed abundantly on these outfall boulders at the 24- to 30-m (80- to
100-ft) depths. This was the only location in the entire study area where
Triphyllozoon was observed. The boulder stacks also form an area of habi
tat space that is far more complex than the broad flat pavement that the
outfall pipe bisects. Several spiny lobsters,PanuZirus spp., were observed
in the shelter of the interstices between the boulders, as wer; several
species of sea urchins, Heterooentrotus mammiZatus, and Eohinothrix spp.
In this sense, the outfall pipe and adjacent boulders are acting as an
"artificial reef" in an area which has extremely low relief and is barren
of extensive settlement.
None of the benthic filter feeders or benthic algae mentioned as being
common around the Sand Island and Kane'ohe Bay outfalls were observed in
the immediate vicinity of the M6kapu outfall. The only abundant filter
feeder that has colonized the area is TriphyZZozoon hirsutum, and this spe
cies is presently limited to the outfall structure itself, appearing in
relatively small clumps with large areas of bare substrata still available.
It is possible that the combination of a relatively low discharge rate
(0.4 m3 js or 9 mgd) and relatively strong and variable current velocities
of up to 1 knot around M6kapu Point (Bathen 1972) disperse the effluent
rapidly enough so that the increase in suspended organic material is not
great enough to support dominating communities of filter feeders or leafy
benthic algae like those found in the proximity of Sand Island or Kane'ohe
Bay. This reasoning is supported by the observation that there is no evi
dence of effluent discharge in the surface waters of Kailua Bay. At Sand
Island the discharge location was readily apparent from the surface, both
by sight and odor.* Also, the discharge effluent at Mokapu has been ob
served to disperse rapidly in the water column; visual effects of impair
ment of water clarity were not apparent except within about 2-3 m of the
discharge exits in the outfall pipe (Plates 2 and 3).
The abundance of the bryozoan, TriphyZZozoon is also significant with
*R. Grigg and M. Palmgren 1978: Personal communication.
22
PLATE 2. DIFFUSER VENT IN MOKAPU OUTFALL PIPE, O'AHU, 1978.PLUME OF SEWAGE CAN BE SEEN PULSING FROM THE VENTIN THE LOWER CENTER OF THE PICTURE. NOTE WATERCLARITY ADJACENT TO OUTFALL VENT, INDICATING THATDISCHARGE AREA IS RAPIDLY FLUSHED. ALSO NOTELACK OF SEDIMENT ACCUMULATION ON BOULDERS ADJACENT TO VENT.
PLATE 3. EXPOSED MOKAPU OUTFALL DIFFUSER PIPE, O'AHU, 1978.NOTE HIGH NUMBERS OF FISH AROUND DIFFUSER.
23
respect to physical parameters of the waters near the diffuser. Soule and
Soule, research fellows at the Alan Hancock Foundation, University of
Southern California, state in an unpublished manuscript that no bryozoans
are found where rapid dispersion of fine sediments is taking place. Grigg
(1978) observed no bryozoans at stations up to 4 miles (6 436 m) from a
sewage discharge off Palos Verdes, Calif. Turbulence is also a controlling
factor for bryozoans. The erect, branching forms are too fragile to toler
ate wave action; and even the tight-clinging, encrusting forms grow only on
the protected surfaces of rocks and coral heads. However, bryozoans do not
tolerate stagnant water well. The polyps must be provided with moving
water for filter-feeding and excretory exchange.*
The Mokapu outfall diffuser seems to be an ideal habitat for TriphyZ
Zozoon hirsutum. The organic load of the surrounding water is high enough
to provide abundant material for filter feeding. However, this load is
presently low enough and water movement is high enough to prevent sedimen
tation that is a limiting factor to bryozoans. Since the diffuser is below
normal wave base of 24 to 30 m (80-100 ft), water movement is not intense
enough to be destructive to the colonies. Therefore, the abundant appear
ance of bryozoans is a good indicator that the physical condition of the
environment is presently not stressed with respect to sewage-induced sedi
mentation.
Another possibility is that the time since discharge commencement has
not yet been long enough for the community to undergo the successional
changes to a climax community of filter feeders or algal domination. It is
possible that the community is now in some preliminary stages where organic
material levels are building to a threshold point, after which filter
feeders and leafy algae will begin to proliferate.
Substrates may also require time to proceed through conditioning
stages, after which new forms will begin to settle and grow. As noted pre
viously, corals have settled on the new substrates immediately adjacent to
the direct flow of the outfall. These colonies are still less than several
centimeters in size; presumably because corals are known to require condi
tioning of their substrate (Harrigan 1972), growth is relatively slow.
However, the clumps of TriphyZZozoon are presently as large as 25 to 30 cm
(Plate 4). It will be interesting to see if the present relationship con
tinues, and, if so, to what degree the bryozoan outcompetes the corals for
*D.F. Soule and J.D. Soule 1978: Personal communication.
24
PLATE 4. BRYOZOAN, TRIPHYLLOZOON HIRSUTUM BUSK, GROWING ONOUTFALL BOULDERS, MOKAPU OUTFALL, OIAHU, 1978.LARGEST COLONIES ARE APPROXIMATELY 30 CM IN DIAMETER. EXPOSED OUTFALL PIPE SURFACE IS AT LEFT.
space on the outfall boulders. It is also possible that other filter feed
ers will begin to settle and proliferate in the outfall area, outcompeting
TriphyUozoon. No "before-after" studies have been conducted previously on
sewage stresses in Hawai'i, so there is no information base with which to
compare the observed pattern of change at M6kapu Point. It may be that the
Sand Island area went through a similar period of small coral and bryozoan
abundance prior to the population explosion of Chaetopterus worms.
Hopefully, the third survey, proposed for the summer of 1979, will
provide additional data to show what the effects of the effluent discharge
are on community successional patterns. If colonization and growth on the
new substrate continue to be mainly corals and other organisms not asso
ciated with severe sewage stress, then it may be concluded that for at
least two years the outfall has had no negative, and even some positive,
effects on coral reef and reef animal growth.
