I. INTRODUCTION

A. Nonindigenous marine species introductions and their impacts on native populations.

The distributions of marine organisms throughout the world have been established with the 
influence of natural physical barriers such as landmasses, temperature or salinity gradients, or 
current systems, which isolated populations and promoted speciation through evolutionary time 
scales.  However, the geographic ranges of formerly separated species populations have always 
been in flux as changing current systems, shifting temperature environments or disappearance of 
land barriers with sea level rise have permitted range extensions of organisms through natural 
means.  What we often traditionally view as a relatively static marine ecosystem with defined 
boundaries dividing distinct populations has in fact been subject to continual transport, invasion, 
competition and changes in dominance at low rates of natural introduction. 

When natural means causing redistribution of species populations in the sea become superceded 
by man-related activities, dramatic changes can occur in the resident communities of the areas 
receiving the introductions.  As stated by Briggs (1974) dominant species can not only succeed 
in colonizing when they manage to migrate across barriers but often show spectacular success 
as the result of man-made introductions, either purposeful or accidental.  Human-mediated 
transport of nonindigenous species is not a new phenomenon.  For thousands of years vessels 
have transported boring organisms and fouling organisms (Carlton 1992; Carlton and Hodder 
1995), and circumnavigation of the world by sailing vessels has been underway since the early 
sixteenth century.  In fact, because classification and identification of marine organisms began 
only in the mid-eighteenth century, and few marine biological surveys were conducted prior to the 
mid-nineteenth century, the natural distributions of many widely distributed marine shallow water 
species remain in question (Carlton 1989).

In the last century, and more especially during the last two decades, man-related redistribution of 
marine shallow water organisms has become more frequent and ever more important in its 
impacts on native communities.  These introductions of nonindigenous species have been 
promoted though five principal vectors (Carlton 1987, Ruiz et al. 1997):

? Transport of planktonic or larval forms in ship ballast water or benthic forms in ballast water 
sediments in large cargo vessels.
? Transport of fouling organisms attached to vessel hulls or of nonattached organisms 
associated with fouling communities
? Intentional or unintentional release or propagation in the natural environment of edible or 
commercially important organisms, or of organisms that are associated with organisms 
brought in for culture purposes.
? Ornamental organisms such as exotic fishes or algae, originally intended for aquarium 
observation, teaching or research but escaping or released into the environment.
? Reconnection of formerly isolated bodies of water by canal construction, the most famous 
example being Lessepsian migration of organisms through the Suez Canal between the 
Red and Mediterranean Seas (Por 1978; Zibrowius 1991).

The primary vector responsible for recent invasions in coastal and inland waters is believed to be 
worldwide commercial and military shipping.  Ships may transport viable organisms within their 
ballast water or on their hulls as fouling organisms. If precautions are not taken, these potential 
invaders may be discharged in harbors where, with no natural predators of controls, they may 
proliferate rapidly. The role of ballast water and ship fouling as transport vectors of nonindigenous 
species has been demonstrated in many parts of the world (Carlton and Geller 1993;Rainer 1995; 
Chu et al. 1997; Smith et al. 1996).  Although these mechanisms for species transport are not 
new, aquatic species introductions may have accelerated in many areas of the world in the last 
decade due to more rapid ship movement and increased traffic to and from some areas such as 
China.

Since the 1970s, a striking surge of exotic species invasions has occurred in harbors, ports, and 
other coastal ecosystems around the world (See Carlton 1985; Carlton and Geller 1993 and Ruiz 
et al. 1997 for reviews).  Introduced species can rapidly monopolize energy resources, act as 
voracious predators, overcome endemic species, or transmit parasites and diseases that can be 
passed to humans through the food chain or direct exposure.  Because of the serious 
consequences that can result from nonindigenous introductions, marine species invasions have 
been ranked among the most serious potential perturbations of marine ecosystems (Carlton 
1994).  As an indication of the growing extent of the problem, more than 75 papers were 
presented at the January, 1999 First National Conference on Marine Bioinvasions on the 
patterns, transport vectors, impacts and control measures for nonindigenous marine species 
throughout the world.  Most introductions of nuisance marine nonindigenous species have 
occurred in the last two decades, such as the fouling mussel Perna perna along the Texas Gulf 
coast in the late 1980s (Hicks and Tennel 1993), the Japanese shore crab Hemigrapsus 
sanguineus on the US Atlantic  coast in 1988 (McDermott 1991, 1999; Brousseau et al, 1999, 
Larson et al. 1999; Seeley 1999; Tyrrell 1999), a variety of molluscs (Zolotarev 1996) and  the 
North American ctenophore Mnemiopsis leidyi (Shushkina et al. 1990) into the Black Sea in the 
early 1980s, the alga Codium fragile tomentosoides (Trowbridge 1996) and a variety of 
invertebrates (Hayward 1997) into New Zealand waters, and Japanese dinoflaggelates 
(Alexandrium spp.) macroalga (Undaria pinnatifida), starfish (Asterias amurensis) and the 
European green crab Carcinus maenus into Australian waters in the late 1980s (Sanderson 1980; 
Hallegroff and Bolch 1991; Buttermore et al. 1994; Rodriguez et al 1999).

San Francisco Bay, which has had a long history of direct commerce and exchange of shipping 
with Hawaii, has been especially impacted by marine species invasions.  Species introductions in 
San Francisco Bay have been so prolific and successful that no shallow water habitat in the Bay 
is free of exotic species, and it is difficult to find any abundant native species (Carlton 1979; 
Cohen and Carlton 1995).  Moreover, species invasions continue in San Francisco Bay (Cohen 
and Carlton 1997; Cohen 1999; Daehler and Strong 1996; Greenberg et al. 1996) which have the 
potential to be ecologically devastating.  The Chinese river clam, Potamocorbula amurensis, has 
reached densities as high as 10,000 per m2 in shallow areas of the bay, sufficient to strip the bay 
of phytoplankton blooms that form the basis of the food chain (Carlton et al. 1990) and alter food 
webs (Thompson and Luoma 1999).  The European green crab Carcinus maenus  consumes a 
wide variety of prey (Grosholz and Ruiz 1999) and is capable of greatly altering the San 
Francisco Bay ecosystem through predatory consumption, competition and indirect effects such 
as hybridization.  This crab has recently extended its range to Oregon coast (Behrens Yamada et 
al. 1999) and Puget Sound (Cohen 1999).  The estuarine Chinese mitten crab Eriocheir sinensis, 
harbors a human parasite, damages levees and shorelines and interferes with fisheries (Cohen 
and Carlton 1998).  Cohen and Carlton (1995, 1998) have estimated that a new invasion occurs 
in San Francisco Bay an average of every 10 to 12 weeks, and new invasions continue to be 
documented (Cohen et al. 1995; Gosliner 1995; Mills and Sommer 1995; Cohen and Carlton 
1998).

B. Marine Nonindigenous Species Introductions in Hawaii

The main Hawaiian Islands are the most isolated land area in the world, lying more than 4300 km 
from the North America and more than 6400 km from Japan, and the native biota of these islands 
have one of the highest rates of endemism in the world (Kay and Palumbi 1987). The Hawaiian 
islands have been a principal port of call for ocean-going ships sailing from San Francisco Bay 
and elsewhere in the Pacific for more than 150 years.  Located at the crossroads of the Pacific, 
Hawaii receives ship traffic from all oceans, principally from the west coast of North America, 
Asia and the South Pacific (Carlton 1987).  Virtually all international shipping passes through 
Honolulu Harbor and Barbers Point Deep Draft Harbor on Oahu, making these the most likely 
entry points for introduced marine species into Hawaii.

Only recently has information become available concerning the abundance of nonindigenous 
species in Hawaiian waters.  Maciolek (1984) listed 19 species of diadromous and marine fishes 
to be present in Hawaiian waters, which was increased to 21 marine species by Randall (1987), 
about 4% of a total of 536 Hawaiian shore fish species (Randall 1992).  Carlton and Eldredge (in 
prep.) reviewed the marine and brackish water invertebrates of Hawaii and determined 
approximately 205 species to be demonstrably or potentially nonindigenous, again about 4% of 
the approximate 5000 marine species estimated for Hawaii (Allison et al. 1995).  Approximately 
18 species of macroalgae have been introduced to Hawaii since 1950 (Russell 1992, Rogers 
1997, 1999), again about 4% of the approximately 430 estimated total macroalgal species for 
Hawaii (G. Smith, pers. comm.).

Baseline studies of Hawaiian nearshore marine biota directed toward the detection of introduced 
species and their impact have shown that nonindigenous introductions vary substantially from 
these average values, depending on the characteristics of the area surveyed.  The most 
comprehensive survey conducted to date, a 1996 survey completed in Pearl Harbor (Coles et al. 
1997; 1999), found 95 known or potentially nonindigenous species, which composed 23% of the 
419 invertebrates, macroalgae and fishes found.  Only three nonindigenous invertebrates and 
one nonidigenous fish occurred at Midway Atoll in 1997 out of a total 444 taxa found (DeFelice et 
al. 1998).  No nonindigenous algae or invertebrates were found in the nearshore and intertidal of 
Kahoolawe Island in 1998 out of a total of 298 species observed or collected (Coles et al. 1998).

Despite the potential importance of Honolulu Harbor or other commercial harbors on Oahu as 
potential gateways for nonindigenous marine species to enter the Hawaiian marine ecosystem, 
no studies have previously been conducted of introduced species in Hawaiian commercial 
harbors, and little information is available for the composition of the marine communities for these 
harbors.  Therefore, we conducted surveys in Honolulu Harbor and Keehi Lagoon in 1997, and in 
Kewalo Basin, the Ala Wai Yacht Harbor and the Barbers Point Deep Draft Harbor in 1998.  The 
results of this study are reported herein and the presence and impact of nonindigenous marine 
introductions in these harbors are evaluated. Studies of nonindigenous introductions in stream 
mouths along Oahus south and west shores and a evaluation of the role of hull fouling, ballast 
water and sediments as vectors for marine introductions were conducted as part of this overall 
project and will be reported in separate reports. 

II. HONOLULU HARBOR AND OTHER SOUTH SHORE HARBORS 

A. Historical Perspective

Although the first contacts between Hawaiians and Europeans occurred on the islands of Hawaii 
and Kauai, Honolulu has played a major role in the history of Hawaii since the Honolulu Harbor 
was first located by British and American ships in the late 18th century (Appendix A).  The first 
European to see Honolulu Harbor may have been a crewman in 1786 from the HMS King 
George, captained by Nathaniel Portlock, which was the third ship to visit the Islands and the first 
to reach Oahu.  However, European discovery of the harbor in 1794 is credited to Captain Brown 
of the British schooner Jackal and Captain Gordon of the sloop tender Prince Lee Boo, later 
joined by the American ship Lady Washington commanded by Captain John Kendrick (Cartwright 
1923).

The harbor was first surveyed by W. R. Broughton of the HMS Providence in 1796, although this 
earliest chart was lost.  Although it was difficult for sailing ships to enter the harbor against the 
usually prevailing northeast tradewinds, mariners soon recognized the value of the harbor as a 
calm berthing spot with relatively deep water, conditions rare in the Hawaiian Islands where few 
natural harbors occur.  With the increased shipping brought to the islands by sandalwood trade 
and whaling in the early nineteenth century, the population and power center of Hawaii shifted to 
the formerly small village on the dusty plain of Honolulu.  It is no exaggeration that Honolulu 
Harbor is the primary factor for Honolulu being the present capital and commerce center of 
Hawaii, since the harbor has historically provided the principal access for goods transported by 
sea to and from the Hawaiian Islands.

As an indication of the increasing importance that Honolulu and Oahu were to play in his recently 
established Kingdom of the Hawaiian Islands, King Kamehameha moved his court to Oahu from 
Kailua-Kona in 1804, establishing one of his three residences along the harbor near the mouth of 
Nuuanu Stream.  It was near this site in 1816 that a fort was built by the Russian trader Georg 
Scheffer and then refurbished by the Hawaiians to become Honolulu Fort, which dominated the 
harbor shoreline until the fort was dismantled in 1856-57.  The fort stood at the intersection of the 
present Queen Street and Fort Street Mall, and its seaward walls were along the shoreline of that 
period.

