Alan M. Friedlander, U.S. Geological Survey and the University of Hawaii

Lisa M. Wedding, NOAA Center for Coastal Monitoring and Assessment Biogeography Branch and the University of Hawaii Manoa

Eric Brown, National Park Service Kalaupapa National Historical Park

Mark E. Monaco, NOAA Center for Coastal Monitoring and Assessment

NOAA Technical Memorandum NOS NCCOS 117
Citation

Friedlander, A.M., L.M. Wedding, E. Brown, M.E. Monaco. 2010. Monitoring Hawaii’s Marine Protected Areas: Examining Spatial and Temporal Trends Using a Seascape Approach. NOAA Technical Memorandum 

NOS NCCOS 117. Prepared by the NCCOS Center for Coastal Monitoring and Assessment Biogeography 

Branch. Silver Spring, MD. 130 pp. 

Acknowledgements 

The completion of this document would not have been possible without the participation of the following 

people: Donna Brown, Chris Caldow, Brent Carmen, Eric Carr, Athline Clark, Alicia Clarke, Mary Donovan, 

Tommy Hagawa, Skippy Hau, Brian Hauk, Jamison Higgins, Sarah D. Hile, Alan Hong, Paul Jokiel, Kevin 

McMahon, Russell Sparks, Kosta Stamoulis, Keoki Stender, Yuko Stender, William Walsh and Darla White. 

Their efforts to help compose, format, review and edit the document are very much appreciated. 

The covers for this document were designed and created by Eric Carr of Consolidated Safety Services. The 

cover photo was provided by Keoki Stender, www.marinelifephotography.com. 

For more information

For more information about this report, please contact CCMA’s Biogeography Branch Acting Chief Chris 

Caldow, at (301) 713-3028 or visit http://ccma.nos.noaa.gov/about/biogeography/.

Mention of trade names or commercial products does not constitute endorsement or recommendation for 

their use by the United States government. 
Monitoring Hawaii’s Marine Protected Areas: Examining 

Spatial and Temporal Trends Using a Seascape Approach

Prepared by the

Biogeography Branch

Center for Coastal Monitoring and Assessment (CCMA)

National Centers for Coastal Ocean Science (NCCOS)

National Oceanic and Atmospheric Association National Ocean Service (NOAA)

1305 East West Highway (SSMC-IV, N/SCI-1)

Silver Spring, MD 20910

USA

August 2010

Alan M. Friedlander 

U.S. Geological Survey and the University of Hawaii

Lisa M. Wedding

NOAA Biogeography Branch and the University of Hawaii at Manoa

Eric Brown

National Park Service, Kalaupapa National Historic Park

Mark E. Monaco

NOAA Center for Coastal Monitoring and Assessment 

NOAA Technical Memorandum NOS NCCOS 117

United States 

Department of 

Commerce

Gary Locke

Secretary

National Oceanic and 

Atmospheric Administration

Jane Lubchenco 

Administrator

National Ocean Service

David Kennedy

Assistant Administrator
TABLE OF CONTENTS

Executive Summary i

Introduction 1

Background 1

Objectives 1

Seascape Ecology Approach 2

Hawaii Marine Protected Areas 2

Methods 5

Study Sites 5

Benthic Habitat Mapping 5

Bathymetric LiDAR Data 11

Sampling Locations 11

Fish Sampling Methodology 12

Fish Sample Size Optimization Analysis 12

Benthic Survey Techniques 13

Rugosity Methods 14

Sample Design 14

Data Analysis 14

Quantifying Seascape Metrics 15

Results 17

Hanauma Bay Marine Life Conservation District (MLCD) and Southeast Oahu 17

 Sample Allocation 17

	 	 Characterization of Spatial Patterns of the Seascape and MLCD Configuration	 17

 Large-scale Benthic Cover 19

 Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes 24

 Spatial and Temporal Patterns in Benthic Cover within the Management Regimes 24

 Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes 34

 Fish Trophic Structure Between Management Regimes and Among Habitats 43

Pupukea MLCD and North Shore Oahu 45

 Sample Allocation 45

	 	 Characterization of Spatial Patterns of the Seascape and MLCD Configuration	 46

 Evaluation of MLCD Boundary Expansion Using a Seascape Perspective 48

 Large-scale Benthic Cover 49

 Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes 49

 Spatial and Temporal Patterns in Benthic Cover within the Management Regimes 52

 Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes 59

	 	 Fish Trophic Structure Between Management Regimes and Among Habitats	 68

Honolua-Mokuleia MLCD and West Maui 70

 Sample Allocation 70

	 	 Characterization of Spatial Patterns of the Seascape and MLCD Configuration	 71

 Large-scale Benthic Cover 71

 Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes 73

	 	 Spatial and Temporal Patterns in Benthic Cover within the Management Regimes	 76

 Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes 83

 Fish Trophic Structure Between Management Regimes and Among Habitats 92
TABLE OF CONTENTS

Kealakekua Bay MLCD and South Kona 94

 Sample Allocation 94

 Characterization of Spatial Patterns of the Seascape within the MLCD 95

 Large-scale Benthic Cover 97

 Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes 102

 Spatial and Temporal Patterns in Benthic Cover within the Management Regimes 105

 Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes 111

 Fish Trophic Structure Between Management Regimes and Among Habitats 121

Discussion 125

Benthic Habitats 125

Seascape Metrics 125

	 Fish Assemblages	 126

Management Implications 127

 

References 129
Executive Summary

Photo courtesy of Keoki Stender, www.marinelifephotography.com
Executive Summary

Monitoring Hawaii’s Marine Protected Areas i

Hawaii’s coastal marine resources have declined dramatically over the past 100 years due to multiple anthropogenic stressors including overfishing, coastal development, pollution, overuse, invasive species and climate 

change. It is now becoming evident that ecosystem-based management, in the form of marine protected areas 

(MPAs), is necessary to conserve biodiversity, maintain viable fisheries, and deliver a broad suite of ecosystem 

services. Over the past four decades, Hawaii has developed a system of MPAs to conserve and replenish marine resources around the state. These Marine Life Conservation Districts (MLCDs) vary in size, habitat quality, 

and management regimes, providing an excellent opportunity to test hypotheses concerning MPA design and 

function using multiple discreet sampling units.

NOAA/NOS/NCCOS/Center for Coastal Monitoring and Assessment’s Biogeography Branch used digital benthic habitat maps coupled with comprehensive ecological studies between 2002 and 2004 to evaluate the 

efficacy of all existing MLCDs using a spatially-explicit stratified random sampling design. The results from 

this work have shown that areas fully protected from fishing had higher fish biomass, larger overall fish size, 

and higher biodiversity than adjacent areas of similar habitat quality. Other key findings demonstrated that top 

predators and other important fisheries species were more abundant and larger in the MPAs, illustrating the 

effectiveness of these closures in conserving these populations. Habitat complexity, protected area size and 

habitat diversity were the major factors in determining effectiveness among MPAs.

In order to more effectively evaluate these MLCDs, a seascape ecology approach was subsequently applied to 

characterize ecosystem patterns at scales that are commensurate with marine resources and their users. Furthermore, a monitoring program in four representative MLCDs (Hanauma Bay and Pupukea, Oahu; HonoluaMokuleia Bay, Maui, and Kealakekua Bay, Hawaii Island) was established and these sites were sampled annually between 2006-2008. These study sites were chosen because they are characterized by a wide range of 

habitats types, representative wave exposures, varying levels of resource protection and human use. NOAA’s 

2007 benthic habitat maps, which utilize a more robust classification scheme, were also integrated to apply a 

stratified random sampling design and to support a seascape ecology approach in order to characterize and 

quantify the seascape within each MLCD and adjacent area open to fishing. General benthic habitat types 

were similar inside and outside MLCDs (Friedlander et al. 2007b) although coral cover tended to be higher inside. Therefore, the focus of this study was to examine trends over time in each of these management regimes. 

This approach allowed for the identification of seascape characteristics that support robust and effective MPAs 

in the state of Hawaii. The major findings of the study are highlighted below:

Benthic Assemblage Characteristics Among Study Sites and Over Time:

• Coral species richness and cover was greater inside the Hanauma, Pupukea and Honolua-Mokuleia Bay 

MLCDs compared to adjacent open areas and the rank order of these metrics among the four MLCDs 

did not change over the study period.

• Coral cover generally increased over time in Hanauma, Pupukea and Honolua-Mokuleia Bay MLCDs 

and did not change within Kealakekua MLCD.

• Macroalgae cover was generally greater outside the MLCDs and increased over time, particularly adjacent to the Hanauma Bay MLCD (Maunalua Bay, Oahu), Honolua Bay MLCD (west Maui) and Kealakekua Bay MLCD (south Kona, Hawaii Island). 

Seascape Characteristics Among Study Sites:

• Seascape metrics derived from the 2007 NOAA benthic habitat maps were used to characterize the 

spatial structure and composition of habitat in the MLCDs, as well as the spatial configuration of the protected area boundary. These metrics included Shannon’s habitat diversity and evenness indices, patch 

fractal dimensions, MLCD perimeter and area and MLCD perimeter/area ratio.
Executive Summary

ii Monitoring Hawaii’s Marine Protected Areas

• Several LiDAR-derived seascape metrics (e.g., depth, rugosity, slope) were developed for each MLCD 

to better describe the three dimensional seascape structure within each protected area.

• The quantification of both two (e.g. habitat diversity) and three dimensional (e.g. habitat complexity) 

structure of the seascape in the MLCDs allowed for the characterization of seascape attributes that support a successful protected area and may inform future marine spatial planning efforts.

• Changes to the size and spatial configuration of the Pupukea MLCD since 2003 have greatly increased 

the habitat quality and ecosystem health within this protected area.

Fish Assemblage Characteristics Among Study Sites and Over Time:

• Rank order of MLCDs by fish assemblage characteristic (species richness, numerical abundance, biomass and diversity) remained the same between the 2002-2004 sampling period and the 2006-2008 

sampling period.

• Fish biomass increased in three of the four MLCDs from the initial sampling period. There was a 14% 

increase in Pupukea, a 7% increase in Kealakekua and a 5% increase in Honolua Bay. Biomass in 

Hanauma Bay did not change appreciably during this same period. 

• Fish biomass declined by 39% outside the Pupukea MLCD on Oahu but increased by 32% outside 

Hanauma Bay on the south shore of Oahu, although the absolute increase (+0.05 t ha-1) was small. 

• Apex predator biomass increased in the Pupukea MLCD and in the Hanauma Bay MLCD as a result of 

increasing numbers and sizes of bluefin trevally jacks (omilu, Caranx melamypgus). Herbivore biomass 

increased by 17% within the Pupukea MLCD and this increase was driven mainly by large parrotfishes 

that are primary fisheries target species.

Combining seascape-scale habitat stratification with in situ ecological data allowed for the development of a 

statistically robust monitoring program of living marine resources (e.g., fishes, corals and other invertebrates, 

algae) within and adjacent to four representative marine protected areas in Hawaii. The results clearly show 

that areas with good habitat quality and management conserve fish populations within their boundaries while 

areas without protection are in poorer ecological health and continue to decline over time. These temporal patterns support the findings of the initial assessment conducted by Friedlander and colleagues (Friedlander et 

al. 2006, 2007a, 2007b) and increase confidence in the effectiveness of marine protected areas as a management tool for the conservation of nearshore marine resources in Hawaii.

Remotely sensed data allowed the examination of these MLCDs at spatial scales that are more appropriate for 

management and provide valuable information on seascape metrics (e.g. rugosity, slope, depth, habitat type) 

that are relevant to the design of future MPAs. Marine protected areas are spatially discrete forms of management and georeferenced data on seascape metrics can be integrated using a spatial framework to ensure 

MPA design scenarios are incorporating an appropriate range in depth, habitat complexity, and a mosaic of 

interconnected habitat types. 

The findings from this study greatly contribute to the understanding of marine protected area design and function in Hawaii and can be useful in the development of comprehensive coastal and marine spatial planning. 

Introduction

Photo courtesy of Kosta Stamoulis
Introduction

Monitoring Hawaii’s Marine Protected Areas 1

Background

Coral reef ecosystems are facing overexploitation and severe depletion on a global scale (Birkeland 2004, 

Bellwood et al. 2004, Pandolfi et al. 2005). Pollution, coastal development, and invasive species all impact 

coral reefs locally, and climate change now is having a global effect on corals and coral reef communities. Fishing, however, has historically exerted the most direct influence on most reefs and other marine ecosystems 

(Jennings and Kaiser 1998, Jackson et al. 2001). Increasing fishing pressure on coral reefs over the last few 

decades has not only led to declines in valued fish stocks, but have also resulted in major impacts to this fragile 

ecosystem as a whole.

Marine resources were important to 

the ancient Hawaiians for subsistence, 

culture, and survival. Today, food, recreation, culture, commerce, aesthetics, 

and shoreline protection are just a few 

of the ecosystem services provided by 

Hawaii’s coral reefs. These reefs also 

have extremely high biodiversity and 

conservation value due to large proportion of species found nowhere else on 

earth. However, the 1.2 million residents 

and nearly seven million tourists each 

year have put increasing pressure on 

Hawaii’s coral reefs, which have been 

valued at over US$10 billion (Figure 1; Cesar and van Beukering 2004, Friedlander et al. 2008a). 

The combination of increased fishing pressure and growing coastal populations has reduced the ability to 

achieve the management goal of sustainable fisheries. In the past, Hawaiians managed fisheries in a sustainable manner based on traditional ecological knowledge and restricted marine resources that were more vulnerable to over harvesting (Poepoe et al. 2007). Currently, the main Hawaiian Islands are faced with the problems 

of increasing anthropogenic impacts that are a result of a growing coastal population and tourism (Friedlander 

et al. 2008a,b). The effects of fishing are evident at the level of individual stocks, as well as throughout the entire ecosystem (Friedlander and DeMartini 2002, Williams et al. 2008). In order to preserve marine biodiversity, 

ecosystem function, and the goods and services provided by resilient systems, marine reserves have been 

increasingly recommended as part of an ecosystem-based approach to management. Benefits derived from 

marine reserves include the enhancement of fisheries, insurance against management failures, the protection 

of essential fish habitat, and increased recruitment of fishes to adjacent open areas (Lubchenco et al. 2003, 

Browman and Stergiou 2004). 

Objectives

The focus of this study was to develop and implement a monitoring program of living marine resources (e.g., 

fishes, corals and other invertebrates, algae) within and adjacent to four representative MLCDs, develop metrics of seascape using remotely sensed data, and organize these data in a GIS to better understand ecosystem processes at the seascape scale.

Specifically:

• Examine temporal variation in fish assemblages and benthic habitat cover across a range of habitat 

types and management strata to determine trends and variability over time. 

• Develop metrics from remotely sensed data (e.g., LiDAR, benthic habitat maps) that provide seascapelevel characterization of the benthic environment.

Figure 1. Honolulu’s crowded coast (left). Photo: A. Clarke. Hawaii’s coral 

reefs provide world famous surfing spots like the Banzai Pipeline on Oahu’s 

north shore (right). Photo: J. Stahl. 
Introduction

2 Monitoring Hawaii’s Marine Protected Areas 

• Integrate remotely sensed data with ecological data into GIS to better understand ecosystem processes 

at broader spatial scales.

• Determine efficacy of marine protected areas using a seascape ecology approach.

Seascape Ecology Approach

Effective management of species and assemblages of concern requires accurate, spatially explicit information that characterizes and quantifies the seascape that supports the resources. Landscape ecology has been 

widely applied in the terrestrial environment to understand ecological patterns and processes, and this information is often used to inform conservation planning and resource management actions (Turner 2005). A 

landscape ecology approach can be integrated within a geospatial framework using Geographic Information 

Systems (GIS) and remote sensing to identify the spatial association of reef fishes with the seascape. Information on the seascape, derived from remote sensing methods (e.g., LiDAR, benthic habitat maps), can provide 

valuable and spatially discrete ecological criteria that can be integrated in GIS, along with social and economic 

considerations, and ultimately provide support for marine spatial planning applications (Pittman et al. 2007a,b, 

Wedding et al. 2008, Wedding and Friedlander 2008, Monaco et al. 2005). This integrated approach aids 

in defining the forces that shape broad-scale community structure, and addresses specific questions about 

particular groups of economically and ecologically important species at the regional scale that management 

decisions are typically implemented across the state of Hawaii. 

