1

A MANUAL FOR MONITORING CORAL REEFS WITH

INDICATOR SPECIES:

Butterflyfishes as Indicators of Change on Indo-Pacific Reefs

Michael P. Crosby, Ph.D

National Research Coordinator

Office of Ocean and Coastal Resource Management

National Oceanic and Atmospheric Administration

Washington, D.C.

and

Ernst S. Reese, Ph.D

Professor of Zoology

Department of Zoology and the

Hawaii Institute of Marine Biology

The University of Hawaii

Honolulu, Hawaii

Sponsored by

the Legacy Resource Management Program, Department of Defense;

the Office of Ocean and Coastal Resource Management, National Oceanic and Atmospheric Administration

and the U.S. Man and the Biosphere Program
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A MANUAL FOR MONITORING CORAL REEFS WITH

INDICATOR SPECIES:

Butterflyfishes as Indicators of Change on Indo-Pacific Reefs

Please cite as: Crosby, M.P. and E.S. Reese. 1996. A Manual for Monitoring Coral Reefs With Indicator

Species: Butterflyfishes as Indicators of Change on Indo Pacific Reefs. Office of Ocean and Coastal Resource Management, National Oceanic and Atmospheric Administration, Silver Spring, MD. 45 pp.

October 1996

Michael P. Crosby, Ph.D

National Research Coordinator

Office of Ocean and Coastal Resource Management

National Oceanic and Atmospheric Administration

Washington, D.C.

and

Ernst S. Reese, Ph.D

Professor of Zoology

Department of Zoology and the

Hawaii Institute of Marine Biology

The University of Hawaii

Honolulu, Hawaii

Sponsored by

the Legacy Resource Management Program, Department of Defense;

the Office of Ocean and Coastal Resource Management, National Oceanic and Atmospheric Administration

and the U.S. Man and the Biosphere Program
3

PREFACE

The importance of coral reef ecosystems may be seen in their numerous ecological,

aesthetic, economic and cultural functions. Atoll and barrier reef islanders recognize that

healthy reefs are essential for the support, creation, and repair of the coral islands upon which

they live. Coral reefs also protect coastlines from shoreline erosion, and serve as a living

pantry for the subsistence harvest and consumption of many reef organisms. The cycle of reef

accretion and erosion maintains beaches, and provides habitat for seagrasses and mangroves.

Coral reefs are important recreational resources for most of the world's people having the

privilege of living near them. An outgrowth of the International Coral Reef Initiative (CRI) is

the early development of national coral reef initiatives. The United States is developing an

inter-agency CRI to create the base for a combined domestic and international effort aimed at

the conservation and effective management of coral reef ecosystems (Crosby et al., 1995,

Crosby and Maragos, 1995).

The ability of coral reef ecosystems to exist in balanced harmony with other naturally

occurring competing/limiting physico-chemical and biological agents has been severely

challenged in the last several decades by the dramatically increased negative and synergistic

impacts from poorly managed anthropogenic activities (Maragos et al., 1996). Globally,

scientists are now working together and with other groups to promote assessment, monitoring,

other research, protection, and restoration of coral reefs. Many non-governmental groups have

been at the forefront in local implementation of various coral reef monitoring efforts. These

grassroots efforts should be applauded and encouraged. However, long-term nationally and

globally coordinated coral reef monitoring programs are essential to manage, archive, translate

and transfer data to scientists, managers and other interest groups. Within the U.S., the National Oceanic and Atmospheric Administration (NOAA) is developing a nationally coordinated coral reef monitoring program to be implemented in 1996, and is actively pursuing

partnerships with other agencies (such as the National Park Service and the Environmental

Protection Agency) and volunteer interest groups (such as American Oceans, The Nature

Conservancy, REEF and Reefkeeper) in this effort.

Numerous coral habitats (particularly in the American Flag Pacific Islands - AFPI) are

under Department of Defense (DOD) management and are sites of military training exercises.

It is, therefore, appropriate that an additional partner in the Coral Reef Initiative is the U.S.
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Department of Defense, and this manual is a direct product of that partnership. This manual

describes a non-invasive and “low-tech” approach to assessing and monitoring coral reef

habitats. It will serve as a reference for individuals with limited technical science background

and expertise. Contents include step by step instructions for determining coral-feeding fish

behavior and how to relate these traits to changes in coral habitat condition, as well as basic

techniques for assessing coral and fish biodiversity and percent cover.

It has become evident that new and innovative techniques are needed that can provide

an “early warning” of stress within coral habitats so that investigations to determine causes of

the stress and implementation of counteractive management strategies to protect these critical

habitats can be initiated. A critical need also exists for relatively low cost, simple (low technology) methods for assessing the overall stress levels (“health”) of coral reef habitats

throughout the world. This need is especially high in regions that do not have readily available “high” technology equipment and expertise for conducting comprehensive “high-tech”

ecological studies. Even within the more “developed” regions of the world, the availability of

low cost methods for coral reef assessments that could be conducted by persons without

extensive technological training and education would be of immense value to monitoring the

stress in these ecosystems.

To date, no DOD studies have been initiated to determine the nature or extent of possible coral reef damage resulting from amphibious exercises. Past actions have excluded

amphibious exercises from training because of a lack of scientific data to prove/disprove

damaging effects on coral reefs. If the U.S. DOD is to maintain its “readiness” capabilities,

training exercises in the direct and/or indirect vicinity of coral reef habitats are critical. If

these exercises are to be conducted with no (or at least minimal) significant negative impacts

to these fragile ecosystems, coral reef habitat assessment and long-term monitoring protocols

must be developed and implemented. It is also desirable that the local indigenous peoples of

the islands where the exercises occur should be significantly involved with DOD in any assessment and monitoring activities.

In a broad sense, the development of the coral reef monitoring approach described in

this manual will provide the basis for a future, more comprehensive effort to provide scientific

evidence of the compatibility of amphibious exercises or highlight potential damages, and

recommendations for mitigating alternatives to minimize effects of these exercises. The

protocols encompassed in this approach will have applicability throughout the Indo-Pacific
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region and will serve to not only assist DOD in being able to conduct military exercises while

also protecting coral habitats, but will also serve to empower local populations in managing

their coral ecosystems in partnership with DOD.