Between 1816 and 1819 three maps were prepared by European visitors which give a good idea 
of the configuration of Honolulu Harbor and the earliest development of the waterfront of the 
village that would become Hawaiis capital city.  The charts show the dimensions of the harbor in 
its natural state, formed from the outflow of Nuuanu Stream through the coral reef, before the 
harbor had been altered by dredging and filling.  The earliest map (Figure 1), drawn in 1816 
under the direction of the Russian commander Otto von Kotzbue, is the most detailed of the three 
in terms of shoreline landmarks, and provides the first data available for depths (missing on the 
English translation version) in the harbor and at its entrance.  It shows the newly built fort, 
fishponds on the Iwilei side of Nuuanu Stream and southeast of the harbor entrance, which is 
Figure 1. English translation of von Kotzebues 1816 chart of Honolulu Harbor.
restricted on both sides by a broad coral reef.  The extent of the village of Honolulu is indicated by 
dwellings along the shoreline north and south of the fort.

Two arrivals to Honolulu Harbor that would significantly affect the history and economy of Hawaii 
were the arrival of two first whaling ships in 1819 and the first company of Christian missionaries 
on the brig Thaddeus in 1820.  A few years after, the first pier in the harbor was improvised in 
1825 by sinking a ships hull near the present Pier 12 site.  During this same year the first harbor 
regulations were published and the fourth existing map of the harbor prepared by C. R. Malden, 
an officer of the HMS Blonde under the command of Lord Byron. (Figure 2).  This rendering 
shows great detail for depth soundings, which indicate the narrowness and relative shallowness 
of the harbor entrance, and the configuration of both the coral reef and substantial intertidal areas 
that became exposed at low tide.  Honolulu had grown from a village to a small town, with the 
beginnings of a grid work of streets.  Two years later, in 1827, two shipwrecked sailors, James 
Robinson and Robert Lawrence formed Robinson and Company and built Honolulus first 
combined wharf and shipyard in the vicinity of the Honolulu Fort.

By 1840, when the fifth existing map of Honolulu Harbor was prepared by the U. S. Exploring 
Expedition under C. R. Wilkes (Figure 3), Honolulu had grown to have a network of streets 
connecting the residences, commercial, and public buildings that made up a town increasingly 
dependent on goods and income derived from ships entering and leaving Honolulu Harbor.  The 
map provides the greatest detail to date for harbor depths and indicates that fishponds formerly 
located southeast of the harbor entrance were no longer present.

By the 1850s development of Honolulu and harbor use had increased to a point where it was 
apparent that alterations of the harbor would be necessary to continue increases in shipping and 
commercial uses.  In 1852 the first steam powered ship, the schooner Constitution, arrived in the 
harbor from San Francisco.  A 940-foot long wall was constructed in the harbor in 1853 to divert 
sediments entering from Nuuanu Stream toward Kalihi Basin.  The seriousness of sedimentation 
that was occurring at that time into the harbor was suggested by a newspaper report that water 
depth in the harbor was decreasing at a rate of 0.6 m (2 ft) per year. In 1854 the first steam tug 
was used to pull sail powered ships into dock against the prevailing tradewinds, a task originally 
performed by Hawaiians in outrigger canoes and later by men or ox teams pulling ropes along the 
reef or intertidal area south of the harbor entrance.

In 1856-57 the most prominent landmark on the Honolulu shoreline, the Honolulu Fort was torn 
down and its materials were used to extend the shoreline seaward of the former fort site, in the 
area of the present Aloha Tower complex.  Dredging was also conducted along Robinsons Pier 
and the Market and Custom House piers in this area, which provided additional fill material as 
well as dock access for ships with deeper drafts. This new area was called the Esplanade, and it 
marked the beginning of extensive dredging and filling activities that would drastically alter the 
size and configuration of the harbor over the next century.  The Esplanade was later expanded in 
1889 by extending the sea wall and filling with dredge material to support the construction of 
wharves and covered storage areas.


Figure 2. Map of Honolulu Harbor prepared by C. R. Malden in 1825.


Figure 3. Map of Honolulu Harbor prepared in 1840 by the U. S. Exploring Expedition under C. R. 
Wilkes.

Honolulu Harbors first lighthouse was first put in use in 1869.  It was located on the north side of 
the entrance channel and was accessible by foot on a boardwalk from Immigration Island, the 
forerunner of Sand Island, which was originally an intertidal area awash at high tide. Mail service 
by steamship to Honolulu began in 1875 and the first marine railway, located at the Pier 2 
location, began service in 1883.

Major alteration of Honolulu from its natural configuration began in 1890 with the dredging of the 
main channel to 60 m (200 ft) width by 9 m (30 ft) deep for about 303 m (1000 ft) through the 
sand bar at the entrance, which formerly restricted ships to those having a draft of less than about 
6 m (20 ft).  Dredging required two years to complete, and the first large vessel to enter the 
harbor though the new entrance channel was the Oceanic, in 1893.  Many dredging and filling 
operations soon followed, and the 1890s and 1900s saw the construction of many new piers and 
channels in the harbor, the dredged material going to create new dry land areas.  In 1898 docking 
areas in the harbor were dredged to provide access to wharves for ships up to 182 m (600 ft) long 
in the vicinity of Nuuanu stream.  The entrance channel was further widened to 121 m (400 ft) 
and deepened to 10.6 m (35 ft) in 1905 and the main harbor enlarged to 364 m (1200 ft) wide in 
1905, and 158,000 cubic yds. of dredged material were deposited on Immigration (Sand) Island in 
1906.  Further dredging was conducted at the base of Alakea Street in 1906 and the dredge 
material used as fill material in the Pier 1 area around Immigration Island.  Piers were constructed 
at the base of Richards Street in 1896, at the site of Piers 17 and 18 in 1901 to accommodate 
sugar loading in 1901, and at Piers 7 and 12 in 1907.  Along with this increased activity in 
Honolulu Harbor during this period, Oahus second harbor was constructed at the site of the Ala 
Wai Yacht Harbor during the 1900s.

By 1900 (Figure 4) it was apparent that Honolulu Harbor was becoming insufficient to 
accommodate the shipping activities of the growing city of Honolulu, and a channel was proposed 
to be dredged form the main harbor to Kalihi.  Dredging commenced in 1915 and was completed 
in 1920, connecting the original harbor and the area that would be Kapalama Basin with a 303 m 
(1000 ft) long by 10.6 m (35 ft) deep and 242 m (800 ft) wide channel.  This allowed the 
construction of additional piers, and the first fuel oil line for oil powered vessels was constructed 
at Pier 16 in 1920.  This year also saw the completion of Hawaiian Electrics Honolulu Generating 
Station at Pier 7.  This facility was the primary source the electricity for the city for the next twenty 
years and is still used today during peak load periods.  At maximum operation it circulated 
200,000 gpm of cooling water from its intake to its discharge basin, increasing the water 
temperature 5-6?C and providing one of the principal sources of water circulation in the harbor.

In 1921 Oahus third harbor was constructed at Kewalo Basin, primarily to provide berthing for 
fishing boats in clean water, to prevent contamination of cleaning fish catches in the polluted 
runoff of Nuuanu Stream.  At about the same time, the Ala Wai Canal was dredged in 1921-28 to 
drain Waikiki marshes.  The original mouth of the canal was not at its present location near the 
Ala Wai Harbor, but was into a channel dredged along the shore to join the recently completed 
Kewalo Basin channel.


Figure 4. Map of Honolulu Harbor around 1900 showing dredged entrance channel, lighthouse, 
marine railway, the Esplanade and new piers.
The 1920s and 1930s were a time of rapid harbor expansion and alteration (Figures 5 and 6).  
The Kapalama channel was widened to 242 m (800 ft) and deepened to 10.6 m (35 ft) in 1922-27 
and further widened in 1931-35 when the entrance channel was dredged to a depth of 12 m (40 
ft) and widened to 152 m (500 ft).  The Aloha Tower and Piers 8 to 11 were completed in 1926 on 
the site of the Esplande, ushering in a new era of tourism that would dominate Hawaiis future 
economy.  Pier construction continued in the enlarged Honolulu Basin as well as in the Kapalama 
Channel.  By 1940 the total harbor area had at least doubled in size from its natural state and had 
a total of 32 piers (Figure).  With the onset of World War II, administration of the harbor was 
assumed by the U. S. Navy, and a massive expansion of the harbor and Keehi Lagoon was 
begun.  Between 1941 and 1945 the Kapalama Channel was widened another 303 m (1000 ft) 
and the Kapalama Basin was dredged to of 10.6 m (35 ft) deep, 303 m (1000 ft) wide by 1030 m 
(3400 ft) long.  A second entrance channel, originally proposed in 1900 to provide two entrances 
to an enlarged harbor, was scheduled to be dredged, but the project was considered 
unnecessary by the end of the war and abandoned.

Considerable dredging was also conducted outside of Honolulu Harbor during this period.  The 
Kewalo Basin and channel were deepened to 4.8-5.2 m (16-17 ft), and three seaplane runways 
and a mooring basin were dredged in Keehi lagoon with more than 16 million cu. yds. of dredged 
material deposited on the Keehi Lagoon shoreline to form the area of the Honolulu airport.

In 1946 a liberty ship, the SS Britain Victory, grounded on the entrance channel reef, blocking the 
harbor for one week.  This renewed interest in opening a second entrance channel to the harbor 
at the west end of Kapalama Basin, which was recommended for construction in 1949 by the 
chief of the U. S. Army Corps of Engineers.  However this project was not completed until 1962 
when the Kalihi entrance channel was opened.  Road access to Sand Island was maintained by 
the use of a Bascule drawbridge, which could be opened to allow ships to pass through the Kalihi 
Channel.  Opening of the Kalihi Channel constituted the last major alteration of Honolulu Harbor, 
which appears today much the same as it did in 1968 (Figure 7).

Keehi Lagoon however, was substantially altered during the 1960s and 1970s. The Keehi Lagoon 
Marina was completed in 1963, and the Honolulu Airport Reef Runway was constructed from 
1972 to 1975.  This involved the filling of an area 666 m (2200 ft) by 3,636 m (12,000 ft) on former 
reef, burying an area of 1,240 acres off shore of the airport outside of Keehi lagoon.  Within the 
lagoon, the project also involved dredging of channels that increased circulation, alleviating the 
stagnant water conditions that had been created by dredging the seaplane runways in the 1940s.  
Elsewhere, the Ala Wai Yacht harbor was enlarged in the early 1970s to accommodate two 
additional basins and berths for more than 350 additional boats.

During the 1980s activity in Honolulu Harbor included dredging of the main entrance channel to 
13.6 m (45 ft) depth and dredging of the main basin, Kapalama Channel and Kapalama Basin to 
12.1 m (40 ft).  Piers 16, 17 and 18 were reconstructed or modified to provide moorings for 
commercial fishing boats in the same area from where they had been removed to Kewalo Basin 


Figure 5. Map of Honolulu Harbor in 1928 showing widened channel entrance and main basin, channel dredged to Kapalama Basin, proposed 
Kalihi Channel and fill areas on Sand Island and on the reef east of the harbor entrance.


Figure 6. Map of Honolulu Harbor in 1940.


Figure 7.  Honolulu Harbor and Keehi Lagoon in 1968.

in the 1920s.  However, the main harbor activity during this period was the creation of a deep 
draft harbor at Barbers Point on Leeward Oahu in 1982-85. This harbor was constructed to divert 
large ship cargo, especially those carrying flammable or explosive cargo, away from the highly 
populated areas around Honolulu Harbor.

The most recent phase of development of Honolulu Harbor was the conversion in 1990-1993 of 
the Aloha Tower complex from its former use as Harbormasters offices to a shopping and visitors 
center.  This has included the moving of the historic four-masted Falls of Clyde museum ship to 
the Pier 7-8 basin, the opening of the Maritime Museum on Pier 7, and the placing of numerous 
restaurants and shops along the historic waterfront of the former Esplanade.  The juxtaposition of 
such facilities adjacent to the docks and wharves of a working harbor has provided an interesting 
opportunity for visitors to view the vital role that Honolulu Harbor continues to play in the lives of 
Oahu residents.