Benthic habitat type is a major determinant of fish assemblage structure and 

can be used as a surrogate to explain 

the distribution of fishes across the seascape (Figure 2). Benthic habitat maps 

provide fundamental information that 

can be used to guide marine conservation and management actions such as 

marine protected area (MPA) design, 

and evaluation (Friedlander et al. 2008 

a,b). These maps allow a better understanding of the seascape, ecosystem 

function, and species habitat utilization 

patterns in a coastal area (Christensen 

et al. 2003, Friedlander et al. 2003, Pittman et al. 2007a). This work evaluated 

four marine protected areas in Hawaii by 

utilizing NOAA’s Biogeography Branch 

benthic habitat maps in order to utilize 

a seascape approach. Further, these maps were used to characterize and quantify the seascape within each 

marine protected area to assist in understanding the important attributes of the seascape that support high 

abundance, diversity and biomass.

Hawaii Marine Protected Areas

Hawaii established its first MPAs over 30 years ago and since that time numerous protected areas have been 

established with varying levels of protection, ranging from complete ‘no-take’ areas to areas that have allowed 

a wide variety of activities to occur within their boundaries (Figure 3). The state of Hawaii, Division of Aquatic 

Resources has developed Marine Life Conservation Districts (MLCDs) to conserve and replenish marine resources around the state. Between 2002 and 2004, NOAA/NOS/NCCOS/Center for Coastal Monitoring and 

Assessment’s Biogeography Branch conducted a comprehensive evaluation of all MLCDs in the state (Friedlander et al. 2006, 2007a, 2007b). Results from this comprehensive assessment showed MLCDs protected 

from fishing, with high habitat complexity and good habitat quality (e.g., high coral cover and low macroalFigure 2. A school of white cheek surgeonfish swim over typical Hawaiian 

reef structure. Photo: K. Stamoulis. 
Introduction

Monitoring Hawaii’s Marine Protected Areas 3

gae cover), had higher values for most 

fish assemblage characteristics. To 

more effectively assess the efficacy 

of these protected areas, a monitoring 

program in four representative MLCDs 

(Hanauma Bay, Honolua-Mokuleia Bay, 

Kealakekua Bay and Pupukea) was 

established and these sites were sampled annually between 2006 and 2008. 

These four sites were chosen because 

they were representative of the diverse 

habitat types, wave exposures, and 

benthic environments across a broad 

geographic area of the main Hawaiian 

Islands. 

Figure 3. Hanauma Bay Marine Life Conservation District (MLCD) was 

designated as the first “no-take” marine protected area (MPA) in Hawaii in 

1967 and encompasses approximately 41 ha. This area receives over one 

million visitors per year and is the most visited MPA in the world. Photo: L. 

Wedding. 
Methods

Photo courtesy of Keoki Stender, www.marinelifephotography.com
Methods

Monitoring Hawaii’s Marine Protected Areas 5

Study Sites

The study areas are located in the 

main Hawaiian Islands at Pupukea and 

Hanauma Bay on the island of Oahu; 

Honolua-Mokuleia Bay, Maui; and Kealakekua Bay, Hawaii (Figure 4 and 

Table 1). These study sites were chosen because they are characterized by 

a wide range of habitats types, representative wave exposures, varying levels of resource protection, and human 

use. Each site has an MPA, referred to 

as MLCDs in Hawaii, and an adjacent 

area of comparable habitat that is open 

to resource extraction. This provides an 

opportunity to examine the relationship 

between relatively intact fish assemblages (MPAs), and their associated 

habitat, as well as the ability to compare 

the strength of the fish-habitat relationships in adjacent open access areas. Figure 4. Map of the Hawaiian Islands and study locations.

Table 1. Summary of Marine Life Conservation Districts at four study sites in the main Hawaiian Islands.

Marine Life 

Conservation 

District

Island Established Area

(km2

)

Use Protection 

from fishing

Permitted activities

Pupukea Oahu 1983* (2003) 0.71 moderate moderate pole-and-line from shore; harvest of seaweed (seasonal)**

Hanauma Bay Oahu 1967 0.41 high high complete no-take

Honolua-Mokuleia 

Bay

Maui 1978 0.19 moderate high complete no-take

Kealakekua Bay Hawaii 1969 1.24 high moderate hook and line; throw net; har- vest of seaweed (seasonal)** 

and crustaceans in 60% of the 

total area

“Use” represents the level of use that was classified by the state of Hawaii, Department of Land and Natural Resources, Division of Aquatic Resources (DAR 1992). Protection from fishing is based on the qualitative ranking of Division 

of Aquatic Resources (DAR) regulations. 

*Pupukea Marine Life Conservation District (MLCD) boundary was expanded and rules were modified in 2003.

**Selar crumenophthalmus (November-December) and Decapterus spp. (August-September)

Benthic Habitat Mapping

NOAA acquired and visually interpreted orthorectified satellite imagery for selected near-shore waters and 

parts of the main Hawaiian Islands. An important step in producing benthic habitat maps was the development 

of a habitat classification scheme. Thirty-two distict benthic habitat types (e.g., four major and 14 detailed 

geomorphological structure classes; eight major and three detailed biological cover types) within 13 attribution zones were digitally mapped in GIS using heads-up visual interpretation of orthorectified satellite imagery 

(Tables 2, 3 and 4). 

The classification scheme was influenced by many factors including requests from the management community, the National Ocean Service's coral reef mapping experience, existing classification schemes, and quantitative habitat data for the area. The hierarchical scheme allows users to expand or collapse the thematic detail 

of the resulting map to suit their needs. For instance, additional hierarchical categories can be added into a 

GIS system by users with more detailed knowledge or data for specific areas. For example, habitat polygons 
Methods

6 Monitoring Hawaii’s Marine Protected Areas 

Table 2. Description of attribute “Zone” in the 2007 NOAA benthic habitat classification scheme. Source: Battista et al. 

2007. 

Zone Description Typical Habitats

Shoreline 

intertidal

Area between the mean high water line and the lowest spring tide 

level.

Sand, seagrass, rocks, reef and man- groves.

Vertical wall Near-vertical slope between the shore and the shelf. Not usually 

distinguishable in remotely sensed imagery. 

Rock, boulder, algae and coral

Lagoon Shallow, between shore and intertidal zone, only present when 

reef crest is present.

Sand, seagrass, algae, hard bottom, 

patch reefs

Back reef Between edge of reef crest and lagoon floor. Only present when 

a reef crest and lagoon exists.

Sand, rubble, seagrass, algae patch 

reef

Reef flat Shallow, between intertidal zone and reef crest. Not usually pres- ent if a lagoon is present.Sand, rubble, seagrass, algae, patch 

reef

Reef crest Flattened emergent or nearly emerged segment of reef between 

back reef and fore reef.

Rubble, aggregated coral

Fore reef Between edges of reef crest sloping deeper to the edge of the 

shelf platform. Also, features not forming emergent reef with a 

slope greater than the shelf.

Spur and groove

Bank/Shelf A flattened platform extending seaward from the fore reef to the 

insular shelf escarpment that drops to deeper water. 

Sand, patch reefs, algae, pavement, 

sand channels

Bank/Shelf 

Escarpment

Edge of shelf where depth increases rapidly to oceanic water. Sand, spur and groove

Channel Naturally occurring channels can cut across other zones. Sand, mud, pavement

Dredged Natural geomorphology disrupted or altered by excavation or 

dredging.

Mud, sand

Unknown Zone, cover, and structure un-interpretable due to turbidity, cloud 

cover, water depth or other interference.

Land Terrestrial features above the spring high tide line.

Table 3. Description of attribute “Geomorphic Structure” in the 2007 NOAA benthic habitat classification scheme. Source: 

Battista et al. 2007. 

Geomorphic Structure Description

Unconsolidated 

sediment

Sand Coarse sediment typically found in exposed areas

Mud Fine sediment, associated with river discharge and sheltered areas

Coral Reef and Hard

bottom

Spur and groove Alternating sand and high relief coral formations, oriented perpendicu- lar to shore

Patch reef Isolated coral formations with no organized structural axis

Individual patch reef Distinctive single patch that are larger or equal to the minimum map- ping unit

Aggregate patch reef Clustered patches that are too small or too close together to map 

separately

Aggregate reef High relief lacking sand channels of spur and groove

Scattered coral/rock in 

unconsolidated sediment

Sand or seagrass bottom with small rocks and coral heads too small to 

delineate individually

Pavement Flat, low-relief, solid carbonate rock with biological cover dense enough 

to obscure surface

Rock/boulder Solid carbonate blocks, boulders, or volcanic rock.

Reef rubble Dead, unstable coral rubble often colonized by biological cover

Pavement with sand 

channels

Alternating low-relief sand/surge channels oriented perpendicular to 

shore. 

Other Delineations Artificial Man-made, including: wrecks, piers, submerged jetties, fish ponds, and 

shoreline created by dredge spoil

Land Terrestrial features above the spring high tide line

Unknown -- Zone, cover, and structure un-interpretable due to turbidity, cloud cover, 

water depth or other interference.
Methods

Monitoring Hawaii’s Marine Protected Areas 7

Table 4. Description of attribute “Biological Cover” in the 2007 NOAA benthic habitat classification scheme. The minimum 

mapping unit (MMU) is 1 acre. Source: Battista et al. 2007. 

Biological Cover 

Live Coral Substrates colonized by live reef building corals and other organisms with at least 10% live coral cover.

Continuous coral Live coral covering > 90% of the substrate. May include areas < 90% coral cover if 

< 10% of the total area is too small to be mapped independently (< 1 MMU).

Patchy coral Discontinuous live coral with breaks in coverage (< 1 MMU) to be mapped as continuous coral. Live coral cover 50% > 90% of the bottom.

Sparse coral Discontinuous live coral with breaks in coverage (< 1 MMU) to be mapped as continuous coral. Live coral cover 10% > 50% of the bottom.

Seagrass Habitat with 10% or more of seagrass.

Continuous seagrassSeagrass > 90% of the substrate. May include blowouts < 10% of the total area that 

are too small to be mapped independently (< 1 MMU).

Patchy seagrass Discontinuous seagrass community with breaks in coverage (< 1 MMU) to be 

mapped as continuous seagrass. Cover 50% > 90% of the bottom.

Sparse seagrass Discontinuous seagrass community with breaks in coverage (< 1 MMU) to be 

mapped as continuous seagrass. Cover 10% > 50% of the bottom.

Macroalgae Substrates with > 10 % coverage of any combination of numerous species of red, green, or brown macroalgae. 

 Continuous

macroalgae

Macroalgae covering > 90 % of the substrate. May include areas < 10% of the total 

area that < 1 MMU.

Patchy macroalgae Discontinuous macroalgae with breaks in coverage (< 1 MMU) to be mapped as 

continuous macroalgae. Cover 50% > 90% of the bottom.

Sparse macroalgae Discontinuous macroalgae with breaks in coverage that are too diffuse or irregular, or result in isolated patches of macroalgae that are too small (smaller than the 

MMU) to be mapped as continuous macroalgae. Overall cover is estimated at 10% 

> 50% of the bottom.

Encrusting/

Coralline Algae

 > 10% coverage of any combination of encrusting or coralline algae. 

 Continuous 

corraline algae

Coralline algae covering > 90% of the substrate. May include areas < 10% of the 

total area that are < 1 MMU.

Patch corraline 

algae

Discontinuous coralline algae with breaks in coverage (< 1 MMU) to be mapped as 

continuous coralline algae. Cover 50% > 90% of the bottom.

Sparse corraline 

algae

Discontinuous coralline algae with breaks in coverage (< 1 MMU) to be mapped as 

continuous coralline algae. Cover 10% > 50% of the bottom.

Turf Algae A community of low lying species of marine algae composed of algal divisions dominated by filamentous 

species lacking upright fleshy macroalgal thali.

Continuous turf Turf algae > 90 %of the substrate. May include areas < 10 % of the total area that 

are< 1 MMU.

Patchy turf Discontinuous Turf algae with breaks in coverage (< 1 MMU) to be mapped as continuous Turf algae. Cover50% > 90% of the bottom.

Sparse turf Discontinuous Turf algae with breaks in coverage (< 1 MMU) to be mapped as continuous Turf algae. Cover 10% < 50% of the bottom.

Emergent 

vegetation

Emergent habitat composed primarily of Rhizophora mangle (red mangrove) and Hibiscus sp (hau) trees. 

Continuous 

vegetation

Emergent vegetation > 90 % of the substrate. May include areas < 10% of the total 

area that are < 1 MMU.

Patchy vegetation Discontinuous Emergent vegetation with breaks in coverage (< 1 MMU) to be 

mapped as continuous Emergent vegetation. Cover 50% < 90% of the bottom.

Sparse vegetation Discontinuous Emergent vegetation with breaks in coverage (< 1 MMU) to be 

mapped as continuous Emergent vegetation. Cover 10% > 50% of the bottom.

Uncolonized Substrates not covered with a min. of 10% of any of the above biological cover types. This habitat is usually on sand or mud structures.

Unknown Zone, Cover, and Structure uninterpretable due to turbidity, cloud cover, water depth, or other interference.
Methods

8 Monitoring Hawaii’s Marine Protected Areas 

smaller than the minimum mapping unit (the minimum mapping unit, or MMU, is 1 acre for NOAA maps) can 

be delineated, such as individual patch reefs, or habitat polygons delineated as colonized pavement using 

this scheme could be further attributed with health information (e.g., bleached, percent live cover) or species 

composition. For more information, visit http://ccma.nos.noaa.gov/products/biogeography/hawaii_cd_07/htm/

overview.html.

The hierarchical scheme was modified throughout the development process based upon feedback provided by 

workshop participants and other contributors. Additional modifications were made during the mapping process 

to ensure that each category definition reflected the intended habitats and zones encountered in the field as 

accurately as possible. For instance, the separation of biological cover and geomorphological structure in the 

present scheme represents a significant evolution of previous versions of the classification schemes developed for mapping of the Caribbean in 1999 and of the Hawaiian Islands in 2000. The major product of this effort 

is a series of GIS-based benthic habitat maps that are characterized by a high degree of spatial and thematic 

accuracy (Figures 5 and 6; Battista et al. 2007, Coyne et al. 2003).
Methods

Monitoring Hawaii’s Marine Protected Areas 9

Figure 5. Example of NOAA’s Biogeography Branch benthic habitat map of geomorphic structure in Kealakekua Bay, 

Hawaii.
Methods

10 Monitoring Hawaii’s Marine Protected Areas 

Figure 6. Example of NOAA’s Biogeography Branch benthic habitat map of biological cover in Kealakekua Bay, Hawaii.
Methods

Monitoring Hawaii’s Marine Protected Areas 11

Bathymetric LiDAR Data

The U.S. Army Corps of Engineers 

Scanning Hydrographic Operational 

Airborne LiDAR Survey (SHOALS) system is an airborne LiDAR bathymeter 

utilized to remotely collect topographic 

and bathymetric measurements using infrared (1,064 nm) and blue-green 

(532 nm) scanning laser pulses (Figure 

7). SHOALS LiDAR data was collected 

in Hawaii between 1999 and 2000. The 

sensor provides a vertical accuracy of ± 

20 cm and a horizontal resolution of ± 

1.5 meters. The minimum depth range 

is typically 0 - 1 meter, with a maximum 

depth range of approximately 40 meters, and a spatial resolution of 4 meters 

(Irish and Lillycrop 1999; Figures 7, 8 

and 9).

Sampling Locations

A field team consisting of two divers 

navigated to waypoints using GPS, they 

then marked the location with a lead 

weight and float and accurately established the location using GPS measurements. Direction of each transect was 

determined randomly along the isobath 

of that point except in cases where that 

direction caused the transect to traverse multiple habitats. In those situations, transects were run within a habitat polygon at a similar isobath strata. 

Divers descend together; with diver 1 

carrying a 25 m transect line and rugosity chain and diver 2 carrying a digital 

camera and a 1 square-meter quadrat. 

Diver 1 began a 25 x 5 m fish transect 

starting at the marked waypoint and 

moved along the depth contour. As the 

fish count diver started his/her count, 

he or she visualized out to the end of 

the transect line and enumerated all individuals that were potentially leaving 

the census area. This partially accounts 

for the behavior that targeted species 

acquire in areas that are frequented by 

spearfishers. The fish count method is 

described in detail below. As diver 1 laid 

out transect line, diver 2 conducted benthic surveys within the quadrat. Once diver 1 completed the fish transect, he/she conducted rugosity measurements as described below. 

Figure 7. Bathymetric grid derived from LiDAR data at Hanauma Bay, Oahu.