As part of the development of this manual and associated video, Drs. Crosby and Reese

conducted pilot classroom and in situ coral reef assessment and monitoring training exercises

with selected individuals from the Commonwealth of the Northern Mariana Islands, American

Samoa and Hawaii. Participants in this training exercise were nominated by the Governors’

Principal Points of Contact for the U.S. Coral Reef Initiative in Hawaii, American Samoa and

the Commonwealth of the Northern Mariana Islands, as well as the Kaho’olawe Island Reserve

Commission. Trainees participated in approximately three days of training at the study sites

that Drs. Crosby and Reese are examining as part of a larger Marine and Coastal Ecosystem

Directorate Core Project of the U.S. Man and the Biosphere Program. Field testing of the

approach described in this manual included 640 man-hours of on-site, in water, data collection

from November, 1992 to June, 1996.

The pilot training exercise was accomplished by demonstrating the use of indicator

species of coral feeding chaetodontid fishes to detect low level, sub-lethal changes in the coral

reef habitat. These techniques serve as an early warning of stress within a coral reef prior to

reaching a “point-of-no-return” (hence remedial actions may be taken). Specifically, correlations of fish feeding preference, abundance, and behavior with coral abundance and health are

determined. Demonstrations of line transect methods for determining coral diversity and

abundance, and fish counting methods were also made. Proper methods for data entry and

analyses were also presented.

Support of this study and the collaborative interplay between DOD, NOAA scientists,

University scientists, as well as the Local Level Coral Reef Initiatives, Coastal Zone and

Natural Resource agencies has produced a monitoring approach important to the future planning and management of sensitive coral reef habitats, and will facilitate transfer of developed

techniques and information to local indigenous peoples in the AFPI and other sites around the

world. Of special utility will be the ability to utilize a low cost, low technology method in

areas of the world that critically need assistance in determining the level and degree of environmental perturbations to coral and hard bottom marine habitats. Such local level monitoring

programs will form the basis of efforts to not only produce the scientific data required to

objectively ascertain impacts of training exercises and recommend mitigation actions, but will
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also serve to demonstrate DOD and NOAA commitment to forming partnerships in training

and information transfer that will support the empowerment of local populations in the ability

to better assess and manage their coral reef ecosystems.

The authors wish to thank the Legacy Resource Management Program and Department

of the Navy - Department of Defense, the National Oceanographic and Atmospheric Administration, the U.S. Man and the Biosphere Program, and the University of Hawaii for their

support in this effort. In addition, they are grateful for the active participation by the Governors’ Principal Points of Contact for the U.S. Coral Reef Initiative (Lelei Peau, American

Samoa; Douglas S.Y. Tom, Hawaii; and Manuel Sablan and Eric L. Gilman, Commonwealth

of the Northern Mariana Islands), the Local Level Coral Reef Initiative Programs in Hawaii,

American Samoa and the Commonwealth of the Northern Mariana Islands, as well as the

Kaho’olawe Island Reserve Commission. Special thanks to Katie Geenen and Kai Wang for

their assistance in preparing this manual and associated video; to Javier Mendez and Randy

Kosaki for their assistance with the field work and in situ training aspects of this project; and

to the trainees of our pilot training sessions.

Cover Photograph: A pair of butterflyfish, Chaetodon multicinctus, the indicator

species being used in Hawaii. Photo by Randall Kosaki using a Nikonos V camera with a 35

mm lens, Nikon SB-103 flash, Kodachrome 200 film at f16.
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There are many methods to monitor coral reefs and the associated biota. Each method

has its strengths and weaknesses, benefits and costs, all of which must be considered carefully.

Each method also varies in how much scientific training is necessary to successfully use it.

The choice of a method depends on the question being asked, the resources available,

and the training of the persons who will do the work. The approach described in this manual is

aimed at answering the question: Is the condition of the coral reef changing?

The Butterflyfish Fish Indicator method is relatively simple and inexpensive. It does

not require highly trained personnel with extensive knowledge of reef organisms. More

importantly, however, it is sensitive to changing conditions on the reef and, we believe, should

provide an “early warning” that change is occurring. This may provide enough time for

remedial actions to be taken before entire coral reef habitats are irreversably lost.

I. INTRODUCTION

The approach described in this manual has at least four important features:

• It does not require that the scientific names of all the coral and fishes are known by the data collectors. It

does require that one or two key coral feeding indicator species be recognized for each geographical

location. Suggested species are listed in Appendix 1.

• It can be used stepwise with each step providing more information. The first step, which is counting

the coral feeding butterflyfishes along the transect, or even just the designated indicator species,

provides useful information. The next step, which is counting the corals species occurring at 0.5 m

points along the transect line, provides further information and so forth. This means that the method

can be used to match the time, resources and personnel available.

• It is best applied where gradual, slow (even sub-lethal) environmental disturbance is suspected, which

would be difficult and expensive to measure by other methods, such as collecting water samples or tissue

samples for analysis. Clearly, the method is not appropriate for catastrophic disturbance such as oil spills,

storms, etc.

• It is an "environmentally friendly" non-destructive or non-consumptive method that can be used by

volunteers, local communities, and individuals with no previous former technical scientific training.
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A. The Indicator Species Concept

Butterflyfishes of the family Chaetodontidae are conspicuous inhabitants of coral reefs

throughout the world (Burgess, 1978; Motta, 1989). Many species are obligate corallivores,

and thus depend on the live tissue of corals for their food. Because their metabolic or energetic demand is so intimately linked to the existence and overall condition or "health" of the

coral substrate, we believe that these species of butterflyfishes are excellent candidates for

indicators of changes in conditions on the coral reef.