Apparent dissimilarities between the corresponding transects of the
1975 and 1978 surveys could have been from several causes. The most obvious
cause for less coral cover at the later date would be adverse effects of
25
sewage introduction to environment. However, this does not seem to be the
case. As stated previously, none of the detrimental effects of sewage-
turbidity, sedimentation, proliferation of algal blooms, or high concentra
tions of benthic filter feeders or algae--are in evidence. Also, the
greatest decrease in coral cover (other than at the outfall station) occurs
at stations A and E. Station E is farthest away from the outfall and would
be expected to exhibit the least change in coral cover if the range of in
fluence of sewage were responsible for the noted decrease.
What does seem to be the cause of the variance is the inexact replica
tion of the transects. It appears that natural variability from patchiness
of bottom topography and coral cover and the variability of transect loca
tion caused by inexact navigation is of a magnitude great enough to mask
any changes caused by sewage stress.
While it would be expected, under random patchy conditions, that as
many transects would have greater coral cover in 1978 as in 1975, this was
not the case. On only one transect (D20) was the coral cover higher in
1978. While this result appears to be caused by some nonrandom phenomena,
examination of the areas involved provides reasons for the unexpected dis
tributions.
In 1975 station B showed coral cover of 29.5, 44.3, and 36.5% at the
6, 12, and 18 m (20, 40, and 60 ft) transects, respectively. In 1978 the
same transects showed coral cover of 5.5, .8, and .5%. Although this
reduction could be the result of outfall construction that proved to be
lethal to corals in the immediate area, it seems that the bottom topography
of the two transects is different enough to conclude that the locations are
different. With the exception of B20-78 which was located in an area en
tirely of basalt substrata, most of the substrate at the 12- and l8-m depths
was unconsolidated sand and rubble, and a flat limestone pavement mostly
devoid of benthic cover. In contrast, the 1975 transects all appear to
have occurred on patches of limestone veneers of active reef growth. It
seems that the small variation in horizontal distance within the study area
reflects large differences in community structure and the factors that
affect this structure. In this case, this proves to be an advantageous
situation since the area where the outfall occurs is mostly barren; and,
initially at least, the outfall is providing settling sites for a more
diverse substrata. On the other hand, the benthic communities in the area
26
surveyed in 1975 may have been more damaged by outfall construction.
Similarly, the percent coral cover at transects E60-75 and E60-78 were
85.4 and 13.4%, respectively. Much of the coral cover in the area of the
1975 survey was one species, Montipora patuZa (62.4%). This species was
not recorded at all on the 1978 transect. The high abundahce in the first
survey was due to the transect location in the center of and parallel to
the depth contours of a small area of steep vertical relief. Seaward of
this small cliff area the substrata were sand and coral rubble, while the
shoreward side was a flat limestone shelf with numerous encrusting species.
It appears that M. patuZa has the adaptive advantage of populating steep
vertical areas by growing in an overlapping plate-like form. This growth
form is rarely observed on flat areas. The steep cliff area encountered in
the 1975 survey area was very narrow, with a change of only about 4 verti
cal meters and did not appear to be a common topographic feature in Kailua
Bay. Since the 1978 transect was not located on this cliff, bottom commu
nity structure was markedly different between the two surveys. It is inter
esting to note that even though the total coral cover at E60 was markedly
different, species diversity (HI) was closely parallel (Fig. 3). While
total coral cover varied considerably, both transects were dominated by one
species, P. Zobata in 1978 (86.3%) and M. patuZa in 1975 (73.1%).
The opposite situation with respect to species cover diversity occurred
at station D. In the 1975 survey transects D40 and D60 were conducted par
allel to the upper and lower edges of a cliff area densely populated with
plates of M. patuZa. In the 1978 transect, the l2-m (40-ft) transect was
conducted on a flat reef veneer that was not clearly dominated by anyone
species. While the coral cover is greater on the 1975 transect (83%) com
pared to 1978 (34%), the diversity relationships are reversed: .89 in 1975
compared to 1.45 in 1978. When coral cover is low, enough bare substrate
is available for a relatively large number of species to settle without com
petitive superiority by one species. In contrast·, in areas where one spe
cies can effectively outcompete other forms to dominate most of the avail
able space, diversity is low.
Variability within very small areas is most clearly shown in station A.
This station was the easiest to relocate since it was positioned directly
off Mokapu Point and landmarks on the point were readily recognizable. How
ever, results show that at the 6-, 12-, and l8-m (20-, 40-, and 60-ft)
27
transects the 1975 survey consistently showed higher total coral cover and
species cover diversity (Fig. 3). This result could be interpreted to re
flect detrimental effects from the outfall causing a reduction of live
coral cover. If this were the case, it would be expected that higher lime
stone cover would be recorded in the later survey since this is the compon
ent including dead coral colonies. However, at all three transects lime
stone cover was higher in the 1975 surveys, conducted before the sewage
outfall was constructed. Again it is concluded that small variation in
locations can reflect relatively large differences in benthic community
structure. This area is directly off a point projecting into the area of
highest wave activity and current energy. Apparently these are the factors
which affect coral community structure in this area since there is no pau
city of hard substrata and the most common species is P. meandrina. This
coral is considered a fugitive species since it is first to colonize areas
of new substrata and occurs in areas too harsh, with respect to wave energy,
for other species. Relatively small changes in depth and distance from the
shoreline cliff around Mokapu Point are apparently enough to reflect changes
in community response to factors of wave stress. This is most clearly seen
at the 6-m (20-ft) transect which showed 23% P. meandrina cover in 1975 and
only 7.6% in 1978.
MICROMOLLUS KS
Methods
The micromollusks described below were obtained primarily on transects
C, D, and E. A single sample from a l2-m depth (40 ft) was obtained on
transect A; and, of the two samples from transect B (excluding the two sam
ples from the diffuser), only the sample from 24 m (80 ft) had sufficient
specimens for analysis.
Micromollusks were analyzed as described previously (Russo et al.
