#### 30 April 2004 {#april-2004 .heading-style1 .unnumbered}

Alan M. Friedlander

NOAA, National Ocean Service,

National Centers for Coastal Ocean Science - Biogeography Program

&

Oceanic Institute

Makapu'u Point/41-202 Kalanianaole Hwy, Waimanalo, Hawai'i 96795.

E-mail: <afriedlander@oceanicinstitute.org>

Herman Cesar

Cesar Environmental Economics Consulting

Kastanjelaan 9\
6828 GH Arnhem\
The Netherlands

E-mail: <herman.cesar@ivm.vu.nl>

Web: www.ceec.nl

Contents

1\. Introduction [3](#introduction)

2\. Marine Protected Area Theory and Empirical Evidence
[4](#marine-protected-area-theory-and-empirical-evidence)

3\. Description of Study Sites [4](#description-of-study-sites)

4\. Empirical Information On Fish Assemblage Characteristics
[4](#empirical-information-on-fish-assemblage-characteristics)

5\. Empirical Information on MPAs in Hawaii -- Overall Comparison of
Protected Areas
[4](#empirical-information-on-mpas-in-hawaii-overall-comparison-of-protected-areas)

6\. Recommendations for Modifications to Existing MPAS and for the
Design and Siting of Future Protected Areas
[4](#recommendations-for-modifications-to-existing-mpas-and-for-the-design-and-siting-of-future-protected-areas)

7\. References [4](#references)

Appendix I. Top ten species observed on transects for the Six Study
Sites
[4](#top-ten-species-observed-on-transects-for-the-six-study-sites)

  -----------------------------------------------------------------------
  ***Colophon***

  This study is part of a larger report with the title "Assessment of
  Economic Benefits and Costs of Marine Managed Areas in Hawaii\" by
  Herman Cesar, Pieter van Beukering and Alan Friedlander. This
  publication is a result of research carried out by Cesar Environmental
  Economics Consulting (CEEC) and funded by the National Oceanic and
  Atmospheric Administration, Coastal Ocean Program, under awards NA
  160A2412 to the University of Hawaii for the Hawaii Coral Reef
  Initiative Research Program. Co-funding was obtained from the Division
  of Aquatic Resources (DAR) and the Department of Business, Economic
  Development & Tourism (DBEDT). This specific study was compiled by Alan
  M. Friedlander, under a subcontract with CEEC.
  -----------------------------------------------------------------------

# Introduction

#### Declining fisheries resources worldwide and in Hawaii 

Fish and other harvested resources provide economic, social, cultural,
and spiritual benefits to people throughout the world. Global fisheries
have undergone enormous changes in the past several decades with
three-quarters of the world's major fisheries now considered to be
harvested at or beyond their maximum capacity (FAO 2000) despite
increased regulation in most fisheries sectors. The U.S. has some of the
most highly regulated fisheries in the world with substantial
investments in science, management plans, monitoring, and enforcement
(Eagle et al. 2003), yet 40% of U.S. fisheries are considered overfished
(NMFS 2002). Examples include the collapse of cod stocks off New England
and Canada, which has led to large-scale ecosystem-wide changes, and
economic collapse of many coastal communities in these areas (Kurlansky
1998). Because selective fishing can affect a number of population
characteristics (e.g. size and age composition, sex ratio, genetic
make-up, and large-scale behavioral phenomena like spawning
aggregations) those fish that remain may pass on less-preferred
characteristics that may confound future fishing efforts (Sladek Nowlis
and Friedlander 2004a). In addition to overexploitation, a variety of
factors such as habitat loss, climate change, and natural variability
have contributed to the collapse of many fisheries. Regardless of the
causes, declining fish stocks have had negative economic, social, and
ecological consequences at an unprecedented scale.

Large marine vertebrates such as whales, sharks, turtles, groupers, and
manatees were once important components of marine ecosystems worldwide
but have been systematically removed from the ocean by humans over the
pat 500 yr (NRC 1995, Jackson et al. 2001, Pitcher 2001). Large
predatory fish have been reduced to one-tenth of their historic
abundance (Myers and Worm 2003) and these top predators have specialized
niches that when depleted can lead to a phase transition of ecosystems
dominated by lower trophic guilds (Pauly et al. 1998, Pinnegar et al.
2000). The 'shifting baseline syndrome' (Pauly 1995, Sheppard 1995)
makes it difficult to determine what constitutes a natural ecosystem and
how to mange these ecosystems accordingly.

As is the case elsewhere throughout the world, coastal fisheries in
Hawai'i are facing unprecedented overexploitation and severe depletion
(Shomura 1987, Smith 1993, Gulko et al. 2000, Friedlander 2003, Lowe
2003). This decline in fish abundance and size, particularly around the
more populated areas of the state, is likely the cumulative result of
years of chronic overfishing (Shomura 1987; Gulko et al. 2000;
Friedlander and DeMartini 2002). Fishing pressure on nearshore resources
in heavily populated areas of the main Hawaiian Islands (MHI) appears to
exceed the capacity of these resources to renew themselves (Smith 1993).
Fish assemblages in the northwestern Hawaiian Islands---a remote area
that experiences only limited fishing activity---are dominated by large
apex predators, such as sharks and jacks that likely have a profound
impact on the structure of the entire coral reef ecosystem (Friedlander
and DeMartini 2002). The near-extripration of apex predators and heavy
exploitation of lowed trophic levels in the MHI from intensive fishing
pressure has resulted in a stressed ecosystem that does not contain the
full complement of species and interrelationships that would normally
prevail (Friedlander and DeMartini 2002) (Figure 1)

![](media/image1.png){width="5.683333333333334in"
height="3.7111111111111112in"}

Figure 1: Lightly fished areas in the Northwestern Hawaiian Islands
contain greater biomass of all trophic guilds than sites in the main
Hawaiian Islands (data from Friedlander and DeMartini 2002). The
difference is especially large for apex predators, which account for the
majority of all biomass in the Northwestern Hawaiian Islands.

Factors contributing to the decline of inshore fisheries in the MHI
include a growing human population, destruction or disturbance to
habitat, introduction of new fishing techniques (inexpensive
monofilament gill nets, SCUBA, spear guns, power boats, sonar fish
finders), and loss of traditional conservation practices (Brock et al.
1985, Lowe 1996, Birkeland and Friedlander 2002, Friedlander et al.
2003). The proliferation of long and inexpensive gill nets has allowed
new fishers to enter the fishery and set nets deeper and in locations
not previously harvested (Clark and Gulko 1999). Intensive fishing
pressure on highly prized and vulnerable species has led to substantial
declines in catch as well as size and has raised concerns about the
long-term sustainability of these stocks (Friedlander and Parrish 1997,
Friedlander and DeMartini 2002, Friedlander and Ziemann 2003, Tissot and
Hallacher 2003, Tissot et al. 2004). Despite the opinion of many
fishermen that overharvesting is one of the major reasons for the
long-term decline in inshore marine resources, there is poor compliance
with state fishing laws and regulations (Harman and Katekaru 1988). The
lack of marine-focused enforcement and minimal fines for those few cases
that have been prosecuted contribute to a lack of incentive by the
population to abide by fisheries management regulations.

Under-reporting by commercial fishers and the existence of a large
number of recreational and subsistence fishers without licensing or
reporting requirements have resulted in uncertainty in actual fisheries
catch statistics for the state (Lowe 1996). The nearshore recreational
and subsistence catch is likely equal to or greater than the nearshore
commercial fisheries catch, with more species taken using a wider range
of fishing gear (Friedlander and Parrish 1997, Gulko et al. 2000,
Everson and Friedlander 2003).

#### Why has conventional management failed? 

Sustainability and biodiversity conservation are cited as objectives of
ecosystem management (National Research Council 1999) yet current
management regimes have failed to meet these objectives. Failures in
management can be linked to uncertainties due to incomplete knowledge of
populations, communities, and ecosystems (Fogarty et al. 2000).
Traditional western concepts of fisheries management (maximum
sustainable yield, growth overfishing, recruitment overfishing, etc.)
have their genesis in single-species population dynamics and stock
assessment (Murawski 2000). This approach is primarily concerned with
the conservation of the parts as opposed to the interrelationships among
them. Many management tools---including size limits, gear limits, quota
systems on effort or total catch, and even temporary closures---are used
frequently but do not create a refuge for populations, habitats, and
ecosystems, nor do they reallocate fishing effort across space (Sladek
Nowlis and Friedlander 2004a). The failure of these more conventional
management tools is apparent in the status of fished populations around
the world. Several challenges contribute to these management failures,
including excessive fishing capacity (FAO 2000), environments degraded
by fishing and other activities (Watling and Norse 1998), and management
systems that require far more information than is available (NRC 1998;
PDT 1990; Sladek Nowlis and Bollermann 2002). Another concern is the
possibility of critical depensation (Allee effect) where the per capita
birth rate declines at low populations because, for example, of the
increased difficulty of finding a mate (Allee 1931).

[]{.mark}

# Marine Protected Area Theory and Empirical Evidence

Because of the generally poor state of fisheries worldwide, marine
resource managers are inspired to consider fresh tools to stem the
decline in global fish stocks (FAO 1999). Marine ecosystems are complex
with highly variable natural replenishment and therefore require more
spatially-based management tools. The Food and Agriculture Organization
(FAO) has established a code of conduct that supports the precautionary
principle, which states should apply to conservation, management, and
exploitation of living aquatic resources in order to protect them and
preserve the aquatic environment (FAO1995). It states that the \'absence
of adequate scientific information should not be used as a reason for
postponing or failing to take conservation and management measures.\'

Theoretical evidence has been available for decades on policies that
scale back fishing rates when abundance drops. One means for achieving a
constant escapement-like policy is the use of marine reserves to protect
part of the stock (Figure 2). What reserves offer that other management
tools cannot is the ability to control fishing rates in a manner that is
relatively easy to enforce (PDT 1990; Sladek Nowlis and Friedlander,
2004a) and requires relatively little scientific information (Sladek
Nowlis and Bollermann 2002). In addition, reserves prevent habitat
disturbance due to fishing, protect non-target organisms, and preserve
biodiversity (Bohnsack 1996, Murray et al. 1999, Fogarty et al. 2000).
In this paper, we will refer to these reserves as Marine Protected Areas
(MPAs) as is common in the fisheries literature. In the accompanying
papers of this study, we will refer more broadly to Marine Managed Areas
(MMAs) to highlight that these areas tend to have other management
aspects than conservation and fisheries, such as tourism.

![](media/image2.jpeg){width="5.7034722222222225in"
height="4.959722222222222in"}

Figure 2 Marine reserves as a tool to achieve constant escapement. The
dotted line represents estimated productivity, and the solid lines
represent policies that link marine reserves with catch quotas. The
centerline represents the use of a reserve encompassing 30% of the
unfished stock biomass and fairly aggressive fishing on the remaining
stock. Line (a) represents a relatively smaller reserve, with a
more-controlled fishing effort but less insurance, while line (b)
represents a larger reserve and even more aggressive fishing pressure
and greater insurance. The optimal policy will depend primarily on the
degree to which the biology, ecology, current abundance, and fishing
mortality rate of the stock or stocks are unknown, and thus amount of
insurance that is desirable or necessary (from Sladek Nowlis and
Bollermann, 2002).