The conceptual ideas are simple. Corallivorous butterflyfishes have coevolved with,

and are intimately related to the corals on which they fed (Reese, 1977, 1981, 1991;

Harmelin-Vivien and Bouchon-Navaro, 1983; and Roberts et al., 1988). The distribution and

abundance of these fishes should be directly correlated with the distribution and abundance of

the corals. If the corals are adversely affected by stressful environmental conditions, such as

chronic low levels of pollution, their health will deteriorate. This deterioration should be

detected by the fishes which feed on them. In many species of organisisms sub-lethal levels of

stress cause re-allocation of available energy. A shifting of energy from production of storage

products such as glycogen or lipids to an increase in respiration would make an organism less

"nutritious" for those consuming that organism. The corals are sessile and cannot avoid the

stress, whereas the fishes are mobile, and can emigrate to healthier regions of the reef.

Chronic low levels of pollutants on the reef and the slowly deteriorating condition of the corals

are difficult to detect by conventional methods (Dahl, 1981; Brown and Howard, 1985; IUCN,

1993; UNEP/AIMS, 1993; Rogers et al., 1994; Jones and Kaly, 1996). In contrast, simply

counting the abundance of the brightly colored, diurnal butterflyfishes using conventional

census techniques is a relatively simple task. However, as will be discussed in a following

section, it is important to focus on coral feeding fishes since plankton-feeding fishes would be

less dependent on the coral. A diver can be taught in a few hours to recognize and census key

indicator species along a transect. The indicator hypothesis can be stated as follows: Coralfeeding butterflyfishes respond to declines in coral quality or abundance by behavioral and

spatial adjustments that can be easily and rapidly observed.

Reese (1977, 1981) first proposed that obligate corallivores, such as many species of

butterflyfishes, could serve as indicator organisms. Subsequent experimental studies have

increased our knowledge of the relationships between butterflyfishes and corals, and support

their potential as indicators to monitor changes in conditions of coral reefs.
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B. Behavioral Ecology of Butterflyfishes

Fishes of the family Chaetodontidae, the butterflyfishes, are found in all tropical seas

of the world. There are 114 species in 10 genera with 90 of the species in the genus

Chaetodon. The systematics of the family is well studied using classical systematic methods

(Burgess, 1978). Butterflyfishes are characterized as diurnally active, brightly colored inhabitants of coral reefs. They belong to three feeding guilds: corallivores, benthic omnivores, and

planktivores. The coral feeders are especially interesting because they are closely associated

with the living coral reef for both food (as described in the previous section) and shelter. Coral

feeding species tend to live in monogamous pairs and are broadly home ranging or territorial.

Pair bonds are established based on size reflecting a division of labor between mates.

Males actively defend the territory allowing the female more time to feed. Larger males are

able to defend larger territories, while larger females produce more eggs. Thus, there is assortative mating and pair bond formation based on size. Butterflyfishes are dusk spawners on

evenings around new and full moon. They are broadcast spawners so there is no parental

care. Coral-feeding pairs of chaetodontids show a very high degree of site fidelity with the

same individual pairs being found on the same territories for seven to eight years (Reese,

1991). Their life span is 10 to 12 years depending on the species. Predation is minimal on

adults but may play a significant role in limiting population sizes during the planktonic larval

dispersal stage in their life history. Their striking color patterns make them ideal for behavioral observation in the field, while their typical life history allow individual paris to be observed for many years. Not only can species be readily identified from a distance and interactions recorded, but variation within each species permits individual recognition in the field.

All of these characteristics make butterflyfishes ideal candidates for indicator species of

ecological conditions on coral reefs.

C. Butterflyfishes as Indicator Species

The concept of using certain key species as indicators of ecological conditions is now

well established (Soule and Kleppel, 1988; Jones and Kaly, 1996). The situation with respect

to butterflyfishes is reviewed by Hourigan et al. (1988).

A number of points must be emphasized. First, sensitive biotic indicators are most
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useful when one wishes to detect low levels of chronic perturbations such as low levels of

chemical pollutants or small changes in temperature or nutrient levels. Over time such low

levels of chronic perturbations can have marked detrimental effects on the ecosystem they are

impacting. Yet it is extremely difficult and expensive to devise a sampling regime to detect

such low levels. It is under such conditions that sensitive biological indicators are most useful.

Clearly, one doesn’t need a sensitive indicator for episodic, catastrophic events like oil spills or

storms. The second point of importance is that not all chaetodontids are candidates for indicator species. The planktivores, in particular, hovering above the reef facing into the current to

intercept plankton are not sensitive to the corals on the reef beneath them. Likewise, the more

omnivorous species, feeding on benthic invertebrates other than corals and on algae, tend to be

opportunistic and they feed on prey in proportion to their abundance. Therefore, as the prey

changes, they change their diets and so do not indicate that a change is occurring in the ecosystem.

In contrast, the coral feeding chaetodontids make ideal indicators because they are

directly dependent on the corals as their principal source of energy. Many species are obligate

corallivores and do not feed on anything else. Furthermore, they show strong preferences for

certain species of corals which provides a further dimension of sensitivity to the system. Since

they are territorial, strongly site attached, and live for many years, they provide a longitudinal

component to the system which has great value. Even if changes occur very slowly over a

number of years in the ecosystem which will eventually make the corals moribund, the same

individual pairs of butterflyfishes will be present to experience the change.

Current efforts to use butterflyfishes as indicators of coral reef diversity in Indonesia

and the Philippines (Nash, 1989; White, 1989) have overlooked this important point. Forty

species are listed on their survey form and many of these are not corallivores. To recognize all

these species is a difficult task for non-specialists charged with making the surveys. In addition, less time and attention can be given to the distribution, abundance and social behavior of

the truly important indicator species, the corallivores. Nevertheless, these efforts are an

important start and make it even more important that the utility of the concept be demonstrated

in the correct way.

A reef which is dead from siltation or from crown-of-thorns or from some other causal

factor no longer has its characteristic assemblage of coral feeding chaetodontids (Reese, 1981;

Bouchon-Navaro et al., 1985; Hourigan et al., 1988). The time course of this change in popu-
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lation structure is unknown at present and currently is being investigated. There appears to be

a threshold level of reef deterioration at which the fishes begin to leave, perhaps related to the

decrease in both abundance and diversity of the corals upon which they are feeding. However,

once a coral reef has died, preventive medicine in the form of monitoring with indicator

species is useless. Although this approach should still be useful in assessing coral reef recovery.