Figure 8. Aerial imagery of Hanauma Bay, Oahu overlaid on LiDAR-derived 

digital elevation model.
Methods

12 Monitoring Hawaii’s Marine Protected Areas 

Fish Sampling Methodology

Fish assemblages at each location 

were assessed using standard underwater visual belt transect survey methods (Brock, 1954; Brock, 1982). A diver 

swam each 25m x 5m transect at a constant speed (approximately 15 min/transect) and identified to the lowest possible taxon, all fishes visible within 2.5 

m to either side of the centerline (125 m2

transect area). Nomenclature followed 

Randall (1996). Total length (TL) of fish 

was estimated to the nearest centimeter. 

Length estimates of fishes from visual 

censuses were converted to weight using the following length-weight conversion: W = aSLβ where the parameters 

a and β are constants for the allometric 

growth equation and SL is standard 

length in mm and W is weight in grams. 

Total length was converted to standard 

length (SL) by multiplying standard length to total length-fitting parameters obtained from FishBase (www.fishbase.org). Length-weight fitting parameters were available for 150 species commonly observed on visual fish 

transects in Hawaii (Hawaii Cooperative Fishery Research Unit, unpublished data). This was supplemented by 

using information from other published and web-based sources. In the cases where length-weight information 

did not exist for a given species, the parameters from similar bodied congeners were used. All biomass estimates were converted to metric tons per hectare (t/ha) to facilitate comparisons with other studies in Hawaii. 

Finally, fish taxa were categorized into three trophic categories (herbivores, secondary consumers and apex 

predators) according to various published sources and FishBase (www.fishbase.org).

Fish Sample Size Optimization Analysis

Optimal sample size was determined for 

number of species and number of individuals per transect among the four major habitat types surveyed in the study 

area (Table 5, Figures 10 and 11). A 

technique developed by Bros and Cowell (1987) using the standard error of the 

mean to resolve statistical power was 

used to determine the number of samples needed using number of species 

and number of individuals. This method 

uses a Monte Carlo simulation procedure to generate a range of sample sizes versus power. The sample size at which further increase in sample 

size does not substantially increase power (decreasing standard error of the mean) is taken as the minimum 

suitable number of samples. For number of species per transect, high and low standard error of the mean 

began to level off and converge at ca. four samples in the colonized hardbottom habitat and unconsolidated 

sediment habitat and ca. eight samples for the macroalgae and uncolonized hardbottom habitats. For number 

of individuals per transect, high and low standard error of the mean began to converge at six samples in the 

unconsolidated sediment habitat and nine samples in the macroalgae habitat. Given these results, nine to 10 

samples per habitat appeared to be adequate to control the standard error of the mean for number of individuals and number of species per transect.

DISTANCE (KM)

DEPTH (M)

Figure 9. Bathymetric profile of Hanauma Bay, Oahu derived from LiDAR 

data.

Table 5. Summary of sampling allocation of transects by location and man- agement regime with a total sample size of n=329 per year. 

Pupukea Hanauma 

Bay

HonoluaMokuleia 

Bay

Kealakekua 

Bay

Total

MLCD 35 33 37 34 139

FRA 29 29

Open areas 38 47 63 13 161

Total 73 80 100 76 329

* MLCD = Marine Life Conservation District; FRA = Fisheries Replenishment Area; 

Open = Area open to resource extraction.
Methods

Monitoring Hawaii’s Marine Protected Areas 13

Benthic Survey Techniques

The four MLCDs and the surrounding 

benthic habitat were surveyed using 

the in situ planar point intercept quadrat method (Reed 1980). Each 25 m 

fish transect was stratified into 5 x 5 m 

segments with quadrat placement randomly allocated within each segment. 

The quadrat grid was 1m2

 in area and 

consisted of 1 inch PVC tubing fitted 

with nylon line spaced 10 centimeters 

apart to form a square grid with 81 intersections. A subset of 25 randomly 

selected intersections were marked 

and used for substrate identification. 

The rationale for the subset was that 25 

points sufficiently represented the habitat with acceptable error and optimized 

sampling time (Friedlander et al. 2006). 

Each intersection was identified using 

substrate categories of sand, coralline 

algae, turf algae, macroalgae, Halimeda

spp and coral. Coral and macroinvertebrates were identified down to species 

using Veron (2000) and Hoover (2002) 

respectively. The macroinvertebrates 

category incorporated echinoderms 

and other large invertebrates (e.g., zooanthids, octocorals) that occupied significant portions of the substrate. Macroinvertebrates were also included in 

the results for comparative purposes, 

but the methodology limited conclusions about distribution and abundance 

for this group of organisms. Limitations of in situ methodology precluded 

taxonomic resolution of algae down to 

the species level so algae were identified to genera using Littler and Littler 

(2003). Percent cover values for each 

substrate category and coral species 

were derived by dividing the number of 

occupied points by the total number of 

intersections (25) within each quadrat.

Figure 10. Four major habitats using NOAA benthic habitat maps to inform 

a stratified random sampling approach types (Maunalua Bay and Hanauma 

Bay, Oahu).

Figure 11. GPS points of sampling locations at (A) Pupukea (B) Hanauma 

Bay (C) Honolua-Mokuleia and (D) Kealakekua Bay MLCDs in the main 

Hawaiian Islands.
Methods

14 Monitoring Hawaii’s Marine Protected Areas 

Rugosity Methods

To measure reef rugosity or surface relief, a chain of small links (1.3 cm per link) was draped along the full 

length of the centerline of each transect (Risk 1972). Care was taken to ensure that the chain followed the 

contour of all natural fixed surfaces directly below the transect centerline. A ratio of distance along the reef 

surface contour to linear horizontal distance gave an index of spatial relief or rugosity. 

Sample Design

A stratified random sampling design was employed (Figure 12). Habitat polygons were attributed into four major habitat types (colonized hard bottom, uncolonized hard bottom, macroalgae and sand). Within each major 

habitat type, sampling was further stratified by management regime (MLCD; fisheries management area, or 

FMA; and open access). All sampling locations were randomly generated in ArcGIS (Figure 13). 

Colonized hard bottom Uncolonized hard bottom Macroalgae Sand

MLCD FMA Open MLCD FMA Open MLCD FMA Open MLCD FMA Open

Habitats

Management Strata

Colonized hard

 bottom

Uncolonized 

hard bottom

Macroalgae Sand

The overall benthic assemblages were generally similar between the MLCDs and the corresponding areas 

open to fishing based on multivariate analysis of similarities (ANOSIM; Friedlander et al. 2007). The pairwise 

ANOSIM R test value comparing benthic habitat composition between MLCDs and open areas was highest in 

Pupukea (R = 0.327), followed by Kealakekua (R = 0.107), Hanauma (R = 0.040) and Honolua (R = 0.002). 

The ANOSIM R statistic ranges from well separated (R>0.75), overlapping but clearly different (R>0.5), or 

barely separable at all (R<0.25). The small R statistics indicated that these benthic assemblages were still 

relatively similar to their adjacent open access areas.

General benthic habitat types were similar inside and outside MLCDs (Friedlander et al. 2007b) although coral 

cover tended to be higher inside. Therefore, the focus of this study was to examine trends over time in each 

of these management regimes. The trends over time tended to be consistent with the level of protection (e.g. 

MLCD).

Figure 12. A stratified random sampling design was employed to assess MLCDs, FMAs and areas open to fishing by 

major habitat types.

Data Analysis

For fish assemblage characteristics, the number of individuals and biomass were ln(x+1) transformed for 

statistical analysis. Numbers of individuals were converted to number/m2

 and biomass was converted to t/

ha for comparisons with other studies throughout the state. Comparisons of fish species richness, biomass, 

and diversity among management strata were conducted using a Nested Analysis of Variance (ANOVA) us-
Methods

Monitoring Hawaii’s Marine Protected Areas 15

ing only the habitat types common to 

all management strata. Significant differences between pairs were examined 

using the Tukey-Kramer HSD (honestly 

significant difference) test for ANOVAs 

(α = 0.05).

Species diversity was calculated from 

the Shannon-Weaver Diversity Index 

(Ludwig and Reynolds 1988): H’=S (pi

ln pi), where pi is the proportion of all 

individuals counted that were of species

i. An index of relative dominance (IRD) 

for each fish taxa was created by multiplying the percent frequency of occurrence of the taxa on each transect by 

the relative percent number of that taxa 

(Greenfield and Johnson 1990). 

Quantifying Seascape Metrics

Bathymetric grids were created from the 

LiDAR data for each study area by interpolation using Inverse Distance Weighting in ArcGIS Spatial Analyst (ESRI) at a 

grid cell size of 4 meters. Slope was derived from the gridded (4 m cell size) LiDAR bathymetry using the ArcGIS Spatial Analyst extension, where the raster 

cell values represented the maximum 

rate of change in elevation between 

neighboring cells and were calculated in 

degrees. Rugosity was derived from the 

gridded (4 m cell size) LiDAR bathymetry using the ‘Benthic Terrain Modeler 

for ArcGIS’, where the raster cell values 

reflected the ratio of the seascape surface area to the planimetric area determined in a neighborhood analysis (Jenness 2004, Lundblad et al. 2006). 

Figure 13. GIS layers used to quantify and characterize the seascape cover 

were compiled for each site by management regime.
Results

Photo courtesy of Keoki Stender, www.marinelifephotography.com
Results

Monitoring Hawaii’s Marine Protected Areas 17

HANAUMA BAY MLCD AND 

SOUTHEAST OAHU 

The south shore Oahu study area extended from Wailupe Peninsula to 

Sand Beach (9.1 km2) and included the 

Hanauma Bay MLCD (Figure 14). 

Sample Allocation

A total of 80 samples were collected in 

2004 and 2006, in 2007 and 2008 only 

hardbottom habitat was sampled (n=59; 

Table 6). The two levels of sampling 

stratification included major habitat 

types (CHB, MAC, UCH and UCS) and 

fisheries management regime (open access and MLCD). Macroalgae habitat 

was not present at the 1 acre minimum 

mapping unit within the MLCD.

Characterization of Spatial Patterns 

of the Seascape and MLCD 

Configuration

The total seascape area within Hanauma Bay MLCD was 0.41 km2

, with a 

perimeter to area ratio of 7.78 and a 

fractal dimension of 1.47. (Tables 7 and 

8). Based on the NOAA Biogeography 

Branch 2007 benthic habitat maps the 

habitat diversity, represented by Shannon’s diversity index and Shannon’s 

evenness index was 0.85 and 0.62 respectively (Table 7). Bathymetric grids 

were created from the LiDAR data within Hanauma Bay MLCD that represented an approximate depth range of 0–27.68 m ( x =8.55; Table 9 and 

Figure 14. Overview of Hanauma Bay, Maunalua Bay and adjacent areas 

with sampling locations.

Table 6. Sample allocation at Hanauma Bay Marine Life Conservation District by habitat (2004-2008).

2004 2006 2007 2008 Total

MLCD 33 33 22 22 110

 CHB 12 12 12 12 48

 UCH 10 10 10 10 40

 UCS 11 11 22

Open 47 47 37 37 168

 CHB 11 11 11 11 44

 MAC 11 11 11 11 44

 UCH 15 15 15 15 60

 UCS 10 10 20

Total 80 80 59 59 278

* MLCD = Marine Life Conservation District; CHB = Colonized hard bottom; UCH = Uncolonized hard bottom; UCS = Unconsolidated sediment; Open = Area open to resource extraction; MAC = Macroalgae.
18 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 15). Percent slope (ranging from 0-92.18, [ x =7.89]) and rugosity (ranging from 0–1.45, [ x =1.01]) were 

also derived from the bathymetric grids.

Table 7. Summary of seascape metrics derived from 2007 

NOAA Biogeography Branch benthic habitat maps. 

Hanauma Bay MLCD*

Shannon's diversity index 0.85

Shannon's evenness index 0.62

Mean patch fractal dimension 1.47

* MLCD = Marine Life Conservation District.

Table 8. Summary of marine reserve boundary analysis.

Hanauma Bay MLCD*

Reserve perimeter (km) 3.19

Reserve area (km2

) 0.41

Perimeter to area ratio 7.78

* MLCD = Marine Life Conservation District.

Table 9. Summary of seascape structure derived from LIDAR bathymetric grids for Hanauma Bay Marine Life Conserva- tion District.

Hanauma Bay MLCD*

Depth (m) Rugosity Slope (percent)

Average depth 8.55 Average rugosity 1.01 Average slope 7.89

Standard

deviation

6.67 Standard

deviation

0.02 Standard

deviation

8.58

Depth range 0 – 27.68 Rugosity range 1 – 1.45 Slope range 0 – 92.18

* MLCD = Marine Life Conservation District.

Figure 15. Aerial imagery of Hanauma Bay, Maunalua Bay and other adjacent open areas with other panels describing 

seascape characteristics (e.g. depth, slope, rugosity) derived from bathymetric LiDAR data.
Results

Monitoring Hawaii’s Marine Protected Areas 19

Large-scale Benthic Cover

Benthic coverage for the Hanauma 

Bay MLCD was derived from the 2007 

NOAA benthic habitat maps with the 

majority (69%) of the geomorphic structure comprised generally of coral reef 

and hard bottom within the MLCD (Table 

10, Figures 16 and 17). The coral reef 

and hard bottom structure consisted 

mostly of pavement (34%), aggregated 

reef (18%) and rock/boulder (15%) with 

only 2% of individual patch reef. The 

remaining portion of the MLCD was 

characterized by sand (22%). Biological cover was dominated by coral, with 

coral 10%-50% cover (48%) followed 

by coral 50%-90% cover (7%). The remaining portion of the biological cover 

in the MLCD consisted of uncolonized 

bottom (22%), turf (8%) and coralline algae (6%; Table 11, Figures 18 and 19).

Table 10. Geomorphic structure for the Hanauma Bay Marine Life Conservation District (MLCD) derived from NOAA benthic habitat maps.

Major Structure Detailed 

Structure

Area 

(km2)

Percentage

Coral Reef and Hard bottom Aggregate 

Reef 0.07 18%

Coral Reef and Hard bottom Individual 

Patch Reef 0.01 2%

Coral Reef and Hard bottom Pavement 0.14 34%

Coral Reef and Hard bottom Rock/Boulder 0.06 15%

Other Delineations Land 0.01 2%

Unconsolidated Sediment Sand 0.09 22%

Unknown Unknown 0.03 7%

Table 11. Biological cover for the Hanauma Bay MLCD derived from 

NOAA benthic habitat maps.

Major Cover Percent Cover Area (km2) Percentage

Coral 10% - 50% 0.20 48%

Coral 50% - 90% 0.03 7%

Coralline Algae 50% - 90% 0.02 6%

Turf 50% - 90% 0.03 8%

Unclassified 0.01 2%

Uncolonized 90% - 100% 0.09 22%

Unknown 0.03 7%
20 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 16. Sampling locations and benthic structure for the Hanauma Bay Marine Life Conservation District, Maunalua 

Bay and other adjacent open areas.
Results

Monitoring Hawaii’s Marine Protected Areas 21

Figure 17. Sampling locations and benthic structure for the Maunalua Bay study area.
22 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 18. Sampling locations and benthic cover for the Hanauma Bay MLCD, Maunalua Bay and other adjacent open 

areas.
Results

Monitoring Hawaii’s Marine Protected Areas 23

Figure 19. Sampling locations and benthic cover for the Maunalua Bay study area.
24 Monitoring Hawaii’s Marine Protected Areas

Results

Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes

Average coral species richness per 

transect for colonized hard bottom 

(CHB) habitat in Hanauma Bay MLCD 

was slightly greater than the adjacent 

open area with grand means of 5.77 

and 4.39, respectively. Average species 

richness in CHB habitat in the MLCD 

ranged from 5.67 (±0.28) in 2007 to 

5.83 (±0.41) in 2004. This compared 

to the adjacent open area where richness ranged from 4.09 (± 0.25) in 2004 

to 4.64 (± 0.62) in 2006 (Table 12 and 

Figure 20). Spatial patterns in coral richness from 2004-2007 increased overall 

in shallow water areas of both Maunalua 

and Hanauma Bays (Figure 20). Coral 

richness decreased over time for several transects, typically located in shallow water. For a large number of 

transects inside and outside the MLCD richness did not change. Generally, over time, coral richness increased 

in both the MLCD and open areas from 2004 to 2007 (Table 12, Figures 21 and 22a). Average coral richness 

was relatively stable across sampling years for both open CHB and MLCD CHB with the marcroalgae (MLCD) 

being greater across all years sampled. Uncolonized hard bottom (UCH) habitat in both areas generally had 

greater variability in coral species richness, possibly due to the patchy distribution of coral cover on the hard 

substrate in these environments. Patterns in coral richness in MAC habitat demonstrated similar variability with 

overall low values over time compared to the other two major habitats (Figure 22a). 