#### Benefits of MPAs

Benefits of MPAs are:

-   Increase stock abundance - Theory supports the ability of marine
    reserves to rebuild overfished fisheries and enhances catches
    (Beverton and Holt 1957; DeMartini 1993; Polacheck 1990; Sladek
    Nowlis and Roberts 1999) and several empirical studies have shown
    increased catch despite 25% to 40% reserve closures (McClanahan and
    Kaunda-Arara 1996, Russ and Alcala 1999, Roberts et al. 2001). Catch
    around Apo Island in the Philippines has increased since the early
    1980's despite the closure of more than 10% of the fishing area
    (Russ and Alcala 1999, Figure 3). In contrast, Sumilon Island (25%
    no-take reserve), also in the Philippines, showed a decline in catch
    following the opening of the reserve to fishing. Despite the
    increased fishing area, both total catch and catch per unit effort
    declined to half of their previous values Subsequent closures showed
    slight increases in catch but these catches were much smaller than
    those observed when a complete closure of 25% was in place;

-   Preserve desirable traits - Selective fishing can affect a number of
    population characteristics---size and age composition, sex ratio,
    genetic make-up, and large-scale behavioral phenomena like spawning
    aggregations (PDT 1990), while reserves have been shown to preserve
    these traits;

-   Provide spillover of adults and juveniles into fished areas --
    Movement can serve as a mechanism for exporting productivity from
    marine reserves to fishing areas. Johnson and colleagues (1999)
    demonstrated that, in addition to a build-up of biomass within a
    reserve off Cape Canaveral, Florida, some fish moved in and out of
    the reserve. Consequently, a number of world record trophy fish were
    caught in the vicinity of the reserve (Roberts et al. 2001);

-   Increase reproductive output and recruitment inside and outside the
    reserve; - The protection of adult biomass greatly increases
    reproductive potential (Polacheck 1990; DeMartini 1993; Guenette and
    Pitcher 1999) and therefore spawning output. Because adult retention
    and population growth rates provide the engine to power all
    population-level benefits, leakiness of adults (spillover effect)
    can have negative consequences to the population (Sladek Nowlis and
    Roberts 1999). Emerging evidence about fish movement suggests that
    even fish with the potential to swim long distances might stay in
    the same area for long periods of time (e.g., Attwood and Bennett
    1994; Holland et al. 1996). In support of this notion is the fact
    that most populations studied in marine reserves responded
    positively to protection, even though many of the reserves were
    small (Halpern 2003);

-   Insurance against uncertainty - Most fisheries, even those that are
    actively managed and well studied, are prone to crashing because
    management reference points have a high likelihood of being off by
    50 percent or more (NRC 1998). By protecting a set amount of fish,
    marine reserves have shown strong potential to protect stocks from
    collapse in varying and uncertain environments (Lauck et al. 1998;
    Mangel 1998; Sladek Nowlis and Bollermann 2002);

-   Reduce overfishing by controlling fishing mortality (Sladek Nowlis
    and Bollermann 2002);

-   Ecosystem management - By reallocate fishing effort in space and
    protect populations, habitats, and ecosystems within their borders,
    marine reserves provide a spatial refuge for the ecological systems
    they contain (Sladek Nowlis and Friedlander 2004a);

-   Maintain system productivity - Destructive fishing practices can
    disturb habitats essential to fisheries production (Watling and
    Norse 1998, Morgan and Chuenpagdee 2003). System productivity can
    also reduce by fishing activity through the disruption of species
    interactions (Jackson et al. 2001);

-   Provide unfished reference areas - distinguish between natural and
    fishery-related changes in marine systems, dramatically limiting a
    manager's ability to explain past events and predict future ones.

![](media/image3.jpeg){width="5.854861111111111in"
height="3.7916666666666665in"}

*Figure 3: Mean number (left columns at each time) and mean biomass
(right columns) of large predatory reef fish (Sub-F. Epinephelinae, F.
Lutjanidae and F. Lethrinidae) per 1000 m2 in the Sumilon and Apo
reserves in the Phillipines from 1983-1993. The Sumilon reserve had been
protected from fishing for almost 10 years in 1983. Solid arrows
indicate when fishing in the reserve began (1984, 1992) and the open
arrow indicates when fishing ceased (1987). (Russ and Alcala 1999).*

#### Biodiversity and conservation benefits 

Biodiversity can be defined as the variability among living organisms
from all sources including terrestrial, marine and other aquatic inter
alia ecosystems and the ecological complexes of which they are part;
this includes diversity within species, between species and of
ecosystems (de Fontaubert et al 1996). All except one of the presently
described phyla (33) occur in the ocean while only about half that occur
on land (Norse, 1993). Furthermore, 15 phyla are exclusively marine.
Consequently, marine organisms display a much larger phyletic diversity
than those on land (Ray, 1988). In addition to its intrinsic value,
maintaining marine biodiversity is important to the success of
humankind. Norse (1993) detailed the importance of marine biodiversity
to humans as follows:

-   Food

-   Medicine and tools for biomedical research

-   Coastal processes and protection

-   Global climate control

-   Recreational resources

-   Knowledge

It has been shown that marine reserves can halt the loss of biodiversity
and changes in species interaction too commonly seen under current
management strategies by conserving habitats and biological communities
(Dayton et al. 1995; Boehlert 1996; Hixon and Carr 1997). Instead of
focusing on a system with representative species, we should focus on
representative habitats or ecosystems. Ecosystem-level management may be
best achieved by acknowledging the ecological phenomena we know in
management decisions while also allowing natural systems to function
naturally in designated marine reserves.

Endemism is a key attribute of biotic communities. One reason of general
biogeographic interest is that speciation and the origin and maintenance
of biodiversity are undoubtedly related to degrees of isolation and
endemism. The faunas of isolated oceanic archipelagos like the Hawaiian
Islands represent species conservation hotspots that have become
increasingly important due to the continual losses of fish and other
organismal biodiversity on coral reefs (Roberts et al 2002). That over
one-half (by numbers and biomass, not merely species-presence) of the
fishes on shallow NWHI reefs are endemic to the Hawaiian Islands is a
noteworthy finding, given our contemporary appreciation of the nature
and magnitude of species loss in the sea (DeMartini and Friedlander in
press).

#### MPA design 

A comprehensive zoning process may be preferable to solely designating
marine reserves because additional zone types can reduce a number of
additional user conflicts while also providing buffers to protect marine
reserves from inevitable edge effects. Marine reserve networks have the
greatest chance of including all species, life stages, and ecological
linkages if they encompass representative portions of all ecologically
relevant habitat types (Ballantine 1997; Friedlander and Parrish 1998;
Murray et al. 1999). Two of the most basic tenets of marine reserve
network design are that all habitats be represented and that the
resulting network be self-sustaining (Ballantine 1997).

##### Stakeholder Driven Process

The input of people from coastal communities will be vital to good
design and function of marine reserves and to maintain public support
for them. Without this support, enforcement may be greatly compromised
and future political changes can lead to the dismantling of reserves
(Alcala and Russ 1990). Participatory planning represents a bottom up
approach to resource management that incorporates local knowledge into
the management process. Incorporating stakeholder group concerns and
interests into the management process will increase the perceived
legitimacy of decisions and make compliance with rules and regulations
easier (Bunce et al. 2000). Stakeholder input provided invaluable
information on resource distribution and their uses (Johannes 1997,
Friedlander et al. 2003b). By airing concerns and priorities,
stakeholders also develop a sense of ownership and a better
understanding of the management process. Where reserves have been
established without broad public support, they are vulnerable to
dismantling when politics shift (e.g., Russ and Alcala 1999) or in
danger of never being created in the first place.

##### Total coverage

Fished populations that fall to less than 30% of their unexploited
population size are legally considered to be overfished in the U.S.
(NMFS Magnuson Stevens). At this spawning potential ratio (the ratio of
total egg production under fishing to total egg production without
fishing), egg survival must be 3.3.times its natural survival to
maintain the population (Bohnsack et al. 2003). Studies of fish movement
and habitat utilization suggest that protecting 20-30% of the habitat
would likely protection 20-30% of the population (Bohnsack et al. 2003,
Sladek Nowlis and Friedlander 2004a). Based on empirical evidence from
temperate and boreal fish populations, a minimum of 20-35% of the
spawning biomass should be preserved to conserve viable population size
(Mace and Sissenwine 1993). The greater the impacts outside the reserve,
the larger the reserve size required.

One means for achieving a constant escapement-like policy is the use of
marine reserves to protect part of the stock. In order to ensure
productive catches into the future, models have predicted that reserves
might have to encompass over half the management area (Lauck et al.
1998; Mangel 1998). Reserve networks need to protect a population
consisting of 30 to 50% of its unfished size to insure against collapses
in the face of large uncertainties (Lauck et al. 1997; Mangel 2000;
Sladek Nowlis and Bollermann 2002).

##### Size and shape

If individual reserves are too few and large, export of production from
reserves to fishing grounds may be limited. If, on the other hand,
reserves are universally small, they may provide only limited protection
for most species. Suggestions of at least 10 km2 per individual reserve
have been proposed to contain viable populations of a wide range of
species (Friedlander et al. 2003b). Reserves should be designed to
minimize the perimeter to area ration in order to reduce the edge
effects outside the reserve (Fogarty et al. 2000). Higher perimeter to
area ratios may be desirable to accrue adult spillover and larval
replenishment (Fogarty et al. 2000). Straight boundaries that run
north-south or east-west are ideal, while complex shapes---like
following a depth contour---are difficult to recognize or enforce.
Replication of reserve units and selection of appropriate replicated
control (i.e., open to fishing) areas will aid greatly in the evaluation
of reserve success.

##### Area selection and habitat types 

Representative proportions of all habitats types should be included in
any reserve (Ballantine 1997). To be effective, it is generally accepted
that MPA networks should be distributed along environmental gradients
and should protect representative species and habitat types (Ballantine
1997; Murray et al. 1999), although rare and vulnerable habitat types
should be represented more fully (Sladek Nowlis and Friedlander 2003a).
It may also be beneficial to focus marine reserve efforts on known
ecological connections among habitat types (Appeldoorn et al. 2003).
Reserves should provide habitat connectivity for ontogenetic migration,
typically from shallow to deeper sites (Parrish 1987, Appeldoorn et al.
2003). Reserve design should also consider diel movement patterns such
as the movement of many species from daytime resting habitat on the reef
to nocturnal foraging in soft bottom habitats (Holland et. al 1996,
Meyer 2003, Friedlander et al. 2002). Choosing areas that are
characterized by diverse habitats may foster these ecological
connections and increase the capacity of even very small reserves to
sustain productive populations within their borders (Appeldoorn et al.
1997).