 The size of corallivorous butterflyfishes territories are determined by the amount of

coral food contained therein. Experimental removal of coral from territories results in pairs of

fish attempting to enlarge their territories at the expense of their neighbors which results in

increased agonistic levels of behavior (Hourigan, 1987; Tricas, 1986, 1989). These changes in

social behavior, in what otherwise is a stable situation, provide a sensitive early indication that

changes are occurring. Furthermore, these events which precede the actual exodus of the fish

from the reef appear to occur at a time when the corals are becoming stressed ("unhealthy")

but before they have become moribund beyond recovery. Since we are interested in detecting

slow changes in the ecosystem, this early warning should provide time for remedial actions to

be taken by persons or agencies charged with management of the reef resources, providing the

changes are due to perturbations caused by human activities which are impacting the area and

could otherwise be controled.

II. APPLICATION OF THE METHOD

A. Assessment of the Problem

The first steps in the application of the indicator species method are to:

• accurately assess the problem

• clearly identify the questions to be asked

• state the goals of the monitoring program

For example, in an ongoing study of the condition of the coral reefs in Hawaii (Crosby,

1994), we assess reefs in near pristine condition, as well as those impacted only by non-point

source runoff high in sediments and those impacted only by recreational diving. In the reef
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impacted by sediments, the sediment came from erosion of the soil due to grazing of the

vegetation by feral goats and the effects of using the island as a military target causing fires

and further destruction of the vegetation.

Since the military stopped bombing the island and removed the goats, and Hawaiian

groups began an extensive replanting program, we asked the question: Will the coral reefs

recover as the amount of sediment impacting them decreases? In order to answer this question, we looked for two coral reefs areas, one which was severely impacted by the sediment

and another which was relatively pristine and less impacted by sediment. Using information

gathered by the Division of Aquatic Resources, Department of Land and Natural Resources,

State of Hawaii (Kanenaka et al., 1993), we identified two coral reef areas: one near Hakioawa

is pristine with little evidence of sediment damage, while the second reef area near Kuheia Bay

is heavily impacted by sediment. We also include as a third study site, the island of Molokini

which has as its only clear human perturbation, excessive recreational diving and boat anchoring. In recent years, mooring buoys have been installed in Molokini and commercial boat

permits are required.

The goal of our monitoring program is to establish long-term monitoring stations to

detect evidence of the coral reefs recovering or declining at Kuheia Bay and Molokini Island,

that is a change in the direction of replenishment of the coral biota gradually over time. In

contract, we postulated that we would see less change in the relatively pristine reefs at

Hakioawa.

Thus we assessed the problem, asked the relevant question, and clearly stated our

goals.

B. Field Observations

All of the data collection described in this manual involves direct observation of

butterflyfishes and corals. Therefore, it is necessary to use scuba. Since butterflyfishes are

diurnal, all work is done during daylight hours. In addition, reef fishes don't demonstrate

"normal" behavior patterns under conditions of high turbulence and surge and poor visibility.

Hence most observations should be collected under normal weather conditions for the particular study site. Because many flourishing coral reefs in the Indo-Pacific region are at depths of

about 5 - 20 m, surface weather conditions directly affect the underwater conditions.
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C. Getting Started: Establishing the Transect Lines

Transect lines are simple to make. Brightly colored polypropylene line about 1/4 to not

more than 1/2 inch in diameter (approximately 1 cm in diameter) works best. We recommend

sections slightly longer than 30 m. The line floats so it does not get tangled in the coral. At

one meter intervals colored cloth or colored surveyor's tape can be knotted through the weave

of the line. Use different colored cloth for different lines. It is convenient to tie a lead weight

at both ends of the line which helps secure the line at either end.

Transect lines are not placed haphazardly (Fig. 1). They are placed in areas where

coral is abundant since the question that is the focus of this Indicator Species Approach is

whether or not the condition of the coral on the reef is changing. If one was interested in

assessing the relative amount of different kinds of bottom substrata such as sand, coral, algae

turf, rubble etc., over the broader area of the study site, then, one would place the lines randomly in the area to be sampled. There is much confusion on this point, but what is important

is to always consider the essential question being asked. Again the approach described in this

manual focuses on the condition of the coral, not its abundance relative to other kinds of

bottom in a broad area.

We recommend that at each study site from one to four 30 m transects be established.

The configuration depends on the reef’s contour. For example, on an expansive reef of fairly

uniform coral cover, a star-burst pattern is used. If the reef is a system of raised coral ridges

("spur and groove") or a narrow reef front, the transects are placed roughly in a parallel

pattern. Since we are interested in change of the living corals and behavior of coral feeding

butterflyfishes, the transects are purposely placed in areas of high coral cover.
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Figure 1. The transect lines are set at 90 degrees to each other in a star-burst pattern. Here only two

lines are visible. The diver on the left is in the process of laying out the 30 meter transect line, while the

diver at the right is counting pairs of butterflyfishes along the transect line.

Data are recorded on underwater paper attached to a regular, inexpensive clipboard

using a soft lead No. 1 pencil. Plastic underwater paper is available under the brand name of

"Nalgene Polypaper".1 Since plastic paper and woodless pencils are relatively expensive, an

inexpensive alternative is to cut pieces of matte acetate to the correct size, place the acetate

sheet on top of the appropriate data sheet, securing it under the clip on the board and with a

rubber band at the bottom of the board. A regular No. 1 or 2 lead pencil writes easily on the

acetate sheet. Upon completion of the dive, the data from the acetate sheet can be copied onto

1 Nalgene Polypaper is available from Technical Service, Nalge Company, Sybron Corp., Box 20365, Rochester,

N.Y. 14602-0365 U.S.A., Telephone (716) 586-8800, Fax (716) 586-8431. Although regular No. 1 or 2 lead

pencils work, we have found that woodless pencils which are available in most art supply stores or from a

manufacturer such as "Pentalic Woodless Pencils", M. Grumbacher, Inc., Cranbury, N.J. 08512 U.S.A., work even

better. The softer grade of woodless pencils work best.
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D. Counting the Numbers of Coral Feeding Butterflyfish

Once the transect lines are established, the numbers of each species of butterflyfish

within five meters of either side of the transect line are counted. This is known as the Belt

Transect Census method (Brock, 1954, 1982; Rogers et al., 1994), and is widely used. Where

one is interested in the numbers of fishes of all kinds, including smaller cryptic species and

larger transitory species, the Stationary Fish Census method (Bohnsack and Bannerot, 1986)

may be preferred.