Spatial and Temporal Patterns in Benthic Cover within the Management Regimes

Average percent coral cover over the sampling period was more than three times greater in Hanauma Bay 

MLCD ( x =18.05) compared to the adjacent area ( x =5.65). Average percent coral cover per transect in 

Hanauma Bay MLCD ranged from 14.50% (±3.72) in 2004 to 25.20% (±4.15) in 2007. This compared to the 

adjacent open area where coral cover ranged from 4.69% (±1.02) in 2004 to 6.62% (±1.57) in 2007 (Figure 23). 

Patterns of percent change of coral cover per transects between 2004-2007 generally increases in Hanauma 

Bay MLCD while only increasing slightly or staying the same in Maunalua Bay (Figure 24). Temporal patterns 

in percent coral cover are generally consistent over the sampling period, with the greatest values consistently 

found within CHB habitat inside the MLCD in all three years (Figure 22b). 

Average percent macroalgal cover over the sampling period was greater in the open area ( x =35.48) compared to the MLCD ( x =4.36). Average percent macroalgal cover per transect in the MLCD ranged from 1.55% 

(±0.39) in 2004 to 4.36% (±0.75) in 2007. This compared to the adjacent open area where macroalgal cover 

ranged from 16.68% (±3.29) in 2004 to 35.48% (±4.14) in 2007 (Figure 25). Patterns of percent change of 

macroalgae cover on transects between 2004-2007 show a general increase in Manualua Bay with small or 

no increases in Hanauma Bay (Figure 26). Temporal patterns in percent macroalgae cover show increases in 

the open MAC but decreases in open UCH habitat across the study site (Figure 22c). Macroalgae cover in the 

MLCD showed small increases across all years in both the CHB and UCH habitat types.

Table 12. Comparison of coral species richness in major habitats and 

among different management regimes at Hanauma Bay, Oahu.

2004 2006 2007

Mean SE Mean SE Mean SE

MLCD

CHB 5.83 0.41 5.83 0.55 5.67 0.28

UCH 1.30 0.21 2.70 0.62 2.80 0.44

Open

CHB 4.09 0.25 4.64 0.62 4.36 0.64

MAC 0.09 0.09 0.82 0.63 0.18 0.18

UCH 1.53 0.24 3.00 0.48 2.53 0.54

* MLCD = Marine Life Conservation District; CHB = Colonized hard bottom; 

UCH = Uncolonized hard bottom; Open = Area open to resource extraction; 

MAC = Macroalgae
Results

Monitoring Hawaii’s Marine Protected Areas 25

Figure 20. Coral species richness by individual transects for southeast Oahu study area including Hanauma Bay MLCD, 

Maunalua Bay and other adjacent open areas from 2004-2007. Classification based on natural breaks.
26 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 21. Change in coral species richness from 2004-2007 for the Hanauma Bay MLCD, Maunalua Bay and other adjacent open areas.
Results

Monitoring Hawaii’s Marine Protected Areas 27

x

x

Figure 22. Temporal change in benthic communities at Hanauma Bay MLCD, Maunalua Bay and other adjacent open ar- eas by habitat type from 2004-2007. (a) Mean species richness by year (b) Mean percent cover of coral (c) Mean percent 

cover of macroalgae (d) Mean percent cover of turf algae.

Average percent turf cover over the sampling period was greater in Hanauma Bay MLCD ( =33.72) compared to the adjacent open area ( =27.54). Average percent turf cover per transect in the MLCD ranged 

from 26.64% (±4.33) in 2006 to 38.93% (±5.61) in 2004. This compared to the adjacent open area where turf 

cover ranged from 15.56% (±2.66) in 2006 to 42.03% (±4.82) in 2004 (Figure 27). Patterns of percent change 

of turf cover on transects between 2004-2007 show a general decrease in cover in Manualua Bay with small 

decreases in Hanauma Bay. A few transects both inside and outside the MLCD had increases in turf cover 

(Figure 28). Temporal patterns in percent turf cover show overall decreases in all management regimes and 

habitat types from 2004-2006. Turf algae is generally much greater in Manualua Bay compared to Hanauma 

Bay MLCD across all years sampled (Figure 22d). 
28 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 23. Mean percent coral cover by individual transects for southeast Oahu study area including Hanauma Bay 

MLCD, Maunalua Bay and other adjacent open areas from 2004-2007. Classification based on natural breaks.
Results

Monitoring Hawaii’s Marine Protected Areas 29

Figure 24. Change in mean coral cover from 2004-2007 for the Hanauma Bay MLCD, Maunalua Bay and other adjacent 

open areas.
30 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 25. Mean percent macroalgal cover by individual transects for southeast Oahu study area including Hanauma Bay 

MLCD, Maunalua Bay and other adjacent open areas from 2004-2007. Classification based on breaks.
Results

Monitoring Hawaii’s Marine Protected Areas 31

Figure 26. Change in mean macroalgal cover from 2004-2007 for the Hanauma Bay MLCD, Maunalua Bay and other 

adjacent open areas.
32 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 27. Mean percent turf algal cover by individual transects for southeast Oahu study area including Hanauma Bay 

MLCD, Maunalua Bay and other adjacent open areas from 2004-2007. 
Results

Monitoring Hawaii’s Marine Protected Areas 33

Figure 28. Change in mean turf algal cover from 2004-2007 for the Hanauma Bay MLCD, Maunalua Bay and other adjacent open areas.
34 Monitoring Hawaii’s Marine Protected Areas

Results

Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes

Average fish species richness per transect in Hanauma Bay MLCD ranged from 16.91 (± 5.08) in 2008 to 19.45 

(± 5.21) in 2004 (Figure 29). This compared to the adjacent open area where richness ranged from 6.84 (± 

6.06) in 2008 to 9.30 (± 8.24) in 2006. Richness averaged 2.54 (± 4.54) fewer species in the MLCD between 

2004 and 2008 while richness declined by 1.16 (± 5.04) species on average in the open area during this same 

time period (Figure 30). The lowest overall richness was observed in the open MAC ( x = 2.68 ± 0.54) and the 

highest was in the MLCD CHB ( x = 19.10 ± 1.73). These trends remained consistent over the five year study 

period (Figure 31a).

Figure 29. Fish species richness by individual transect for southeast Oahu study area including Hanauma Bay, Maunalua 

Bay and other adjacent open areas from 2004-2008. Classification based on natural breaks.
Results

Monitoring Hawaii’s Marine Protected Areas 35

Figure 30. Change in fish species richness from 2004-2008 for the Hanauma Bay MLCD, Maunalua Bay and other adjacent open areas.
36 Monitoring Hawaii’s Marine Protected Areas

Results

x x

x

x

x x

x x

x

Figure 31. Temporal change in fish assemblages at Hanauma Bay MLCD, Maunalua Bay and other adjacent open areas 

by habitat type from 2004-2008. (a) Species Richness (b) Numerical Abundance (c) Biomass (d) Diversity.

Average number of individuals m2 in the MLCD ranged from a high of 0.69 (± 0.33) in 2004 to a low of 0.50 (± 

0.27) in 2007 (Figure 32). The highest average number of individuals per m2

 in the open area was recorded 

in 2006 ( = 0.38 ± 0.54) and the lowest occurred in 2008 ( =0.18 ± 0.27). Average number of individuals 

declined by 0.17 m-2 (± 0.32) in the MLCD and 0.05 (± 0.27) in the open area between 2004 and 2008 (Figure 

33). The lowest overall numerical abundance was observed in the open MAC ( =0.08 ± 0.01) and the highest 

was in the MLCD CHB ( =0.69 ± 0.11). Trends were fairly consistent over time except for an increase in open 

CHB in 2006 due to the presence of several large schools (50-200+) of blackfin chromis (Chromis vanderbilti; 

Figure 31b). 

Biomass (t ha-1) in the MLCD ranged from 0.82 (± 0.50) in 2007 to 0.95 (± 0.81) in 2004 (Figure 34). In contrast, 

biomass in the open area was lowest in 2004 ( = 0.11 ± 0.14) and highest in 2006 ( = 0.17 ± 0.29). Between 

2004 and 2008, biomass in the MLCD declined by 0.05 (± 0.68) and increased by approximately the same 

amount in the open area ( = 0.05 ± 0.30; Figure 35). Biomass was lowest in the open MAC ( = 0.02 ± 0.01) 

and highest in the MLCD UCH ( = 0.89 ± 0.20). Within the open area, biomass varied little among years while 

both UCH and CHB habitats within the MLCD tended to vary inversely with one another (Figure 31c). 
Results

Monitoring Hawaii’s Marine Protected Areas 37

Figure 32. Fish abundance by individual transect for southeast Oahu study area including Hanauma Bay, Maunalua Bay 

and other adjacent open areas from 2004-2008. Classification based on natural breaks.

Diversity varied little among years and between management strata. The highest diversity within the MLCD 

was observed in 2004 ( x = 2.26 ± 0.34) and lowest in 2006 ( x = 2.22 ± 0.42; Figure 36). In the open area, 

diversity ranged from 1.26 (± 0.82) in 2008 to 1.40 (± 0.69) in 2004. Diversity decreased by 0.03 (± 0.35) in 

the MLCD and 0.13 (± 0.59) in the open area between 2004 and 2008 (Figure 37). These trends remained 

consistent over the five year study period (Figure 31d).
38 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 33. Change in fish abundance from 2004-2008 for the Hanauma Bay MLCD, Maunalua Bay and other adjacent 

open areas.
Results

Monitoring Hawaii’s Marine Protected Areas 39

Figure 34. Fish biomass by individual transect for southeast Oahu study area including Hanauma Bay, Maunalua Bay and 

other adjacent open areas from 2004-2008. Classification based on natural breaks.
40 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 35. Change in fish biomass from 2004-2008 for the Hanauma Bay MLCD, Maunalua Bay and other adjacent open 

areas.
Results

Monitoring Hawaii’s Marine Protected Areas 41

Figure 36. Fish diversity by individual transect for southeast Oahu study area including Hanauma Bay, Maunalua Bay and 

other adjacent open areas from 2004-2008. Classification based on natural breaks.
42 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 37. Change in fish diversity from 2004-2008 for the Hanauma Bay MLCD, Maunalua Bay and other adjacent open 

areas.
Results

Monitoring Hawaii’s Marine Protected Areas 43

Fish Trophic Structure Between Management Regimes and Among Habitats

Overall, apex predators accounted for 0.9% of the numerical abundance in Hanauma Bay MLCD yet they were 

more than 23 times more abundance than in the adjacent open area where they only accounted for 0.08% of 

the individuals in that location. Numbers of apex predators were lower in the MLCD CHB ( x = 0.004 ± 0.006) 

compared with the MLCD UCH ( x = 0.006 ± 0.016) but they were much more variable in this habitat (Figure 

38a). Apex predators accounted for 11.4% of the total biomass in the MLCD but 0.3% in the open area. The difference in biomass of apex predators between the MLCD and open areas was more than 220%. Apex predator 

biomass was highest in the MLCD UCH and values in 2006 were more than fivefold higher than the average of 

the other three years in that habitat due to the presence of several schools of large (55-65 cm) bluefin trevally 

(Caranx melampygus; Figure 39a).

Benthic carnivores comprised 33% of the numerical abundance in the MLCD and 49% in the open area although density was 1.5 times higher in the MLCD. Numbers of benthic carnivores were similar between the 

MLCD CHB ( x = 0.24 ± 0.14) and the open CHB ( x = 0.26 ± 0.18) habitat types. Little variation was observed 

in benthic carnivore numerical abundance among habitat types or management strata over time (Figure 38b). 

Benthic carnivores comprised 19% of the biomass in the MLCD and 52% in the open area. Despite this difference, biomass of benthic carnivores was more than two times higher in the MLCD. The highest biomass 

of benthic carnivores was observed in the MLCD CHB ( x = 0.21 ± 0.25) and the open CHB ( x = 0.18 ± 0.32) 

habitat types. High biomass of benthic carnivores in the MLCD CHB in 2004 was due to schools of yellowfin goatfishes (Mulloidichthys vanicolensis) and bonefish (Albula glossodonta). Large schools of yellowstripe 

goatfishes (M. flavolineatus) accounted for the high biomass in the open CHB in 2007 and 2008 (Figure 39b). 

Planktivores accounted for 9% of the numerical density in the MLCD and 26% in the open area with density 

about 34% higher in the open area. Planktivore density was highest in the open CHB habitat ( x = 0.19 ± 0.36) 

and ranged from a high of 0.39 (± 0.54) to a low of 0.04 (± 0.08) in 2008 (Figure 38c). The high value in 2006 

was associated with large schools of blackfin chromis. Planktivores comprised 10% of the biomass in the open 

area, which was 68% higher than their biomass in the MLCD where they comprised 3% of the total biomass. 

High biomass of planktivores in the MLCD in 2004 resulted from high abundance of the endemic Hawaiian 

sergeant (Abudefduf abdominalis; Figure 39c). 

Herbivores were the most conspicuous trophic group by number in the MLCD accounting for 58% of total 

abundance. In the open area, this trophic group was five times less abundance compared with the MLCD 

and comprised 25% of total numerical density in this management stratum. The highest densities were in the 

MLCD CHB ( x = 0.36 ± 0.30) and UCH ( x = 0.29 ± 0.15) habitats and showed little variation among years 

(Figure 38d). Herbivores comprised 67% of the biomass in the MLCD and had 11 times greater biomass than 

the open area where herbivores accounted for 37% of the biomass. The highest densities of herbivores were 

in the MLCD UCH ( x = 0.63 ± 0.39) and CHB ( x = 0.60 ± 0.66) habitats and showed little variation among 

years (Figure 39d).
44 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 38. Temporal change in fish abundance by trophic level at Hanauma Bay MLCD, Maunalua Bay and other adjacent 

open areas by habitat type from 2004-2008. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.

Figure 39. Temporal change in fish biomass by trophic level at Hanauma Bay MLCD, Maunalua Bay and other adjacent 

open areas by habitat type from 2004-2008. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.
Results

Monitoring Hawaii’s Marine Protected Areas 45

PUPUKEA MLCD AND NORTH SHORE OAHU

The north shore Oahu study area extended from Sunset Beach to Kawailoa Beach (ca. 6.5 km2) and included 

the Pupukea MLCD (Figure 40).

Sample Allocation

A total of 80 samples were collected in 2004 and 2006, in 2007 and 2008 only hardbottom habitat was sampled 

(n=51; Figure 41 and Table 13). The two levels of sampling stratification included major habitat types (CHB, 

MAC, UCH and sand) and fisheries management regime (open access and MLCD). Macroalgae habitat was 

not present at the one-acre minimum mapping unit within the MLCD, and CHB habitat was likewise not present 

in the open area at the one-acre minimum mapping unit. 

Figure 40. Overview of Pupukea MLCD and adjacent areas 

with sampling locations.

Figure 41. Sampling locations and benthic structure for the 

Pupukea MLCD and adjacent areas.

Table 13. Sample allocation at Pupukea MLCD by habitat (2002-2008).

MLCD CHB UCH UCS Open MAC UCH UCS Total

2003 35 9 15 11 38 12 15 11 73

2006 35 9 15 11 38 12 15 11 73

2007 24 9 15 27 12 15 51

2008 24 9 15 27 12 15 51

Total 118 36 60 22 130 48 60 22

* MLCD = Marine Life Conservation District; CHB = Colonized hard bottom; UCH = Uncolonized hard bottom; UCS = Unconsolidated sediment; Open = Area open to resource extraction; MAC = Macroalgae.
46 Monitoring Hawaii’s Marine Protected Areas

Results

Characterization of Spatial Patterns of the Seascape and MLCD Configuration

The total seascape area within the Pupukea MLCD was 0.11 km2

 in 1983 and 

was expanded to 0.71 km2

 in 2003, with 

the perimeter to area ratio decreasing 

from 27.9 to 7.34 and the fractal dimension also increasing from 1.35 to 1.79. 

(Tables 14 and 15, Figure 42). Based on 

the NOAA Biogeography Branch benthic habitat maps the habitat diversity, 

represented by Shannon’s diversity index and Shannon’s evenness index was 

0.91 and 0.65 respectively in 1983 and 

1.45 and 0.81 respectively in 2003 (Table 14). Bathymetric grids were created 

from the LiDAR data within Hanauma 

Bay MLCD that represented an approximate depth range of 0 – 12.07 meters 

( x = 3.59) in 1983 and in increased to 

include a depth range up to 16.98 ( x

= 8.09) in 2003. The average slope ( x

= 8.55) and average rugosity ( x = 1.02) 

both increased slightly between the 

1983 and 2003 boundary respectively 

(Table 16 and Figure 43).