B. Environmental Settings

1. Honolulu Harbor

Honolulu Harbor originally was a deep embayment formed by the outflow of Nuuanu Stream 
creating an opening in the shallow coral reef that lies along the south shore of Oahu.  It was first 
described scientifically by Agassiz (1889) as nothing but a channel kept open by the flow of the 
Nuuanu River, whichhas killed the corals in its path, scouring at the same in freshets the whole 
harbor and the adjacent limestone forming the channel.  The river forming the Honolulu Harbor 
brings down a large amount of volcanic mud in its short course, and has deposited this in the 
harbor and channel, so that there appears to be nothing but dark volcanic mud for a considerable 
distance out towards the entrance to the channel, where the coral limestone reappears.

In its natural state the harbor consisted only of this river-formed main basin, which was only 6 m 
deep at its entrance.  Its perimeter was enclosed by shallow reef and intertidal areas that were 
exposed at low tide. A small white sand beach extended along the eastern shoreline from the 
present Aloha Tower complex to the Pier 1 area.  The reef extended across the present 
Kapalama Channel continuous with the area that is now Sand Island.  Formerly this was a much 
smaller island (Immigration Island) surrounded by a large shallow reef flat  (Figures 4 and 5).

Honolulu Harbor now consists of a main basin which has been substantially enlarged and 
deepened from the original natural embayment, Kapalama Channel, which was first dredged 
through the reef west of the main basin in 1915-20, and Kapalama Basin, first dredged to 10.6 m 
(35 ft) depth in 1941-45 (Figure 8).  The harbor receives the runoff of two major fresh water 
sources, Nuuanu Stream at the head of the original harbor between Piers 15 and 16, and 
Kapalama Canal which empties into Kapalama Basin between Piers 38 and 39. The harbor 
originally had only one opening to the sea until the Kalihi Channel was completed in 1962, and 
the presence of this channel at the west end of the harbor has undoubtedly increased circulation 
and  water  quality.   Limited  salinity  data (Oceanit 1990)  suggests that surface salinities can be


Figure 8. Current map of Honolulu Harbor and Keehi Lagoon showing Pier Locations.

reduced in the harbor by freshwate runoff by as much as one third, but subsurface salinities remain at an 
oceanic 35 o/oo.  Overall average salinities in the harbor average 34 o/oo (Buske and McCain 1972).

The present harbor ranges in depth down to 13.5 m (45 ft), maintained by periodic dredging.  Very little 
natural substrata remain in the harbor.  Extensive modifications by dredging and filling have greatly 
enlarged the deeper areas of the harbor and reduced the reef flats that enclosed the original main basin.  
More than 50 piers compose most of the shoreline throughout the harbor, and the original entrance 
channel is lined and reinforced with large basalt boulders.  Natural coral reef substratum occurs only in 
two places in the harbor, between Piers 29 and 30 on the landward side of Kapalama Channel and on 
both sides of Kalihi channel between Snug Harbor and the Bascule bridge.  Elsewhere the benthic 
substratum above the silt or sand bottom is composed of concrete abutments or pilings supporting docks 
and piers, many of which jut out 10-25 m (33-82 ft) from the dredged shoreline.  The bottom of most of 
the harbor is composed primarily of flocculent loose silt, which becomes finer as the mouths of Nuuanu 
Stream and Kapalama Canal are approached.  However, with approach to the harbor entrance at Piers 1 
and 2 the bottom sediments become fine, white calcareous sand, as described by Agassiz (1989) over a 
century ago.

Honolulu Harbor (Figure 8) remains the primary shipping port for commercial goods entering Honolulu or 
being trans-shipped to the neighbor islands, and port activity is dominated by container ships unloading at 
the Matson and Sealand Terminals at Pier 52 on Sand Island.  Just eastward Pier 53 provides berthing 
for U.S. Coast Guard ships and a cable repair ship, and the University of Hawaii berths its fleet of 
research vessels at Snug Harbor, near the Kalihi Channel entrance.  Pier 2 is the foreign trade zone 
docking area, and cruise ships that transport thousand of passengers utilize Piers 10 and 11.  
Commercial fishing boats moor at Piers 16-18, and Piers 19-27 are berths for harbor and inter-island 
tugs.  The Clean Islands Council oil spill emergency response vessels dock at Pier 35, Young Brothers 
interisland tugs and barges utilize Piers 38-40, and a floating dry dock is in place at Pier 41.  Although 
wastes from the pineapple canneries were originally discharged into Kapalama Canal until the early 
1970s, resulting some of the highest bacterial concentrations measured in the state waters at that time 
(Cox and Gordon 1970), the only significant industrial use of harbor water at the present time is for once-
through cooling of the Hawaiian Electric Generating Station.  This facility has, in the past, raised the 
temperature of up to 200,000 gpm cooling water 5-6?C circulating from its intake by Pier 7 to its discharge 
at Pier 5, but usage has decreased in recent decades as generation load has been shift to more efficient 
newer power stations.

2. Keehi Lagoon

Keehi Lagoon (Figure 8) was originally a large shallow reef and subtidal area no more than 1-2 m deep 
that extended more than two miles off the mouths of Kalihi and Moanalua Streams.  Its present eastern 
boundary is formed by Kalihi Channel, which was originally a shallow channel across the reef through 
which the combined outflow of Kalihi and Moanalua Streams reached the sea.  Much of the present land 
for Honolulu International Airport was originally reef, Keehi Lagoon shoreline, ponds or marshes.

Dredge and fill activities in the 1940s and the 1970s drastically altered Keehi Lagoon from its original 
state.  A mooring basin and three seaplane runways two to three miles long by 30.3 m (100 ft) wide and 3 
m (10 ft) deep were dredged in the lagoon in 1941-45 and the dredged material placed along the shore.  
Because these channels essentially trapped water that otherwise would have moved on and off shore 
with tidal exchange and wide movement, stagnant conditions and lowered water quality resulted, retaining 
pollutants in the deeper water in the runways.

Further alteration of the lagoon resulted from the construction of the Honolulu International Airport Reef 
Runway, constructed in 1972-75.  This effectively divided the lagoon into an eastern portion extending 
from the western end of the runway to the Kalihi channel entrance, and a western portion adjoining the 
Hickam small boat harbor.  In the process of constructing the runway, some 1,240 acres of former reef 
and shallow flats were buried under 2.7 m (9 ft) of fill material.  Also, to increase circulation and provide 
boat access, channels were dredged around the eastern end of the runway to the seaplane runways and 
to Hickam Harbor.  Monitoring conducted prior to and following completion of the runway construction 
indicated a substantial improvement in water quality due to the increased circulation provided by these 
channels (Environmental Consultants 1977, 1979; OI Consultants 1986; Noda & Assoc. 1978; Chapman, 
1979).

The eastern portion of Keehi Lagoon sampled in this study consists of a shallow reef flat enclosed by the 
three seaplane runways, the Kahili Entrance Channel to Honolulu Harbor and the access channel east of 
the reef runway that was dredged in 1971-75.  The lagoon receives the combined drainage of Kalihi and 
Moanalua Streams on its north apex, which is completely lined with a dense growth of red mangrove 
(Rhizophora mangle).  A series of small islands line the northeast-southwest directed seaplane runway 
and more are forming on the central reef flat where mangroves have begun to grow and accumulate 
sediments.

3. Kewalo Basin

Kewalo Basin (Figure 9) is located between Honolulu Harbor and Ala Moana Park on Oahus south 
shoreline.  It was dredged from coral reef along the shoreline in the 1920s to provide mooring for fishing 
boats away from the pollution of Nuuanu Stream entering Honolulu Harbor.  The basin is approximately 
242 m (800 ft) wide by 303 m (1000 ft) long and 6.1 m (20 ft) deep, with a 61 m (200 ft) wide entrance 
channel at its southwest side.  The basin entrance is highly exposed to south swell waves, and popular 
surf sites exist both east (Shark Hole) and west (Point Panic) of the entrance.  No freshwater streams 
or industrial discharges enter the basin, although at one time it did receive wastes from a tuna cannery 
that is no longer in operation.  However, the bottom of the basin is highly littered with chunks of concrete, 
coral, bottles, tires and other debris that has been discarded from moored boats through the years.

A total of more than 120 berths accommodate boats of various sizes and uses in Kewalo Basin, including 
cruise, charter fishing and research vessels.  The largest of these are commercial tour boats, which take 
on passengers daily at a dock on the basins northwest side for trips to Waikiki, Honolulu Harbor and 
Pearl Harbor.  Adjacent to this dock is Honolulu Marine ship repair. Other smaller commercial tour boats 

Figure 9. Location maps for Kewalo Basin, Ala Wai Yacht Harbor and Barbers Deep Draft Harbor
are berthed along the northeast (Ala Moana Boulevard) side of the basin.  Two research facilities are 
located at Kewalo Basin, the University of Hawaii Pacific Biological Research Center located along the 
west of the entrance, and the Kewalo Basin Marine Mammal Laboratory near the basins southeast 
corner.

4. Ala Wai Yacht Harbor

The Ala Wai Yacht Harbor (Figure 9), located on the south shore of Oahu between Ala Moana Park and 
Waikiki Beach, was first constructed from a barge channel in the 1900s and has been periodically 
enlarged to where it presently provides more than 1000 berths for pleasure craft.  It is surrounded by 
resort and commercial properties, the largest being the Ilikai and Prince Hotels.  A small man-made salt-
water pond (Hilton Lagoon) adjacent to the harbor discharges into its southeast corner.  Along with 
public berthing of power and sailing boats, the harbor accommodates two yacht clubs, the larger Waikiki 
Yacht Club located on the northeast side of the harbor, and the Honolulu Yacht Club located at the end of 
the central groin.

The Ala Wai Canal discharges into the northwest corner of the harbor near the Waikiki Yacht Club docks 
and is a source of reduced salinity and increased sedimentation, turbidity and pollution from canal runoff.  
The canal, which was dredged in 1921-1928, provides an outlet to the sea for Palolo and Manoa 
Streams, which formerly drained into the Waikiki marshes that were eliminated by construction of the 
canal.  However, the discharge of the original canal did not empty into the Yacht Harbor but flowed into a 
channel dredged along the present Ala Moana shoreline and joined the Kewalo Basin entrance channel.

5. Barbers Point Deep Draft Harbor

Barbers Point Deep Draft Harbor (Figure 9) is the newest harbor on Oahu and is located near Oahus 
industrial park at the southwest end of Oahu.  It was dredged entirely from coral rock above the shoreline 
and receives no stream inflow.  A smaller version of the harbor was enlarged in 1982-1985 to form its 
present configuration of about 121m (700 ft) by 182 m (600 ft) with an entrance channel 85 m (280 ft) 
wide.  Design depth of the harbor is 13.6 m (45 ft) to accommodate the largest vessels arriving in 
Honolulu.  The harbor is the Oahu offloading point for large bulk carriers and oil tankers, and one docking 
area along the south pier is used exclusively for offloading coal which is carried from the pier by a 
conveyor belt system for use in power generation.  A large floating dry dock is moored along the harbors 
southwest pier, and a shallow channel on the northwest side connects the harbor with a private marina 
controlled by the Ko Olina resort area.

C. Previous Biological Studies

1. Honolulu Harbor

Despite its early discovery and the importance that it has played in Hawaiis history and economy since 
European contact, biological sampling has been sporadic in Honolulu Harbor and little was known about 
its marine biota prior to the present study.  Only 205 records of marine invertebrates collected from 
Honolulu harbor are listed in Bishop Museums collection catalog, the earliest a being a decapod, Saron 
marmoratus, collected by W. A. Bryan in 1915.  Other than 38 decapods collected by T. Dranga in the 
early 1920s, most of the collections were made Dr. C. H. Edmondson between 1940 and 1956.

The first systematic surveys of the biota of Honolulu Harbor were conducted by staff and contractors of 
the Environmental Department of the Hawaiian Electric Co. during the early 1970s (McCain and Peck 
1972; McCain and Coles 1973; McCain et al. 1975; Environmental Consultants and Hawaiian Electric 
1974).  These studies were made in the area of Piers 5 to 7, in the vicinity of Hawaiian Electrics Honolulu 
Power Station intake and discharge, and focused on plankton, reef corals and fish.  Later studies made in 
the 1990s in the same area (Brock 1991, 1992, 1993, 1994, 1995, 1997) have described benthic 
invertebrates and fish encountered on annual monitoring surveys in this area.