Since coral feeding butterflyfishes are home ranging and often territorial and are not

preyed on by large transient predators, they usually do not hide or flee when a diver swims

along the transect line (Fig. 1). An exception to this may be in areas where spear fishing has

occurred. Furthermore, since they usually occur in female and male pairs, the observer places

a “2” on the data sheet (Fig. 2) when a pair is counted, a “1” for a solitary individual, and a “3,

4, 5, etc.” if a larger group is observed. An experienced observer may wish to place the letters

“J, S, M, L” above the numbers to indicate relative size such as juvenile, small, medium, and

large. Please note: This additional size designation method should only be used by a thoroughly experienced individual who is familiar with the size categories of the species in question.

Note that on the Fish Transect Data sheet (Fig. 2) the designated indicator species, C.

multicinctus, is shown both as an adult and as a juvenile. C. ornatissimus and C. trifasciatus

are obligate coral feeders and they could be used as indicator species as well. Because C.

multicinctus is more abundant and has smaller territories, it is a more convenient indicator

species in Hawaii. C. unimaculatus is a facultative coral feeder. It is, therefore, not as good a

choice for an indicator species. C. fremblii is an omnivore and feeds on a variety of small

invertebrates. It is, therefore, not as closely dependent on coral as the other species, and may

serve as a sort of control. If a pair is seen, the number “2” is placed to the right of the fish, if a

single fish is seen, the number “1”, if a group, the number designating the numbers of fish in

the group is noted, for example “3”. Monitoring trainees often ask the following two important

regular paper or duplicated with a copier if one is available. The acetate sheet can then be

scrubbed clean using a kitchen scouring pad and reused many times. Matte acetate comes in

sheets and is available in art supply and most stationary stores.
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questions: How much time should I take counting the fish along the 30 m transect? and

Should I count all the butterflyfishes along the 30 m transect line?

Slowly swimming a 30 m transect line, visually scanning the reef 5 m to either side of

the line should take about 5 minutes. Every effort must be made not to count any individuals

twice and swimming at this rate decreases the chance that pairs of fish swim back into the

observers’ view.

The second question is more difficult to answer, because it depends on the experience

of the observer and the amount of circumstantial data she or he wishes to collect. Usually only

one or two, at the most three, species of butterflyfishes are designated as the indicator species

for a given study site. In Hawaii, we are using only one species, the brown-barred

butterflyfish Chaetodon multicinctus. It is abundant and easily observed. In most tropical

locations where diversity is greater, the decision may be made to use two or more species as

the local indicator species. Candidate species are listed in Appendix 1.

The following is a step-wise progression of observations that data collectors should

employ as they become more experienced over time and more comfortable with their knowledge of the local fish assemblages.

Below are listed the species which observers may wish to count along the transect:

1. At the minimum, the numbers of pairs and single individuals of the designated focal indicator species must be counted.

2. Next, all the known coral feeding species of butterflyfishes should be counted.

3. If the observer knows the fish fauna well, she or he may wish to count other species of

butterflyfishes as well. For example, species which feed on plankton or are benthic omnivores feeding on algae and small invertebrates like crabs, shrimps, tube worms, etc. may be

counted. These may serve as a “control” since their abundance and distribution should be

less immediately or directly affected by the changing condition of the coral than those

species which depend on the coral directly for their food.
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Figure 2. This is an example of the Fish Transect Data sheet being used in Hawaii. Sample data entries are

shown on the data sheet.

OBSERVER: _

OATE: _

SITE: _

TIME: _ DEPTH: _

IFish Transect Data: Record the number of ea. spp. seen within 5m on either side of 30m line

species

C. multicinctus

, I

2..2.2..

number of individuals

C. mlliticinctus Juveniles «4 em.)

C. ornatissimus

•

C. trifasciatus

C. unimaculatus

•

C. fremblii

other species:

2

2.

32.
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E. Estimating the Percent Coral Cover under the Transect Line

Prior to beginning the estimation of coral cover along the 30 m transect line, the observer should make note of the most abundant or apparent species under the transect line. This

familiarization period usually focuses on six to perhaps 12 species of corals. The six species

of coral shown on the Coral Transect Data sheet (Fig. 3) are those which are most abundant at

the Kaho’olawe and Molokini study sites in Hawaii.

If the scientific names of the coral species are known, these can be entered on the data

sheet. If the scientific names are not known, common local names, such as “finger coral” or

“brain coral”, can be used. Even a simple description is useful, such as “brown lumpy coral”

or “reddish brown plate coral with white polyp tips.” The latter description for example would

be adequate to recognize Montipora verrucosa in Hawaii.

After the data collector feels comfortable with his or her coral identification skills, he

or she swims along the same 30 m transect line used for the fish counts and records the coral

directly beneath the line at 0.5 or 1.0 m intervals. If a 30 m transect line is in use, then this

results in either 60 or 30 data points, each data point being a type of coral. This is known as

the Line and Point Intercept Transect method (Ohlhorst et al., 1988; Rogers et al., 1994).

These authors also discuss other widely used methods for estimating coral cover such as the

Quadrat Grid Transect method. However, the Line and Point Intercept Transect method is

quicker, somewhat easier for novices, provides reliable data and is the preferred when applying

our Indicator Species Monitoring method.

A shortfall of both the Line and Point Intercept Transect and the Quadrat Grid

Transect methods is that they do not take into account the rugosity or uneven surface contours of many coral reefs. A more accurate estimation of the surface coral cover, taking into

account the rugosity of the reef, can be obtained using the Chain Transect method. This

method also has its limitations, is time consuming, and can be destructive to fragile corals.