##### Networking

An effective reserve design, i.e., reserves that sustain protected
populations and enhance non-protected populations, will benefit from our
understanding of patterns of larval dispersal and how species attributes
(e.g., larval behavior, developmental traits) and environmental features
(e.g., hydrographic patterns, geomorphological features) influence such
patterns (Cowen et al. 2000, Jones et al. 1999, Roberts 1997, Swearer et
al. 1999). Self-replenishment can be achieved by reserves of sufficient
size to contain a substantial amount of larval dispersal, or by
networking reserves at suitable distances such that propagules produced
by populations in one reserve replenish populations in other reserves.
Individual reserves are likely to be most effective at supplying
productivity to fished areas if there are several small, networked sites
(Sladek Nowlis and Roberts 1999). Marginal decreases in the perimeter to
area ratio with increasing reserve area declines at larger sizes
suggesting diminishing returns at larger reserve sizes (Fogarty et al.
2000). This suggests that several smaller reserves may be more
beneficial than one large one particularly when one is concerned with
limiting the exposure at the boundaries.

Sources are areas that produce a net gain of fish, some of which move at
some point in their life cycle out to other areas. By contrast, sinks
are areas that experience a net loss of fish and are only sustained
because they are supplemented from sources. Crowder and colleagues
(2000) showed that managers could theoretically do more harm than good
if they created a marine reserve network that encompassed more sinks
than sources. A major flaw with their reasoning, though, was the
assumption that sources and sinks are static---that they do not change
with the establishment of marine reserves (Sladek Nowlis and Friedlander
2004b). In fact, strong evidence supports the fact that reserves become
sources most of the time they are established by creating an area with
many fish producing lots of offspring (Appeldoorn et al., 2003).

Dispersal distances for marine larvae ranges from meters to thousands of
kilometers with time in the plankton ranging from minutes to months.
Examination of a wide range of taxa found dispersal to follow a bimodal
distribution with one group of propagules dispersing \< 1 km propagules
and another group that dispersed \> 20 km. (Shanks et al. 2003). These
authors suggested that reserves be designed large enough to contain the
short distance dispersing propagules and spaced far enough apart that
long distance dispersing propagules released from one reserve can settle
in adjacent reserves. A reserve 2 km in diameter, would contain all the
larval types that disperse \<1 km/yr while reserves 4 to 6 km in
diameter would be sufficient for maintenance of larger adult
populations. A reserve 4 to 6 km in diameter should be large enough to
contain the larvae of short distance dispersers and reserves spaced 10
to 20 km apart should be close enough to capture propagules released
from adjacent reserves.

One can manage using a quasi-MSY approach by establishing a network of
marine reserves likely to encompass the proportion of the biomass of
populations of concern necessary to sustain maximum yields (Sladek
Nowlis and Bollerman 2003). With this approach, one uses marine reserve
to provide insurance for future catches, even if the biology of the
system is poorly understood. This approach may be particularly helpful
when traditional management will not work due to gaps in knowledge or
inability to enforce more subtle management measures, situations when
insurance will be of greatest value.

# Description of Study Sites 

#### Hanauma Bay, Oahu

*General description:* Hanauma Bay is located on the southeast tip of
Oahu and comprises an area of ca. 101 acres. It is an enclosed embayment
formed through the partial collapse of two volcanic craters and
subsequent erosional processes. High cliffs surround the sandy beach
that is partially composed of olivine. The back-reef is composed largely
of sand and coral rubble. The reef flat is composed of carbonate with
low coral cover and receives heavily impacted by human use. The deeper
reefs with more extensive coral cover extend out to the mouth of the bay
to 30-meter depths. The bay has a southern exposure but is sheltered
from most major oceanic swells. Hanauma Bay was popular with Hawaiian
royalty and a favorite fishing spot for King Kamahameha V. It is one of
the most heavily used marine preserves in the world, drawing over one
million visitors per year.

Approximately 25% of the marine habitat of Hanauma Bay consists of
colonized pavement (\>10% live coral cover) with an additional 24%
consisting of sand (Table 1). Linear reef (12%) and uncolonized pavement
with less than 10% live coral cover (11%) also comprise important
components of the benthic habitat.

Table 1: Habitat types contained within Hanauma Bay MLCD based on NOAA
benthic habitat maps. (Coyne et al. 2001)

  ---------------------------------------- ------------ -------- ------------
  Habitat Type                             Square Meter Acres    Proportion
                                                                 of total

  Reef/Colonized Pavement                  100937.02    24.94    0.25

  Sand                                     98402.37     24.32    0.24

  Reef/Linear Reef                         47234.20     11.67    0.12

  Hardbottom/Uncolonized Pavement          43881.07     10.84    0.11

  Unknown                                  36382.74     8.99     0.09

  Reef/Colonized Volcanic Rock/Boulders    29280.45     7.24     0.07

  Hardbottom/Uncolonized Volcanic          29280.45     7.24     0.07
  Rock/Boulders                                                  

  Reef/Aggregate Coral                     27300.26     6.75     0.07

  Reef/Patch Reef (Individual)             17135.27     4.23     0.04

  Total                                    7128.69      1.76     0.02
  ---------------------------------------- ------------ -------- ------------

*Fisheries Regulations and Use:* Hawaii Administrative Rules Title 13,
DLNR, Subtitle 4 Fisheries, Part 1, MLCDs, Chapter 28 Hanauma Bay MLCD,
Oahu. §13-28-2 Prohibited activities. (1) Fish for, catch, take, injure,
kill, possess, or remove any finfish, crustacean, mollusk including sea
shell and opihi, live coral, algae or limu, or other marine life, or
eggs thereof; (2) Take, alter, deface, destroy, possess, or remove any
sand, coral, rock, or other geological feature, or specimen;(3) Have or
possess any fishing gear or device, including but not limited to any
hook-and-line, rod, reel, spear, trap, net, crowbar, or other device, or
noxious chemical that may be used for the taking, injuring, or killing
of marine life, or the altering of geological feature or specimen, the
possession of which shall be considered prima facie evidence in
violation of this rule; or (4) Introduce any food, substance, or
chemical into the water, to feed or attract marine life.

#### Waikiki MLCD, Oahu

*General Description:* The Waikiki MLCD, established in 1988, is located
on the south shore of Oahu and comprises ca. 76 acres. The MLCD includes
the waters offshore of Kapiolani Beach Park beginning at the high water
mark on the shoreline to a seaward distance of five-hundred yards (457.2
meters) or to the seaward edge of the fringing reef if one occurs beyond
five-hundred yards (457.2 meters), between the western boundary
delineated by a straight line drawn seaward extending the ewa
(western-most) edge of the groin located seaward of the
Kalakaua-Kapahulu avenues junction and the eastern boundary delineated
by a straight line seaward extending the ewa (western-most) edge of the
wall of the Waikiki War Memorial Natatorium.

A reef flat, consisting mostly of rubble and coralline algae with some
patches of live coral, extends out from the Waikiki Aquarium seawall for
35 yards to a dredged channel that is about 8 feet deep. The reef flat
(3-4 feet water depth) has little topographic relief and is covered with
an alien alga, Gracilaria salicornia. At the outer edge of the reef
flat, a sharp edge with numerous arches and crevices descends down to
about 15-20 feet of water depth. Uncolonized pavement accounts for 43%
of the habitat within the MLCD, followed by 50-90% macroalgae at 38%,
with an additional 14% consisting of sand (Table 2).

Table 2: Habitat types contained within Waikiki MLCD based on NOAA
benthic habitat maps. (Coyne et al. 2001)

  ---------------------------------------- ------------ -------- ------------
  Habitat Type                             Square Meter Acres    Proportion
                                                                 of total

  Hardbottom/Uncolonized Pavement          134751.73    33.30    0.43

  Macroalgae/50-90%                        119123.30    29.44    0.38

  Sand                                     42717.69     10.56    0.14

  Unknown                                  15628.42     3.86     0.05

  Artificial/Other Man Made Structures     694.60       0.17     \>0.01

  Total                                    312915.73    77.32    1.00
  ---------------------------------------- ------------ -------- ------------

*Fisheries Regulations and Use:* Hawaii Administrative Rules Title 13,
DLNR, Subtitle 4 Fisheries, Part 1, MLCDs, Chapter 36 Waikiki MLCD,
Oahu.§13-36-2 Prohibited activities. No person shall engage in the
following activities in the Waikiki Marine Life Conservation District:
(1) Fish for, catch, take, injure, kill, possess, or remove any finfish,
crustacean, mollusk including sea shell and opihi, live coral, algae or
limu, or other marine life, or eggs thereof; (2) Take, alter, deface,
destroy, possess, or remove any sand, coral, rock, or other geological
feature, or specimen; or (3) Have or possess in the water, any spear,
trap, net, crowbar, or any other device that may be used for the taking
or altering of marine life, geological feature, or specimen.

#### Waikiki-Diamond Head Shoreline Fisheries Management Area

*General Description:* The Waikiki-Diamond Head Shoreline Fisheries
Management Area extends from the ewa wall of the Waikiki War Memorial
Natatorium to the Diamond Head Lighthouse, from the highwater mark out
to a minimum seaward distance of 500 yards, or to the seaward edge of
the fringing reef if one occurs beyond 500 yards.

Uncolonized hardbottom pavement accounts for 55% of the habitat within
the FMA (Table 3). Another 18% consisted of macroalgae-dominated
substrate, with greater than 10% live coral reef accounting for an
additional 15%.

*Fisheries Regulations and Use:* The Waikiki-Diamond Head Shoreline
Fisheries Management Area (FMA) was established in 1978 as a rotating
closed area. From 1978 to 1988, management was on a four year cycle with
the entire area closed to fishing for two years, then open to hook and
line fishing only for one year, followed by one year open to all fishing
methods (Brock and Kamm 1993). From July 1998 onward, the management
regime was changed to one year closure and one year open to all fishing
except gillnets and night spearfishing.

Table 3: Habitat types contained within Waikiki Diamond Head MFA based
on NOAA benthic habitat maps. (Coyne et al. 2001)

  ---------------------------------------- ------------ -------- ------------
  Habitat Type                             Square Meter Acres    Proportion
                                                                 of total

  Hardbottom/Uncolonized Pavement          523378.48    129.33   0.55

  Reef/Patch Reef (Individual)             143434.18    35.44    0.15

  Macroalgae/10-50%                        112871.93    27.89    0.12

  Macroalgae/50-90%                        54873.13     13.56    0.06

  Sand                                     39939.30     9.87     0.04

  Unknown                                  30214.95     7.47     0.03

  Hardbottom/Uncolonized Pavem. with       22921.69     5.66     0.02
  Channels                                                       

  Hardbottom/Reef Rubble                   18406.81     4.55     0.02

  Reef/Spur and Groove Reef                12155.44     3.00     0.01

  Artificial/Other Man Made Structures     1389.19      0.34     0.00

  Total                                    959585.09    237.11   1.00
  ---------------------------------------- ------------ -------- ------------

It is open to fishing from January 1 to December 31 of even-numbered
years (2000, 2002, etc.). And closed to fishing from January 1 to
December 31 of odd-numbered years (2001, 2003, etc.). It is permitted to
fish for, take or possess any legal size marine life in season during
the "open to fishing" period, provided that only hook-and-line,
thrownet, handnet to land hooked fish, and spear fishing and hand
harvesting methods are employed.