With the exception of a masters thesis conducted on the benthic community associated with the 
introduced octocoral Carijoa (=Telesto) riisei (Thomas 1979), all remaining biological information for 
marine organisms in Honolulu Harbor has been derived from project related environmental baseline or 
impact assessment surveys.  These have been conducted in the vicinity of Sand island (U. S. Army Corps 
of Engineers 1978), the commercial fishing vessel berthing area at Pier 16 (AECOS 1982), Piers 12 to 15 
(AECOS 1982), Pier 1 (AECOS 1988), and the waterfront at Aloha Tower (Oceanit 1990).  All of these 
surveys were based on field observations with little or no laboratory analysis of sampled organisms, and 
therefore were likely to report only the most abundant and prominent of the organisms present.

2. Keehi Lagoon

All biological reports for Keehi Lagoon are from environmental baseline or impact assessment surveys, 
and most these are from studies associated with the construction of the Honolulu Airport Reef Runway.  
The earliest available biological information for the lagoon comes from a report by Harvey (1970), which 
listed an assortment of macroalgae and micromolluscs found in the sediments throughout the lagoon.  An 
environmental impact assessment prepared in 1975 for disposal of solid waste bales in Keehi Lagoon 
(Parsons 1975) listed a number of invertebrates by their common name but did not identify them by 
genus or species.  Post construction monitoring surveys for the reef runway (Environmental Consultants 
Inc. 1977, 1978, 1979) were more specific and provide information on the composition of the benthic and 
fish biota in the lagoon, along with a ten-year follow up survey conducted in 1988 (Guinther 1988).  A brief 
survey in the mangrove areas at the mouths of Moanalua and Kalihi Steams (Environmental Consultants 
Inc. 1978) provided limited information about biota in this sector of the lagoon.  Like past studies in 
Honolulu Harbor, the previous surveys in Keehi Lagoon were field studies and reported only the most 
obvious organisms.

3. Kewalo Basin, Ala Wai Yacht Harbor and Barbers Point Deep Draft Harbor.

Of the three reports available for Kewalo Basin (Appendix B) only one (Harbors Division 1984) mentions 
biological conditions, stating that no coral was found in the basin channel and that the bottom in the basin 
ranged from soft silt and clay sediments to coarse shells, sand and coral blocks. Because of extensive 
environmental studies conducted in the 1970s (Gonzalez 1971; Shultz 1971; Harris 1972; Luoma 1974; 
Miller 1975) and the 1990s (Laws et al. 1993; Beach et al. 1995; Glenn and McMurtry 1995;Glenn et al 
1995; McMurtry et al. 1995; Raine et al. 1995; Resig et al. 1995; Spencer et al. 1995) much more 
information is available for the Ala Wai Canal, but these studies were concerned with conditions above 
the Ala Moan Boulevard bridge, and no information has existed for the biota in the Yacht Harbor itself.  
The only reference to biological conditions in the harbor is a brief mention of fishes, using popular names, 
as part of a 1974 draft environmental statement for harbor expansion (Department of the Army 1974).

Extensive biological surveys have been conducted on the reef areas off the entrance of the Barbers Point 
Deep Draft Harbor (Environmental Consultants 1975; Bienfang and Brock 1980; OI Consultants 1990).  
However, the only information related to biological conditions inside the harbor can be derived form two 
stations located in the small barge harbor that was expanded in 1978-1980 to the present deep draft 
harbor (Environmental Consultants 1975). 

III. METHODS

A. Literature Search

A variety of sources of information on the environmental conditions and biological communities of the 
harbors on the south shore of Oahu were examined.  Literature consulted included published papers in 
the open scientific literature, taxonomy-based monographs and books, unpublished reports for 
environmental studies in the harbors, and newspaper and magazine articles that were concerned with the 
development or environmental and biological communities of the harbors.  Resources that were consulted 
in this search were the libraries of Bishop Museum, the University of Hawaii, and the Pacific Maritime 
Center.  Environmental reports and Environmental Impact Statements and Assessments were reviewed 
from the University of Hawaii Environmental Center, the Hawaiian Electric Co. Environmental Department 
and AECOS Inc.  An annotated bibliography of all the literature assembled is presented in Appendix B.

B. Bernice P. Bishop Museum Collections

Bishop Museum collections databases for algae, invertebrates, malacology and ichthyology were 
reviewed for all marine or estuarine organisms indicated to have been collected in Honolulu Harbor, 
Keehi Lagoon, Kewalo Basin, the Ala Wai Yacht Harbor or the Barbers Point Deep Draft Harbor. The 
retrieved data were assembled into a combined database for Oahu south shore harbors (other than Pearl 
Harbor) containing taxa identity, taxonomic authority, collection location and date, collector and collectors 
notes, when available.  This information is included with the general listing of all taxa for the study 
developed from all sources and presented in Appendix C.

C. Field Surveys

Benthic fouling and sediment biota were sampled and observations of fishes were made at 15 stations in 
Honolulu Harbor and five stations in Keehi Lagoon in 1997, and four stations in Kewalo Basin, five 
stations in Ala Wai Yacht Harbor and three stations in Barbers Point Deep Draft Harbor in 1998.  Station 
locations, coordinates and dates of sampling are given in Table 1, and station locations are shown in 
Figures 10-12.

Table 1. Station locations and sampling dates for Oahu south and west shore harbors. Longitude and 
latitude coodinates are in WGS84 datum.

Figure 10. Locations of sampling stations in Honolulu Harbor and Keehi Lagoon.
Figure 11. Locations of sampling stations in Kewalo Basin and Ala Wai Yacht Harbor.
Figure 12. Locations of sampling stations at Barbers Point Deep Draft Harbor.

The sampling and analysis process is summarized in Figure 13. Collections and observations were made 
by two experienced investigators sampling as large a variety of habitats as possible at each station while 
snorkeling or using Scuba.  One diver sampled fouling organisms growing on hard surfaces from the 
intertidal zone to the bottom by scraping three samples of approximately 0.1 m2 each.  The other diver 
observed fishes swimming in the area, recorded their identities and also noted the presence of abundant 
invertebrate megafauna and macroalgae.  Collected organisms, which range 4-8 liters in total volume for 
each station were inspected on site and selected hydroids and tunicates were removed to be relaxed in a 
solution of Epsom salts and seawater before preserving in 5% formalin.  The remaining organisms were  
preserved on site in 70% alcohol before returning the samples to the laboratory for sorting and 
identification of organisms.  

Sediment-dwelling organisms and their substratum were collected by inserting a 12.5 cm diameter 
cylinder 15 cm into the sediment, closing off the bottom and top with lids and then transporting the sample 
to the laboratory where it was sieved through a 0.5 mm mesh size screen.  A subsample of 10 to 25 cm3 
was retained from each sample for determination of micromollusc populations.

Figure 13. Summary of field and laboratory methods. 


Specimens collected were sorted and identified to species or the lowest practicable taxa, using 
dissecting or compound microscope magnification when necessary.  Identifications were made 
using descriptions available in Reef and Shore Fauna of Hawaii Sections 1 to 4 (published), 5 
and 6 (unpublished), various taxonomic references, and voucher specimens in the Bishop 
Museum collections.  Specimens from various groups were sent to taxonomic experts for 
verification of preliminary identifications (see Acknowledgments).

D. Data Analysis

All organisms identified from the field study were entered on an Access database relational with 
the databases for previous literature reports and museum collections of organisms from Pearl 
Harbor.  The combined information was used to track the occurrence of species chronologically 
as they were reported from Pearl Harbor.

The Sorensons Index of percent similarity, based on presence-absence of species at station 
pairs, was used to measure the degree of association between stations.  By this index, the more 
species two stations share relative to their total species complements, the greater their ecological 
similarity.  Based on a matrix of Sorensen Index values, cluster analysis was used to arrange 
stations into groups or clusters.  Intercluster distances were calculated using an unweighted pair 
group average method.  In this analysis, similar stations will form clusters distinct from other 
stations.  These clusters are arranged in a hierarchical, treelike structure called a dendrogram 
(see Figures 14 and 15).  Calculation of the similarity measures and cluster analysis were 
performed using the Multi-Variate Statistical Package, ver. 3.1 (Kovach 1993).

IV. RESULTS

A. Station Locations and Descriptions

Locations of station sampled are shown in Figures 10-12, and details for sampling dates and 
coordinates in Latitude-Longitude (WGS84 Datum) are given in Table 1.

Station 1. (Latitude 21? 18 13.6N, Longitude 157? 51 49.8W)
Concrete pier walls and pilings and basin bottom along Pier 3, Ala Moana Boulevard and Pier 4 
used as a landing for Coast Guard shuttle boats.  Walls and pilings support abundant red algae 
Galaxaura acuminata, a few corals and some bryozoans, primarily Schizoporella sp. A.  The 
bottom depth along the walls is about 3 m, deepening in the center of the basin to 10 m (33 ft) 
maximum, with numerous trash such as tires, pipes sheet plastic and bottles along the piers.  
Going deeper, the bottom is coarse white sand, coral rubble and pebbles, with some hard coral 
substratum from the middle to the end of Pier 4.

Station 2. (Latitude 21? 18 18.1N, Longitude 157? 51 52.4W)
This area is a rectangular basin about 40 m (132 ft) x 100 m (330 ft) between Piers 5 and 6, 
which receives the thermal discharge from the Hawaiian Electric Honolulu Generating Station.  
The perimeters of the basin are concrete walls about 3 m deep constructed on top of fringing reef 
that has been dredged to about 10 m (33 ft) depth.  The wall along Pier 5 that is directly in the 
path of the thermal discharge has abundant oysters and limpets, and many fish were observed in 
the discharge plume which increases turbidity in this part of the discharge basin.  Reef corals are 
very abundant along the Ala Moana Boulevard wall, where occurs a large colony of Pocillopora 
eydouxi with a diameter of ca. 1 m, the largest of this species that has been reported in Hawaii.  
The west side of the basin is a dock which juts out from Pier 6 and is supported by concrete 
pilings, and addition hard surface is formed by two concrete abutments in the middle of the basin 
which formerly support a parking lot road.  These abutments formerly supported an abundant 
growth (Thomas 1979) of the introduced octocoral Carijoa (=Telesto) riisei, but only a few 
colonies were found at the time of our survey.  The bottom of the basin is primarily soft silt and 
fine sand ranging down to 10 m (33 ft) except along the sides in depths of 4-6 m (13-20 ft) where 
pieces of coral rubble range up to boulder size.

Station 3. (Latitude 21? 18 21.6N, Longitude 157? 51 53.9W)
This basin is the location of the intake for the Hawaiian Electric power station cooling water along
 the Ala Moana Boulevard wall, where a concrete wall extends to 2.5 m (8 ft) depth to the top of 
the dredged reef.  Reef corals are abundant along this wall, similar to the Ala Moana side of 
Station 2. The east side of the basin at Pier 7 and the west side at Pier 8 are wooden piers 
supported by concrete pilings which were been constructed with the development of the Hawaii 
Maritime Center Museum and the Aloha Tower Market Place within the last ten years.  Pier 7, 
which is also the mooring place for the museum ship Falls of Clyde, was formerly a parking lot 
with concrete walls that supported abundant corals (McCain and Coles 1973; McCain et al 1975), 
but corals are now scare because of the decks shading of the pier walls.  The pillars now have 
an abundant fouling community of suspension feeders.  Also, the iron hull of the Falls of Clyde, 
which is moored at Pier 7, provides additional surface for benthic organisms and was the only 
location in Honolulu Harbor where the pearl oyster Pinctada margaretifera was found.  The depth 
of the basin away from the walls is about 10 m (33 ft) and is composed of loose fine silt with 
intermittent coral rubble and a fine coating of algae.