Simulation studies of the different methods indicate that the best estimates of percent coral

cover and diversity are obtained by doing more Line and Point Intercept Transects rather than

fewer and more time consuming samples taken with the other methods (Kinzie and Snyder,

1978).

The percent coral cover is calculated by dividing the number of observations of a

certain species of coral by the number of data points along the transect line. For example, if an

observation was made at each 0.5 m interval along the 30 m transect, and the coral Porites

compressa was recorded at 12 of the 60 data points along the transect, then P. compressa

accounts for 20% of the coral cover along the transect line.
19

Figure 3. This is an example of the Coral Transect Data sheet being used in Hawaii. Sample data entries are

shown.

DATE: _ TIME: _ DEPTH: _

SITE: OBSERVER: _

oral Transect Data: Record the type of substratum lying directly under the transect line

at O.5m intervals

number of points

Porites compressa

Finger Coral

8

-····· . -.

. ~7

Porites /obara

Lobe Coral

~w

MOnJipora verrucosa

Brown plate coral

..

f.~fC~ •. l ... , J

',l!

Monlipora patula

Crown cain coral

IIllIlIl/lli

/lJJI

1I1/1I/flJI/1I1

,,,,,,,11/111111/1

II/II

II'

Pocil/opora meandrina

Caulinower coral

PaVOlIll

-

varians

Meandering polyp coral

aTHER CORAL SPP. II

NON-CORAL (dead coral. sand, algae, etc.) J
20

F. Marking the Territory Boundary and Measuring Chasing Behavior

Marking of obligate coral feeding Butterflyfish territory boundaries and observations of

chasing behavior are usually done together in a single dive using the data sheet shown in

Figure 4. First the diver locates along the transect line the focal pair of fish whose territory

will be marked. Once this pair is recognized and the first color tagged nail is placed where the

pair of fish is first seen, the diver begins the first 10 minute period of observation and marking

territory. It is hightly recommended that a diver watch with a timer and audio alarm feature be

used for all timed observations. It is advised to look at your watch and enter the time you start

above the “0-10 min.” designation, for example 0930-0940, the next “10-20 min.” designation,

0940-0950 etc. This helps the diver remember in which 10 minute observation period she/he

should be noting the chasing observations.

If you are observing Chaetodon multicinctus, the indicator we are using in Hawaii, and

one fish in your focal pair chases a fish in an adjacent pair of C. multicinctus, you would

record “Agg Cm” in the appropriate 10 min. block on the data sheet. This indicates that one

fish of the pair you are watching was the aggressor and chased a fish of the neighboring pair of

C. multicinctus. If both fish of the pair you are observing chase both fish of the neighboring

pair, then you should note “Agg Cm 2x.” This indicates that two chases occurred and it is

entered as two aggressive chases initiated by the pair you are watching in the computer entry.

If the neighboring pair of C. multicinctus is the aggressor and the pair you are observing is submissive, that is, they flee, then you would record “Sub Cm” in the appropriate time

interval on the data sheet. If all four fish of the two pairs are involved, you would enter “Sub

Cm2x”. Please note: Any shorthand that one wishes to use, such as "+" and "-" instead of

"agg" and "sub" respectively is perfectly acceptable as long as the shorthand is clearly defined.

These behavioral encounters often occur in bouts where one pair chases another pair

out of the territory, but then when the pair being chased enters its own territory, it turns and

becomes the aggressor. Thus your entry might look like, “Sub Cm, Agg Cm” or if all four fish

are involved, “Sub Cm 2x, Agg Cm2x.” It is well known that animals tend to be more aggressive when they are in their home territory but they become submissive when they are outside

of their territory. Thus, you may observe sort of a “see-saw battle” as they chase each other

back and forth between their territories.

Note that another species, Plectroglyphidodon johnstonianus, a damselfish, is included
21

on the Chase Data sheet. This is because this little fish often lives within the territory of pairs

of C. multicinctus where it feeds on coral. It also lays its eggs on the coral. It is aggressive

and often chases C. multicinctus. In contrast, C. fremblii which is not a coral feeder is rarely

chased.

The diver observes the pair of fish for 50 minutes as shown on the data sheet (Fig. 4).

This is not an arbitrary time period. Please note: While it is advisable to complete all 50

minutes of observations in a single dive, it is not essential. However, if a second dive is

required, it should be done as quickly as safety requirements allow. Previous studies have

shown that for coral feeding butterflyfishes with relatively small territories, 50 minutes of

observation and placing nails on the perimeter of the territory provides an accurate measure of

the boundary of the territory.

The process of marking the territory proceeds throughout this 50 minute period after

the first nail is placed on the reef where the pair of fish is noted. In all probability this first nail

is inside the territory and subsequently will be relocated as the boundary of the territory becomes more clearly defined (see Fig. 5 in which the color tagged nails on the boundaries of the

territories are connected by a dotted line).

As the diver observes the pair of fish, she/he places a color tagged nail where the fishes

hesitate and turn back into their territory. This hesitating and turning behavior is repeated

when the fishes reach the boundary of their territory. Most agonistic chasing behavior occurs

at the boundaries as well. With experience the diver learns to recognize these movements and

becomes very good at placing the nails along the territory boundary. The diver places most of

the nails during the first two or three ten minutes observation periods as the pair of fish forage

through their territory. By the fourth and fifth ten minutes periods very few nails are placed

because the boundary of the territory has already been fairly accurately marked. Throughout

this process nails placed during the first few observations periods may be moved to more

accurately delineate the boundary of the territory.

It is often useful to know the sizes of the focal pair of obligate coral feeding

Butterflyfish one is observing. Since the method is unobtrusive, we do not catch the fish

to measure them. Rather when the fish stops to feed, you note the spot behind its snout

and the tip of its tail on the substratum. Then when the fish swims away you measure this

distance with the ruler glued to the side of your clipboard and note it on the data sheet.

This can be done while you are marking the territory and observing chasing behavior and
22

also when you are recording feeding behavior. With a little practice this method of estimating fish length is quite accurate. Nevertheless, for the sake of accuracy it is good to

take at least five to six measurements and average them. It is useful to try to measure the

lengths of both members of the pairs even though we know that the sizes of the two fish of

a pair are about the same.