It is prohibited to fish for, take or injure any marine life (including
eggs), or to possess in the water any fishing gear during the "closed to
fishing" period. To use any spear between the hours of 6:00 pm to 6:00
am, or have or possess in the water any trap or net except thrownet or
handnet to land hooked fish during the "open to fishing" period.

#### Molokini Shoal MLCD

*General Description:* The Molokini shoal marine life conservation
district is the southern rim of an extinct volcanic crater, 3 miles off
Maui. It encompasses 200 acres and was established 1977. The shallow
inner cove is the crater\'s submerged floor. The cove area slopes off
from the shoreline to a depth of about 100 feet before dropping off. The
bottom consists of sand patches, coral and basaltic boulders. A shallow
reef in less than thirty feet of water extends from the shoreline
northward at the islet\'s northwestern point. The back (southern) side
of the islet has a steep face that drops off to depths of over 200 feet.
Small patches of coral are scattered across the wall. Crevices and
outcroppings harbor large populations of fishes.

*Fisheries Regulations and Uses*: §13-31-3 Prohibited activities. No
person shall engage in the following activities in the Molokini shoal
marine life conservation district: (1) Fish for, catch, take, injure,
kill, possess, or remove any finfish, crustacean, mollusk including sea
shell and opihi, live coral, algae or limu, or other marine life, or
eggs thereof except as provided for in section 13-31-4(1); (2) Have or
possess in the water, any spear, trap net, crowbar, or any other device
that may be used for the taking or altering of marine life, geological
feature, or specimen; (3) Take, alter, deface, destroy, possess, or
remove any sand coral, rock, or other geological feature, or specimen;
31-2 §13-31-5(4) Feed or deliberately introduce any food material,
substance, or attractant, directly to or in the vicinity of any aquatic
organism, by any means for any purpose except as provided in section
13-31-4(1); (5) Moor boats for commercial activities except as provided
for in section 13-31-5; or (6) Anchor a boat when a day use mooring
system and management plan is established by this department.

#### Honolua-Mokule'ia Bay MLCD

*General Description*: Honolua-Mokule'ia are semi-enclosed bays bordered
by north and south basalt cliffs. The surrounding area are for
conservation and agriculture use with intermittent stream input.. North
and south bay reef flats crest and slope to central 8 to 13 m deep
coarse sand channel with beds of Halimeda incrassata. The south reef is
composed of coral colonized basalt. The north reef is more developed
with higher coral coverage compared to the south. Coverage on both reefs
increases with depth. Sand comprises 45% of the total habitat, followed
by uncolonized hardbottom (35%), and colonized reef habitat (19%, Table
4). The north reef is sheltered except from winter north Pacific swells.
This area has high human use: water recreation, boat traffic, tourism.
Fish feeding, anchoring, trampling, sedimentation.

Table 4: Habitat types contained within Honolua-Mokuleia MLCD based on
NOAA benthic habitat maps. (Coyne et al. 2001)

  ---------------------------------------- ------------ -------- ------------
  Habitat Type                             Square Meter Acres    Proportion
                                                                 of total

  Sand                                     82448.28     20.37    0.45

  Hardbottom/Uncolonized Volcanic          62975.71     15.56    0.35
  Rock/Boulders                                                  

  Reef/Colonized Volcanic Rock/Boulders    13350.31     3.30     0.07

  Reef/Aggregate Coral                     11947.85     2.95     0.07

  Reef/Colonized Pavement with Channels    9412.64      2.33     0.05

  Hardbottom/Uncolonized Pavement          1132.75      0.28     0.01

  Total                                    181267.54    44.79    1.00
  ---------------------------------------- ------------ -------- ------------

*Fisheries Regulations and Uses:* Prohibited activities. No person shall
engage in the following activities in the Honolua- Mokuleia Bay Marine
Life Conservation District: (1) Fish for, catch, take, injure, kill,
possess, or remove any finfish, crustacean, mollusk including sea shell
and opihi, live coral, algae or limu, or other marine life, or eggs
thereof; (2) Take, alter, deface, destroy, possess, or remove any sand,
coral, rock, or other geological feature, or specimen; or (3) Have or
possess in the water, any spear, trap, net, crowbar, or other device
that may be used for the taking or altering of marine life, geological
feature or specimen.

#### Kahaluu Beach Park, Hawai'i

*General Description:* Kahalu'u is the largest sand beach between Kailua
and Keauhou on the kona coast of the big island. In the days of the
Hawaiian kings, with many of the islands\' beaches having dangerous surf
and riptides, King Kamehameha wanted a safe place for his family to
enjoy the ocean. He had his workers construct a seawall in the surf to
protect a small cove on the sunny side of the Big Island. This cove
today is known as Kahaluu Beach Park and is one of the most popular
swimming and best snorkeling sites in the Kona district.This site is
shallow, mostly less than 1 m, with only a few patches of live coral.

*Fisheries Regulations and Uses:* Kahalu'u Beach Park lies within the
Kailua-Keauhou Fish Replenishment Area (FRA) which prohibits the taking
of aquatic life for either commerical or non-commercial aquarium
purposes or to engage in or attempt to engage in fish feeding..

#### Wai Opae MLCD

*General Description:* A shallow basalt/coralline algae platform on the
seaward margin of the tide pools causes waves to break, thereby
dissipating most of their energy before they enter the pools. The tide
pools are one of the few areas on the entire windward coast of the Big
Island that are easily accessible almost all of the time. People can
drive to within a few yards of the nearest pools and except for during
major storms, it is possible to swim in the tide pools under most
weather conditions. This makes the pools particularly vulnerable to
potential negative impacts from human perturbation.

The MLCD area has a total coral cover of approximately 47% that is
dominated by Porites lobata (Hallacher, Tissot, and Walsh, unpublished
data).

*Fisheries Regulations and Uses*: Prohibited activities -- No person
shall engage in the following activities in the district: (1) take,
injure, kill, possess, or remove any marine life; (2) take, alter,
deface, destroy, possess, or remove any sand, coral, rock, or other
geological feature or specimen; (3) anchor or moor any vessel; (4)
conduct commercial activities, such as, but not limited to, commercial
tours, dive groups, sightseeing tours, hikes, or guided services.

# Empirical Information On Fish Assemblage Characteristics

#### Fish Assemblage Characteristics

Fish assemblage characteristic data were derived from CRAMP data
(Friedlander et al. 2003), NOAA fish habitat utilization and MPA
efficacy study (Friedlander and Brown 2003), and Wai Opae tidepool
investigation (Hallacher, Tissot, and Walsh, unpub.). High species
richness, number of individuals, biomass, and diversity in observed at
Hanauma Bay, Honolua-Mokule'ia, and Molokini Shoal (Table 5).

[Table 5: Fish assemblage characteristics among various MPAs in
Hawaii]{.mark}

+----------------+----------+----------+---------+----------+--------+
| Location       | Species  | Num/ha   | Biomass | D        | Ev     |
|                |          |          | (t/ha)  | iversity | enness |
|                |          | x (1000) |         |          |        |
+----------------+----------+----------+---------+----------+--------+
| Kahalu'u Beach | 15.00    | 3.28     | 0.39    | 2.31     | 0.87   |
| Park1          |          |          |         |          |        |
|                | (7.07)   | (2.38)   | (0.12)  | (0.35)   | (0.03) |
+----------------+----------+----------+---------+----------+--------+
| Hon            | 22.78    | 12.04    | 0.91    | 2.32     | 0.76   |
| olua-Mokule'ia |          |          |         |          |        |
| Bay MLCD^1^    | (6.18)   | (6.51)   | (0.77)  | (0.35)   | (0.09) |
+----------------+----------+----------+---------+----------+--------+
| Waikiki        | 11.00    | 3.98     | 0.41    | 1.78     | 0.79   |
| MLCD^1^        |          |          |         |          |        |
|                | (6.16)   | (3.38)   | (0.56)  | (0.63)   | (0.20) |
+----------------+----------+----------+---------+----------+--------+
| Waikiki FMA^1^ | 10.35    | 3.67     | 0.22    | 1.75     | 0.81   |
|                |          |          |         |          |        |
|                | (5.91)   | (2.94)   | (0.30)  | (0.42)   | (0.10) |
+----------------+----------+----------+---------+----------+--------+
| Hanauma Bay    | 27.57    | 16.71    | 1.38    | 2.44     | 0.74   |
| MLCD^2^        |          |          |         |          |        |
|                | (5.35)   | (4.86)   | (0.30)  | (0.31)   | (0.06) |
+----------------+----------+----------+---------+----------+--------+
| Molokini Shoal | 20.50    | 6.86     | 0.98    | 2.52     | 0.84   |
| MLCD^2^        |          |          |         |          |        |
|                | (5.07)   | (2.44)   | (0.80)  | (0.35)   | (0.09) |
+----------------+----------+----------+---------+----------+--------+
| Wai Opae^3^    |          | 5.6      |         |          |        |
+----------------+----------+----------+---------+----------+--------+
| Totals         | 17.00    | 10.35    | 0.54    | 1.98     | 0.73   |
| excluding      |          |          |         |          |        |
| MLCDs          | (7.16)   | (8.03)   | (0.48)  | (0.51)   | (0.16) |
+----------------+----------+----------+---------+----------+--------+

Sources: 1 -- NOAA/NOS fish habitat utilization study (Friedlander and
Brown 2003). 2 -- CRAMP (Friedlander et al. 2003a), and 3 -- UH Hilo/DAR
long-term data (Hallacher, Tissot, and Walsh). Totals are statewide and
based on 208 transects, not including MLCDs.

The lowest values for species richness and biomass were observed at the
two Waikiki locations with low biomass also observed at the shallow
protected Kahalu'u Beach Park. Habitat and level of protection seem to
be the reasons for these observed differences. These areas with greater
protection for fishing and greater habitat quality has fish assemblage
values higher than the average values observed on hardbottom habitats
around the state while the areas with limited protection from fishing
and or poor habitat quality had values below those average values.

#### Species Composition among Sites

Surgeonfishes dominate the Honolua-Mokule'ia Bay MLCD based on weight
and index of relative dominance (IRD) and (Table A-1 in the Appendix).
Seven of the top ten species were surgeonfishes accounting for 58% of
total fish biomass at this location. Fishers target many of the species
and the abundance and size of these species in the MLCD are evidence
that these species are being protected from excessive exploitation.