1977). Sediment samples obtained by hand by scuba divers at depths of 6 to
24 m (20 to 80 ft) were picked for shells under a binocular dissecting
microscope from volumes of 25 cm~ The micromollusks were then separated
to species, counted, and analyzed for species composition and species diver
sity. In this report the term "abundance" is substituted for "standing
crop" used in the previous report (Russo et al. 1977); it was determined as
28
was standing crop in the earlier report, by dividing the number of shells by
sediment volume. Figures for abundance are minimal in that they do not in
clude fragments and juvenile shells too small for identification. Species
diversity indices were calculated from the Shannon-Wiener function, HI =rpilog~Pi (Pielou 1969). Species composition is represented by relative
abundance values for various groups and species.
Results
Abundance, species diversity, and species composition of the micro
molluscan assemblages are shown in Table 4, and the distribution of these
parameters on the transect lines is shown in Figures 10 to 13. In the
following discussion, the samples are separated into two groups of stations,
at 6- to l2-m (20- to 40-ft) depths and 18- to 30-m (60- to 100-ft) depths.
This separation was suggested by differences in species composition which
were confirmed by analysis utilizing the Sorenson coefficient of similarity
which demonstrated two clusters of assemblages, a shallow water assemblage
at 6 to 12 m and a deeper water grouping at 18 to 30 m.
As in the 1975 samples, gastropods comprise between 86 and 100% of the
micromolluscan assemblages on transects C to E. Most of the gastropods are
epifaunal; hence, the assemblage is largely a weed- or rubble-associated
assemblage rather than one associated with infauna.
At depths of 6 to 12 m (20 to 40 ft) on transects C through E, average
abundance is 7.1 shells/cm 3• Four groups of micromollusks comprise about
50% of the assemblages on each transect: the archaeogastropod TpicoZia
vaPiabiZis, the rissoids Rissoina ambigua and R. miZtozona, the cerithids
Bittium paPcum and B. zebPum, and species of the family Triphoridae. At
depths of 18 to 30 m (60 to 100 ft), average abundance is slightly higher,
9.1 shells/cm 3, than at the 6- to 12-m (20- to 40-ft) depths. The dominant
species at the deep stations are the rissoid VitPieithna marmopata, the
dialid CePithidium peppaPVuZum, species of the family Triphoridae, and
TPicoZia.
At the 6- to 12-m depth interval abundance, species composition, and
distribution are in the main similar to those described in 1975; but some
differences, especially with respect to the proportions of species present,
are apparent. Overall abundance at this depth interval was little changed
with respect to the 1975 abundance figure (7.1 shells/cm 3 vs. 7.6 shells/
TABL
E4.
ABUN
DANC
E,SP
ECIE
SD
IVER
SITY
,AN
DSP
ECIE
SCO
MPO
SITI
ON
OFM
ICRO
MOL
LUSK
SAT
TRAN
SECT
STA
TIO
NS,
MOK
APU
OU
TFA
LL,
OIA
HU
,19
78
Stl
ltlo
n*C2
002
0E2
0A4
0Cl
tO04
0E4
0C6
006
0E6
0B8
008
0D
lff.
At
Dlf
f.Bt
~o.
mlc
rom
ollu
sks
6241
425
515
145
125
188
3310
567
349
446
269
319
No.
/cm
!2.
516
.610
.26.
01.
85.
07.
51.
34.
22.
714
.017
.810
.812
.8H
'(s
pec
ies
div
ersi
ty)
3.6
4.4
2.3
4.8
4.1
3.6
4.1
1.2
3.4
4.3
3.4
4.1
2.8
3.4
No.
gas
tro
po
ds
6140
725
414
745
125
184
3210
258
323
407
267
316
%C
ompo
sIt
Ion
tT
rio
oti
ava
ria
bit
is25
1864
89
1417
----
147
4§
4
Ris
80in
aam
bigu
aJ3
4--
II4
--I
Ris
soin
aep
ham
ilta
--§
--I
2--
§--
----
I4
32
Ri8
soin
am
itto
sona
1514
§5
2223
23--
37
--4
I§
Vit
rio
ith
na
mar
mor
ota
37
--4
9--
5--
37
832
I4
Bit
tiw
np
ar=
lzeb
rum
6II
§5
----
----
----
--§
ceri
thid
iwn
perp
arvu
lwn
--§
----
2--
§--
234
4218
2332
Dia
tava
ria
----
--5
--§
§3
I--
5I
4724
Tri
ph
ori
dae
53
--9
910
5--
--14
25
46
*ln
dlc
ates
tran
sect
(A,
B,C
,0,
E)an
dde
pth
(20,
40,
60,
80ft
).tS
amp
les
at
dif
fuse
r.T
Per
cent
of
gas
tro
po
ds.
§Am
ount
too
smal
lto
beco
unte
d,
N 1.0
30
5.0
30 B
204.0
10
0
15 3.0
10
,'a,5 0 ..4.0
...... -J:, \, ,,., 'o-~-o 0'
E ~\
u 0 \
"-.... \
ci 30 Diii \
Z IX: ,~
UJ 3.0UJ ><.> 20 0 Dz<l ,0 (J)
0 IUJ
Z U:::> 10 IlD
, UJ
<l 'o----d, 0-
(J) 4.0 ,0
,\ ,,
50 E0-...... ,
"-0
40 3.0
E
30
20 4.0
1975
10 1978 1975
1978
0 3.020 40 60 80 100 120 20 40 60 80 100 120
DEPTH, ft DEPTH, ft
NOTE: Ft x 0.304 8 = m.
FIGURE 10. ABUNDANCE AND SPECIES DIVERSITY OF MICROMOLLUSKS ONTRANSECTS B THROUGH E, 1975 AND 1978, MOKAPU OUTFALL,O'AHU
31
25
1975 \1 \----- 1978 21 1 \
1 \I \
1 \~ 17 1 \0 1 \40 z I \0 13
I \.... I \
30 ::l 1CD I.. 0:: 9 I, .... I.. Cf) 0
20, , I'0 0 ,
1.. 5,
I, ,, I10
, ,,, ,I, 0
0'0
DiDID vtlf'iD17 48
13 44
9 40~0
Z 5 0, 360.... ' ..,Cf) ' ..0 I '0------0----_0 320-~
I")
12 rrit:(J/iD WlriDbili6 E0 u0 ~ 280z
8ILl, 0 24, z
4,
~, 0, z,, ,,-- ::l 20CD
0.. ".