Another method of estimating fish sizes underwater without catching them is to practice observing fish shapes of known size at different distances underwater (Bell et al., 1985).

This method is useful where the diver does not have time to observe a fish for a long period of

time, for example when counting fish along a belt transect census or when counting fish using

the stationary census method.

The first method described above is recommended when using the indicator species

monitoring method.

At the completion of this dive, the boundaries of the territory have been marked by the

color tagged nails and agonistic chasing behavior recorded on the data sheet. The rate of

chasing behavior should be higher where the reef is undergoing change. The next step is to

measure feeding behavior.
23

Figure 4. This is an example of the Chase Data sheet being used in Hawaii.

OBSERVER:, __

DATE:. _

SITE: _

TIME: _ DEPTH:. _

time period

0-10 min.

10·20 min.

20·30 min.

30-40 min.

40·50 min.

hase Data: Record each event when the focal pair:

• chases another fish .- Agg Iso· name)

• is chased b another fish .• Sub _(sp. name)

chases.

Co mu/ticinctus C. OTnlIlissimus C. tri!asciatus C. unimacu/atus C. fremblii P. johnslonianus
24

G. Measuring Feeding Behavior and Measuring the Size of Territory

These two activities can usually done in a single dive using the data sheets shown in

Figures 6 and 7. The two activities are depicted in Figure 5. Unlike marking the territory

boundary and observing chasing behavior which are done simultanously over the entire dive,

measuring feeding behavior and measuring the territory size are done sequentially. First

feeding behavior is observed for three 10-minute observation periods, then the territory size is

measured during the remainder of the dive or on a subsequent dive. Please note: Observations

of feeding behavior will require very focused attention by the data collector.

Figure 5. Observations are made on territory size, chasing and feeding behavior. The two divers on the

left are measuring a territory. The diver at the right is observing the focal pair of butterflyfish in the

territory.
25

1. Measuring Feeding Behavior

The data collector should look for any individual identifying marks on one fish of the

focal pair. Variations in the marking pattern of the fish enable an experienced diver to recognize individual fish in the pairs, and this fish is then used to measure feeding behavior. The

Feeding Data sheet (Fig. 6) shows three 10-minute time periods in which the number of bites

on a particular species of coral is recorded. At the conclusion of the dive the numbers of bites

per 10 minutes for each species of coral can be calculated. This gives an indication of which

coral is preferred as food, but one must take into consideration the relative abundance of the

corals (Fig. 3). A species of coral which is preferred as food may be rare in the area of reef

and so the rate at which it is fed on may be relatively low. Feeding bites may occur as a single

bite and be designated in the proper space as “1”, or they may occur in bouts of two or more

bites and be designated as “2”, “5”, etc. Since feeding bites usually occur in bouts of two or

three bites with often a brief pause in between each bite, a typical data entry may look like “| |

|” indicating three bites on the species of coral. Sometimes bites may be grouped more closely

in rapid sequence. The observer counts five bites and then records “5” in the appropriate

square on the data sheet.

On the left side of the data sheet are shown the six most common species of corals in

our study sites at Kaho’olawe and Molokini Islands in Hawaii. Note that a sketch of the coral

to help an inexperienced diver identify it is shown together with its common and scientific

name. Please note that at the bottom there is space for bites taken on “other coral” and on

“Non-coral” substrata.

It must be emphasized that the scientific names of the corals are not necessary to be

known for the successful application of the indicator species method described in this manual.

It is extremely important, however, that descriptive names are given to the most common

species of corals in the study area and those which are fed on by the species of butterflyfishes

being used as indicators. These names and perhaps sketches should be used on the data sheet.

They must be used consistently by all data collectors. Consistency is the key to the data

quality control required for successful application of the method. The reason that scientific

names of the corals are not important is that we are looking for changes in feeding rates as an

indication of changing conditions on the reef. Therefore, it is important to know that the

indicator species of fish is feeding more or less on the “brown lumpy” coral. It is not necessary to know the scientific name of the coral. Please note: With an accurate and thorough

description of the physical appearance of the coral, scientific names to at least the genus level

can usually be determined at a later date.
26

Figure 6. This is an example of the Feeding Data sheet being used in Hawaii. Sample data entries are shown

for the first 0-10 min. time period.

OBSERVER: _

DATE: _

SITE: _

TIME: _ DEPTH: _

IFeeding Data: Record no. at bites taken (per 10 min. period) on each spp. of coral by one focal fish

0-10 min 10·20 min. 20-30 min

//312

Porites compressa

Finger Coral

i 1.5'11

. )-~

Porites /obata

Lobe Coral

1/£2.//3 J.' ;'

. oi: .

. ;. ~" . .':

:- . . Momipora verrucosa

Brown olate coral

.,~~ /112.3/

.~.;-~~ .~rl!N!:'~,

Montipora patula

Crown calvx coral

///25/~

*

..

Pocillopora meandrina

Caulinower coral

•

/1

Pavolla varians

Meanderinll oolvn coral

OTHER CORAL 1/

NON-CORAL I
27

2. Measuring the Territory Size

The boundary of the territory is marked with color-tagged nails. The shape of territories is usually more or less circular or oval, and the longest axis across the territory is evident

to the diver (Fig. 5). It is possible for one diver to measure a territory, but it is much easier and

faster if the diver’s buddy or assistant helps to measure the territory. First, a fiberglass measuring tape2

 is stretched across the longest axis of the territory and anchored at both ends with a

small weight. Then, while one diver holds one end of the second tape at each meter interval

along the first tape, the second diver measures the distance to the boundary of the territory and

enters the distance on the data sheet (Fig. 7). Note that in Figure 7 the line down the center

represents the tape measured line down the longest dimension of the territory (Fig. 5). The

length of the line is indicated at the appropriate meter mark, in this example it is 15 meters.