Table 14. Summary of seascape metrics derived from 2007 NOAA Biogeography Branch benthic habitat maps.

1983 Boundary

MLCD*

2003 Boundary

MLCD*

Shannon's diversity index 0.91 1.45

Shannon's evenness index 0.65 0.81

Mean patch fractal dimension 1.35 1.79

* MLCD = Marine Life Conservation District.

Table 15. Summary of marine reserve boundary analysis for Pupukea 

MLCD.

1983 Boundary

MLCD*

2003 Boundary

MLCD*

Reserve perimeter (km) 2.97 5.21

Reserve area (km2

) 0.11 0.71

Perimeter to area ratio 27.9 7.34

* MLCD = Marine Life Conservation District.

Figure 42. Aerial imagery of Pupukea MLCD with original (1983) and extended boundaries (2000).
Results

Monitoring Hawaii’s Marine Protected Areas 47

Table 16. Summary of seascape structure derived from LiDAR bathymetric grids for Pupukea MLCD.

Depth Rugosity Slope (percent)

Average Standard 

deviation

Depth

range

Average Standard 

deviation

Rugosity

range

Average Standard 

deviation

Deviation

range

1983 Boundary MLCD* 3.59 2.62 0-12.07 1.02 0.04 1-1.52 8.55 9.65 0.02-94.04

2003 Boundary MLCD* 8.09 4.23 0-16.98 1.02 0.04 1-1.52 9.28 10.63 0.01 - 94.04

* MLCD = Marine Life Conservation District.

Figure 43. Aerial imagery of Pupukea and surrounding study region with other panels 

describing seascape characteristics (e.g. depth, slope, rugosity) derived from bathymetric 

LiDAR data.
48 Monitoring Hawaii’s Marine Protected Areas

Results

Evaluation of MLCD Boundary Expansion Using a Seascape Perspective

The total area within the Pupukea MLCD 

increased by 646% after the expansion 

of the boundaries (Table 15). Prior to expansion, the MLCD was dominated by 

rock/boulder habitat (74%). After boundary expansion, the geomorphic structure 

consisted of a mix of pavement (37%), 

sand (33%) and rock/boulder (26%). 

Before expansion the biological cover 

was dominated by macroalgae (63%), 

followed by uncolonized hard bottom 

(24%), and turf algae (11%; Figure 44). 

Following expansion, the biological cover consisted of a mixture of uncolonized 

hard bottom (33%), macroalgae (25%), 

turf algae (25%) and coral (12%; Figure 44). Habitat diversity and evenness 

increased by 59% and 25%, respectively, after boundary expansion. The 

mean perimeter-area ratio among habitat polygons increased by 24% and the 

mean patch fractal dimension increased 

by 33%. The total perimeter-area ratio 

decreased by 66%. 

Figure 44: Sampling locations and benthic cover for the Pupukea MLCD 

and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 49

Large-scale Benthic Cover

Benthic coverage for the Pupukea 

MLCD was derived from the NOAA 

benthic habitat maps with coral reef and 

hard bottom major structure accounting 

for 64% of the total habitat within the 

MLCD, followed by unconsolidated sediment (33%). The coral reef and hard 

bottom structure consisted mostly of 

pavement (37%), rock/boulder (26%), 

and with only 1% of scattered coral and 

rock. The remaining portion of the MLCD 

was characterized by sand (33%; Table 

17). Biological cover was dominated 

by macroalgae 10-50% (26%). The remaining portion of the biological cover 

in the MLCD consisted of uncolonized 

(33%) and turf (26%; Table 18).

Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes

Average coral species richness per 

transect in Pupkea MLCD was 1.6 

times greater than the adjacent open 

area with a grand mean of 4.45(± 0.43) 

and 2.79 (± 0.49), respectively. Average species richness in CHB habitat in 

the MLCD ranged from 4.56 (± 0.50) in 

2003 to 5.89 (± 0.35) in 2006 (Table 19). 

Patterns in coral richness from 2003-

2007 increased slightly inside and outside of the MLCD (Figure 45). Patterns 

of change over time among management and habitat regimes were generally consistent over time with coral richness in the MLCD CHB habitat being 

the greatest overall, with all categories increasing in 2006. CHB habitat was not present outside of the MLCD 

(Figure 46). Coral richness outside of the MLCD was nearly equal among habitats in 2003 and 2006, although 

richness was slightly greater in the UCH habitat compared to the MAC habitat in 2007 (Figure 47a).

Table 17. Geomorphic structure for the Pupukea MLCD derived from 

NOAA benthic habitat maps.

Major Structure Detailed 

Structure

Area 

(km2)

Percentage

Coral Reef and Hardbottom Pavement 0.26 37%

Coral Reef and Hardbottom Rock/Boulder 0.18 26%

Coral Reef and Hardbottom Scattered 

Coral/Rock 0.00 <1%

Other Delineations Land 0.03 4%

Unconsolidated Sediment Sand 0.24 33%

Unknown Unknown 0.00 <1%

Table 18. Biological cover for the Pupukea MLCD derived from NOAA 

benthic habitat maps.

Major Cover Percent Cover Area (km2) Percentage

Coral 10% - 50% 0.09 12%

Macroalgae 10% - 50% 0.18 26%

Turf 10% - 50% 0.003 <1%

Turf 50% - 90% 0.18 25%

Unclassified 0.03 4%

Uncolonized 90%-100% 0.24 33%

Unknown Unknown 0.002 <1%

Table 19. Comparison of coral species richness in major habitats and 

among different management regimes at Pupukea MLCD.

2003 2006 2007

Mean SE Mean SE Mean SE

MLCD

CHB 4.56 0.50 5.89 0.35 4.56 0.50

UCH 2.60 0.38 5.47 0.39 3.67 0.46

Open

MAC 1.83 0.32 3.67 0.51 2.42 0.50

UCH 1.87 0.46 3.47 0.53 3.53 0.60

* MLCD = Marine Life Conservation District; CHB = Colonized hard bottom; 

UCH = Uncolonized hard bottom; Open = Area open to resource extraction; 

MAC = Macroalgae.
50 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 45. Coral species richness by individual transects for north shore Oahu study area including Pupukea Bay MLCD 

from 2003-2007.
Results

Monitoring Hawaii’s Marine Protected Areas 51

Figure 46. Change in coral species richness from 2003-2007 for the Pupukea MLCD and adjacent areas.
52 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 47. Temporal change in benthic communities at Pupukea MLCD and surrounding areas by habitat type from 2003-

2007. (a) Mean species richness by year (b) Mean percent cover of coral (c) Mean percent cover of macroalgae (d) Mean 

percent cover of turf algae.

Spatial and Temporal Patterns in Benthic Cover within the Management Regimes

Average percent coral cover over the sampling period was greater in Pupukea MLCD ( x =10.21) compared 

to the adjacent area ( x =7.09). Average percent coral cover per transect in the MLCD ranged from 6.24% (± 

1.30) in 2003 to 12.64% (± 1.79) in 2006. This compared to the adjacent open area where coral cover ranged 

from 3.26% (± 0.71) in 2003 to 9.95% (± 1.58) in 2007 (Figure 48). Patterns of percent change of coral cover on 

transects between 2004-2007 show an overall increase both inside Pupukea MLCD and in the adjacent open 

areas (Figure 49). Patterns of change among management and habitat regimes were generally consistent over 

time with MLCD CHB covering the greatest percentage in all years (Figure 47b). 

Average percent macroalgal cover over the sampling period was greater in the open area ( x =6.32) compared 

to the MLCD ( x =2.94). Average percent macroalgal cover per transect in the MLCD ranged from 1.55% (± 

0.30) in 2003 to 5.97% (± 1.15) in 2007. This compared to the adjacent open area where cover ranged from 

5.66% (± 1.13) in 2006 to 7.11% (± 1.25) in 2007 (Figure 50). Patterns of percent change of macroalgal cover 

on transects between 2004-2007 show a general increase to the north of Pupukea MLCD in hard bottom habitat with small decreases in adjacent open areas (Figure 51). Macroalgae is generally much greater in the open 

areas compared the MLCD, and higher on hard bottom habitats when examined over time among management and habitat regimes. Macroalgal cover is similar for both habitat types in the MLCD and follow consistent 

trends over time (Figure 47c). 
Results

Monitoring Hawaii’s Marine Protected Areas 53

Figure 48. Mean percent coral cover by individual transects for north shore Oahu study area including Pupukea MLCD 

from 2003-2007.

Average percent turf cover over the sampling period was slightly greater in the open area ( x =47.74) compared to the MLCD ( x =43.00). Average percent turf cover per transect in the MLCD ranged from 30.70% (± 

4.01) in 2006 to 59.10% (± 2.03) in 2007. This compared to the adjacent open area where cover ranged from 

37.92% (± 4.38) in 2006 to 54.43% (± 2.62) in 2007 (Figure 52). Spatial patterns of percent change of turf cover 

on transects between 2004-2007 show a general decrease in turf cover inside and outside of the MLCD (Figure 53). Patterns of change over time among management and habitat regimes were generally consistent over 

time with all categories being relatively similar and decreasing slightly over the sampling period (Figure 47d).
54 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 49. Change in mean coral cover from 2003-2007 for the Pupukea MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 55

Figure 50. Mean percent macroalgal cover by individual transects for north shore Oahu study area including Pupukea 

MLCD from 2003-2007. 
56 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 51. Change in mean macroalgal cover from 2003-2007 for the Pupkea MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 57

Figure 52. Mean percent turf algal cover by individual transects for north shore Oahu study area including Pupukea MLCD 

from 2004-2007.
58 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 53. Change in mean turf algal cover from 2003-2007 for the Pupkea MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 59

Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes

Average fish species richness per transect in Pupukea MLCD ranged from 19.96 (± 5.45) in 2007 to 21.33 

(± 5.58) in 2003 (Figure 54). This compared to the adjacent open area where richness ranged from 14.51 (± 

6.22) in 2007 to 15.89 (± 6.28) in 2003. Richness averaged 3.04 (± 7.23) fewer species in the MLCD between 

2003 and 2008 while richness declined by 0.96 (± 6.43) species on average in the open area during this same 

time period (Figure 55). The lowest overall richness was observed in the open MAC ( x = 13.94 ± 1.04) and 

the highest was in the MLCD CHB ( x = 22.56 ± 2.54). Richness remained consistent over the six year study 

period within the open area and declined slightly in the MLCD (Figure 56a).

Figure 54. Fish species richness by individual transect for north shore Oahu study area 

including Pupukea from 2003-2008.
60 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 55. Change in fish species richness from 2003-2008 for the Pupukea MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 61

Figure 56. Temporal change in fish assemblages at Pupukea MLCD and surrounding areas by habitat type from 2003-

2008. (a) Species Richness (b) Numerical Abundance (c) Biomass (d) Diversity.

Average number of individuals m2 in the MLCD ranged from 0.75 (± 0.45) in 2007 to 1.18 (± 0.53) in 2006 (Figure 57). The highest average number of individuals per m2

 in the open area was recorded in 2006 ( x = 0.73 ± 

0.43) and the lowest occurred in 2007 ( x = 0.56 ± 0.39). Average number of individuals increased by 0.28 m-2

(± 0.62) in the MLCD and 0.08 (± 0.52) in the open area between 2003 and 2008 (Figure 58). The lowest overall numerical abundance was observed in the open MAC ( x = 0.57 ± 0.08) and the highest was in the MLCD 

CHB ( x = 1.08 ± 0.22). Trends were fairly consistent over time except for a large decrease across all habitat 

types in 2007 due to a reduced number of small planktivores during that year, primarily blackfin chromis and 

oval chromis (Chromis ovalis; Figure 56b).

Biomass (t ha-1) in the MLCD ranged from 0.75 (± 0.54) in 2003 to 0.86 (± 0.62) in 2008 (Figure 59). In the open 

area, biomass was lowest in 2008 ( x = 0.18 ± 0.18) and highest in 2007 ( x = 0.27 ± 0.39). Between 2003 and 

2008, biomass in the MLCD increased by 0.11 (± 0.57) but decreased in the open area ( = 0.07 ± 0.40) during 

that same time period (Figure 60). Biomass was lowest in the open MAC ( x = 0.18 ± 0.05) and highest in the 

MLCD CHB ( x = 0.81 ± 0.10). Within the open area, biomass declined within UCH and varied little within MAC 

(Figure 56c). In the MLCD, the increase in biomass was most pronounced in CHB. 

The highest diversity within the MLCD was observed in 2003 ( x = 2.41 ± 0.40) and lowest in 2008 ( x = 1.92 ± 

0.48; Figure 61). In the open area, diversity ranged from 1.90 (± 0.44) in 2008 to 2.16 (± 0.41) in 2003. Diversity 

decreased by 0.49 (± 0.45) in the MLCD and 0.26 (± 0.51) in the open area between 2003 and 2008 (Figure 

62). Diversity declined sharply between 2003 and 2006 across all habitat types and management strata (Figure 56d). 
62 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 57. Fish abundance by individual transect for north shore Oahu study area including Pupukea from 2003-2008.
Results

Monitoring Hawaii’s Marine Protected Areas 63

Figure 58. Change in fish abundance from 2003-2008 for the Pupukea MLCD and adjacent areas.
64 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 59. Fish biomass by individual transect for north shore Oahu study area including Pupukea from 2003-2008.
Results

Monitoring Hawaii’s Marine Protected Areas 65

Figure 60. Change in fish biomass from 2003-2008 for the Pupukea MLCD and adjacent areas.
66 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 61. Fish diversity by individual transect for north shore Oahu study area including Pupukea from 2003-2008.
Results

Monitoring Hawaii’s Marine Protected Areas 67

Figure 62. Change in fish diversity from 2003-2008 for the Pupukea MLCD and adjacent areas.
68 Monitoring Hawaii’s Marine Protected Areas

Results

Fish Trophic Structure Between Management Regimes and Among Habitats

Apex predators accounted for only 0.2% of the numerical abundance in Pupukea MLCD but were 3.3 times 

more abundance than in the adjacent open area where they accounted for 0.1% of the individuals in that location. Numbers of apex predators were low overall and varied greatly without trend among the management and 

habitat strata (Figure 63a). Apex predators accounted for 1.8% of the total biomass in the MLCD and 3.1% in 

the open area yet they comprised more than twice as much total biomass in the MLCD compared with the open 

area. Apex predator biomass was similar in the MLCD Open UCH ( x = 0.014 ± 0.050) and CHB ( x = 0.014 ± 

0.028) habitats with slightly lower biomass in the open UCH ( x = 0.012 ± 0.062). High biomass in 2007 in the 

MLCD and UCH habitats was associated with bluefin trevally (Caranx melampygus; Figure 64a).

Benthic carnivores comprised 30% of the numerical abundance in the MLCD and 48% in the open area with 

densities nearly equal. Numbers of benthic carnivores were similar among habitat and management combinations. Abundance declined across all strata from 2006 to 2007 but increased again in 2008 (Figure 63b). 

Benthic carnivores comprised 24% of the biomass in the MLCD and 43% in the open area. Biomass of benthic 

carnivores was nearly twice as high in the MLCD compared to the open area. The highest biomass of benthic 

carnivores was observed in the MLCD UCH ( x = 0.22 ± 0.22) and this trend was consistent over time (Figure 

64b). The other strata fluctuated without trend over the study period.

Planktivores accounted for 47% of the numerical density in the MLCD and 33% in the open area with density 

more than two times higher in the MLCD. Planktivore density was highest in the MLCD CHB habitat ( x = 0.53 

± 0.52) and MLCD UCH ( x = 0.46 ± 0.51) habitats. Abundance across all strata (except open MAC) declined 

from 2006 to 2007 but increased again in 2008 (Figure 63c). Planktivores comprised 5% of the biomass in 

the MLCD and 6% in the open area although biomass was nearly three times higher in the MLCD. Planktivore 

biomass varied without trend among management and habitat strata (Figure 64c). 

Herbivores accounted for 23% of total numerical abundance in the MLCD and 19% in the open area. Herbivore 

biomass was more than two times higher in the MLCD compared to the open area. The highest densities were 

in the MLCD CHB ( x = 0.26 ± 0.12) and UCH ( x = 0.23 ± 0.18) habitats and showed modest declines over 

time (Figure 63d). Herbivores comprised 70% of the biomass in the MLCD and had five times greater biomass 

than the open area where herbivores accounted for 48% of the biomass. The highest densities were in the 

MLCD CHB ( x = 0.63 ± 0.55) and UCH ( x = 0.52 ± 0.52) habitats and showed a modest increase over time 

(Figure 64d). Most of this increase was due to the increase in number and size of the ember parrotfish (Scarus 

rubroviolaceus). 
Results

Monitoring Hawaii’s Marine Protected Areas 69

Figure 63. Temporal change in fish abundance by trophic level at Pupukea MLCD and surrounding areas by habitat type 

from 2004-2008. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.