~
16 Trlpllorldoe 16
12 12
/a...B 8 / '\
I ,I ,
I,
4I "4 I '0
0 ..
0.........
0A 8 C 0 E A B C 0 E
TRANSECT TRANSECT
NOTE: ft x 0.304 8 = m.
FIGURE 11. SUMMARY OF MICROMOLLUSCAN SPECIES DISTRIBUTION AND ABUNDANCEAT DEPTHS OF 60 TO 100 FT, 1975 AND 1978, MOKAPU OUTFALL, O'AHU'
32
TRANSECT C TRANSECT 0 TRANSECT E30
064%@20'
\\ \
20 \ 'a.. "\
\"."'0\
10
~" #,1)...
0Hi~~oino millozono,0 H/~~oino mlliozono
20 ' \,, \0' \
\10 \
\\\
0Vifricirhntl Vifricifhna Vifricithna
0~ 8 '\0 , \ ..0- " \
.. 'Z , \ 0'0 4 ~. \ ,... ~.
,• ,(/) \0 \ ~
\ ,Q. 0:E a. porcum /8. ztllJrum 8.parclllll /8. z_rum0U
8
4
0Trip horldo. Triphorido. Triphorldoe
15
10 ,0, \
,/ \\
5 d \\\
\
0
DEPTH,ft
NOTE: Ft x 0.304 8 = m.
FIGURE 12. MICROMOLLUSCAN SPECIES COMPOSITION AT DEPTHS OF 20 TO 100FEET, MOKAPU OUTFALL, 1975 AND 1978
33
30Tricolio yorlobilis
20
10
1975
- ---- 1978
---0,"' ...
'0
Bittfum porcum
Rlssoino mll/ozono
,,~" ," ," ,,," ,"cI ", ,,,
O~I:::::.-~~-----==::::::::~H
4
01---------=---------;8
cf?-.i 20oIen 10o0..:E8 0 r..-------------------l
EoC
TRANSECT
BA
24 r--r----,.---,------r---....,-......
4
O~..L___~.:..___..L.____.J. ~_J
20
~ 12z<toz 8:JCD<t
toE~ 16oz
EoC
TRANSECT
B
_........-o-----~,--- ""
'0
0 _
--------0' .. ........
0 .. ....' ....
A
4
8
Vllricithno mormor%
8
4
O~~----L----'------'--_--L---'
01-----------........---......-;12
Note: Ft x 0.304 8 = m.
FIGURE 13. SUMMARY OF MICROMOLLUSCAN SPECIES COMPOSITION AND ABUNDANCEAT DEPTHS OF 20 TO 40 FT, MOKAPU OUTFALL, 0 I AHU, 1975 AND 1978
34
cm 3). However, in 1978 there were proportionately more TricoZia, Bittium
parcum, and Vitricithna and lesser proportions of Triphora spp. in the sam
ples than were recorded in 1975. Rankings of the most abundant species are
shown in Table 5, and the changes in rank as reflected in the general dis
tribution of species are shown in Figure 13.
TABLE 5. MEAN PERCENT COMPOSITION AND RANKS OF SIX DOMINANT SPECIESOF MICROMOLLUSKS AT DEPTHS OF 20-40 FEET*, 1975 AND 1978,MOKAPU OUTFALL, O'AHU
1975 1978x % of x %ofSample Rank Sample Rank
Rissoina miZtozona 19 1 14.7 2
Triphoridae 9.7 2 4 5
TricoZia variabiZis 8 3 26.3
Rissoina anibigua 5.7 4 4.3 4
Bittium parcum 4.3 5 5.7 3
Vitricithna marmorata 1.6 6 4 5
*Ft x 0.304 8 = m.
Differences in species composition at the 18- to 30-m depth level are
not so noticeable as at the lesser depths, but there is a noticeable differ
ence in abundance, from an average of 23.8 shells/cm 3 at these stations in
1975 to an average of 9.1 she11s/cm 3 in 1978. Vitricithna replaces Cerithi
dium perparvuZum as the most abundant mollusk at these depths in the 1978
samples, and TricoZia variabiZis occurs in greater abundance (Table 6).
TABLE 6. MEAN PERCENT COMPOSITION AND RANKS OF FIVE DOMINANT SPECIESOF MICROMOLLUSKS AT DEPTHS OF 60-100 FEET*, 1975 AND 1978,MOKAPU OUTFALL, O'AHU
1975 1978- % of x % ofxSample Rank Sample Rank
Cerithidium perparvuZum 30 1 10.5 2
Diala varia 9 2 1 5
Vitricithna marmorata 7.3 3 15 I
Triphoridae 5 4 8 3
Tricolia variabilis 4.7 5 7.5 4
*Ft x 0.304 8 = m.
35
Comparison of the 36.6-m (120-ft) depth level samples on transect B
with the two samples from the diffuser area in 1978 show little change in
species composition but a major difference in average abundance from 22.7
she11s/cm 3 in 1975 to 12.8 shells/cm3 in 1978. Cerithidium perparvuZum was
the dominant species in 1975 and again in 1978; the ranking of the five
dominant species was about the same in 1978 as in 1975.
Discussion
Differences between the 1975 and 1978 micromolluscan assemblages in
clude:
1. A generally lower abundance (number of shells/cm3 of sediment)
2. Increased proportions of Bittium parcum, TricoZia variabiZis, and
Vitricithna marmorata at the 6- to l2-m (20- to 40-ft) depth
levels and concomitant decrease in the proportions of Triphoridae
3. Increased proportions of Vitricithna at the 18- to 30-m (60- to
lOO-ft) depth levels.
The differences may in part be ascribed to the effects of the diffuser
system on the biota and may in part be associated with the patchy nature of
the substrate.