In contrast to the fish assemblage structure in the Honolua-Mokule'ia
Bay MLCD, fish assemblage structure in the Waikiki MLCD had a greater
prevalence of species that are of little extractive value such as the
Sargent Major (Abudefduf abdominalis, mamo), the endemic saddle wrasse
(Thalassoma duperrey, hi¯na¯lea lauwili), the reef triggerfish
(Rhinecanthus rectangulus, humuhumunukunukuapua\'a), and belted wrasse
(Stethojulis balteata, o¯maka) (Table A-2 in the Appendix). This shift
in fish assemblage structure is even more pronounced at the Waikiki FMA
where the saddle wrasse is the dominant species based on index of
relative dominance (IRD - biomass x frequency of occurrence) (Table A-3
in the Appendix). The brown surgeonfish (Acanthurus nigrofuscus,
ma¯i\'i\'i ) was the second most important species in the Waikiki FMA
based on IRD. Although this species is consumed locally, it has very low
consumptive value.

Dominant species within the Hanauma Bay MLCD included the goldring
surgeonfish (Ctenochaetus strigosus, kole), the introduced bluestripe
snapper (Lutjanus kasmira, ta\'ape), the Achilles tang (Acanthurus
Achilles, pa¯ku'iku'i), and the endemic spectacled parrotfish (Chlorurus
perspicillatus, uhu uliuli) (Table A-4 in the Appendix). The presence of
large parrotfishes, particularly large terminal phase males, in Hanauma
Bay is an indication of what more natural assemblages of fishes would
look like without overexploitation.

Molokini Shoal MLCD contains a diverse assemblage of fishes that is
represented by five different families within the top ten dominant
species (Table A-5 in the Appendix). Black (Melichthys niger,
humuhumu\'el\'ele) and pink durgon (Melichthys vidua, humuhumuhi\'ukole)
are the number one and four species based on IRD. These species feed
mainly on plankton and are more common on offshore islets. Resource
species of interest included one 80 cm giant trevally (Caranx ignobilis,
\'ulua aukea) and several large bullethead parrotfish (Chlorurus
sordidus, uhu) were observed at Molokini.

The Kahalu'u Beach Park was dominated by juvenile parrotfishes (Table
A-6 in the Appendix). Palenose (Scarus psittacus, uhu) and bullethead
(Chlorurus sordidus, uhu ) accounted for more than 32% of total fish
biomass. This shallow rubble location provided optimal habitat for these
species.

The endemic saddle wrasse (Thalassoma duperrey, hi¯na¯lea lauwili) was
the dominant species observed in the Wai Opae tidepool MLCD, followed by
two species of parrotfishes (Scarus psittacus and S. sordidus), the
majority of which were juveniles (Hallacher, Tissot, and Walsh,
unpublished data) (Figure 4). Families of fishes representing the top
ten species included: parrotfishes (Scaridae), damselfishes
(Pomacentridae), wrasses (Labridae), surgeonfishes (Acanthuridae),
puffers (Tetraodontidae), and butterflyfishes (Chaetodontidae).

![](media/image4.png){width="4.983333333333333in"
height="2.527083333333333in"}

Figure 4: Numerical abundance of top tens species in Wai Opae MLCD prior
to MLCD creation (Hallacher, Tissot, and Walsh, unpublished data).

#### Comparisons of Waikiki MLCD, FMA, and areas open to fishing 

##### Fish assemblage characteristics among fisheries management regimes in Waikiki

Fish assemblage characteristics (species, number, and biomass) differed
significantly among fisheries management regimes (MLCD, FMA, and open
access) in the uncolonized hardbottom habitat type (\<10% live coral
cover) in the Waikiki-Diamondhead area (Friedlander and Brown 2003).
Although number of species and number of individuals was higher in the
MLCD, it was not significantly different from the FMA and open access
areas in the uncolonized habitat (Table 6).

Table 6: Comparison of fish assemblage characteristics among various
management regimes in uncolonized hardbottom habitats (\<10% live coral
cover) in the Waikiki-Diamondhead area. MLCD -- Marine Life Conservation
District, Open -- open to all fishing, FMA -- Fisheries Management Area
(Friedlander and Brown 2003).

  ----------------- -------------- ------------- ------------ ------- ---------
  Species           Management     Mean (SD)     Statistics   P       Comp

                    MLCD           13.00 (6.94)  0.008        0.92    A

                    FMA            12.27 (6.92)                       A

                    Open           11.94 (6.01)                       A

  Number/ha x       Management     Mean (SD)     Statistic    P       Comp
  (1000)                                                              

                    MLCD           5.05 (3.96)   0.11         0.9     A

                    Open           4.76 (3.98)                        A

                    FMA            4.47 (3.45)                        A

  Biomass (t/ha)    Management     Mean (SD)     Statistics   P       Comp

                    MLCD           0.61 (0.73)   9.32         0.009   A

                    FMA            0.31 (0.37)                        AB

                    Open           0.16 (0.14)                        C
  ----------------- -------------- ------------- ------------ ------- ---------

Table 7: Comparison of fish assemblage characteristics among various
management regimes in macroalage habitats in the Waikiki-Diamondhead
area.

  ------------------ ------------- ------------- ------------ --------- ---------
  Species            Management    Mean (SD)     Statistics   P         Comp

                     MLCD          9.18 (5.00)   10.06        \<0.001   A

                     FMA           8.00 (3.46)                          A

                     Open          3.18 (2.81)                          B

  Number/ha x (1000) Management    Mean (SD)     Statistics   P         Comp

                     MLCD          3.01 (2.56)   7.42         0.002     A

                     FMA           2.69 (1.94)                          A

                     Open          0.85 (1.12)                          B

  Biomass (t/ha)     Management    Mean (SD)     Statistics   P         Comp

                     MLCD          0.22 (0.25)   12.05        0.003     A

                     FMA           0.11 (0.11)                          A

                     Open          0.03 (0.02)                          B
  ------------------ ------------- ------------- ------------ --------- ---------

Fish biomass in uncolonized hardbottom habitats was four times higher in
the MLCD compared to the open access areas and more than twice as high
as in the FMA. In the macroalgae habitats, the number of species, number
of individuals, and biomass were all highest in the MLCD, followed by
the FMA and lowest in the open access areas (Table 7). Fish biomass was
twice as high in the MLCD compared to the areas open to fishing.

##### Fish assemblage characteristics among fisheries management regimes in West Maui

Fish assemblage characteristics differed among fisheries management
regimes (MLCD and open access) in the colonized hardbottom (\>10% live
coral cover), uncolonized hardbottom (\<10% love coral cover), and
unconsolidated sediments habitat types along the west Maui coast (Table
8). Although number of species was higher in the MLCD, it was not
significantly different from the open access areas except for the
uncolonized hardbottom habitats (Table 8). Number of individuals was
also higher in the MLCD compared with the open assess areas but only
significantly different in the colonized hardbottom habitat (Table 8).
Fish biomass was significantly higher in the Honolua-Mokule'ia MLCD
compared to open assess areas for all three habitat types (Friedlander
and Brown 2003).

Table 8: Fish assemblage characteristics between the Honolua-Mokule'ia
MLCD and areas open to fishing among different habitat types
(Friedlander and Brown 2003).

  --------- ------------------ -------------------- ----------------------
  Species   Colonized          Uncolonized          Unconsolidated
            Hardbottom         Hardbottom           sediments

  MLCD      24.87 (1.2)        20.17 (2.0)          12.00 (2.24)

  Open      20.61(2.0)         14.87 (1.55)         9.47(1.44)

  T         1.90               2.03                 0.99

  P         0.07               0.05                 0.32

  Number    Colonized          Uncolonized          Unconsolidated
            Hardbottom         Hardbottom           sediments

  MLCD      15.14 (1.53)       8.18 (1.47)          4.63 (1.16)

  Open      10.41 (1.53)       6.05 (1.09)          3.70 (0.81)

  T         2.08               1.29                 1.69

  P         0.047              0.20                 0.10

  Biomass   Colonized          Uncolonized          Unconsolidated
            Hardbottom         Hardbottom           sediments

  MLCD      1.37 (0.40)        0.88 (0.25)          0.49 (0.16)

  Open      0.41 (0.08)        0.20 (0.04)          0.12 (0.03)

  T         3.05               3.71                 2.80

  P         0.005              \<0.001              0.007
  --------- ------------------ -------------------- ----------------------

##### Fish trophic structure between management regimes and among habitats

Herbivores were the dominant trophic guild by weight over all habitat
types, accounting for over 67% of the total fish biomass observed on
transects. These were followed by mobile invertebrate feeders (21%),
planktivores (6%), and piscivores (3.4%). Biomass for all trophic guilds
was higher in the MLCD compared to areas open to fishing (Figure 5).
Piscivores showed the largest difference (1226%) in biomass between
areas open to fishing and those closed to fishing. Herbivores showed a
740% difference followed by planktivores (108%), and mobile invertebrate
feeders (46%).

![](media/image5.wmf){width="4.865972222222222in"
height="2.8159722222222223in"}

Figure 5 Mean fish biomass (t/ha) among trophic guilds for the
Honolua-Mokule'ia MLCD compared to open access areas (Friedlander and
Brown 2003).

##### Comparison of fish assemblage characteristics between Kahalu'u Beach Park and similar habitat types within Kailua-Keauhou FRA. 

Number of species, number of individuals, biomass, and diversity were
all lower at the Kahaluu Beach Park compared with similar uncolonlized
(\<10% live coral cover) habitat within the Kailua-Keauhou FRA (Table
9). The low habitat heterogeneity in the backreef habitat of the beach
park resulted an assemblage dominated by juvenile parrotfishes.

Table 9: Comparison of fish assemblage characteristics between Kahaluu
Beach Park and similar habitat types within Kailua-Keauhou FRA.

  ------------------------------ ----------------- -----------------------
  Assemblage Characteristic      FMA               Kahaluu Beach Park

  Species richness               18.00 (5.06)      15.00 (7.07)

  Number of individuals          6.65 (4.14)       3.28 (2.38)

  Biomass                        0.52 (0.32)       0.39 (0.12)

  Diversity                      2.25 (0.25)       2.31 (0.35)
  ------------------------------ ----------------- -----------------------

# Empirical Information on MPAs in Hawaii -- Overall Comparison of Protected Areas 

#### Hawaii Coral Reef Assessment and Monitoring Program (CRAMP)

As part of the Hawaii Coral Reef Assessment and Monitoring Program
(CRAMP), fish assemblage structure and habitat utilization patterns were
examined at 60 sites around the main Hawaiian Islands (Friedlander et
al. 2003a). Areas completely protected from fishing had distinct fish
assemblages with higher standing stock and diversity than areas where
fishing was permitted or areas that were partially protected from
fishing (Table 10, Friedlander et al. 2003a). Hanauma Bay on Oahu and
Honolua Bay on Maui, two no-take areas with the highest levels of
protection from fishing in the main Hawaiian Islands, had the highest
values for most fish assemblage characteristics.