Station 4. (Latitude 21? 18 32.9N, Longitude 157? 51 52.2W)
This site is the basin located between Pier 11 and the former Pier 12, which was the location of 
the first pier that was made in Honolulu Harbor in 1825 by sinking a ships hull at the foot of 
Nuuanu Avenue and building a dock around the hull (Rush 1957).  When the Honolulu Fort was 
torn down in 1857 and its materials were used in filling the nearshore reef and subtidal area 
which make up the present Aloha Tower area, formerly known as the Esplanade (Anon. 1936; 
Judd 1975).  Pier 12 was built in 1907 (Wilson, Okamoto & Assoc. 1968) and is no longer 
present, but a small area used for docking small boats exists at the former Pier 12 site.  The 
perimeter of this area is made of cut coral blocks, stated in Oceanit (1990) to be taken from the 
Honolulu Fort, but this is unverified.  The blocks extend down to about 6 m (20 ft) depth in a 
mixed silt-rubble bottom which has abundant trash such as wire, old appliances and fishing lines.  
Despite generally poor water clarity in this area, live reef corals are abundant along the pier, 
especially Porites lobata and Montipora capitata (=verrucosa) at the edge of the dredged reef.  
The zoanthid Zoanthus pacificus  and the coral Pocillopora damicornis  are also abundant on the 
coral blocks right up to the pilings which support Nimitz Highway.  These pilings stand in front 
storm drainage culverts and have a rich coverage of oysters, bryozoans, hydrozoans and a few 
corals.  The Pier 11 side of the basin is a pier that extends about 10 m (33 ft) from the shore and 
is supported by concrete pilings that have an abundant fouling community.  Depth in the basin 
ranges up to 10 m (33 ft), with a bottom of soft muddy sediments and abundant trash.

Station 5. (Latitude 21? 18 38.9N, Longitude 157? 51 54.4W)
Located along Pier 14, this site is one of the two closest to the mouth of Nuuanu Stream and its 
freshwater runoff.  The pier extends 10 m  (33 ft) from the shore and is supported on pilings that 
have an abundant coverage of fouling organisms such as fanworms, tunicates and the coral 
Montipora capitata (=verrucosa).  The depth along the pier is 8 m (26 ft), with a bottom of fine 
muddy sediments

Station 6. (Latitude 21? 18 38.2N, Longitude 157? 52 2.2W)
This site is at Pier 20, which is the docking area for numerous harbor and interisland tugs, and is 
just outside of the mooring areas for commercial fishing vessels at Piers 16-18.  It is also the area 
most directly exposed to runoff from Nuuanu Stream which discharges into the harbor about 91 m 
(300 ft) northeast of the site.  The dock is supported by concrete piers extending up to 10 m (33 
ft) out from the shoreline in dredged coral substratum, and depths along the shore and outside 
the dock range 2-10.5 m (6.6-35 ft).  Fouling is abundant on the dock pilings and the fine 
sediments are composed mostly of terrigenous material.


Station 7. (Latitude 21? 18 33.2N, Longitude 157? 52 10.6W)
This station lies at the end of Pier 27, which is the approximate location of the northwest edge of 
the original Honolulu Harbor that existed before the Honolulu Basin was enlarged and the 
Kapalama Channel dredged in the early part of this century.  All characteristics of this site are 
otherwise similar to those at Station 6.

Station 8. (Latitude 21? 18 39.4N, Longitude 157? 52 24.3W)
This site lies between Piers 29 and 30 along the Kapalama channel, and it represents a relatively 
natural environment compared to other areas this far into the harbor.  Although the area was 
formed from dredging the channel through a former reef flat, it has the appearance of reef slope 
outside of a narrow fringing reef which extends about 5 m (16.5 ft) from the shoreline.  Although 
this 1-2 m flat area is quite barren, the slope outside the reef has a variety of coral species with 
moderate coverage and numerous fishes, which are probably attracted to the rugose habitat 
provided by the numerous small holes and ledges on the slope. The reef extends to nearly 10 m 
(33 ft) depth where the bottom levels off to a fine silt substratum.  Just southeast of this site at 
Pier 29, the concrete pilings of the pier and the reef substratum below have heavy fouling and 
abundant sponges with a heavy coating of silt.

Station 9. (Latitude 21? 19 0.4N, Longitude 157? 52 37.2W)
This site is in the most interior section of the harbor at Pier 36, across from the mooring area 
where the Clean Island Council docks its vessels used for rapid oil spill response.  Pier 39 itself is 
used for docking large fishing boats.  The area is at the end of a basin that has been dredged 
from fossil reef and is quite isolated from the more open parts of the harbor, suggesting relatively 
stagnant conditions.  The pier supports of concrete pilings extend to 8.5 m (28 ft) depth along Pier 
39 to a shallow area at the head of the basin about 1 m 3.3 ft) deep.  The pilings are heavily 
fouled with suspension feeders, while the algae Acanthophora spicifera and Galaxaura acuminata 
are abundant in the shallows.  A school of tilapia (Oreochromis mossambicus) was observed 
along the Nimitz Highway end of the basin, the only place this fish was observed to be abundant 
in the harbor.

Station 10. (Latitude 21? 19 0.5N, Longitude 157? 52 50.4W)
Located at the end of Pier 39, where the pier is a grooved concrete wall extending down 9 m (30 
ft) to a deep soft silt bottom.  Fouling on the pier wall was relatively sparse, and surfaces were 
heavily sedimented, probably from runoff from Kapalama Canal, which reaches the harbor at the 
head of this basin about 400 m (1320 ft) landward of this site.  The bottom depth is about 9.5 m 
(31 ft) with a substratum of fine muddy silt.

Station 11. (Latitude 21? 19 2.4N, Longitude 157? 52 57.7W)
Sampling for this station was done from the surface of the main dry-dock operating in Honolulu 
Harbor, located at the end of Pier 41.  The dry-dock is on jack-up legs standing in 11 m (36 ft) of 
water on a fine sediment bottom.  The dry-dock is heavily fouled with tunicates and hydrozoans.


Station 12. (Latitude 21? 18 55.5N, Longitude 157? 53 12.3W)
This site is just outside of the Snug Harbor mooring area for ships operated by the University of 
Hawaii. The water at the site is shallow, ranging 2-5 m (6.6-16.5) and the shoreline has a row of 
wooden piers that are the main hard surface at the site.  The sediment has a greater component 
of calcareous material than occurs further into the harbor, but biotic coverage on the available 
surface is relatively sparse.

Station 13. (Latitude 21? 18 24.7N, Longitude 157? 52 19.5W)
Located at the east end of the Coast Guard Station Docking area.  Concrete pilings extend to 9.5 
m (31 ft) depth in a turbid area with numerous coils of cable, tires, rope and other trash next to 
the dock.  Fouling was moderate, with virtually no corals and few fish present.

Station 14. (Latitude 21? 18 8.0N, Longitude 157? 52 9.4W)
Located at the border of Anuenue Fisheries Center and Sand Island Park near the beginning of 
the harbor entrance channel.  Substratum is a steep slope dredge from the reef and small 
boulders from 1-2 m (3.3-6.6 ft) to the fine sediment harbor bottom at 9 m depth.  Corals and 
associated invertebrates were moderately abundant and a variety of fish species were present.

Station 15. (Latitude 21? 18 50.7N, Longitude 157? 53 9.5W)
Located west of the Sea Land Pier and adjacent to the entrance of the Kalihi entrance channel.  
Substratum is a rock seawall made of large basalt boulders on top of the reef, which has been 
dredged to 11 m (36 ft) depth.  Boulder and trash are abundant on the silt bottom.  Corals were 
common and reef fish were abundant.

Station 16. (Latitude 21? 17 58.6N, Longitude 157? 54 2.2W)
This site is the most exposed to open ocean conditions of any in the study, being located on a 
reef along the west side of the Kalihi Channel, approximately 1.75 km the entrance, in 2-3 m (6.6-
10 ft) depth.  The substratum is consolidated reef with numerous sand channels and large pieces 
of the iron hull of a wrecked boat or barge.  Abundant filamentous algae and macroalgae 
dominate the benthos.

Station 17. (Latitude 21? 18 42.2N, Longitude 157? 55 9.3W)
The site is the docking area for the Department of Transportation Fire Rescue boat hanger 
located at the Reef Runway end of the seaplane runway that runs along Lagoon Drive.  This dock 
was constructed as part of the completion of the Reef Runway in 1972-1975 and therefore can be 
utilized as a dated surface no more than 25 years old.  The sites surfaces are concrete pilings in 
3 m (10 ft) of water which have an abundant fouling community, and the bottom substratum is 
primarily medium to fine grained calcareous sediments.

Station 18. (Latitude 21? 19 11.0N, Longitude 157? 53 39.8W)
Keehi Lagoon Marina floating docks located midway between Honolulu Harbors Kalihi Channel 
and mouths of Kalihi and Moanalua Streams.  The dock surfaces are very heavily fouled and are 
anchored in 3 m (10 ft) of turbid water over a muddy sediment bottom.

Station 19. (Latitude 21? 19 5.2N, Longitude 157? 54 26.8W)
Located midway along the reef side of the Lagoon Drive seaplane runway, the site was an iron 
barge hull stranded on the reef edge.  The barge has since been removed.  Depth on the runway 
side of the barge was 4.5 m (15 ft) and decreased to 1 m (3.3 ft) on the reef side of the barge.  
The hull had only moderate fouling with a heavy sediment coating, and the bottom substratum 
was fine sand to silt.

Station 20. (Latitude 21? 19 54.6N, Longitude 157? 53 35.2W)
The site was at the mouth of Moanalua Stream where abundant red mangrove (Rhizophora 
mangle) roots provide the only solid substratum in the muddy bottom.  Samples were taken from 
the roots at 0-0.5 m (0-1.7 ft) depth.

Station 21. (Latitude 21? 17 36.4N, Longitude 157? 51 30.1W)
Located in Kewalo Basin, along Honolulu Marine Dock on north side of the basin just inside the 
entrance.  Along with vessel repair activities conducted by Honolulu Marine this area is used for 
docking tourist ferries that conduct daily tours of Waikiki, Honolulu Harbor and Pearl Harbor.  
Depths along the dock range down to 6 m, and the fouling community on hard surfaces is 
dominated by macroalgae.  The fine sand bottom has a great deal of litter and trash, especially 
old tires.

Station 22. (Latitude 21? 17 29.5N, Longitude 157? 51 56.9W)
This is the docking area for small boats at the southwest corner of Kewalo Basin, across the road 
from the offices and training pools of the Hawaii Marine Mammal Center.  The area is shallow, 
ranging down to only 3 m (20 ft), and both the hard surfaces on the docks and the sandy bottom 
are dominated by a heavy growth of the algal-like bryozoan Zoobotryon verticillatum.

Station 23. (Latitude 21? 17 38.9N, Longitude 157? 51 26.6W)
Docking area for large fishing boats along Fishermans Wharf at the northwest corner of Kewalo 
basin.  The docks extend about 10 m (33 ft) from the share on cement piers that extend down to 
a maximum of 4 m (13 ft) over a mixed sand/silt/shell bottom.  The area can receive storm water 
runoff from drainage culverts located along Ala Moana Boulevard, which runs along the north side 
of Kewalo Basin. 

Station 24. (Latitude 21? 17 34.7N, Longitude 157? 51 19.5W)
Small boat docks near site of former McWaynes Marine Center at the northeast corner of Kewalo 
Basin.  Sampling site was a concrete wall along the shoreline extending down to 3 m (10 ft) and a 
fine sand-shell bottom.

Station 25. (Latitude 21? 17 15.6N, Longitude 157? 50 26.8W)
Ala Wai Canal Bridge where the canal empties into the Ala Wai Yacht Harbor.  Samples were 
taken only from the concrete base of the a bridge abutment in the intertidal zone at 0-0.5 m (0-1.7 
ft) depth

Station 26. (Latitude 21? 17 16.3N, Longitude 157? 50 30.8W)
Located along the wall in front of the Waikiki Yacht Club on the northwest side of the Ala Wai 
Yacht Harbor and by on adjacent cement pilings and floating docks.  Depth is shallow, maximum 
2 m (6.6 ft) over a muddy, soft silt bottom.

Station 27. (Latitude 21? 17 6.5N, Longitude 157? 50 26.8W)
Floating docks in Ala Wai Yacht Harbors most shoreward  channel, in front of end of Hobron 
Lane.  Depth was 3.5 m (12 ft) over a muddy bottom with abundant trash, water was very turbid 
with a surface oil sheen.

Station 28. (Latitude 21? 17 5.6N, Longitude 157? 50 36.8W)
Along the Texaco fueling area at the end of the most seaward dock in the Ala Wai Yacht Harbor, 
adjacent to the harbor entrance.  Substratum was cement pilings, floating docks and walls along 
the front of the fueling area.  The fine sand bottom is up to 6 m (20 ft) deep along the sloping 
coral rubble side of the basin.