At the 6- to l2-m depth levels the three species which increased in
abundance between 1975 and 1978 were those associated with frondose algae
and/or increasingly soft substrata rather than hard substrates. High pro
portions of all three are characteristic of micromolluscan assemblages
elsewhere on O'ahu where some stress has been identified (Kay 1978). It is
interesting that there are increases in abundance for all three species at
all the stations and concomitant decreases in sponge-associated species
(Triphoridae) at all three stations. But, while it is tempting to speculate
that the relative increases in abundance of the three species are associated
with increased nutrient content of the water, it is also possible to suggest
that the samples reflect a variety of substrates in the area. Dollar sug
gests, for example, that the generally lower coral cover he recorded for this
report are the result of patchy distribution of the benthos and types of sub
strates rather than biotic responses to outfall stress. Thus, the samples
reported here may also reflect the patchy nature of the substrate rather
than an actual change in abundance and species composition associated with
the diffuser system. It should be noted, however, that decrease in abund-
36
ance of shells at depths of 18-30 m (60-100 ft) parallels the finding in
Mamala Bay where decrease in abundance was noted following the opening of
the diffuser system in 1977 (Kay 1978).
CONCLUSION
From the results of this study, the following conclusions are drawn:
1. The fish population study showed no significant difference in fish
abundance, composition, or diversity at stations A, C, D, or E between
1975 and 1978.
2. Fish abundance and diversity increased significantly at the outfall
station (B) from 1975 (before outfall construction) to after outfall
operation in 1978. The formation of new substrate by the outfall con
struction seems to have provided substrate for microscopic algal growth.
Coincident with this was a large increase in herbivorous fishes.
3. The biomass of algae did not change significantly at any of the stations
between studies.
4. Coral cover was lower on all but one transect in 1978. This trend does
not appear to be correlated with outfall stress since some of the
greatest differences occurred at stations farthest from the source of
discharge. Differences may reflect the patchy distribution of the
substrate.
5. The greatest differences in coral species-cover diversity of all species
occurred at the outfall.
6. Filter feeders (bryozoans) were abundant at the outfall site, probably
due to new hard substrate and an increase in suspended organic material
in the water column.
7. Similiarity coefficient and dendrograph analysis revealed a separation
of two clusters of micromolluscan assemblages, a shallow water assem
blage at 6 to 12 m (20-40 ft) and a deeper water grouping at 18 to 30 m
(60-100 ft).
8. Differences between the 1975 and 1978 micromo1luscan assemblages include
(a) a generally lower abundance (no. of she1ls/cm 3 of sediment),
(b) increased proportions of Bittium parcum, Tricolia variabilis, and
Vitricithna marmorata at the 6- to l2-m (20- to 40-ft) depths and a con
comitant decrease in Triphoridae, and (c) increased proportions of
37
Vitricithna at 18- to 30-m (60- to 100-ft) depths. These differences
may in part be associated with patchy substrate.
REFER~NCES
Bathen, K.H. 1972. A descriptive study of the circulation and water quality in Kailua Bay, Oahu, Hawaii during 1971 and 1972. Tech. Rep. No. 29,Look Laboratory, University of Hawaii.
Davis, W.P., and Birdsong, R.S. 1973. Coral reef fishes which forage inthe water column. HelgoLander Wiss. MUresunters 24:292-306.
Dollar, S.J. 1975a. "Zonation of reef corals off the Kona Coast of Hawaii."Master's thesis, University of Hawaii.
1975b. Ecology of West Hawaii coastal waters: A preliminary reporton the relationship between community structure and environmental stressin three Kona Coast coral communities. Report to Water Resources ResearchCenter, University of Hawaii.
EPA and Sanitation District of Los Angeles County. 1976. EISjEIR FinalJoint Outfall System Plan: Vol. I. U.S. Environmental Protection Agency,C-06-1051-010, SCH-740-506-05.
Grigg, R.W. 1972. Some ecological effects of discharged sugar mill wasteon marine life along the Hamakua coast, Hawaii. Report submitted to C.Brewer and Co. and Theo. H. Davies and Co., Ltd., Honolulu, Hawai'i.
1978. Rocky bottom communities: Long term changes at Palos Verdes.In press.
, and Kiwala, R.S. 1970. Some ecological effects of discharged wastes---on marine life. Calif. Fish and Game 56(3):145-55.
Harrigan, J.F. 1972. "The planula larva of PociUopora damicornis: Lunarperiodicity of swarming and substratum selection behavior." PhD. dissertation, University of Hawaii.
Hobson, E.S. 1974. Feeding relationships of teleostean fishes on coralreefs in Kona, Hawaii. Fishery Bull. 72:915.
Kay, E.A. 1978. Micromolluscan assemblages in Mamala Bay, 1977. WaterResources Research Center, Progress Report for the Department of PublicWorks, City and County of Honolulu, Hawaii.
; Reed, S.A.; and Russo, A.R. 1973. A baseline survey of benthic-'b-;-i-ot"-a in Kailua Bay. In The Quality of Coastal Waters: Second Annual
Progress Report, Tech. Rep. No. 77, Water Resources Research Center,University of Hawaii.
Maragos, J.E. 1972. "A study of the ecology of Hawaiian reef corals."PhD. dissertation, University of Hawaii.
Mueller-Dombois, D., and Ellenberg, H. 1974. Vegetation ecology. New York:John Wiley and Sons.
Patrick, R., and Strawbridge, D. 1963. Methods for studying diatom popUlations. J. Water Poll. Control Fed. 35:151.
38
Pielou, E.C. 1969. An introduation to mathematiaaL eaology. New York:Wiley-Interscience.
Russo, A.R.; Dollar, S.J.; and Kay, E.A. 1977. An inventory of benthicorganisms and plankton at Mokapu, O'ahu. Tech. Rep. No. 101, WaterResources ResBarch Center, University of Hawaii.
Smith, S.V. 1977. A preliminary report on the responses of a coral reef/estuary ecosystem to relaxation of sewage stress. In Proa. Third IntL.CoraL Reef Symposium, Miami, ,Florida, pp. 578-83.
Stein, J.E., and Denison, J.G. 1963. Limitations of Indicator organisms.In PoLLution and Marine EaoLogy, ed. T.A. Olson and F. Burgess, p. 323.New York: John Wiley and Sons.