The State of Hawaii has been encouraging community-based management of
subsistence fishing areas since 1994, and a number of community-managed
areas are now being established. Locations influenced by customary
stewardship harbored fish biomass that was equal to or greater than that
of no-take protected areas. The remoteness of these locations combined
with the light fishing pressure (on-island consumption only) and
community oversight has resulted in high standing stock of reef fishes
compared to other locations in Hawaii.

Marine protected areas in the main Hawaiian Islands with high habitat
complexity, moderate wave disturbance, a high percentage of branching
and/or lobate coral coupled with legal protection from fishing pressure
had higher values for most fish assemblage characteristics. Habitats
with these optimal characteristics should possess fish assemblages with
high species richness, abundance, biomass, and diversity (Friedlander et
al. 2003a).

Table 10: Fish assemblage characteristics among various management
regimes at 60 sites in the main Hawaiian Islands (Friedlander et al.
2003a).

  --------------------------------- ---------------------- ---------------
  Management                        Species Richness       Comp

  No-take                           24.99                  A

  Customary stewardship             21.29                  AB

  Partial protection                19.13                  AB

  Open access                       17.54                  B

                                    Number of individuals  

  No-take                           146.26                 A

  Customary stewardship             142.50                 A

  Partial protection                113.81                 A

  Open access                       105.39                 A

                                    Biomass                

  No-take                           15.89                  A

  Customary stewardship             16.27                  A

  Partial protection                6.07                   AB

  Open access                       6.16                   B

                                    Diversity              

  No-take                           2.52                   A

  Customary stewardship             2.33                   AB

  Partial protection                2.22                   AB

  Open access                       2.14                   B
  --------------------------------- ---------------------- ---------------

#### NOAA/NOS Fish Habitat Utilization Study

Coupling the distribution of habitats and species habitat affinities
using NOS digital benthic habitat maps has proven to be powerful tool to
examine the efficacy of MPAs in Hawaii. Habitat quality and size were
important determinates of the effectiveness of the MPAs examined with
respect to their fish assemblages (Friedlander and Brown 2003). In most
cases, when compared within habitats, fish assemblage characteristics
were higher in protected areas compared with areas open to fishing,
illustrating the effectiveness of these areas in conserving coral reef
fish assemblages (Table 11).

Table 11: Mean fish biomass (t/ha) in areas under various management
regimes. MLCD -- Marine Life Conservation District, Open -- open to all
fishing, FMA -- Fisheries Management Area. Hard bottom habitats only
(Friedlander and Brown 2003).

  -------------------- ---------- ---------- ---------- ------------------
  LOCATION             MLCD       Open       FMA        Comparisons

  West Maui            0.68       0.16                  MLCD\>Open

  Kaneohe Bay, Oahu    0.65       0.32                  MLCD\>Open

  South coast, Lanai   0.63       0.44                  MLCD\>Open

  Kailua Kona, Hawaii  1.52       0.94       0.46       MLCD\>Open\>FMA

  Waikiki, Oahu        0.41       0.13       0.22       MLCD\>FMA\>Open
  -------------------- ---------- ---------- ---------- ------------------

##### Size of fishes among management regimes

Mean size of fishes among management regimes at 449 sampling locations
around the main Hawaiian Islands showed significantly different (Table
12) with fish in the MLCDs being significantly larger than in open
access and slightly larger (but not significantly) than those fishes in
the FMAs.

Table 12: Mean size of all fishes among various management regimes.
Results of One-way ANOVA: F~2,5823~ = 85.2, P \< 0.001. Tukey HSD
multiple comparisons test. Management regimes with the same latter are
not significantly different at = 0.05 (Friedlander and Brown 2003).

  -------------------------- --------------- -------------- --------------
  Management regime          Mean size (cm)  Standard Error Multiple
                                                            comparison

  MLCD                       12.3            0.16           A

  Fisheries Management Area  11.9            0.24           A

  Open access                9.97            0.11           B
  -------------------------- --------------- -------------- --------------

##### 

##### Comparison of parrotfish size frequency distributions between fished and unfished areas

Adult parrotfish are highly prized by fishers and large adult parrotfish
(uhu) are rare in locations that are heavily fished. Size frequency
distributions of parrotfishes in the Moku o Loe (Coconut Island) Hawai'i
Marine Laboratory Refuge are significantly larger than parrotfishes
observed in areas open to fishing (P \< 0.001). Examination of the size
spectra of parrotfishes greater than 20 cm in length show a
significantly (P \< 0.05) smaller number of the larger size classes
represented in the areas open to fishing compared with Moku o Loe
(Coconut Island) Hawai'i Marine Laboratory Refuge when harvest is
prohibited (Figure 6). These larger size classes targeted by fishers
also contain a disproportionately greater number of terminal phase males
that are important for reproductive success and therefore, population
replenishment.

![](media/image6.wmf){width="5.854166666666667in"
height="3.386111111111111in"}

Figure 6: Size spectrum of parrotfishes greater than 20cm around Moku o
Loe (Coconut Islands Marine Laboratory Reserve) and patch reefs open to
fishing around Kaneohe Bay, Oahu (Friedlander and Brown, 2003).

# Recommendations for Modifications to Existing MPAS and for the Design and Siting of Future Protected Areas

There are a variety of marine areas in Hawai*'*i that have some type of
protected status. These include Marine Life Conservation Districts
(MLCDs), Fisheries Management Areas (FMAs), Fisheries Replenishment
Areas (FRAs), a Marine Laboratory Refuge, Natural Area Reserve (NARs),
Kahoolawe Island Reserve (KIR), National Wildlife Refuges, the Hawaiian
Islands Humpback Whale Sanctuary (Clark and Gulko, 1999) and the
Northwestern Hawaiian Islands Coral Reef Ecosystem Reserve. In almost
all cases examines, protected areas had healthier more diverse fish
assemblages with higher standing stock compared with similar adjacent
areas. However, many of these reserves are either too small, lack
suitable habitats, or are not fully protected from fishing and therefore
do not function effectively as refuges (Friedlander 2001, Friedlander et
al. 2003a, Meyer 2003). This is the most critical issue to address if
reserves are to be an effective fisheries and conservation management
tool in Hawai'i.

The value of Hawai'i's commercial coastal fisheries was approximately
2.5 million dollars in 2001 (DeMello 2003). However, 80% of catch
consists of two species of coastal pelagics (akule ands opelu). The
recreational catch and artisanal catch combined likely exceeds the
commercial catches, excluding akule and opelu, and these fishers take a
wider variety of species (Friedlander and Parrish 1997, Everson and
Friedlander 2003). Income derived from recreational use of the fishery
through the sale of fishing tackle, bait, license fees, fuel, etc. is
also an important component of the local economy in many areas around
the state. For the reasons stated above, MPAs in Hawai'i should focus
more on improving recreational and subsistence catch versus enhancing
commercial yield. Although recreational fishers can tolerate a much
lower yield than commercials fishers, subsistence fishers need to
maintain yields for sustenance and for cultural survival. Although this
need does not always translate to a dollar value, its importance to the
people of Hawai'i cannot be neglected. MPAs will also provide insurance
against other management inaction as well as providing insurance against
environmental disasters. Fishing tourism is important to many coastal
states and if Hawaii chooses to pursue this sector, MPAs will be needed
to reduce the risk of collapse and rebuild stocks.

#### Waikiki MLCD

Although the Waikiki MLCD has been a no-take area since 1988, low
habitat heterogeneity, degraded reef environment, and small size have
resulted in this area having a very low standing stock of fish. However,
the size and number of fishes is greater within the MLCD compared to
adjacent habitats (Friedlander and Brown 2003, Meyer 2003). Meyer (2003)
noted that abundance and size of both target and non-target species was
greater in the Waikiki MLCD compared to adjacent fished areas,
suggesting that fishing is not the only factor determining patterns in
abundance and size. The author noted that despite having generally poor
habitat quality, the habitat within the MLCD had greater complexity
compared to the adjacent areas. Based on tracking data and the
distribution of critical habitat, Meyer (2003) also determined that the
area of the Waikiki MLCD (0.32 km^2^) would need to be at least tripled
in size (1 km^2^) to begin to effectively protect more mobile species
such as jacks and goatfishes. This modest increase in size along with
the inclusion of essential softbottom habitats would greatly improve the
effectiveness of the MLCD in conserving reef assemblage structure,
however, a much larger area or areas would be necessary to achieve any
type of fisheries benefits.

#### Waikiki-Diamond Head FMA

Visual census data of fishes conducted by DAR since 1978 has shown that
fish biomass was higher within the Waikiki-Diamondhead FMA when the area
was closed to fishing or open to hook and line only (Brock and Kamm
1993). Despite these closures, standing crop of fishes never exceeded 50
g/m2 and Brock and Kam (1993) attributed this to the lack of adequate
shelter habitat. The benefits derived from the closure were quickly lost
when the area was open to all types of fishing, but hook-and-line
fishing appeared to have little impact on fish standing stock. (Brock
and Kamm 1993). Current regulations prohibit fishing on odd-numbered
years and prohibit trap and net fishing, except throw nets, and
nighttime spearfishing during even-numbered open years. Holland and
Meyer (2003) found that alternating closures is less important than the
fact that no nighttime spearfishing or gillnetting was allowed in the
FMA during open years. Recent analysis of DAR survey data (1985-2000)
from the Waikiki-Diamond Head FMA revealed a recovery for most trophic
and taxonomic groupings in closed years with declines occurring in open
years (DAR unpub. data). The most disturbing trend from this analysis
was the overall decline in most trophic guilds over time despite annual
closures.

#### Habitat

In Hawaii, habitats with low spatial relief and limited shelter were
found to be associated with low biomass and diversity of reef fishes
while highly complex habitats harbored high fish biomass and diversity
(Friedlander and Parrish 1998a). Ideally, essential fish habitat in the
main Hawaiian Islands should consist of an area with high rugosity or
relief with moderate wave exposure that has a high percentage of
branching and/or lobate coral coupled with legal protection from fish
pressure. Habitats with these optimal characteristics should possess
fish assemblages with high species richness, abundance, biomass, and
diversity (Friedlander at al. 2003a). If protective areas are to be
effective, they must include the diversity of habitats necessary to
accommodate the wide range of fish species.

Coupling the distribution of habitats and species habitat affinities
using GIS technology enables the elucidation of species habitat
utilization patterns for a single species and/or assemblages of animals.
This integrated approach is useful in quantitatively defining essential
fish habitat and defining biologically relevant boundaries of marine
protected areas. By integrating spatial data into the biological
sampling design, significant progress can be made towards identifying
and quantifying spatial dependencies in habitat utilization by reef
fishes. This design also lends itself to elucidating factors that might
suggest cause for differential patterns in ontogenetic habitat
selection, ergo distribution, within the available landscape. Such
patterns in population and community structure are necessary and
fundamental components of any intent to understand and maximize the
benefits derived from a Marine Protected Area.