Station 29. (Latitude 21? 16 58.2N, Longitude 157? 50 7.4W)
Southeast interior of the Ala Wai Yacht Harbor adjacent to the Hilton Lagoon, which discharges 
effluent circulated from the lagoon to this site.  Depth ranges down to 4 m (13 ft) to a bottom of 
coarse shells in fine sand.

Station 30. (Latitude 21? 19 18.5N, Longitude 158? 7 11.2W)
Located at the principal cargo area along the south side of the Barbers Point Deep Draft Harbor, 
which presently is used primarily for unloading coal transported by a conveyor belt from docked 
bulk carriers.  Concrete pilings, many almost completely covered with the nonindgenous 
bryozoan Amathia distans, stand along the dock front in about 10 m (33 ft) depth over a silt-mud 
bottom.

Station 31. (Latitude 21? 19 18.9N, Longitude 158? 7 14.9W)
This site is a floating dry-dock, which is moored about 10 from the shore on the west side of 
Barbers Point Deep Draft Harbor in 11.5 (38 ft) m water.  All fouling samples were taken from the 
dry-docks hull, which was in approximately 9.5 m (31 ft) deep along the ships bottom.

Station 32. (Latitude 21? 19 19.8N, Longitude 157? 7 16.7W)
Barge Pier along south side of the Barbers Point Deep Draft Harbor channel, near the entrance.  
The hard substratum was a concrete wall extending to 7 m (23 ft) depth on a mixed pebble-sand 
bottom with abundant twisted metal and wrecked car debris along the wall.

B. Benthos and Fish Surveys

A total of 1122 taxa with 852 named species algae, invertebrates and fishes, listed in Appendix C, 
have been reported in all studies in the five south and west shore Oahu harbors. Of these, 728 
taxa including 585 named species were observed or collected in the five harbors or basins.  Table 
2 shows the distribution of the total taxa and named species among major taxonomic groups for 
each of the five harbors.

Table 2. Numbers of total taxa and named species in major groups observed or collected in the 
five harbor areas.



Figure 14. Percent similarity cluster analysis of all invertebrate taxa by station.

Figure 15. Percent similarity cluster analysis of amphipod crustacean taxa by station.

Although levels of association within any station groups were not very high (maximum 68%), the 
overall pattern of station clusters corresponds to environmental conditions and locations among 
and within the harbors (Figures 10-12).  Four clusters of stations are suggested, with three 
outliers.  Station 16, along the Kalihi channel outside of Keehi Lagoon, and Station 20, in 
Moanalua Stream mouth showed low association with other stations, indicating the uniqueness of 
the communities at those locations.  Also, Station 32, at the Barbers Point barge pier, although 
lying within Cluster B, was separated from all other stations in that cluster.  Cluster A was 
composed of nine stations all within Kewalo basin and the Ala Wai Yacht Harbor, with two 
stations, 25 at the Ala Wai Bridge, and 27, at the end of Hobron Lane forming a subcluster.  
Environmental conditions at these two stations were noted at the time of sampling to be the 
poorest of any in the harbor, with abundant floating garbage, turbid conditons and an oil sheen on 
the water at Station 27.

The stations at the coal loading dock (30) and the floating dry-dock (31) at Barbers Point 
composed Cluster B, while Cluster C was made up of three stations in Keehi Lagoon.  The 
remaining Cluster D was composed of the 15 Honolulu Harbor stations arranged into two groups 
and one outlier. The larger of the groups contained Stations 4 to 13 and 15 from the interior of the 
harbor, and the other group was composed of three stations near the harbors main entrance 
(Stations 2, 3 and 14).  Station 1, also near the entrance, was the final station within Cluster D.

Of the analyses done for the major invertebrate taxonomic groups and the fishes, only the 
similarity indices and dendrographs for the Amphipoda (Figure 15) showed a discernable pattern.  
The station groupings are similar to those for the total invertebrates (Figure 14) and have a 
generally higher lever of association within clusters, suggesting that much of the pattern shown 
for the total invertebrates was accounted for by distributions of amphipods.  The Kalihi Channel, 
Moanalua Stream mouth and Barbers Point barge pier stations remain as outliers, while Cluster 
A contains Kewalo Basin and Ala Wai Harbor stations plus a single Keehi Lagoon station, 
grouped in two subclusters.  Cluster B contains all Honolulu Harbor stations except for two from 
Keehi Lagoon, arranged in four subclusters.  Subclusters B1 and B2 are from the interior of 
Honolulu Harbor, B3 has Station 11 from the Honolulu Pier 41 dry-dock and Station 18 from 
Keehi Lagoon.  Subcluster B4 contains the three stations near the Honolulu Harbor main channel 
and one form the reef edge in Keehi Lagoon.  Station 1 is missing from this analysis because the 
amphipods from this station were apparently misplaced before identification. 

The distribution of total numbers of taxa of invertebrates and fishes among the stations is shown 
in Figure 16.  Total taxa ranged from a low of 24 at the mouth of Moanalua Stream (Station 20) 
where the only hard substratum to sample was on mangrove prop roots, to highs of more than 
180 taxa at Station 2, in the discharge basin of Hawaiian Electrics Honolulu Plant, and at Station 
14, across the harbor on the Sand Island side of the entrance. Numbers of taxa for eight major 
groups of invertebrates and fishes are shown for each station in Figure 17.  Although the patterns 
vary, the highest or second highest numbers for seven of the nine groups were at stations near 
the Honolulu Harbor main entrance, with maximum numbers occurring at Station 14, on the Sand 

Figure 16. Total numbers of invertebrates and fish taxa observed or collected at the stations.

Island side of the main channel for Cnidaria, Mollusca and fishes.  Echinoderms also showed 
maximal numbers in this area at Stations 1 and 3.  For the remaining two groups, Crustacea were 
most abundant at Station 16 along the Kalihi Channel outside of Keehi Lagoon, where amphipods 
and isopods accounted for 20 of the 61 taxa that occurred.  Highest numbers of ascidian taxa 
were at piers well within Honolulu Harbor (Station 10) and Keehi Lagoon (Station 18) and at the 
Honolulu Marine Pier (Station 21) in Kewalo Basin.

Taxonomic richness was highest among the Crustacea, with a maximum number at Station 16 
along the Kalihi Channel outside of Keehi Lagoon, where 26 of the 61 crustaceans were 
amphipod or isopod species occurring only at that site.  Other groups with high species richness 
were the Polychaeta, which had 36 taxa at Station 2 near the Honolulu Harbor entrance, and 
fishes, which had 33 taxa at Station 14, on the other side of the harbor entrance along Sand 
Island.  The distribution of the Echinodermata was highly restricted, with more than one or two 
species occurring only near the Honolulu Harbor main entrance at Stations 1, 2, 3 and 14, or at 
Station 16, in open ocean water along the Kalihi Channel outside of Keehi Lagoon. 

The patterns of abundance for total taxa are shown on the dot maps for each harbor in Figures 18 
to 20.  Except for Barbers Point, every location shows a distinct increase in numbers of taxa with 
proximity to a harbor entrance.  This is shown most clearly for Honolulu Harbor, where three of 
the four stations near the entrance had more than 150 taxa, but also held for Kewalo Basin and 
the Ala Wai, where taxa numbers near the entrance exceeded 100 while 50 to 100 were found at 
more interior stations.   Lowest numbers of taxa occurred at the mouths of Moanalua Stream in 
Keehi Lagoon (Station 20) and the Ala Wai Canal (Station 25).  This is due in part to the low 

Figure 18. Total invertebrate and fish taxa at Honolulu Harbor and Keehi Lagoon stations.
Figure 19. Total invertebrate and fish taxa at Kewalo Basin and Ala Wai Yacht Harbor stations.

Figure 20. Total invertebrate and fish taxa at Barbers Point Deep Draft Harbor stations.

water quality and high turbidity at these sites preventing diving observations, consequently 
sampling was conducted only near the waters surface.  However, the lack of suitable substrate 
high siltation from stream outflow, and the likelihood of considerable salinity fluctuations probably 
result in a limited marine biota at these locations.

C. Comparison with Previous Reports

A total of 728 taxa including 585 named species (Appendix D) were collected or observed in the 
present study at all sites  Tables 3 and 4 summarize the numbers of total taxa and named 
species for major taxonomic groups in each harbor area in for the present study, along with the 
corresponding numbers for all previous surveys combined.  The greatest numbers of taxa and 
species were in Honolulu Harbor, where the total taxa for the 15 stations of the present study of 
598 is about 1.4 times the number of taxa from all previous reports from this harbor, and is about 
three times the numbers for each of the other harbors.  With the exception of Keehi Lagoon, 
where previous numbers of taxa reported were slightly more than found in the present study, the 
numbers of taxa found in the present study exceeded by many times those for previous reports.  
These greater numbers applied across all major taxonomic groups except for the Keehi Lagoon 
results, where the greater number of taxa from previously reported studies were due primarily to a 
single survey (Harvey 1970), which listed 70 of the 74 molluscs noted in Appendix D.

Comparing values of Table 3 with Table 4 indicates that most taxonomic identifications were 
made to the species level.  Named species as a percentage of total taxa ranged from 73% for 

Table 3. Numbers of total taxa in present study compared with totals of previous studies.


Table 4. Numbers of named species in present study compared with totals of previous studies

Honolulu Harbor to 87% for Keehi Lagoon.  As suggested in Table 3 for total taxa, Honolulu 
Harbor had the most species, and numbers of species found in the present study were 
substantially greater than numbers for previous studies at all harbors except at Keehi Lagoon, 
where the values were about equal.

D. Nonindigenous and Cryptogenic Species

Species previously reported in Hawaii were categorized as native, nonindigenous or cryptogenic 
(i. e. of uncertain origin but with indications of being introduced) according to the designations in 
Carlton and Eldredge (in prep) and Coles et al. (1997, 1999). For species new to Hawaii, status 
was assigned using the criteria presented by Chapman (1988) and Chapman and Carlton (1991) 
and described in Coles et al. (1997).  These criteria include new appearances in the region, 
association with known dispersal mechanisms or other introduced species and disjunct 
geographic distributions.  Taxonomic specialists were also consulted for their input in assessing 
the status of newly reported species.

Of the total 585 named species found in the five harbors in this study, 27 are considered to be 
cryptogenic and 73 are designated nonindigenous (Appendix E), for a total of 100 non-native 
species, or 17% of the total species overall.  Tables 5 and 6 list the numbers of designated 
nonindigenous and cryptogenic species found in the present study and in all previous surveys for 
each harbor area.  As was the case for total taxa and named species (Tables 3 and 4) Honolulu 
Harbor had the greatest numbers, with a maximum of 51 nonindigenous and 23 cryptogenic 
species in the present study.  Past surveys in Honolulu Harbor also reported a substantial 
number of nonindigenous and cryptogenic species, in contrast to the other harbors, where limited 
sampling did not detect introductions at Barbers Point or Kewalo Basin, and only one 
nonindigenous organism was found in the Ala Wai Harbor.  Similarly, only 13 nonindigenous and 
two cryptogenic species were previously reported in Keehi Lagoon.

Totals for introduced and cryptogenic species in each harbor are shown in Table 7, which 
indicates that Honolulu Harbor has had the greatest numbers of nonindigenous and cryptogenic 
species and therefore has been the most impacted by marine introductions.  However, when 
these data are compared with the total numbers of named species in each harbor, a different 
pattern emerges.  Expressing the introduced species as a percentage of the total named species 
found in each harbor in 1997-98 (Table 4), Honolulu Harbor has the lowest value of 15%, while 
the remaining harbors have values ranging from 27% to 33%, with the highest value occurring in 
Keehi Lagoon.

The distribution of the nonindigenous and cryptogenic species among the sites sampled 
(Appendix E) indicates that introductions were not concentrated in any harbor. Most species 
occurred at multiple sites, with only 13 of the 100 species unique to any station (three at Barbers 
Point, three at Keehi lagoon, three at the Ala Wai and four in Honolulu Harbor).  This indicates 
that most introduced species were frequently encountered and well distributed among the five 

Table 5. Introduced species in Oahu south and west shore harbors.

Table 6. Cryptogenic species in Oahu south and west shore harbors.