Strasberg, D.W. 1966. Observations on the ecology of four apongonid fishes.Paa. Sai. 20:338-41.
Turner, C.H. 1965. The marine environment in the vicinity of the OrangeCounty Sanitation District's ocean outfall. Calif· Fish and Game 52:28.
____~; Ebert, E.E.; and Given, R.R. 1964. An ecological survey of amarine environment prior to installation of a submarine outfall. CaLif·Fish and Game 50:176-89.
39
APPENDICES
APPENDIX A. FISH SPECIES LIST, MOKAPU OUTFALLFOLLOW-UP STUDY, 1978. . . . . . . . . . . . . . . . . . . 41
APPENDIX B. DESCRIPTION OF CORAL STUDY SITES,MOKAPU OUTFALL, O'AHU, 1978. . . . . . . . . . . . . . . . 46
41
APPENDIX A. FISH SPECIES LIST, MOKAPU OUTFALL FOLLOW-UP STUDY, 1978(HI =Diversity Index J =Equitability Component [HI /H ' max])
Station A20
ThaZassoma baZZieui 8T. duperreyi 18Acanthurus nigrofusaus 37A. Zeuoopareius 7A. triostegus (sandvicensis) 2Ctenochaetus strigosus 2Chaetodon muZticinctus 4C. quadrimacuZatus 1Chromis Zeucurus 2PZectrogZyphidodon imparipennis* 1P. johnstonianus 4Stegastes fascioZatus t 2SuffZamen bursa 3Pervagor spiZosoma 3Scarus spp. 2Naso Zi turatus 2Parupeneus muZtifasciatus 2
Total Species: 17 TOTAL 100HI = 3.6J = .878*old name - Abudefduf irrrparipennistold name - Pomacentrus jenkinsi
Station A40
ThaZassoma duperreyi 30T. baZZieui 2Chaetodon quadrimacuZatus 3Acanthurus nigrofuscus 5A. oZivaceus 2SuffZamen bursa 7Rhinecanthus rectanguZus 1PZectrogZyphidodon johnstonianus 5Pervagor spiZosoma 4Parupeneus muZtifasciatus 1
Total Species: 10 TOTAL 60HI = 2.47J = .74
Station A60
ThaZassoma duperreyi 21Coris gaimardi 2Acanthurus nigrofuscus 8Naso Zituratus 2Ctenochaetus strigosus 7Chromis verator 2DascyZZus aZbiseZZa 2Chronis Zeuourus 8Stegastes fascioZatus 2Chaetodon ZunuZa 1C. quadPimacuZatus 7C. muZticinctus 6C. mi Ziaris 1Forcipiger f1avissimus 3C. coraZZicoZa 3Scarus spp. 1Parupeneus muZtifasciatus 6Pervagor spi Zosoma 3Canthigaster janthinopterus* 1Paracirrhites arcatus 1SuffZamen bursa 2MUZZoidichthys fZavoZineatus t 1Centropyge potteri 6
Total Species: 23 TOTAL 96HI = 3.97J = .874*old name - C. jactatortold name - M. sandvicensis
42
APPENDIX A--Continued.
Station A80
ThaZaBsoma duperreyi 20Aaanthurus nigrofusauB 7Naso Zituratus 1Zebrasoma fZavesaens 2Ctenoahaetus strigosus 21Chaetodon fremblii 3C. multiainatus 5Centropyge potteri 11SuffZamen bursa 2Pervagor spilosoma 1Anampses auvieri 1Foraipiger fZavissimus 2Saarus spp. 2Chromis leuaurus 7Canthigaster janthinopterus 4Paraairrhites araatus 2Aetobatus narinari 1Saorpaena aoniorta 1Parupeneus multifasaiatus 7
Total Species: 19 TOTAL 100H' = 3.56J = .83
Station 820 on Outfall
Chaetodon miliaris >30Chromis verator 5Abudefduf abominalis 15Kyphosus cinerascens 30MUlloidichthys aurifZa~a 10Ctenochaetus strigosus 6Naso unicornis 3N. literatus 2Monotaxis grandoculis 1Scarus spp. 2Parupeneus porphyreus 5
Total Species: 11 >109H' = 2.78J = .799
Station 820 10 m off Outfall
Thalassoma duperreyi 29Aaanthurus nigrofusaus 20A. leucopareiuB 3A. triostegus 4Plectroglyphidodon imparipennis 5P. johnstonianus 4Pervagor spi Zosoma 3Haliahoeres ornatissimus 1Coris fZavovittata 2Paracirrhites araatus 1P. forsteri 1Anampses spp. 2
Total Species: 13 TOTAL 75H' = 2.90J = .781
Station 840 on Outfall
Mulloidiahthys aurifZamma >20Chaetodon mi liaris >20Kyphosus ainerasaens >20Abudefduf abominalis >20Acanthurus triostegus 15A. olivaaeus 5A. nigrofusaus 5Naso li teratus 3N. uniaornis 2Chaetodon fremblii 2Thalassoma duperreyi 4T. baUieui 5Chromis verator 2Bodianus bilunulatus 2Scarus spp. 4Parupeneus multifasciatus 3
Total Species: 16 TOTAL >132H' = 3.48J = .866
Station 840 10 m off Outfall
Thalassoma duperreyi 3Chaetodon fremblii 1SuffZamen bursa 2Parupeneus multifasciatus 2MUlloidichthys auriflamma 2
Total Species: 5 TOTAL 10H' = 2.25J = .97
TOTAL
APPENDIX A--Continued.