#### Movement

Short and long-term movement patterns of omilu (blue trevally, *Caranx
melampygus*) were monitored around Coconut Island (Moku o Lo'e) (Holland
et al. 1996). The limited range of dispersal of recaptured (75.5% within
0.5 km of the release site) and strong site fidelity observed from
sonically tagged fish suggest that dispersal is much less than might be
predicted for a highly mobile, piscivorous species. The authors suggest
that small refugia (e.g. 5 km of reef face) could provide significant
protection for this species despite its potential for long-range
movements. *Kumu* (whitesaddle goatfish, *Parupeneus porphyreus*), an
endemic goatfish and important fisheries species, were acoustically
tracked around the Coconut Island refuge for periods up to 93 h (Meyer
et al. 2000). The home ranges of all fish were within the boundaries of
the Coconut Island reserve. This small reserve (\< 1 km^2^) was capable
of protecting both large juveniles and some spawning size individuals
(Meyer et al. 2000). *Kala* (blue spined unicornfish, *Naso unicornis*)
were acoustically tracked for periods of up to 22 days in the shallow
high-energy fringing reef habitat in the Waikiki Marine Life
Conservation District (Meyer and Holland 2001). The home ranges of all
of the *kala* tracked were completely encompassed by the boundaries of
the 0.32 km^2^ Waikiki Marine Life Conservation District.

#### Monitoring

Assessments of fished vs. protected areas must consider habitat and
environmental variables when designing assessment programs in order to
properly examine MPA success. Once reserves are established, long-term
monitoring programs can be implemented to help determine the
effectiveness of the zoning plan and to guide future modifications to
either the fishing regulations or the reserve boundaries.

The openness and dynamic nature of ocean environments make it extremely
difficult to prove fisheries enhancements statistically because of the
difficulty separating reserve effects from other changes that might have
occurred (PDT 1990; Sladek Nowlis and Friedlander, 2004b). Marine
reserves are as well supported by this sort of evidence as any other
fishery management technique in use today and should not be held to a
higher standard than these other management measures (Sladek Nowlis and
Friedlander, 2004a).

Most commonly, marine reserve monitoring programs compare inside to
outside of the reserve). These sorts of comparisons can provide useful
information about how fishing is affecting the outside areas with the
reserves serving as a control. In this manner, a marine reserve
monitoring program can follow a standard ecological
before-after-control-impact-pairs design (Sladek Nowlis and Friedlander
2004c). These samples are paired in the sense that the control (reserve)
and impact (fishing) sites are examined more or less concurrently.
Replication comes from collecting such paired samples at a number of
times both before and after the reserve designation. The approach is to
test whether the differences between the control and impact sites
changed after the reserve was established.

#### Fisheries yields

In 1998, the state legislature passed Act 306, which established a West
Hawai'i Regional Fisheries Management Area to provide for effective
management of marine resources. The West Hawai'i Fisheries Council,
composed of stakeholders and government representatives, developed a
network of nine Fish Replenishment Areas (FRAs) encompassing 35.2%
(including existing protected areas) of the coastline (Walsh 1999).

Preliminary analysis (Tissot et al., 2004) indicates that three years
after closure of the FRAs there have been significant increases in the
overall abundance of fishes targeted by collectors. Two species, the
yellow tang and Potter's angelfish (*Centropyge potteri*), showed
significant (74-80%) increases in FRAs relative to previously protected
reference areas (Walsh et al. 2003). Furthermore, no aquarium fishes
declined in abundance in open areas as might be expected if the
intensity of harvesting increased outside of the FRAs. In fact, two
species displayed significant increases in abundance in the open areas.

![](media/image7.wmf){width="5.488194444444445in"
height="2.7534722222222223in"}

Despite a closure of more than 35% of the west Hawaii coast to aquarium
fish collecting, the value of fish caught has increased since the
closure went into effect in 2000 (Walsh et al. 2003). Of special note is
the fact that the dollar value of each yellow tang has increased in the
past two years (Despite a closure of more than 35% of the west Hawaii
coast to aquarium fish collecting, the value of fish caught has
increased since the closure went into effect in 2000 (Walsh et al.
2003). Of special note is the fact that the dollar value of each yellow
tang has increased in the past two years (Despite a closure of more than
35% of the west Hawaii coast to aquarium fish collecting, the value of
fish caught has increased since the closure went into effect in 2000
(Walsh et al. 2003). Of special note is the fact that the dollar value
of each yellow tang has increased in the past two years (Despite a
closure of more than 35% of the west Hawaii coast to aquarium fish
collecting, the value of fish caught has increased since the closure
went into effect in 2000 (Walsh et al. 2003). Of special note is the
fact that the dollar value of each yellow tang has increased in the past
two years (). Indeed, the overall value of the West Hawai'i aquarium
fishery in FY 2003 is the highest it has ever been (). ). Indeed, the
overall value of the West Hawai'i aquarium fishery in FY 2003 is the
highest it has ever been (Figure 8). ). Indeed, the overall value of the
West Hawai'i aquarium fishery in FY 2003 is the highest it has ever been
(Figure 8). ). Indeed, the overall value of the West Hawai'i aquarium
fishery in FY 2003 is the highest it has ever been (Figure 8).

Figure 7: Number and value (adjusted for inflation) of yellow tangs
caught in West Hawai'i per fiscal year (Walsh et al. 2003).

![](media/image8.wmf){width="5.602083333333334in"
height="2.959722222222222in"}

Figure 8: Number caught of top 2nd-5th West Hawai'i species per fiscal
year (Walsh et al. 2003).

#### Basic recommendations for reserves in Hawaii

-   Larger in size -- individual reserves should cover 5-10 km^2^

-   Incorporate essential fish habitat -- areas of high habitat
    complexity, spawning locations, and essential feeding areas need to
    be represented fully in reserve design

-   More connection between shallow and deep habitats -- identify
    juvenile and adult habitats and provide connections for ontogenetic
    movement

-   More connections between resting and feeding habitats. Identify
    feeding corridors to connect habitats.

-   Reserves should be networked. Several smaller reserves may be more
    beneficial than one large one particularly when one is concerned
    with limiting the exposure at the boundaries. A reserve 4 to 6 km in
    diameter should be large enough to contain the larvae of short
    distance dispersers and reserves spaced 10 to 20 km apart should be
    close enough to capture propagules released from adjacent reserves.

-   Develop a more ecosystem-based ahupua'a concept that includes
    shoreline and upland ecosystems and uses

-   Recognize that the baseline has shifted and manage to rebuild not to
    sustain current levels of abundance.

-   Develop monitoring programs that incorporate habitat types and
    examine human use patterns.

-   Involvement of local communities is invaluable to the creation and
    implementation of reserves or reserve networks because they play an
    important role in the enforceability and social acceptability of
    reserves.

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###### Top ten species observed on transects for the Six Study Sites

Table A-1: Top ten species observed on transects at Honolua-Mokule'ia
Bay MLCD (N = 27)

  ----------------- --------------- ------------- --------- --------- ------ --------- -------- ---------
  TaxonName         Hawaiian Name   Common Name   Number    Biomass   Freq   IRD^\*^   \%       \%
                                                  (No./ha x (t/ha)                     number   biomass
                                                  1000)                                         

  *Naso unicornis*  kala            Bluespine     0.20      0.16      0.52   0.082     1.62     17.24
                                    Unicornfish                                                 

  *Acanthurus       ma¯i\'i\'i      Brown         1.43      0.06      0.93   0.059     11.83    7.02
  nigrofuscus*                      Surgeonfish                                                 

  *Acanthurus       manini          Convict Tang  0.75      0.10      0.41   0.040     6.20     10.60
  triostegus*                                                                                   

  *Naso lituratus*  umaumalei       Orangespine   0.16      0.06      0.56   0.034     1.33     6.62
                                    Unicornfish                                                 

  *Acanthurus       ma¯ikoiko       Whitebar      0.24      0.06      0.48   0.028     1.97     6.30
  leucopareius*                     Surgeonfish                                                 

  *Acanthurus       pualu           Ringtail      0.17      0.07      0.33   0.024     1.43     7.88
  blochii*                          Surgeonfish                                                 

  *Thalassoma       hi¯na¯lea       Saddle Wrasse 1.05      0.02      0.96   0.022     8.71     2.51
  duperrey*         lauwili                                                                     

  *Scarus           pa¯lukaluka     Redlip        0.23      0.03      0.56   0.015     1.87     2.90
  rubroviolaceus*                   Parrotfish                                                  

  *Parupeneus       moano           Manybar       0.22      0.02      0.78   0.014     1.80     1.97
  multifasciatus*                   Goatfish                                                    

  *Acanthurus       na\'ena\'e      Orangeband    0.08      0.02      0.41   0.009     0.64     2.51
  olivaceus*                        Surgeonfish                                                 
  ----------------- --------------- ------------- --------- --------- ------ --------- -------- ---------

Table A-2: Top ten species observed on transects at Waikiki MLCD (N =
21)

  --------------- ------------------------- ------------- --------- --------- ------ --------- -------- ---------
  Taxon Name      Hawaiian Name             Common Name   Number    Biomass   Freq   IRD^\*^   \%       \%
                                                          (No./ha x (t/ha)                     number   biomass
                                                          1000)                                         

  *Acanthurus     ma¯i\'i\'i                Brown         0.50      0.06      0.67   0.042     12.44    15.50
  nigrofuscus*                              Surgeonfish                                                 

  *Naso           kala                      Bluespine     0.16      0.06      0.57   0.035     4.11     15.02
  unicornis*                                Unicornfish                                                 

  *Acanthurus     ma¯ikoiko                 Whitebar      0.18      0.05      0.29   0.014     4.40     11.75
  leucopareius*                             Surgeonfish                                                 

  *Thalassoma     hi¯na¯lea lauwili         Saddle Wrasse 0.73      0.01      0.95   0.013     18.28    3.23
  duperrey*                                                                                             

  *Acanthurus     manini                    Convict Tang  0.32      0.02      0.43   0.010     8.13     5.55
  triostegus*                                                                                           

  *Rhinecanthus   humuhumunukunukuapua\'a   Reef          0.08      0.01      0.67   0.008     2.11     2.93
  rectangulus*                              Triggerfish                                                 

  *Abudefduf      mamo                      Sargent Major 0.27      0.02      0.33   0.006     6.70     4.10
  abdominalis*                                                                                          

  *Stethojulis    \'o¯maka                  Belted Wrasse 0.30      0.01      0.81   0.004     7.46     1.25
  balteata*                                                                                             

  *Acanthurus     pualu                     Ringtail      0.12      0.02      0.19   0.004     2.97     5.28
  blochii*                                  Surgeonfish                                                 

  *Naso           umaumalei                 Orangespine   0.06      0.01      0.24   0.003     1.53     3.37
  lituratus*                                Unicornfish                                                 
  --------------- ------------------------- ------------- --------- --------- ------ --------- -------- ---------

*^\*^* IRD = index of relative dominance (biomass x frequency of
occurrence).