Figure 64. Temporal change in fish biomass by trophic level at Pupukea MLCD and surrounding areas by habitat type 

from 2004-2008. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.
70 Monitoring Hawaii’s Marine Protected Areas

Results

HONOLUA-MOKULEIA MLCD AND WEST MAUI

The west Maui study area extended from Kapalua Bay north to Honolua Bay (ca. 6.2 km2) and included the 

Honolua-Mokuleia Bays MLCD (Figure 65).

Sample Allocation

A total of 80 samples were collected in 2004 and 2006, in 2007 and 2008 only hardbottom habitat was sampled 

(n=75; Figure 66 and Table 20). The two levels of sampling stratification included major habitat types (CHB, 

MAC, UCH and unconsolidated sediment [UCS]) and fisheries management regime (open access and MLCD). 

Macroalgae habitat was not present at the one-acre minimum mapping unit within the MLCD. 

Figure 65. Overview of Honolua-Mokuleia MLCD and adjacent areas with sampling locations.

Figure 66. Sampling locations and benthic structure for the 

Honolua-Mokuleia MLCD and adjacent areas.

Table 20. Sample allocation at Honolua-Mokuleia Bay MLCD by habitat (2002-2008).

MLCD CHB UCH UCS Open CHB MAC UCH UCS Total

2003 37 15 12 10 63 13 12 23 15 100

2006 37 15 12 10 63 13 12 23 15 100

2007 27 15 12 48 13 12 23 75

2008 27 15 12 48 13 12 23 75

Total 128 60 48 20 222 52 48 92 30

* MLCD = Marine Life Conservation District; CHB = Colonized hard bottom; UCH = Uncolonized hard bottom; UCS = Unconsolidated 

sediment; Open = Area open to resource extraction; MAC = Macroalgae.
Results

Monitoring Hawaii’s Marine Protected Areas 71

Characterization of Spatial Patterns of the Seascape and MLCD Configuration

The total seascape area within Honolua Bay was 0.11 km2

 and 0.08 km2 in 

Mokuleia Bay, with a perimeter to area 

ratio of 12.23 and 16.79, respectively, 

with an overall mean fractal dimension 

of 1.43 (Tables 21). Based on the 2007 

NOAA Biogeography Branch benthic 

habitat maps the habitat diversity in 

Honolua-Mokuleia MLCD, represented 

by Shannon’s diversity index and Shannon’s evenness index was 0.83 and 

0.76 respectively (Table 22). Bathymetric grids were created from the LiDAR data within Mokuleia Bay MLCD 

that represented an approximate depth 

range of 0 – 22.82 meters ( x = 4.27; 

Table 23 and Figure 67). Percent slope 

(ranging from 0 – 226.43, x =6.08) and 

rugosity (ranging from 0 –2.79, x =1.01) 

were also derived from the bathymetric 

grids. Bathymetric grids within Honolua 

Bay MLCD represented an approximate 

depth range of 0 – 14.01 meters ( x

= 5.27; Table 23 and Figure 67). Percent slope (ranging from 0 – 226.43, 

x =6.99) and rugosity (ranging from 0 

–2.79, x =1.01) were also derived from 

the bathymetric grids. 

Large-scale Benthic Cover

Geomorphic structure for HonoluaMokuleia Bay MLCD, derived from the 

NOAA benthic habitat maps, was characterized by sand (36%) with the hardbottom areas dominated by pavement 

(29%) and rock/boulder structure (17%; 

Table 24). Biological cover in the MLCD 

consisted primarily of uncolonized volcanic rock 90-100% (36%), followed by 

turf 50-90% (35%) and coral 50-90% 

(11%; Table 25).

Table 21. Summary of seascape metrics derived from 2007 NOAA Biogeography Branch benthic habitat maps. 

Honolua-Mokuleia MLCD*

Shannon's diversity index 0.83

Shannon's evenness index 0.76

Mean patch fractal dimension 1.43

* MLCD = Marine Life Conservation District.

Table 22. Summary of marine reserve boundary analysis for HonoluaMokuleia MLCD.

Honolua Bay

MLCD*

Mokuleia Bay 

MLCD*

Reserve perimeter (km) 1.30 1.26

Reserve area (km2

) 0.11 0.08

Perimeter to area ratio 12.23 16.79

* MLCD = Marine Life Conservation District.

Table 23. Summary of seascape structure derived from LiDAR bathymetric grids for Honolua-Mokuleia MLCD.

Honolua Bay MLCD* Mokuleia Bay MLCD*

Depth (m)

Average 5.27 4.27

Standard deviation 3.56 2.18

Depth range 0-14.01 0 – 22.82

Rugosity

Average rugosity 1.00 1.01

Standard deviation 0.03 0.07

Rugosity range 0 – 1.36 0 – 2.79

Slope (percent)

Average slope 6.99 6.08

Standard deviation 8.83 10.20

Slope range 0 – 81.01 0 – 226.43

* MLCD = Marine Life Conservation District.

Table 24. Geomorphic structure for the Honolua-Mokuleia MLCD derived 

from NOAA benthic habitat maps.

Major 

Structure

Detailed 

Structure

Area (km2

) Percentage

Coral Reef & Hard bottom Aggregate Reef 0.02 10%

Coral Reef & Hard bottom Pavement 0.05 27%

Coral Reef & Hard bottom Pavement with 

Sand Channels 0.01 5%

Coral Reef & Hard bottom Rock/Boulder 0.03 17%

Other Delineations Land 0.01 5%

Unconsolidated Sediment Sand 0.06 36%
72 Monitoring Hawaii’s Marine Protected Areas

Results

Table 25. Biological cover for the Honolua-Mokuleia MLCD derived from 

NOAA benthic habitat maps.

Major Cover Percent Cover Area (km2) Percentage

Coral 10% - 50% 0.01 8%

Coral 50% - 90% 0.02 11%

Turf 10% - 50% 0.01 5%

Turf 50% - 90% 0.06 35%

Unclassified 0.01 5%

Uncolonized 90%-100% 0.06 36%

Figure 67. Aerial imagery of Honolua-Mokuleia Bay MLCD and surrounding study region with other panels describing 

seascape characteristics (e.g. depth, slope, rugosity) derived from bathymetric LiDAR data.
Results

Monitoring Hawaii’s Marine Protected Areas 73

Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes

Average coral species richness per 

transect for CHB Honolua-Mokuleia 

MLCD was greater than the adjacent 

open area with a grand mean of 4.51 

and 2.29, respectively (Table 26). Average species richness in CHB habitat 

in the MLCD ranged from 5.53 (± 0.29) 

in 2002 to 6.13 (± 0.49) in 2007. This 

compared to the adjacent open area 

where richness ranged from 3.38 (± 

0.55) in 2006 to 4.77 (± 0.39) in 2007 

(Figure 68). Overall, patterns of change 

in coral richness from 2002-2007 were 

variable, with relatively small increases 

to the south of the MLCD, and comparatively larger increases inside the MLCD 

(Figure 69). Average coral richness over time was generally stable among management regimes and habitat 

types. Coral cover in the MLCD CHB was consistently greater than the other strata. Coral richness increased 

slightly from 2006-2007 across the study area (Figure 70a).

Table 26. Comparison of coral species richness in major habitats and 

among different management regimes at Honolua-Mokuleia Bay, Maui.

2002 2006 2007

Mean SE Mean SE Mean SE

MLCD

CHB 5.53 0.29 5.67 0.36 6.13 0.49

UCH 2.75 0.35 3.42 0.45 3.58 0.58

Open

CHB 3.92 0.31 3.38 0.55 4.77 0.39

MAC 0.42 0.23 0.50 0.26 0.75 0.41

UCH 2.13 0.35 1.96 0.34 2.74 0.36

* MLCD = Marine Life Conservation District; CHB = Colonized hard bottom;

UCH = Uncolonized hard bottom; Open = Area open to resource extraction; 

MAC = Macroalgae.
74 Monitoring Hawaii’s Marine Protected Areas

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Figure 68. Coral species richness by individual transects for west Maui study area including Honolua-Mokuleia MLCD 

from 2002-2007.
Results

Monitoring Hawaii’s Marine Protected Areas 75

Figure 69. Change in coral species richness from 2002-2007 for the Honolua-Mokuleia and adjacent areas.
76 Monitoring Hawaii’s Marine Protected Areas

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x x

x

x

Figure 70. Temporal change in benthic communities at Honolua-Mokuleia MLCD and surrounding areas by habitat type 

from 2002-2007. (a) Mean species richness by year (b) Mean percent cover of coral (c) Mean percent cover of macroal- gae (d) Mean percent cover of turf algae.

Spatial and Temporal Patterns in Benthic Cover within the Management Regimes 

Average percent coral cover over the sampling period was about two times greater in Honolua-Mokuleia 

MLCD ( =15.98) compared to the adjacent area ( =8.09). Average percent coral cover per transect in the 

MLCD ranged from 11.70% (± 2.15) in 2002 to 22.37% (± 3.07) in 2007. This compared to the adjacent open 

area where coral cover ranged from 5.21% (± 0.98) in 2006 to 12.57% (± 2.26) in 2007 (Figure 71). Patterns of 

change in coral richness between 2002 and 2007 generally increased inside the MLCD, with a few transects 

decreasing over time. The areas adjacent to the MLCD followed similar trends with areas of increase, as well 

as decrease over the study period (Figure 72). Temporal patterns in percent coral cover across all management regimes and habitat types generally followed similar trends with the exception of open CHB which decreased slightly in 2006 and then increased again 2007 (Figure 70b). 

Average percent macroalgal cover over the sampling period was about two times greater in the open area (

=14.14) compared to the Honolua-Mokuleia MLCD ( =7.37). Average percent macroalgal cover per transect 

in the MLCD ranged from 6.05% (± 1.36) in 2006 to 8.93% (± 1.82) in 2002. This compared to the adjacent 

open area where macroalgae cover ranged from 12.58% (± 1.62) in 2002 to 17.60% (± 2.25) in 2007 (Figure 

73). Patterns of change in macroalgal cover between 2002 and 2007 generally decreased throughout the 

study area with the exception of transects furthest to the south which showed relatively large increases over 

the sampling period. Marcoalgal cover decreased in most hard bottom habitats in the open access areas, and 

in several adjacent open areas in deeper sea grass bed habitats (Figure 74). Temporal patterns in percent 

macroalgae cover show increases in the open MAC habitat in all years sampled with the largest increase in 

2007. Patterns across all habitats and sampling regimes indicate macroalgae is generally much greater in the 

open areas compared the MLCD, and higher on hard bottom habitats (Figure 70c). 
Results

Monitoring Hawaii’s Marine Protected Areas 77

Figure 71. Mean percent coral cover by individual transects for west Maui study area including Honolua-Mokuleia MLCD 

from 2002-2007.
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Figure 72. Change in percent coral cover from 2002-2007 for the Honolua-Mokuleia and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 79

Figure 73. Mean percent macroalgal cover by individual transects for west Maui study area including Honolua-Mokuleia 

MLCD from 2002-2007.
80 Monitoring Hawaii’s Marine Protected Areas

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Figure 74. Change in mean percent macroalgal cover from 2002-2007 for the Honolua-Mokuleia and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 81

Average percent turf cover over the sampling period was about the same in the open area ( =37.19) compared to the Honolua-Mokuleia MLCD ( =39.99). Average percent turf cover per transect in the MLCD ranged 

from 32.80% (± 3.76) in 2006 to 48.74% (± 2.86) in 2007. This compared to the adjacent open area where turf 

cover ranged from 32.01% (± 3.12) in 2006 to 42.67% (± 2.52) in 2007 (Figure 75). Patterns of percent change 

of turf algae cover on transects between 2004-2007 show an overall decrease in turf algae cover throughout 

the study site with particular areas both inside and outside of the MLCD demonstrating increases in turf cover 

(Figure 76). Temporal patterns in percent turf cover were consistent among all management regimes and habitat types with slight decreases from 2004-2006, and a slight increases from 2006-2007. Open MAC habitat 

consistently had the lowest percentage of turf cover, while the other categories were similar (Figure 70d). 

Figure 75. Mean percent turf algal cover by individual transects for west Maui study area 

including Honolua-Mokuleia MLCD from 2002-2007.

x

x
82 Monitoring Hawaii’s Marine Protected Areas

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Figure 76. Change in mean percent turf algal cover from 2002-2007 for the Honolua-Mokuleia and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 83

Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes

Average species richness per transect in Honolua-Mokuleia MLCD ranged from 19.18 (± 3.89) in 2007 to 

22.78 (± 6.18) in 2002 (Figure 77). This compared to the adjacent open area where richness ranged from 

11.94 (± 6.66) in 2007 to 14.19 (± 8.36) in 2003. Richness declined by 3.59 (± 7.52) species in the MLCD between 2002 and 2007 while richness declined by 2.25 (± 8.08) species on average in the open area during this 

same time period (Figure 78). The lowest overall richness was observed in the open MAC ( = 6.44 ± 0.91) 

and the highest was in the MLCD CHB ( = 21.78 ± 2.78). The decline in richness was most pronounced in 

the MLCD CHB (Figure 79a).

Figure 77. Fish species richness by individual transect for west Maui study area including Honolua-Mokuleia from 2002-

2007.

x

x
84 Monitoring Hawaii’s Marine Protected Areas

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Figure 78. Change in fish species richness from 2002-2007 for the Honolua-Mokuleia MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 85

Figure 79. Temporal change in fish assemblages at Honolua-Mokuleia MLCD and surrounding areas by habitat type from 

2002-2007. (a) Species Richness (b) Numerical Abundance (c) Biomass (d) Diversity.

Average number of individuals per m2 in the MLCD ranged from 1.07 (± 0.64) in 2006 to 1.20 (± 0.65) in 2002 

(Figure 80). The highest average number of individuals per m2 in the open area was recorded in 2002 ( x = 

0.61 ± 0.56) and the lowest occurred in 2007 ( x = 0.48 ± 0.45). Average number of individuals decreased by 

0.05 m-2 (± 0.86) in the MLCD and 0.12 (± 0.48) in the open area between 2002 and 2007 (Figure 81). The 

lowest overall numerical abundance was observed in the open MAC ( x = 0.16 ± 0.03) and the highest was in 

the MLCD CHB ( x = 1.33 ± 0.17). Trends were fairly consistent over time and habitat types except for a slight 

decline in 2007 in open UCH and CHB (Figure 79b).

Biomass (t ha-1) in the MLCD ranged from 0.88 (± 0.79) in 2006 to 1.06 (± 0.80) in 2007 (Figure 82). In the open 

area, biomass was lowest in 2002 ( x = 0.21 ± 0.22) and highest in 2006 ( x = 0.26 ± 0.36). Between 2002 and 

2007, biomass in the MLCD increased by 0.05 (± 1.16) and ( x = 0.03 ± 0.37) in the open area during that same 

time period (Figure 83). Biomass was lowest in the open MAC ( x = 0.11 ± 0.04) and highest in the MLCD CHB 

( x = 1.02 ± 0.15). Within the open area, biomass varied little (Figure 79c). In the MLCD, biomass increased 

sharply in the UCH between 2006 and 2007. 