Station 860 on Outfall
Kyphosus ainerasaensChaetodon mil iarisMUlloidiahthys aurif1ammaCtenoahaetus strigosusChromis veratorAbudefcluf abominalis
Total Species: 6 TOTALHI = 2.33J = .89
Station 860 10 m off Outfall
Thalassoma cluperreyi
Station B80 on Outfall
Kyphosus ainerasaensChaetodon miliarisAaanthurus olivaaeusMUlloidiahthys aUPif1ammaThalassoma cluperreyiBodianus biZunuZatusLutjanus kasmiraChaetodon frembZiiStegastes fasaioZatusHaliahoeres ornatissimusDasayZlus albisaeZZaCoris balZieuiAulostomus ahinensis
Total Species: 13HI = 2.96J = .797
>20>20>20
53
10
>78
4
>20>20
18>20
55
10221311
>108
43
Station 8 at 27.4 m (90 ft)on the Diffuser
MUZloidiahthys f1avolineatus >20Aaanthurus oZivaaeus >15Lutjanus kasmira >15Abudefduf abominalis >10Chromis verator 6Aaanthurus triostegus 4A. nigoris 2A. nigrofusaus 9Naso li turatus 4Chaetodonmiliaris 8C. frembZii 2Thalassoma duperreyi 3Parupeneus porphyreus 2P. muZtifasaiatus 4Ctenoahaetus strigosus 4Meliahthys vidua 2Anampses spp. 1
Total Species: 17 TOTAL >111HI = 2.96J = .71
Station C20
Aaanthurus triostegus(sandvwensisJ 7
ThaZassoma cluperreyi 2Suff1amen bzaosa 1Pervagor spi Zosoma 2
Total Species: 4 TOTAL 12HI = 1.62J = .81
Station C40
Aaanthurus nigrofusausThalassoma duperreyiPervagor spi losomaCtenoahaetus strigosusPZeatrogZyphidodon johnstonianusSuff1amen bursa
Total Species: 6HI = 2.02J = .78
TOTAL
1463311
28
44
APPENDIX A--Continued.
Station C60
Parupeneu8 muZtifa8aiatu8Da8ayZZus aZbiseZZesPervagor SpiZo8omaCanthigaster janthinopterusCentropyge potteriChaetodon miZiaris
Station 020
Aaanthurus nigrofusausA. triostegusA. mataThaZassoma duperreyiCtenoahaetus strigosusParupeneus muZtifasaiatusA. oUvaaeusSuffZamen bursaChaetodon quadrimaauZatusC. unimaauZatusC. frembUiC. muUiainatusCentropyge potteriNaso U turatusBodianus biZunuZatusStegastes fasaioZatusThalassoma balZieuiRhineaanthus aauZeatusCanthigaster janthinopterusPZeatrogZyphidodon johnstonianus
Total Species: 20 TOTALH' = 3.52J = .81
Total Species: 6H' = 2.37J = .91
TOTAL
421113
12
156
1318
5213122111111111
77
Station 040
ThaZas80ma duperreyi 15Stegastes fasaioZatus 9Ctenoahaetus strigosus 8Aaanthurus nigrofusaus 5Centropyge potteri 7Zebrasoma !Zavesaens 1Chaetodon unimaauZatus 6C. multiainatus 8Pervagor spiZosoma 3Saarus spp. 4Chromis spp. 2StethojuZis baZteata 1Canthigaster janthinopterus 1Paraairrhites forsteri 1Parupeneus bifasaiatus 1BaZistes spp. 1
Total Species: 16 TOTAL 73H' = 3.52J = .87
Station 060Lutjanus kasmira 41Chromis verator* 29Thalassoma duperreyi 9Ctenoahaetus strigosus 19Chaetodon unimaauZatus 13Chromis spp. 18Chromis leuaurus 8Aaanthurus triostegus 8A. nigrofusaus 2Zebrasoma fZavesaens 1ThaZassoma baUieui 2Parupeneus multifasaiatus 3Stegastes fasaioZatus 2Pleatroglyphidodon johnstonianus 2Naso U turatus 1Centropyge potteri 3Abudefduf abominalis 2Chaetodon ornatissimus 1Foraipiger fZavissimus 1Bodianus bilunulatus 1Saarus spp. 1Sufflamen bursa 1Pervagor spi losoma 1Parupeneus porphyreus 1
Total Species: 24 TOTAL 170H' = 3.54J = .77*Misidentified as Dasayllus trimaaulatus in Russo et al. (1977) study.
APPENDIX A-Continued.
45
Station D80
Chpomis ve~ato~* 26Ctenoahaetus swigosus 17Thalassoma duperreyi 10Sufflamen b~sa 4Pleawoglyphidodon johnstonianus 3Chae todon mi l iaris 2Fo~aipige~ flavissimus 2Chaetodon unimaaulatus 2Parupeneus muUifasaiatus 3MUlloidiahthys flavolineatus 3Chaetodon multiainatus 1Canthigaste~ janthinoptepus 1Cent~pyge potteri 1Saapus spp. 2COl'is gaimardi 1
Total Species: 15 TOTAL 78HI = 3.03J = .77*Misidentified as Dasayllus trimaaulatus in Russo et al. (1977) study.
Station E40
Thalassoma dupe~~eyi 17Ctenoahaetus strigosus 21Aaanthur>us nig~ofusaus 16Chaetodon unimaaulatus 7
. Stegastes fasaiolatus 6~omis leuauPUs 3Chaetodon multiainatus 3C. f~embUi 1Cenwopyge potteri 1Aaanthupus t~iostegus 2A. olivaaeus 2Pleat~oglyphidodon johnstonianus 2Chaetodon miUaris 1Thalassoma ballieui 2Canthiga8te~ janthinoptepus 1Fo~aipige~ flavissimus 1Pervago~ spi losoma 1Par>upeneus muUifasaiatus 1Anampses spp. 1Paraai~~hites araatus 1
Station E20
Thalassoma dupe~~eyi
Ctenoahaetus strigosusAaanthupus mataStegastes fasaiolatusAaanth~ leuaopareiusAbudefduf so~didus
Chaetodon miliarisC. quadrimaaulatusCanthigaste~ ainatusOst~aaion lentiginosum
Total Species: 10 TOTALHI = 2.83J = .85
12774222111
39
Total Species: 20HI = 2.85J = .66
Station E60
Ctenoahaetus strigosusAaanth~ wiostegusThalassoma duperreyiT. baUieuiChaetodon multiainatusC. miUarisC. lunulaC. f~embUi
Sufflamen b~sa
Bodianus bilunulatus, Paraai~~hi tes fo~ste~i
P. araatusOswaaion meleag~is
Co~is flavovi ttata
Total Species: 14HI = 3.12J = .81
TOTAL 90
186454311221111
TOTAL SO
46
APPENDIX B. DESCRIPTION OF CORAL STUDY SITES,M5KAPU OUTFALL, O'AHU, 1978
Station A20
Large basaltic boulders and wave-scoured basaltic bottom with encrust
ing calcareous algae characterized this transect. Dominant coral cover con
sisted of numerous colonies of PocilZopora meandrina; some flat encrusta
tions of Porites Zobata; and, occasionally, small patches of Montipora spp.
and Pavona varians. Dead colonies of P. meandrina covered with algal
growth were also numerous. The only sea urchins observed were Echinothrix
diadema.