Table 7. Introduced + cryptogenic species in Oahu south and west shore harbors.


harbors.  Honolulu Harbor, with more than twice as many total species than any other harbor, did 
not have a corresponding increase in nonindigenous or cryptogenic forms. Instead the 
percentage of the total species that were nonindigenous or cryptogenic was around half the 
values for other harbors.  This suggests that, although the discovery of rare or introduced species 
will always be a function of sampling effort, the greater effort in Honolulu Harbor did not yield 
increased numbers of introduced species relative to the total community.  The five harbors 
therefore can be considered as a single entity regarding nonindigenous and cryptogenic species.

E. Recent Introductions


Table 8. Nonindigenous species first noted in Oahu south and west shore harbors (including 
Pearl Harbor) since 1995.

Table 8 lists all the nonindigenous species from the present study that are first reports for Hawaii, 
along with similar first reports that were determined by the Pearl Harbor Study in 1996 (Coles et 
al. 1997, 1999).  Thirteen new nonindigenous species were found for the five harbors of the 
present study, compared to 12 new species for Pearl Harbor.  Only two of the species, the 
sponge Gelliodes fibrosa and the barnacle Chthamalus proteus, both from the Caribbean, 
occurred at all stations in the present study and in Pearl Harbor.  Two more species, the sponge 
Sigmadocia caerulea and the pycnogonid Pigrogromitus timsanus, previously found in Pearl 
Harbor, also occurred in one or two harbors in the present study, while the eight remaining Pearl 
Harbor were not found elsewhere.  Three of the four bryozoans (Bugula dentata, B. robusta and 
Cailibugula dendrograpta), the ascidians Bottryloides simodensis and Eusynstela aliena and the 
sponge Neofolitipsa ungiculata were quite widely distributed in two or three of the harbors in the 
present study, but the remaining seven species occurred only at a one or two stations in a single 
harbor.  

Three of the unique reports were from the Ala Wai Yacht Harbor.  The corallimorpharian 
Actinodoscus nummiformis was first found near Station 28 in December 1997 by staff of the 
Waikiki Aquarium.  The organism zooids were artificially attached to a piece of coral rubble, 
indicating that the organisms had been discarded from a salt-water aquarium.  Three small 
clumps of A. nummiformis were still growing at the site in April 1999, but the organism has not 
been observed elsewhere in or outside of the harbor.  Another cnidarian found only in the Ala Wai 
was Diadumene franciscana, previously known only from San Francisco Bay (D. Fautin, pers. 
comm.) which only occurred at Station 26, near the Waikiki Yacht Club, and at Station 28, near 
outflow into the harbor from Hilton Lagoon.  The only occurrence of the Lemon Peel fish 
Centropyge flavissima in this study was an observation near the harbor boat ramp.  However, R. 
Pyle (pers. comm.) has sighted this species infrequently over at least the past ten years at Keehi 
Lagoon, Kewalo Basin, the Ala Wai and Kaneohe Bay.

The remaining single reports were of the pycnogonid Anoplodactylus arescus, the amphipods 
Jassa falcata and Leucothoe micronesiae and the bryozoan Caulibugula caliculata. 
Anoplodactylus arescus is a sediment dwelling pycnogonid occurring in this study only at the 
Barbers Point coal loading dock and previously found throughout the Indo-Pacific as far west as 
East Africa.  Jassa falcata is a common circumtropical and temperate species with unknown 
origins, reported from harbors around the world.  Leucothoe micronesiae was originally described 
from the Caroline Islands and has also been reported from Madagascar. Caulibugula caliculata 
and C. dendrograpta, which occurred here only in Honolulu Harbor, have previous known 
distributions in the western Indo-Pacific (C. Zabin, pers. comm.).  These bryozoans, along with 
the more widely distributed B. dentata and B. robusta, although first reported here for Hawaii, are 
very likely to also occur in Pearl Harbor, which wil be verified by a review of the samples taken for 
the 1996 study.

F. Introductions with Time

Of the 100 nonindigenous or cryptogenic species found in the present study, all but four have a 
recorded date of first appearance in Hawaiian waters, shown in Appendix F along with data on 
the location of the first Hawaiian report, origins or previous ranges of the introduced species, and 
sources of information.  These first Hawaiian reports are grouped by decade and shown in Figure 
21.  The graph can be interpreted to suggest that substantial increases in first reports of 
introduced species occurred in the 1970s and 1990s when 17 to 21 nonindigenous and 7 to 12 
cryptogenic species were first reported in Hawaii.  Figure 22 shows the decades when the same 

Figure 21. First Hawaiian reports of nonindigenous and cryptogenic species found in Oahu south 
and west shore harbors in 1997-98.
Figure 22. First occurrences of nonindigenous and cryptogenic species found in Oahu south and 
west shore harbors in 1997-98.


 
Figure 23. Ratio of new reports of nonindigenous and cryptogenic species to first reports for all 
species by decade (bars) and total new species reports by decade for Oahu south and 
west shore harbors (line).

species were first reported in one of the five harbors of the present study.  An even greater recent 
increase is suggested by this graph, where 69 of the 100 nonindigenous and cryptogenic species 
were first observed or collected in these harbors in the 1990s, most of them during the two years 
of the present study. 

A comparison of the two figures suggests that in decades when sampling effort has been low, it is 
likely that a relatively rare species, introduced or otherwise, may be present but undiscovered 
until a threshold effort has been made which will result in collection of the species.  Prior to the 
1990s the only substantial sampling that had been done in Honolulu Harbor or Keehi Lagoon was 
in the 1940s and 1970s.  These decades were also the only periods of substantial introductions 
introduction into these harbors previous to the 1990s (Figure 22) despite numerous first reports of 
introduced species elsewhere (Figure 21).

Further indication that the apparent recent increases in nonindigenous and cryptogenic species in 
these harbors is sampling effort related is shown in Figure 23, which plots the ratio of new reports 
in the five harbors to the total number of species newly reported in the same decade, along with 
the numbers of total new reports.  In contrast to the relatively high numbers of introductions 
suggested for the 1970s and 1990s in Figures 21 and 22, the ratio of introductions to total 
species is lower for the 1990s than for the 1930s or 1940s, and the value for the 1970s is the 
lowest for any decade in which introduced species were first reported.  This data presentation is 
highly sensitive to low numbers, as in the case of the 1930s when only one nonindigenous out a 
total of five newly reported species occurred, resulting in the highest ratio value for any decade.  
However, the results support a conclusion that increased numbers of newly detected introduced 
species may correlate strongly with total numbers of species found, with both increases probably 
related to sampling effort.

G. Origins and Previous Distributions of Nonindigenous Species.

 
Figure 24. Origins or previous distributions of nonindigenous and cryptogenic species collected or 
observed in Oahu south and west shore harbors in 1997-98.  EA: Eastern Atlantic, CA: 
Caribbean, WA: Western Atlantic, EP: Eastern Pacific, IP: Indo-Pacific, WIP: Western 
Indo-Pacific, RS: Red Sea, WW: Tropical or Temperate World Wide.

The origins or previous recognized distributions of the nonindigenous and cryptogenic species 
collected or observed in the present study are listed in Appendix E, and the totals are 
summarized in Figure 24.  Information was insufficient to assign an origin or distribution to 21 of 
the 99 species. Twenty-nine species, or 38% of the remainder, were considered to have tropical 
or temperate worldwide distributions with unknown origins.  The Indo-Pacific accounted for 42% 
of the species with recognized origins or previous distributions, with 22% coming from the central 
Indo-Pacific, 19% from the western Indo-Pacific regions, 1% from the Red Sea, and only 4% from 
the Eastern Pacific.  The Atlantic accounted for only 16% of the total, with most of this (10%) 
coming from the Caribbean Sea.

V. DISCUSSION

These surveys conducted in 1997-98 detected 13 nonindigenous and two cryptogenic marine 
species not previously reported in Hawaii.  These introductions are in addition to the 12 
nonindigenous and 12 cryptogenic species first found for Hawaii in Pearl Harbor in 1996 surveys 
(Coles et al. 1999).  Only two of these species, the sponge Gelliodes fibrosa and the barnacle  
Chthamalus proteus, occur in all the harbors, while many of the other newly reported species 
occurred at only a single station within a given harbor.

The discovery of 25 nonindigenous species and 14 cryptogenic species within a three-year period 
is a substantial increase in introduction reports above previous periods.  At least three factors 
may be responsible for such an apparent increase in marine introductions.

? An increase in transport, survival and propagation of introduced species has occurred from 
various vectors such as hull fouling, ballast water transport, aquaculture or fisheries 
introductions and aquarium releases.  These factors have undoubted been responsible for 
the recent appearance of the Gelloides fibrosa and other sponges, the barnacle Chthamalus 
proteus, (fouling), the cnidarian Actinodiscus nummiformis and the fish Centropyge 
flavissimus (aquarium releases).
? Increased sampling effort in the 1990s has increased the probability of discovering and rare 
introduced species that may have either recently arrived or may have been in Hawaii for 
many years, but in low abundance or in rarely sampled localities that prevented their 
discovery.  The present study was the first comprehensive biological sampling that had been 
conducted throughout Honolulu Harbor and Keehi Lagoon, and the only substantial sampling 
that has ever been done in Kewalo Basin, the Ala Wai Harbor or Barbers Point Deep Draft 
Harbor.  It is logical that this increased effort would detect a substantial number of introduced 
species that may have been present but previously missed.  An analysis of the 
nonindigenous and cryptogenic introductions by decade in Pearl Harbor indicated a strong 
relationship with total species reported, considered to be an indicator of sampling effort 
(Coles et al. 1999).
? Increased attention to a taxonomic group, especially one with species likely to have been 
transported and introduced to a new area, can produce an apparent increase in introductions 
when these species may have been resident in an area for an undetermined period of time.  
For example the ascidian Botrylloides simodensis, although first reported here, has probably 
been in Hawaii for a very long time (G. Lambert, pers. comm.).

Insufficient information exists to determine which of these factors is most responsible for the 
increased incidence of nonindigenous species reports in the 1990s, but all three undoubtedly 
contribute.  Some species, such as Chthamalus proteus and Actinodiscus nummiformis are 
verifiable recent introductions with identifiable vectors, while others like the ascidians Botrylloides 
simodensis and Eusynstyela aliena are probably the result of increased sampling and taxonomic 
effort.  After review of the 25 newly reported nonindigenous species listed in Table 8, we 
conclude that approximately eight are verifiable recent arrivals.  These are Gelliodes fibrosa, 
Sigmadocia caerulea, Actinodiscus nummiformis, Chama elatensis, Chthamulus proteus, 
Nanosesarma minutum, Symplegma reptans and Centropyge flavissimus

The results of this study are very similar to those obtained from the comprehensive survey of 
Pearl Harbor in 1996 (Coles et al. 1997, 1999).  Both studies found around 100 nonindigenous 
and cryptogenic species, each with about 12 new nonindigenous reports.  Both studies found 
numbers of new introductions by decade to be strongly related with the total numbers of newly 
reported species, indicating that high values for introductions in the 1970s and the 1990s were 
sampling effort related.  In Pearl Harbor during the 1910s and 1940s, the ratio of introductions to 
total species were substantially higher than in all other decades, corresponding to periods of 
harbor opening and construction after 1911, and intense increase in shipping activity during and 
following WW II (Coles et al. 1999).  The increase for this ratio shown during the 1940s in Figure 
23 may also have been related to increased shipping activities in Honolulu and other harbors 
during the decade of WW II when harbor construction and activity increased (Sec. II.A).

Although there was substantial overlap in the nonindigenous and cryptogenic populations of the 
five south and west shore harbors of the present study with Pearl Harbor, this overlap was not 
complete. Seventy-one of the 100 species in the five harbors also occurred in Pearl Harbor, 30 
species which occurred in the present studys five harbors were not in Pearl Harbor, and 27 in 
Pearl Harbor were not in the five harbors.  Even less overlap occurred for the new introductions, 
where only five of the 25 species found for the two studies occurred in Pearl Harbor and at least 
one other harbor and only two occurred in all harbors (Table 8).  This suggests that most 
introduced species have remained highly restricted in their habitat, achieving only low abundance 
in limited locations.  The sampling effort of the present study was equivalent or greater to that 
conducted in Pearl Harbor, and sampling and most identifications were completed by the same 
researchers.  It is therefore unlikely that sampling or analytic inconsistencies contributed 
substantially to finding different assortments of introduced or cryptogenic species in the two 
studies.