The highest diversity within the MLCD was observed in 2002 ( x = 2.32 ± 0.35) and lowest in 2006 ( x = 2.02 

± 0.49; Figure 84). In the open area, diversity ranged from 1.77 (± 0.50) in 2006 to 1.92 (± 0.55) in 2002. Diversity decreased by 0.25 (± 0.45) in the MLCD and 0.07 (± 0.58) in the open area between 2002 and 2007 

(Figure 85). Diversity was lowest in the open MAC ( x = 1.43 ± 0.08) and highest in the MLCD UCH ( x = 2.14 

± 0.18). Diversity declined slightly between 2003 and 2006 across all habitat types and management strata but 

increased slightly in 2007 (Figure 79d). 
86 Monitoring Hawaii’s Marine Protected Areas

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Figure 80. Fish abundance by individual transect for west Maui study area including Honolua-Mokuleia from 2002-2007.
Results

Monitoring Hawaii’s Marine Protected Areas 87

Figure 81. Change in fish abundance from 2002-2007 for the Honolua-Mokuleia MLCD and adjacent areas.
88 Monitoring Hawaii’s Marine Protected Areas

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Figure 82. Fish biomass by individual transect for west Maui study area including Honolua-Mokuleia from 2002-2007.
Results

Monitoring Hawaii’s Marine Protected Areas 89

Figure 83. Change in fish biomass from 2002-2007 for the Honolua-Mokuleia MLCD and adjacent areas.
90 Monitoring Hawaii’s Marine Protected Areas

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Figure 84. Fish diversity by individual transect for west Maui study area including Honolua-Mokuleia from 2002-2007.
Results

Monitoring Hawaii’s Marine Protected Areas 91

Figure 85. Change in fish diversity from 2002-2007 for the Honolua-Mokuleia MLCD and adjacent areas.
92 Monitoring Hawaii’s Marine Protected Areas

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Fish Trophic Structure Between Management Regimes and Among Habitats

Apex predators accounted for only 0.6% of the numerical abundance in Honolua MLCD but were nearly 19 

times more abundance than in the adjacent open area where they accounted for 0.07% of the individuals in 

that location. Numbers of apex predators varied little during the study period among all management and 

habitat strata (Figure 86a). Apex predators accounted for 4.7% of the total biomass in the MLCD and 1.7% in 

the open area and comprised nearly 12 times more biomass in the MLCD compared with the open area. Apex 

predator biomass was similar in the MLCD CHB ( x = 0.047 ± 0.072) and UCH ( x = 0.046 ± 0.011) habitats and 

remained constant over time except for an increase in the MLCD UCH in 2007 due to bluefin trevally (Figure 

87a).

Benthic carnivores comprised 29% of the numerical abundance in the MLCD and 42% in the open area with 

densities about 40% higher in the MLCD. Numbers of benthic carnivores varied little among years and management strata (Figure 86b). Benthic carnivores comprised 24% of the biomass in the MLCD and 43% in the 

open area. Biomass of benthic carnivores was more than two times higher in the MLCD compared to the open 

area. The highest biomass of benthic carnivores was observed in the MLCD UCH ( x = 0.27 ± 0.29), followed 

by the MLCD CHB ( x = 0.22 ± 0.29). Trend was consistent over time except for an increase in benthic carnivores in the MLCD CHB in 2007 due to a mix of species including bigeye emperor (Monotaxis grandoculis) and 

yellowfin goatfish (Figure 87b). 

Planktivores accounted for 37% of the numerical density in both the MLCD and open area although the overall density was more than two times greater in the MLCD. Planktivore density was highest in the MLCD CHB 

habitat ( x = 0.50 ± 0.57) and open CHB ( x = 0.42 ± 0.56) habitats. Abundance across all strata varied little 

among years (Figure 86c). Planktivores comprised 3% of the biomass in the MLCD but 13% in the open area 

although absolute biomass was nearly equal. Planktivore biomass varied little among years except for an 

increase in 2006 in the open CHB due to a school of 85 sleek unicornfish (Naso hexacanthus) that ranged in 

size from 20-30 cm (Figure 87c). 

Herbivores accounted for 33% of total numerical abundance in the MLCD and 21% in the open area. Herbivore 

density was more than three times higher in the MLCD and the highest densities were in the MLCD CHB ( x = 

0.47 ± 0.33) and UCH ( x = 0.27 ± 0.18) habitats. Modest declines in numbers were observed between 2002 

and 2007 (Figure 86d). Herbivores comprised 68% of the biomass in the MLCD and had 6.5 times greater 

biomass than the open area where herbivores accounted for 43% of the biomass. The highest densities were 

in the MLCD CHB ( x = 0.71 ± 0.76) and UCH ( x = 0.61 ± 0.79). Although the MLCD CHB showed a slight 

downward trend, the MLCD UCH trended upward slight (Figure 87d).
Results

Monitoring Hawaii’s Marine Protected Areas 93

Figure 86. Temporal change in fish abundance by trophic level at Honolua-Mokuleia MLCD and surrounding areas by 

habitat type from 2002-2007. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.

Figure 87. Temporal change in fish biomass by trophic level at Honolua-Mokuleia MLCD and surrounding areas by habitat type from 2002-2007. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.
94 Monitoring Hawaii’s Marine Protected Areas

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KEALAKEKUA BAY MLCD AND SOUTH KONA

The south Kona study area extended 

from Nenue Point to Honaunau Bay 

(11.8 km2

) and included the Kealakekua 

Bay MLCD (Figure 88). 

Sample allocation

A total of 80 samples were collected in 

2004 and 2006, in 2007 and 2008 only 

hardbottom habitat was sampled (n=64; 

Table 27). The two levels of sampling 

stratification included major habitat 

types (CHB, UCH and UCS) and fisheries management regimes (open access; 

fisheries replenishment area, or FRA; 

and MLCD). West Hawaii has regional 

fisheries management area (FMA) on 

the Kona Coast that is made up of nine 

FRAs that were developed to prohibit 

specific activities (e.g., aquarium fish 

collection, gill net).

Figure 88. Overview of Kealakekua Bay MLCD and adjacent areas with 

sampling locations.

Table 27. Sample allocation at Kealakekua Bay MLCD by habitat (2002-

2008).

2004 2006 2007 2008 Total

FRA 29 29 29 29 116

CHB 29 29 29 29 116

MLCD 34 34 22 22 112

CHB 22 22 22 22 86

UCS 12 12 24

Open 13 13 13 13 52

CHB 13 13 13 13 52

Total 76 76 64 64

* FRA = Fisheries Replenishment Area; CHB = Colonized hard bottom; 

MLCD = Marine Life Conservation District; UCH = Uncolonized hard bottom; 

UCS = Unconsolidated sediment; Open = Area open to resource extraction.
Results

Monitoring Hawaii’s Marine Protected Areas 95

Characterization of Spatial Patterns of the Seascape within the MLCD 

The total seascape area within Kealakekua Bay was 1.24 km2

, with a perimeter to area ratio of 4.30 and fractal 

dimension of 1.50 (Tables 28 and 29). 

Based on the 2007 NOAA Biogeography 

Branch benthic habitat maps the habitat 

diversity in Kealakekua Bay MLCD, represented by Shannon’s diversity index 

and Shannon’s evenness index was 

1.05 and 0.76 respectively (Table 28). 

Bathymetric grids were created from 

the LiDAR data within Kealakekua Bay 

MLCD that represented an approximate 

depth range of 0 – -37.70 meters ( x = 

15.71; Table 30 and Figure 89). Percent 

slope (ranging from 0 – 1.79, x =9.94) 

and rugosity (ranging from 0 – 146.58, 

x =1.01) were also derived from the 

bathymetric grids.

Table 28. Summary of seascape metrics derived from NOAA Biogeography Branch benthic habitat maps for Kealakekua Bay and adjacent areas.

Kealakekua Bay MLCD*

Shannon's diversity index 1.05

Shannon's evenness index 0.76

Mean patch fractal dimension 1.50

* MLCD = Marine Life Conservation District.

Table 29. Summary of marine reserve boundary analysis for Kealakekua 

Bay MLCD.

Kealakekua Bay MLCD*

Reserve perimeter (km) 5.33

Reserve area (km2

) 1.24 

Perimeter to area ratio 4.30

* MLCD = Marine Life Conservation District.

Table 30. Summary of seascape structure derived from LiDAR bathymet- ric grids for Kealakekua Bay MLCD.

Kealakekua Bay MLCD*

Depth (m)

Average 15.71

Standard deviation 9.98

Depth range 0 - 37.79

Rugosity

Average rugosity 1.01

Standard deviation 0.04

Rugosity range 1 - 1.79

Slope (percent)

Average slope 9.94

Standard deviation 11.61

Slope range 0 - 146.59

* MLCD = Marine Life Conservation District.
96 Monitoring Hawaii’s Marine Protected Areas

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Figure 89. Aerial imagery of Kealakekua Bay and surrounding study region with other panels describing seascape characteristics (e.g. depth, slope, rugosity) derived from bathymetric LiDAR data.
Results

Monitoring Hawaii’s Marine Protected Areas 97

Large-scale Benthic Cover

Geomorphic structure for Kealakekua 

Bay MLCD derived from the NOAA benthic habitat maps consisted primarily of 

sand (29%) followed by rock/boulder 

(20%) and aggregate Reef (18%; Table 

31, Figures 90 and 91). Biological cover 

in the MLCD was characterized dominated by coral cover with a total of 34% 

and uncolonized bottom 90%-100% 

(29%), with only 4% biological cover of 

turf 50%-90% (Table 32, Figures 92 and 

93). Table 32. Biological cover for the Kealakekua Bay MLCD derived from 

NOAA benthic habitat maps.

Table 31. Geomorphic structure for the Kealakekua Bay MLCD derived 

from NOAA benthic habitat maps.

Major Structure Detailed 

Structure

Area 

(km2)

Percentage

Coral Reef and Hardbottom Aggregate 

Reef 0.22 18%

Coral Reef and Hardbottom Rock/Boulder 0.24 20%

Other Delineations Land 0.01 1%

Unconsolidated Sediment Sand 0.36 29%

Unknown Unknown 0.40 32%

Major Cover Percent 

Cover

Area 

(km2

)

Percentage

Coral 10% - 50% 0.14 12%

Coral 50% - 90% 0.10 8%

Coral 90% - 100% 0.18 14%

Turf 50% - 90% 0.04 4%

Unclassified 0.01 1%

Uncolonized 90%-100% 0.36 29%

Unknown Unknown 0.40 32%
98 Monitoring Hawaii’s Marine Protected Areas

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Figure 90. Sampling locations and benthic structure for the Kealakekua Bay MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 99

Figure 91. Sampling locations and benthic structure for Honaunau Bay and adjacent areas.
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Figure 92. Sampling locations and benthic cover for the Kealakekua Bay MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 101

Figure 93. Sampling locations and benthic cover for Honaunau Bay and adjacent areas.
102 Monitoring Hawaii’s Marine Protected Areas

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Spatial and Temporal Patterns in Coral Species Richness within the Management Regimes

Average coral species richness was 

similar Kealakekua Bay MLCD ( x

=4.80), the FRA ( x =4.72), and the adjacent open area with ( x =4.99; Table 

33). Average species richness in CHB 

habitat in the MLCD ranged from 3.86 

(± 0.24) in 2004 to 5.36 (± 0.37) in 2006. 

Richness in CHB habitat in the FRA 

ranged from 4.00 (± 0.23) in 2004 to 

5.41 (± 0.27) in 2006. This compared 

to the adjacent open area where richness ranged from 3.92 (± 0.38) in 2004 

to 6.15 (± 0.55) in 2007 (Figure 94). 

Overall, spatial patterns of change in 

richness from 2004 to 2007 increased throughout the study area, with only a few transects declining slightly 

(Figure 95). When examined among management regimes and habitat types, coral richness was similar and 

trend the same in the MLCD and FRA. Coral richness measured in the open CHB area was similar, but increased in 2007 (Figure 96a).

Table 33. Comparison of coral species richness in major habitats and 

among different management regimes at Kealakekua Bay, Hawaii.

2004 2006 2007

Mean SE Mean SE Mean SE

FRA 

CHB 4.00 0.23 5.41 0.27 5.00 0.28

MLCD

CHB 3.86 0.24 5.36 0.37 4.95 0.24

Open

CHB 3.92 0.38 4.92 0.33 6.15 0.55

* FRA=Fisheries Replenishment Area; CHB = Colonized hard bottom; MLCD = 

Marine Life Conservation District; Open = Area open to resource extraction.

Figure 94. Coral species richness by individual transects for south Kona study area including Kealakekua Bay MLCD 

from 2004-2007.
Results

Monitoring Hawaii’s Marine Protected Areas 103

Figure 95. Change in coral species richness from 2004-2007 for the Kealakekua Bay MLCD and adjacent areas.
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Figure 96. Temporal change in benthic communities at Kealakekua Bay MLCD and surrounding areas by habitat type 

from 2004-2007. (a) Mean species richness by year (b) Mean percent cover of coral (c) Mean percent cover of macroal- gae (d) Mean percent cover of turf algae.
Results

Monitoring Hawaii’s Marine Protected Areas 105

Spatial and Temporal Patterns in Benthic Cover within the Management Regimes 

Average percent coral cover was similar for Kealakekua Bay MLCD ( x =32.99), the FRA ( x =35.12), and the 

adjacent open area with ( x =31.57). Average percent coral cover in the MLCD ranged from 29.01% (± 4.45) 

in 2004 to 44.58% (± 3.55) in 2007. Coral cover in the FRA ranged from 30.81% (± 2.78) in 2006 to 37.48% 

(± 3.55) in 2004. This compared to the adjacent open area where coral cover ranged from 30.28% (± 3.20) in 

2004 to 32.92% (± 3.66) in 2007 (Figure 97). Spatial patterns of percent change of coral cover on transects 

demonstrated an overall greater coral cover inside the protected embayments of Kealakekua and Honaunau. 

Overall, between 2004 and 2007 coral cover decreased in parts of Kealakekua Bay with northwest exposures 

and increased in the more protected shallow regions in the southeast of the Bay (Figure 98). Temporal patterns 

in percent coral cover across all management regimes and habitat types generally remained consistent from 

2004-2006, with a small decrease from 2004-2006 in the FRA (Figure 96b). 

Figure 97. Mean percent coral cover by individual transects for south Kona study area including Kealakekua Bay MLCD 

from 2004-2007.
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Figure 98. Change in mean percent coral cover from 2004-2007 for the Kealakekua Bay MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 107

Average percent macroalgal cover was highest in the open area ( x =5.54), and lowest in Kealakekua Bay 

MLCD ( x =1.11) and moderate in the FRA ( x =3.46). Average percent macroalgal cover in the MLCD ranged 

from 0.45% (± 0.20) in 2004 to 1.67% (± 0.59) in 2007. Macroalgal cover in the FRA ranged from 2.23% (± 

0.60) in 2004 to 4.17% (± 1.09) in 2007. This compared to the adjacent open area where macroalgal cover 

ranged from 2.28% (± 0.58) in 2004 to 7.38% (± 1.49) in 2007 (Figure 99). Patterns of percent change of macroalgae cover on transects between 2004-2007 show a general increase in macroalgae cover in the open sites 

north of Kealakekua Bay MLCD and in Honaunau (Figure 100). Percent macroalgae cover increased over time 

for all management regimes with a slight decrease in the MLCD in 2007 (Figure 96c). 

Figure 99. Mean percent macroalgal cover by individual transects for south Kona study area including Kealakekua Bay 

MLCD from 2004-2007.
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Figure 100. Change in mean percent macroalgal cover from 2004-2007 for the Kealakekua Bay MLCD and adjacent 

areas.
Results

Monitoring Hawaii’s Marine Protected Areas 109

Average percent turf cover was highest in the open area ( x =41.25), and lowest in Kealakekua Bay MLCD (

x =20.28) and moderate in the FRA ( x =39.09). Average percent turf cover in the MLCD ranged from 11.88% 

(± 2.20) in 2006 to 26.36% (± 2.59) in 2007. Turf cover in the FRA ranged from 32.14% (± 2.56) in 2007 to 

46.43% (± 3.73) in 2004. This compared to the adjacent open area where turf cover ranged from 33.17% (± 

3.17) in 2006 to 55.45% (± 3.04) in 2004 (Figure 101). Patterns of percent change of turf cover on transects 

between 2004-2007 show a general decrease throughout the study area, with a few transects increasing inside the MLCD (Figure 102). Temporal patterns in percent turf cover decreases overall among all management 

regimes and habitat types, with the exception of the MLCD where there was a slight increase from 2006 to 

2007 (Figure 96d). 

Figure 101. Mean percent turf algal cover by individual transects for south Kona study area including Kealakekua Bay 

MLCD from 2004-2007.
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Figure 102. Change in mean percent turf algal cover from 2004-2007 for the Kealakekua Bay MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 111

Fish Assemblage Characteristics Among Habitat Types and Between Management Regimes

Average species richness per transect in Kealakekua MLCD ranged from 17.95 (± 5.07) in 2008 to 19.73 (± 

5.08) in 2004 (Figure 103). In the adjacent open area richness ranged from 20.54 (± 2.93) in 2004 to 24.08 (± 

3.07) in 2007 while in the FRA, richness ranged from 18.59 (± 4.60) in 2008 to 21.28 (± 4.52) in 2006. Richness declined by 1.77 (± 4.61) species in the MLCD between 2004 and 2008 while it declined by 1.14 (± 5.24) 

species on average in the FRA during this same time period (Figure 104). Richness increased in the open area 

by 0.62 (± 4.39) between 2004 and 2008. The lowest overall richness was observed in the MLCD CHB ( x = 

19.09 ± 0.80) and the highest was in the open CHB ( x = 22.25 ± 1.68). There was an increase in richness in 

the open CHB between 2004 and 2007 with all three management strata showing declines between 2007 and 

2008 (Figure 105a).