Station A40
Basaltic pavement and boulders were the major topographic features of
this area. Numerous large P. meandrina colonies, both alive and dead, and
small encrustations of P. lobata were common. Sea urchins observed were
Echinometra mathaei, Tripneustes gratiZla, and Echinothrix diadema. The
black sponge, Chondrosia chucaZZa,was observed on many of the dead coral
colonies.
Station A60
Basaltic pavement with less boulder cover than shallower transects at
Station A were found at the 18-m (60-ft) depth. Fewer P. meandrina colo
nies were observed with the dominant coral species being encrusting P.
lobata. Limestone substrata occupied by boring Echinometra mathaei and
numerous small patches of the zoanthid PaZythoa tubercuZosa were common.
The sea urchins Tripneustes gratiZZa and Echinothrix diadema were occa
sionally observed.
Station 820
Basaltic boulders covered with a filamentous algal turf and small coral
colonies, mainly P. meandrina, were observed at the area of the outfall.
The soft coral PaZythoa tubercuZosa and the black sponge Chondrosia chucalla
were observed occasionally as were the sea urchins Echinometra mathaei,
Tripneustes gratiZZa, and Echinothrix diadema.
47
Station 840
This transect was all coarse sand, limestone rubble fragments, and
basaltic pavement covered with sediment. Very few corals were observed on
the pavement, and these were limited to small patches of P. lobata, usually
less than 5 em in diameter. Abundant stands of Codium sp. and other fila
mentous algae were observed in the sandy areas. No sea urchins were ob
served in this area.
Station 860
Coarse sandy rubble and areas of hard pavement with short algal turf
were the main features of this area. Several areas of basaltic boulders
with small encrusting colonies of Montipora verrucosa and Porites lobata
were observed. No sea urchins were observed in this area.
Station C20
A flat, limestone-covered basaltic pavement with very sporadic macro
benthos characterized transect C20. Corals present were mostly small, flat,
encrusting florms with the exception of some large well-formed hemispherical
heads of P. meandrina. The calcareous alga Porolithon sp. was noted on some
of the boulders as was the sea urchin Tripneustes gratilla.
Station C40
Limestone pavement and a fairly well-developed reef framework charac
terized this area. Extensive lobata, flat, encrusting P. lobata, and hemis
pherical P. meandrina colonies were numerous with small clumps of short
fingered P. compressa. Most of this area consisted of a relatively complex
substrata in areas of higher coral cover with interstitial areas between
and under colonies. Several small patches of sand and coral rubble were
also observed. Black sponges were observed on many dead P. meandrina colo
nies, and Palythoa tuberculosa colonies were occasionally present. Echino
thrix diadema was the only sea urchin observed.
Station C60
This site featured flat limestone veneer with very little substrate
relief. Flat, encrusting species of P. lobata and large, flattened, hemis-
48
pherical clumps of P. meandrina were the only abundant corals observed.
Clumps of Porolithon and numerous sea urchings of the species Tripneustes
gratilla were also observed on the flat pavement.
Station 020
Here, a flat, limestone-veneer pavement with extensive encrusting spe
cies, mainly P. lobata, M. verrucosa and M. patula, was found. Many of
these colonies had long narrow cracks produced by commensal alpheid shrimps,
Alpheus deuteropus. Pocillopora meandrina heads were sporadic as were
clumps of Porolithon sp. and some calcareous algae, Halimeda discoidea.
Echinometra matheai was the only sea urchin present, and these were observed
in holes bored in the limestone veneer.
Station D40
The topography of this transect was generally limestone reef structure.
The dominant corals were encrusting P. Zobata and M. ontipora sp. The coral
Pavona explanulata was very abundant at this transect, the only place in
this study where such concentrations of this species occurred. Palythos
tuberculosa and Porolithon sp. were also occasionally observed.
Station 060
This transect occurred on a sloping bottom (approximately a 30° grade)
that was almost entirely covered with flat overlapping plates of Montipora
spp. and Porites lobata. The remaining bottom cover was nonliving lime
stone composed of dead coral skeletal material. Occasional E. matheai were
observed in this limestone.
Station 080
The substratum at this transect was predominantly sandy limestone
rubble with large, flat colonies of Montipora spp., Porites lobata, and
Porites (Synaraea) convexa. Clumps of P. compressa with long thin branches
were also present. No P. meandrina or urchins were encountered in this
area.
49
Station E20
Most of this transect consisted of irregular limestone outcrops
covered with a thin algal turf. Very few corals were encountered, and these
were predominantly small encrustations of Porites Zobata. Colonies of the
soft coral PaZythoa tubercuZosa were fairly abundant. Several sand patches
and sand channels intersected the boulder platform. High wave energy
appeared to be a consistent condition in this area. No sea urchins were
observed on any parts of this transect.
Station E40
This transect consisted of a relatively flat limestone pavement, with
the main coral species being large, well-developed colonies of P. meandrina;
flat, encrusting P. Zobata; and short-fingered clumps of P. compressa.
Pavona expZanuZata was also common. Urchins observed were Echinometra
mathaei in interstitial spaces of the limestone pavement, Tripneustes
gratiZZa, and Echinothrix diadema.
Station E60
A flat limestone pavement mostly devoid of corals was the major topo
graphic feature at this transect. The only common species observed was
P. Zobata. Numerous Tripneustes gratilla, as numerous as 9 per quadrat were
observed, as were Echinometra mathaei, which were common in holes in the
substrata.