Despite this degree of difference between the introduced species determined for the two studies, 
similar results were found for their probable origins or previous distributions.  The central and 
west Indo-Pacific and Red Sea accounted for 42% of the introduced species in the present study, 
compared to 39% for Pearl Harbor (Coles et al. 1997).  Only 4% were from the eastern Pacific in 
the present study compared to 3% for Pearl Harbor, while the Atlantic and Caribbean accounted 
for 16% in the present study and 13% for Pearl Harbor.  Both surveys therefore suggest a 
dominance of marine introductions in Hawaii by organisms originating in the waters of the tropical 
Indo-Pacific, with a large component coming from south and east Asia.

Both the Pearl Harbor and the present study indicate that nonindigenous and cryptogenic species 
are a major component of the total marine biota of the harbors and suggest that introductions 
have been occurring for a long time.  Nonindigenous and cryptogenic species composed 17% of 
the total biota in the present study and 22-23% of the total in Pearl Harbor (Coles et al. 1997, 
1999).  Based on first reports, many of these have been in Hawaii since early in the century. In 
Pearl Harbor, where sufficient sampling has been conducted throughout the century to support a 
conclusion, introductions have been estimated to have occurred at a rate of about 5-10% of the 
total population except during wartime activity (Coles et al. 1999).  This is similar to the 
introduction rates for the present study in the 1970s and 1990s when sufficient data are available 
to sustain such an estimate (Figure 22).

Although nonindigenous species are a major component of the total biota in these harbors, there 
is little evidence of recent introductions that have proliferated to a degree that they constitute a 
nuisance situation, monopolize habitats or negatively impact native organisms or commercially 
important species.  The small intertidal barnacle Chthamalus proteus is the only newly introduced 
organism that is abundant throughout all of the harbors and elsewhere in Hawaii (Southward et 
al. 1998).  Of the remaining 25 nonindigenous species that have been newly detected since 1995 
(Table 8) only the sponge Gelliodes fibrosa, the amphipod Leucothoe micronesiae and the 
bryozoan Bugula dentata occurred at multiple stations throughout most of the five harbors.  Many 
other nonindigenous species such as the octocoral Carijoa (=Telesto) riisei, the hydrozoan 
Halocordyle disticha and the bryozoans Amathia distans and Schizoporella sp. A (previously 
called S. errata or S. unicornis but presently under review, C. Zabin pers. comm.) are frequently 
found and abundant components of harbor fauna throughout the Hawaiian Islands.  However, 
these are introductions dating back many decades which have apparently had minimal success 
invading environments outside of harbor areas.  This contrasts strongly with numerous 
circumstances in temperate regions where nonindigenous marine species have invaded multiple 
habitats causing a variety of negative impacts on local species and commercial fisheries (see 
Ruiz et al. 1997 for review). 

There is very limited information on nonindigenous species introductions into tropical marine 
systems to compare with these Oahu studies.  However, a recent comprehensive survey of 
twelve commercial ports in North Queensland, Australia, and comparison of the results of this 
study with findings by the Centre for Research on Introduced Marine pests (CRIMP) at temperate 
Australian ports provides valuable information for comparison (Hilliard et al. 1997).  The 
Queensland ports are in a prototypical tropical environment ranging 12-25?S latitude along the 
northeast Australian coastline inshore of the Great Barrier Reef.  Substantial export of bulk cargo 
occurs from these harbors, therefore the harbors receive large volumes of ballast water 
discharge, exceeding the volumes discharged at temperate southern Australian ports (Hilliard et 
al. 1997).

Comprehensive surveys of the twelve Queensland harbors detected a total of only 15 
nonindigenous and 15 cryptogenic marine species, with none of these considered being of 
nuisance status or potentially damaging to the resident communities.  By contrast, nonindigenous 
species reported for all Australian ports and harbors by CRIMP surveys totaled 139, with an 
additional 31 species designated as cryptogenic.  These 170 nonindigenous and cryptogenic 
species included nine found in the present study.  The number of nonindigenous species 
occurring in each of the Australian states is shown in Table 9, along with comparative data for the 
two Oahu studies and information available for temperate systems throughout the world.

Table 9. Numbers of marine nonindigenous species found in various world locations 

Greater abundance of nonindigenous species in temperate areas is suggested, both worldwide 
and within Australia.  The Hawaiian reports of around 70 nonindigenous species are in the middle 
of the range of findings, and between those of New South Wales and Victoria.  Although Hawaii 
is located within the tropics, the physical environment may be considered subtropical in regard to 
seasonality and prevailing water temperatures, conditions which also apply to much of New South 
Wales.  The Hawaiian environment is therefore likely to be amenable to introduced species from 
temperate zones, and temperate organisms are a substantial component of the nonindigenous 
species reported in Oahu harbors.  By contrast, the Queensland marine environment is entirely 
tropical, and only nine of the fifteen nonindigenous species found in Queensland occur in 
temperate areas elsewhere in Australia (Hilliard et al. 1997).

The limited available information therefore tentatively supports conclusions that tropical areas are 
less receptive to marine invasions, and that nonindigenous species that do survive in tropical 
environments are less likely to cause disruptions in the ecology or economy of the receptor 
environment.  A number of hypotheses are considered in Hilliard et al (1997) to explain a 
proposed paradigm of greater resistance by tropical marine systems to invasions, which will not 
be considered here.  It should be noted, however, that this model does not consider the serious 
impacts that have resulted from the massive proliferation in Hawaii of the introduced red algae 
Hypnea musciformis, Kappaphycus alvarezii, K. striatum and Gracilaria salicornia (Abbott 1987; 
Russell 1983,1992; Rodgers 1997; Rodgers and Cox, in press).  Such outbreaks of nonidigenous 
algae are unreported elsewhere in the tropics, but are similar to algal invasion in the 
Mediterranean Sea where the green algae Caulerpa taxifolia has proliferated unrestricted since 
its introduction in the early 1980s (Meinesz et al. 1993; Siguan 1996).

It is apparent that more data on nonindigenous species are needed from other tropical areas to 
confirm or deny a paradigm of resistance to invasions by tropical marine communities.  A study 
nearing completion on nonindigenous species in the harbors of Guam (Paulay et al. in prep.)  and 
surveys proposed for Lautoka and Suva Harbors in Fiji hopefully will provide information usable 
for this purpose.

VI. CONCLUSIONS AND MANAGEMENT CONSIDERATIONS

The present study is the first to comprehensively examine the benthic and fish biota of the five 
main, non-military harbors of Oahu, which are the major points of entry to the Hawaiian Islands 
for commercial shipping, fishing vessels and long-range sailing craft that might transport 
nonindigenous species into Hawaiian waters.  The present studys surveys were conducted within 
one year of a similar comprehensive study of Pearl Harbor, the center of U.S. Naval ship 
movement in the Pacific.  The combined information from these studies provides a complete 
picture of the present conditions of nonindigenous and cryptogenic species in major Oahu 
harbors, and the impacts that these introductions have had on resident biota within these harbors.

The findings of the present study are remarkably similar to those of the Pearl Harbor surveys 
conducted 1-2 years previously, with a total of about 100 nonindigenous and cryptogenic species 
and about 12 new nonindigenous species found in both studies.  Contrary to many areas in 
temperate regions in the world, there is little indication of recent increases in nonindigenous 
species introductions into Oahu harbors, nor of proliferation of species impacting native 
populations or commercially important species that would constitute a nuisance invasion.  
Although nonindigenous and cryptogenic species make up a larger component of the total biota 
of harbors than has been found in other tropical areas, most of the more common introduced 
species have been in Hawaiian waters for much of the century, and most recent introductions 
have remained restricted in the occurrence and are infrequently found.

Although these findings provide encouraging information about the limited impact that 
nonindigenous species have had in Oahus harbors, it should not be construed that no potential 
exists for marine introductions that would be damaging to Hawaiis marine communities.  The 
example of the barnacle Chthamalus proteus, which has arrived in Hawaiian waters in the last 25 
years and now dominates the shallow intertidal in harbors and embayments throughout the 
Hawaiian chain, represents a warning that a more damaging nonindigenous species is capable of 
invading Hawaii at any time.  Nor can the information found for these harbors be extrapolated to 
other marine environments in Hawaii such as coral reefs, rocky shores or sandy beach 
communities.  The serious impact that nonindigenous algae have had on reef areas and 
shorelines on the south and west shores of Maui, along Ewa Beach on Oahu, and in Kaneohe 
Bay is perhaps the best example of how each environment is subject to a variety of 
nonindigenous species invasions.  In order to assure that invasions are detected early enough 
that action can be taken to reduce impacts, baseline conditions must be determined for specific 
locations, and monitoring programs should incorporate methodologies for nonindigenous species 
detection.  Pursuant to this need, nonindigenous species baseline assessment surveys have 
been proposed for Kaneohe Bay, Kuapa Pond and Waikiki as areas of transition between harbors 
and open ocean coral reefs.

As an example of the need for continued vigilance and an active program for assessing 
nonindigenous marine introductions, a recent occurrence in the area of Darwin Harbor, Northern 
Territory, Australia is instructive.  In March, 1999, during a resurvey of a marina being conducted 
as part of a baseline assessment program, CRIMP researchers discovered a proliferation of the 
nonindigenous mussel identified as Mytilopsis (=Congeria) sallei.  This species is similar to the 
infamous freshwater Zebra Mussel Dreissena polymorpha which has infested all the waterways of 
the eastern and central United States, causing millions of dollars in economic costs and 
enormous ecological damage.  It is known that the introduction of M. sallei was recent, since it 
had not occurred in the area during a survey conducted six months earlier, and further 
investigation found that it had already spread to two other marinas in the area.  Fortunately, 
because the mussels had been detected early, it was possible to conduct an eradication program 
using sodium hypochlorite and copper sulfate in the marinas which apparently was affective in 
eliminating the invading organisms. 

Detection of this nonindigenous mussel was possible only because an active program conducted 
by knowledgeable personnel was underway, and because knowledge of baseline conditions had 
been established.  It is important for the protection of Hawaiis marine communities from 
nonindigenous invasions that surveys be made of the baseline conditions in critical areas, and 
that efforts continue in the detection and assessment of introduced marine organisms.

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ACKNOWLEDGMENTS

This study was conducted with the financial support of the David and Lucile Packard Foundation 
and Dingle-Johnson Act Funds administered through the State of Hawaii Department of Natural 
Resources Division of Aquatic Resources.  Special thanks to the administrations of those 
organizations for making these funds available. The assistance of the management and staffs of 
the Library and Department of Natural Sciences at Bishop Museum is most appreciated, 
especially the efforts of Richard Pyle, who developed the database and assisted in analyses, 
Arnold Suzumoto, who managed the specimen collections, Pakki Naugle and Jonathan Taylor, 
who provided sorting assistance, and Jeremy Park, who assisted in crab identifications. The 
University of Hawaii Hamilton Library, the Pacific Maritime Center, Donn Fukuda of Hawaiian 
Electrics Environmental Department and Eric Guinther , president of AECOS Inc., provided 
access to unpublished reports and other valuable information from their respective libraries.  Dr. 
James Carlton provided continued input and stimulation. Kim Beasley of Clean Island Council, 
Honolulu generously provided the aerial photos used on the cover.

Taxonomic expertise for identifying organisms was provided by the following individuals, and their 
efforts and contributions to this project are gratefully acknowledged.
Algae: Mr. Jack Fisher, Bishop Museum 
Zoantharians:  Dr. Daphne Fautin, University of Kansas
Pycnogonids: Dr. C. Allan Child, U.S. National Museum of Natural History
Isopods and Tanaids Dr. Brian Kensley, U.S. National Museum of Natural History
Caprellid Amphipods: Dr. Ichiro Tekeuchi, University of Tokyo
Barnacles: Dr. William Newman, Scripps Institution of Oceanography
Molluscs: Ms. Regie Kawamoto, Bishop Museum
Bryozoa: Ms. Chela Zabin, University of Hawaii
Ascidians: Dr. Gretchen Lambert, California State University at Fullerton and Mr. Scott 
Godwin, Bishop Museum
Fish: Mr. Arnold Suzamoto, Bishop Museum



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