Table A-3: Top ten species observed on transects at Waikiki FMA (N = 20)

+--------------+--------------+-------------+----+---+---+---+---+---+
| TaxonName    | Hawaiian     | Common Name | Nu | B | F | I | \ | \ |
|              | Name         |             | mb | i | r | R | % | % |
|              |              |             | er | o | e | D | n | b |
|              |              |             | (  | m | q | ^ | u | i |
|              |              |             | No | a |   | \ | m | o |
|              |              |             | ./ | s |   | * | b | m |
|              |              |             | ha | s |   | ^ | e | a |
|              |              |             |    | ( |   |   | r | s |
|              |              |             | x  | t |   |   |   | s |
|              |              |             | 1  | / |   |   |   |   |
|              |              |             | 00 | h |   |   |   |   |
|              |              |             | 0) | a |   |   |   |   |
|              |              |             |    | ) |   |   |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Thalassoma  | hi¯na¯lea    | Saddle      | 0. | 0 | 0 | 0 | 2 | 8 |
| duperrey*    | lauwili      | Wrasse      | 91 | . | . | . | 4 | . |
|              |              |             |    | 0 | 9 | 0 | . | 5 |
|              |              |             |    | 2 | 5 | 1 | 7 | 4 |
|              |              |             |    |   |   | 8 | 5 |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Acanthurus  | ma¯i\'i\'i   | Brown       | 0. | 0 | 0 | 0 | 9 | 1 |
| nigrofuscus* |              | Surgeonfish | 36 | . | . | . | . | 2 |
|              |              |             |    | 0 | 6 | 0 | 8 | . |
|              |              |             |    | 3 | 0 | 1 | 1 | 2 |
|              |              |             |    |   |   | 6 |   | 2 |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Stethojulis | \'o¯maka     | Belted      | 0. | 0 | 1 | 0 | 1 | 3 |
| balteata*    |              | Wrasse      | 60 | . | . | . | 6 | . |
|              |              |             |    | 0 | 0 | 0 | . | 8 |
|              |              |             |    | 1 | 0 | 0 | 2 | 2 |
|              |              |             |    |   |   | 8 | 5 |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *            | humuhumunuk  | Reef        | 0. | 0 | 0 | 0 | 2 | 5 |
| Rhinecanthus | unukuapua\'a | Triggerfish | 08 | . | . | . | . | . |
| rectangulus* |              |             |    | 0 | 6 | 0 | 2 | 2 |
|              |              |             |    | 1 | 0 | 0 | 9 | 7 |
|              |              |             |    |   |   | 7 |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Acanthurus  | manini       | Convict     | 0. | 0 | 0 | 0 | 5 | 5 |
| triostegus*  |              | Tang        | 21 | . | . | . | . | . |
|              |              |             |    | 0 | 4 | 0 | 6 | 9 |
|              |              |             |    | 1 | 0 | 0 | 7 | 5 |
|              |              |             |    |   |   | 5 |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Abudefduf   | mamo         | Sargent     | 0. | 0 | 0 | 0 | 6 | 7 |
| abdominalis* |              | Major       | 24 | . | . | . | . | . |
|              |              |             |    | 0 | 2 | 0 | 6 | 3 |
|              |              |             |    | 2 | 5 | 0 | 5 | 6 |
|              |              |             |    |   |   | 4 |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Naso        | umaumalei    | Orangespine | 0. | 0 | 0 | 0 | 1 | 5 |
| lituratus*   |              | Unicornfish | 04 | . | . | . | . | . |
|              |              |             |    | 0 | 3 | 0 | 2 | 4 |
|              |              |             |    | 1 | 0 | 0 | 0 | 6 |
|              |              |             |    |   |   | 4 |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Naso        | kala         | Bluespine   | 0. | 0 | 0 | 0 | 1 | 7 |
| unicornis*   |              | Unicornfish | 05 | . | . | . | . | . |
|              |              |             |    | 0 | 1 | 0 | 3 | 3 |
|              |              |             |    | 2 | 5 | 0 | 1 | 1 |
|              |              |             |    |   |   | 2 |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Chlorurus   | uhu          | Bullethead  | 0. | 0 | 0 | 0 | 1 | 2 |
| sordidus*    |              | Parrotfish  | 06 | . | . | . | . | . |
|              |              |             |    | 0 | 3 | 0 | 7 | 9 |
|              |              |             |    | 1 | 0 | 0 | 4 | 5 |
|              |              |             |    |   |   | 2 |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+
| *Parupeneus  | moano        | Manybar     | 0. | 0 | 0 | 0 | 1 | 1 |
| mul          |              | Goatfish    | 05 | . | . | . | . | . |
| tifasciatus* |              |             |    | 0 | 3 | 0 | 3 | 8 |
|              |              |             |    | 0 | 5 | 0 | 1 | 7 |
|              |              |             |    |   |   | 1 |   |   |
+--------------+--------------+-------------+----+---+---+---+---+---+

Table A-4: Top ten species observed on transects at Hanauma Bay MLCD (N
= 7)

  ----------------- ------------------- --------------- --------- --------- ------ --------- -------- ---------
  TaxonName         Hawaiian Name       Common Name     Number    Biomass   Freq   IRD^\*^   \%       \%
                                                        (No./ha x (t/ha)                     number   biomass
                                                        1000)                                         

  *Ctenochaetus     kole                Goldring        0.64      0.32      1.00   0.318     28.66    22.34
  strigosus*                            Surgeonfish                                                   

  *Lutjanus         ta\'ape             Bluestripe      0.08      0.09      0.86   0.078     3.56     6.43
  kasmira*                              Snapper                                                       

  *Acanthurus       pa¯ku'iku'i         Achilles Tang   0.07      0.06      1.00   0.057     3.01     3.98
  achilles*                                                                                           

  *Chlorurus        uhu uliuli          Spectacled      0.01      0.11      0.43   0.047     0.41     7.67
  perspicillatus*                       Parrotfish                                                    

  *Melichthys       humuhumuhi\'ukole   Pinktail Durgon 0.02      0.05      0.86   0.041     0.96     3.35
  vidua*                                                                                              

  *Zebrasoma        lau\'i¯pala         Yellow Tang     0.07      0.05      0.86   0.040     3.08     3.25
  flavescens*                                                                                         

  *Abudefduf        mamo                Sargent Major   1.36      0.09      0.43   0.039     8.14     6.41
  abdominalis*                                                                                        

  *Scarus           uhu                 Palenose        0.09      0.04      0.71   0.029     3.69     2.82
  psittacus*                            Parrotfish                                                    

  *Chromis ovalis*                      Oval Chromis    0.26      0.07      0.43   0.028     10.94    4.60

  *Chaetodon        lauwiliwili         Milletseed      0.11      0.05      0.57   0.026     4.65     3.25
  miliaris*                             Butterflyfish                                                 
  ----------------- ------------------- --------------- --------- --------- ------ --------- -------- ---------

*^\*^* IRD = index of relative dominance (biomass x frequency of
occurrence).

Table A-5: Top ten species observed on transects at Molokini Shoal MLCD
(N = 8)

  --------------- ------------------- -------------- --------- --------- ------ --------- -------- ---------
  Taxon Name      Hawaiian Name       Common Name    Number    Biomass   Freq   IRD^\*^   \%       \%
                                                     (No./ha x (t/ha)                     number   biomass
                                                     1000)                                         

  *Melichthys     humuhumu\'el\'ele   Black Durgon   0.69      0.31      0.88   0.270     10.06    29.15
  niger*                                                                                           

  *Naso           umaumalei           Orangespine    1.05      0.18      0.88   0.160     15.31    17.30
  lituratus*                          Unicornfish                                                  

  *Lutjanus       ta\'ape             Bluestripe     0.18      0.04      0.75   0.033     2.62     4.14
  kasmira*                            Snapper                                                      

  *Melichthys     humuhumuhi\'ukole   Pinktail       0.07      0.02      0.63   0.013     1.02     2.04
  vidua*                              Durgon                                                       

  *Caranx         \'ulua aukea        Giant White    0.01      0.09      0.13   0.012     \<0.01   8.76
  ignobilis*                          Trevally                                                     

  *Ctenochaetus   kole                Goldring       0.56      0.01      0.88   0.011     8.16     1.22
  strigosus*                          Surgeonfish                                                  

  *Cantherhines   \'o¯\'ili           Barred         0.09      0.02      0.50   0.011     1.31     2.12
  dumerilii*                          Filefish                                                     

  *Chlorurus      uhu                 Bullethead     0.07      0.02      0.63   0.009     1.02     1.42
  sordidus*                           Parrotfish                                                   

  *Acanthurus     ma¯i\'i\'i          Brown          0.51      0.01      1.00   0.009     7.43     0.84
  nigrofuscus*                        Surgeonfish                                                  

  *Acanthurus     na\'ena\'e          Orangeband     0.08      0.02      0.38   0.008     1.17     1.94
  olivaceus*                          Surgeonfish                                                  
  --------------- ------------------- -------------- --------- --------- ------ --------- -------- ---------

Table A-6: Top ten species observed on transects at Kahalu'u Beach Park
(N = 2)

  ----------------- ------------------------- ------------- --------- --------- ------ --------- -------- ---------
  TaxonName         Hawaiian Name             Common Name   Number    Biomass   Freq   IRD^\*^   \%       \%
                                                            (No./ha x (t/ha)                     number   biomass
                                                            1000)                                         

  *Scarus           uhu                       Palenose      0.32      0.07      1.00   0.066     9.76     17.24
  psittacus*                                  Parrotfish                                                  

  *Chlorurus        uhu                       Bullethead    0.08      0.05      1.00   0.054     2.44     14.10
  sordidus*                                   Parrotfish                                                  

  *Zebrasoma        lau\'i¯pala               Yellow Tang   0.28      0.03      1.00   0.034     8.54     8.93
  flavescens*                                                                                             

  *Acanthurus       na\'ena\'e                Orangeband    0.16      0.03      1.00   0.031     4.88     8.12
  olivaceus*                                  Surgeonfish                                                 

  *Naso lituratus*  umaumalei                 Orangespine   0.12      0.03      1.00   0.026     3.66     6.85
                                              Unicornfish                                                 

  *Acanthurus       manini                    Convict Tang  0.44      0.04      0.50   0.018     13.41    9.35
  triostegus*                                                                                             

  *Melichthys       humuhumu\'el\'ele         Black Durgon  0.08      0.03      0.50   0.013     2.44     6.74
  niger*                                                                                                  

  *Rhinecanthus     humuhumunukunukuapua\'a   Reef          0.08      0.01      1.00   0.012     2.44     3.01
  rectangulus*                                Triggerfish                                                 

  *Novaculichthys                             Rockmover     0.24      0.01      0.50   0.007     7.32     3.87
  taeniourus*                                                                                             

  *Calotomus                                  Stareye       0.04      0.01      0.50   0.007     1.22     3.82
  carolinus*                                  Parrotfish                                                  
  ----------------- ------------------------- ------------- --------- --------- ------ --------- -------- ---------

*^\*^* IRD = index of relative dominance (biomass x frequency of
occurrence).