Average number of individuals m-2 in the MLCD ranged from 0.68 (± 0.26) in 2008 to 0.92 (± 0.65) in 2007 

(Figure 106). The highest average number of individuals per m-2 in the open area was recorded in 2006 ( x = 

1.42 ± 0.52) and the lowest occurred in 2004 ( x = 1.24 ± 0.50). Within the FRA, numerical abundance ranged 

from 0.88 (± 0.38) in 2004 to 1.10 (± 0.59) in 2007. Between 2002 and 2008, average number of individuals 

decreased by 0.14 m-2 (± 0.36) in the MLCD, but increase by 0.07 (± 0.43) in the FRA and 0.03 (± 0.33) in the 

open area (Figure 107). The lowest overall numerical abundance was observed in the MLCD CHB ( x = 0.82 ± 

0.10) and the highest was in the open CHB ( x = 1.33 ± 0.09). Numerical abundance was fairly level between 

2002 and 2007 for all management strata but all three showed slight declines between 2007 and 2008. (Figure 

105b).

Biomass (t ha-1) in the MLCD ranged from 0.82 (± 0.48) in 2004 to 0.97 (± 0.86) in 2007 (Figure 108). In the 

open area, biomass was lowest in 2002 ( x = 0.54 ± 0.20) and highest in 2007 ( x = 0.84 ± 0.25). Similar patterns and numbers were observed in the FRA with low biomass in 2004 ( x = 0.54 ± 0.32) and high biomass 

in 2007 ( x = 0.84 ± 0.58). Between 2002 and 2008, biomass increased across all management strata. The 

MLCD saw an increase of 0.06 (± 0.67), while the FRA and open areas was increases of 0.15 (± 0.34) and 0.13 

(± 0.27), respectively (Figure 109). Overall, biomass was lowest in the open CHB ( x = 0.70 ± 0.13) and highest 

in the MLCD CHB ( x = 0.87 ± 0.07). All management strata showed increases in biomass between 2002 and 

2007, with slight declines between 2007 and 2008 (Figure 105c). 

Diversity within the MLCD varied little among years with the highest diversity observed in 2006 ( x = 2.17 ± 

0.26) and lowest in 2008 ( x = 2.13 ± 0.35; Figure 110). Greater fluctuations were observed in the open area 

with a high in 2007 of 2.27 (± 0.32) and a low in 2004 of 2.01 (± 0.37). The FRA showed intermediate variability 

with a high of 2.25 (± 0.32) in 2006 and a low of 2.12 (± 0.33) in 2008. Diversity decreased in the FRA by 0.11 

(± 0.33), by 0.32 (± 0.28) in the open area and 0.28 (± 0.35) in the MLCD between 2002 and 2007 (Figure 

111). Lowest overall diversity was found in the open CHB ( x = 2.08 ± 0.13) while the highest was observed in 

the FRA CHB ( x = 2.20 ± 0.06). Diversity was stable among management strata between 2004 and 2007 with 

open CHB showing a slight decline in 2008 (Figure 105d). 
112 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 103. Fish species richness by individual transect for south Kona study area including Kealakekua Bay from 2004-

2008.
Results

Monitoring Hawaii’s Marine Protected Areas 113

Figure 104. Change in fish species richness from 2004-2008 for the Kealakekua Bay MLCD and adjacent areas.
114 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 105. Temporal change in fish assemblages at Kealakekua Bay MLCD and surrounding areas by habitat type from 

2004-2008. (a) Species Richness (b) Numerical Abundance (c) Biomass (d) Diversity.
Results

Monitoring Hawaii’s Marine Protected Areas 115

Figure 106. Fish abundance by individual transect for south Kona study area including Kealakekua Bay from 2004-2008.
116 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 107. Change in fish abundance from 2004-2008 for the Kealakekua Bay MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 117

Figure 108. Fish biomass by individual transect for south Kona study area including Kealakekua Bay from 2004-2008.
118 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 109. Change in fish biomass from 2004-2008 for the Kealakekua Bay MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 119

Figure 110. Fish diversity by individual transect for south Kona study area including Kealakekua Bay from 2004-2008.
120 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 111. Change in fish diversity from 2004-2008 for the Kealakekua Bay MLCD and adjacent areas.
Results

Monitoring Hawaii’s Marine Protected Areas 121

Fish Trophic Structure Between Management Regimes and Among Habitats

Apex predators accounted for 0.8% of the numerical abundance in Kealakekua MLCD, 0.4% in the FRA, and 

0.3% in the open area. Apex predator abundance was 68% greater in the MLCD compared to the FRA and 

70% greater compared to the open area. Numbers of apex predators declined in all three management strata 

between 2004 and 2008 (Figure 112a). Apex predators accounted for 8.6% of the total biomass in the MLCD, 

2.8% in the FRA, and 3.1% in the open area. Apex predator biomass was 3.8 times greater the MLCD compared to the FRA and 3.5 times greater compared to the open area. Apex predator biomass was similar over 

time in the FRA and open area but high biomass in 2007 in the MLCD was associated with one whitetip reef 

shark (Triaenodon obesus; Figure 113a).

Benthic carnivores comprised 39% of the numerical abundance in Kealakekua MLCD, 37% in the FRA, and 

31% in the open area. Numbers of benthic carnivores were similar between the MLCD, FRA, and open areas. 

All management areas showed a slight increase in benthic carnivore abundance in 2007 followed by a drop 

in 2008 (Figure 112b). Benthic carnivores comprised 38% of the biomass in the MLCD, 39% in the FRA, and 

37% in the open area. Biomass of benthic carnivores was 19% higher in the MLCD compared to the FRA and 

25% than the open area. There was a slight upper trend in biomass between 2004 and 2008 with the greatest 

increase occurring in the MLCD (Figure 113b). 

Planktivores accounted for 27% of the numerical density in the MLCD, 32% in the FRA, and 50% in the open 

area. Planktivore abundance was 28% lower in the MLCD compared to the FRA and 66% lower compared 

to the open area. Trends were fairly consistent over time (Figure 112c). Planktivores comprised 3% of the 

biomass in the MLCD, 7% in the FRA and 10% in the open area. Planktivore biomass was 52% lower in the 

MLCD compared to the FRA and 66% lower than the open area. There were slight upward trends over time 

in the MLCD and FRA while the open area showed a large spike in 2007 associated with several schools of 

sleek unicornfish (Figure 113c). 

Herbivores accounted for 33% of total numerical abundance in the MLCD, 30% in the FRA, and 19% in the 

open area. Herbivore densities were similar among the three management areas. All three management areas 

showed large increases in numerical abundance of herbivores between 2004 and 2006 with subsequent declines, which were most pronounced in the MLCD and open areas (Figure 112d). Herbivores comprised 51% 

of the biomass in the MLCD, 51% in the FRA, and 50% in the open area. Herbivore biomass was 23% higher 

in the MLCD compared to the FMA and 27% higher compared to the open area. Biomass increased slightly 

between 2004 and 2007 and then declined slightly in 2008 (Figure 113d).
122 Monitoring Hawaii’s Marine Protected Areas

Results

Figure 112. Temporal change in fish abundance by trophic level at Kealakekua Bay MLCD and surrounding areas by 

habitat type from 2004-2008. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.

Figure 113. Temporal change in fish biomass by trophic level at Kealakekua Bay MLCD and surrounding areas by habitat 

type from 2004-2008. (a) Apex predators (b) Benthic invertivores (c) Planktivores (d) Herbivores.

Discussion

Photo courtesy of Keoki Stender, www.marinelifephotography.com
Discussion

Monitoring Hawaii’s Marine Protected Areas 125

The focus of this study was to 1) develop and implement a monitoring program of living marine resources (e.g., 

fishes, corals and other invertebrates, algae) within and adjacent to four representative MLCDs in Hawaii, 2) 

quantify the seascape using remotely sensed data to characterize the protected areas at a broader spatial 

scale, and 3) organize all of the associated spatial data sets in a GIS to better understand spatially-explicit 

ecosystem processes at the scale of the seascape. 

Benthic Habitats

General benthic habitat types were similar inside and outside MLCDs (Friedlander et al. 2007b) although coral 

cover tended to be higher inside. Therefore, the focus of this study was to examine trends over time in each 

of these management regimes. The trends over time tended to be consistent with the level of protection (e.g. 

MLCD).

Coral species richness and coral cover were greater inside the Hanauma, 

Pupukea and Honolua-Mokuleia Bay 

MLCDs compared to adjacent open areas and the rank order in these metrics 

among these MLCDs did not change 

over the study period (Figure 114). 

Coral cover generally increased over 

time inside Hanauma, Pupukea and 

Honolua-Mokuleia Bay MLCDs and did 

not change within Kealakekua MLCD. 

These results highlight the benefits of 

protection from fishing to benthic communities. 

Macroalgae cover was generally greater 

outside the MLCDs and increased over 

time, particularly adjacent to the Hanauma Bay MLCD (Maunalua Bay, Oahu), 

Honolua Bay MLCD (west Maui) and 

Kealakekua Bay MLCD (south Kona, 

Hawaii Island). Areas adjacent to the Hanauma, Honolua and Kealakekua MLCDs all have varying degrees of 

human population and runoff associated with stream beds or freshwater springs that may facilitate increased 

macroalgae growth. 

Seascape Metrics

Seascape metrics derived from NOAA benthic habitat maps were used to characterize the MLCDs. The NOAA 

benthic habitat maps released in 2007 were improved to include a hierarchical classification scheme and allowed for the quantification of seascape metrics based on zone, geomorphic structure, and biological cover 

classes. The seascape metrics used to characterize the MLCDs included: Shannon’s habitat diversity and 

evenness indices, patch fractal dimensions, MLCD perimeter and area, and MLCD perimeter/area ratio. An 

increase in the area of the Pupukea MLCD from 0.11 km2 in 1983 to 0.71 km2 in 2003, due to changes in regulations, resulted in a broader diversity of habitats within the new reserve boundaries. Sand habitat was absent 

from the 1983 protected area and the geomorphic structure within the new boundary was characterized by 

large sand habitats adjacent to rock/boulder habitats. The inclusion of these sand habitats within the reserve 

provides a corridor between hard bottom habitats for mobile organisms and also provides important feeding habitats for a number of organisms. In addition, fish and other mobile organisms that transit these sandy 

habitats benefit from the protection from fishing that was not provided in the older boundary configuration. 

Hanauma Bay and Kealakekua Bay MLCDs had small perimeter to area ratios (P/A), which are preferable for 

marine reserve design since fish and mobile invertebrates are less likely to spill over into open access areas 

Figure 114. MLCDs in Hawaii show a diverse mosaic of habitats such as 

this shallow, coral rich area. Photo: K. Stender, www.marinelifephotography.com
Discussion

126 Monitoring Hawaii’s Marine Protected Areas 

where they are vulnerable to fishing pressure. This small P/A ratio also increases the core area within the protected area and reduces the amount of boundary perimeter that is available for fishing. 

Several LiDAR-derived seascape metrics (e.g., depth, rugosity and slope) were quantified for each MLCD to 

better characterize the seascape within each protected area. In order for fisheries enhancement goals to be 

reached, marine protected areas should protect a range of structural complexities and habitat types in (SladekNowlis and Friedlander, 2004). The Kealakekua Bay MLCD showed the highest overall seascape diversity 

among all protected areas. Hanauma Bay and Honolua-Mokuleia Bay MLCDs demonstrated comparable seascape-level diversity. Biological cover within the 1983 Pupukea MLCD was dominated mostly by macroalgae 

and turf algae in a shallow depth range, but following boundary expansion, biological cover showed a more 

representative diversity of cover types. Further, the boundary was expanded to included deeper habitats that 

were less impacted by wave disturbance, particularity during the winter months, and as a result the new 

boundary now protects approximately 8.5 hectares of coral habitat (10-50% coral cover) that was not included 

in the older design. With the inclusion of coral habitats, a greater depth range, and a broader diversity of habitats in the reserve, there was a greater diversity and biomass of fishes protected within this new reserve area 

(see discussion below). The application of landscape pattern analysis used in this study represents an effective way for scientists and resource managers to evaluate potential or current protected area design scenarios 

in the marine environment at spatial scales that are appropriate for conservation.

Fish Assemblages

Rank order of MLCDs for various fish 

assemblage characteristic (species 

richness, numerical abundance, biomass, and diversity) remained the 

same between the 2002-2004 sampling 

period and the 2006-2008 sampling period. Species richness and diversity did 

not change appreciably in any of the 

MLCDs over the monitoring period. Numerical abundance was highly variable 

and fluctuated without pattern among 

MLCDs and areas open to fishing. 

Biomass increased in three of the four 

MLCDs from the initial sampling period. 

There was a 14% increase in Pupukea, 

a 7% increase Kealakekua, and a 5% 

increase in Honolua Bay. Biomass within the Hanauma Bay MLCD declined by 

5% during this same period. Hanauma 

Bay has been fully protected since 1967 

and the small decline in biomass over 

the monitoring period is not statistically significant and is within the margin of sampling variability. Although 

overall biomass declined slightly, apex predator biomass in the Hanauma Bay MLCD increased as a result 

of increasing numbers and sizes of bluefin trevally jacks (omilu, Caranx melamypgus; Figure 115). A similar 

increase in apex predator biomass was observed in the Honolua Bay MLCD over the same time period. Herbivore biomass increased by 17% within the Pupukea MLCD and this increase was driven mainly by large 

parrotfishes that are prized fisheries species and likely responded to the increased protection provided by the 

expansion of the protected area in 2003. The significant increase in apex predator biomass within the Pupukea 

MLCD is also likely a result of the increase in protected area size. 

Figure 115. Protected areas in Hawaii harbor large apex predators, like this 

jack. Photo: K. Stender, www.marinelifephotography.com
Discussion

Monitoring Hawaii’s Marine Protected Areas 127

Biomass declined by 39% outside the Pupukea MLCD and may reflect increased fishing pressure outside the 

MLCD over time. Biomass outside the Hanauma Bay MLCD increased by 32% although the absolute increase 

(+0.05 t ha-1) was small. Because biomass was so low in this area, the presence of a few schools of larger 

fishes resulted in the large relative increase in biomass although the absolute biomass was still extremely low. 

Management Implications

Combining seascape-scale habitat stratification with in situ ecological data allowed for the development of a 

statistically robust monitoring program of living marine resources (e.g., fishes, corals and other invertebrates, 

algae) within and adjacent to four representative MLCDs of marine protected areas in Hawaii. The results 

clearly show that areas with good habitat quality and management conserve fish populations and benthic communities within their boundaries while areas without protection are in poorer ecological health and continue to 

decline over time. The consistency of these patterns over time support the findings of the initial assessment 

conducted by Friedlander and colleagues (Friedlander et al. 2006, 2007a, 2007b) and increase confidence in 

the effectiveness of marine protected areas as a management tool for the conservation of nearshore marine 

resources in Hawaii.

Marine protected areas are spatially discrete forms of management and georeferenced data on seascape 

metrics can be integrated within a spatial framework to ensure MPA design scenarios are incorporating an 

appropriate range in depth, habitat complexity, and an interconnected mosaic of benthic habitat types. In this 

study, remotely sensed data allowed for the examination of MLCDs at spatial scales that are appropriate for 

management and provided valuable information on seascape metrics (e.g. rugosity, slope, depth, habitat type). 

Using a seascape approach controlled for differences in habitat by utilizing the NOAA benthic habitat maps 

and supported the development of a robust, spatially explicit monitoring program. This approach allows for the 

identification of spatial and temporal trends across the seascape to be highlighted and supports subsequent 

spatial planning and management actions. The findings from this study greatly contribute to the understanding 

of marine protected area design and function in Hawaii and are imperative to the development of comprehensive coastal and marine spatial planning.
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Photo courtesy of Keoki Stender, www.marinelifephotography.com
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U.S. Department of Commerce

Gary Locke, Secretary

National Oceanic and Atmospheric Administration

Jane Lubchenco, Ph.D., Under Secretary for Oceans and Atmosphere

National Ocean Service

David Kennedy, Acting Assistant Administrator for Ocean Service and Coastal Zone 

Management

The mission of the Center for Coastal Monitoring and Assessment’s Biogeography Branch is to develop information and 

analytical capabilities through research, monitoring, and assessment of the distribution and ecology of living marine 

resources and their associated habitats for improved ecosystem-based management. For more information, visit:

http://www.ccma.nos.noaa.gov/about/biogeography/welcome.html