Then the distance to the boundary of the territory is measured at each meter to the right and

left sides of the line, and the distances are recorded to the right and left sides of the line on the

data sheet. The territory size (in square meters) is estimated by simply adding up all the values

on the data sheet. Sample entries in Figure 7 are shown for the first five meters on the right

side. The last data entry on the right side would be at 15 meters. The process is then repeated

for the left side of the territory.

As with all aspects of this and other activities described in this manual, every effort

should be made to avoid contact with the coral during the measurement. This is best done if

the divers are slightly positively buoyant.

2 Fiberglass Measuring Tapes are available from companies such as Ben Meadows Co., P.O.Box 80549,

Chamblee, GA 30366, U.S.A., Fax: U.S. and Canada 1-800-628-2068, International 1-770-457-1841, or Forestry

Supplies, Inc., P.O.Box 8397, Jackson, MS 39284, U.S.A., Fax: U.S. and Canada 1-800-543-4203, International

1-601-355-5126. They are available in both English and Metric units of measure, in varying lengths. Two

excellent brand names are Keson and Lufkin. For our research we prefer the 30m open reel model. Current 1996

prices are about U.S. $ 25-30.
28

Figure 7. This is an example of the Measuring Territory Data sheet being used in Hawaii.

OBSERVER: _

DATE: _

SITE: _

TIME: _ DEPTH: _

Territory Measurement: Measure and record the distance from the center line

to the territorial boundary at 1 m intervals

o -a5

-+------/.0

-+-------/.3

---------·2./

-+-------1,.3

5 -r~-------I.g

10

15

20
29

H. Picking-up the Color-tagged Nails, the Fiberglass Measuring Tapes, and the Polypropylene Transect Lines

After the final territory measurements have been collected, the color-tagged nails and

the fiberglass measuring tapes are removed from the reef. The polypropylene transect lines

can be left in place for several days until the work is completed in the study area. Then they

too are removed from the reef. At the conclusion of the study, nothing should be left on the

reef in order to leave the area as undisturbed as when the study began. Providing that the fish

are not collected by spear or hand nets, they habituate to the presence of a diver by the end of

the first dive. By the third or fourth dive the focal pairs have become so tame that a diver may

approach within a meter before their normal feeding/foraging behavior is affected. In order to

ensure that monitoring can continue at the same site for many months and years, it is advisable

to record accurate site coordinates, note any shoreside landmarks, and/or leave a buoy marker

(sub-surface or surface) as the particular situation allows. Please note: Ensure that any buoy

markers deployed meet all federal, state and local regulations, and are as environmentally

"friendly" as possible.

III. ANALYSIS OF DATA

A. The Design of the Monitoring Program

The correct analysis and interpretation of data depends on an understanding of the

design of the monitoring program, and the design is based on the question being asked. For

this monitoring approach we are relying on the behavioral ecology of the coral feeding

butterflyfish to provide the answer to the question: Is the condition of the coral reef changing?

The design of the monitoring program must include at least two study sites. One site

should have no (or limited and known) environmental perturbations or chronic anthropogenic

disturbances. Little or no change in the behavioral ecology of the indicator species of

butterflyfishes is postulated for this site which can then serve as a control or reference reef.

Additional sites are selected due to concern that disturbance is either occurring or anticipated.

Here the hypothesis is that changes in the behavioral ecology of the indicator species will

provide an “early warning” that sub-lethal and/or potentially lethal change is occurring on the

reef, providing time for remedial action to be taken before the corals on the reef become
30

moribund and die. Although comparisons between the two sites for any sampling period may

be interesting, it is the comparison of the data at the same site over time which is essential.

B. Step-wise Use of the Monitoring Program

The Indicator Species Monitoring approach has the advantage of providing useful

information in a step-wise or incremental fashion. For example, in its simplest form or first

step, the numbers of coral feeding butterflyfishes and especially the species selected as the

indicator species are counted along the 30 meter Belt Transect (see Section II. D. above).

Since two important criteria for the indicator species are (1) being long-lived and (2) being

strongly site attached, home ranging or territorial, then within-site changes in their abundance

and distribution suggest that changes are occurring on the reef where disturbance is anticipated. This evidence of change is strengthened if similar within-site changes do not occur on

the control or reference reef.

The next incremental step is the estimation of coral cover along the transect (see

Section II. E. above). Again the within-site comparisons are important and the reasoning is the

same as for the abundance and distribution of fishes with one very important difference: the

fish are mobile and are able to leave the area as conditions deteriorate whereas the corals are

attached and therefore are unable to avoid the changing conditions of the water around the

reef. They must tolerate and adapt to the conditions or become moribund and die. Needless to

say, when dead corals begin to appear and the estimation of coral cover decreases, evidence

that the reef is being impacted by detrimental environmental factors is strong. Unfortunately it

may be too late to enact remedial management policies at this stage of manifestation to lethal

perturbations.

It is important to note that the overall abundance of corals, as estimated by the percent

coral cover, may not change, but the distribution of the different species, the composition of

the coral community may change. This is because the different species of corals compete with

one another for space on the reef. Alterations in environmental conditions may alter competitive advantages between corals. For example, in Hawaii corals of the genus Montipora are

able to kill and grow over corals of the genus Porites, but the presence of coral feeding

butterflyfishes, especially Chaetodon unimaculatus, which selectively feeds on Montipora

corals prevents this from happening (Cox, 1986). Nevertheless, if adverse conditions affect
31

C. Data Management

All too often environmental monitoring (especially volunteer efforts at local levels)

ends with the collection of the raw data. This is unfortunate because without careful data entry

and statistical analysis the significance of the results cannot be determined. Without a careful

the reef, and the numbers of coral feeding butterflyfishes decline, then changes in the species

composition of the corals on the reef may occur. These changes occur slowly, yet are within

the time-frame of a long-term monitoring program of a few years duration.

The next steps in the monitoring program involve the behavioral observations of

agonistic behavior, as indicated by aggressive behavior, and territory size (see II. F. above) and

feeding behavior (II. G. above). Within-site changes in any of these measure provide evidence

that change is occurring on the reef. For example if low level chronic inputs of pollutants are

gradually causing the healthy conditions of the coral on the reef to deteriorate, the food

"value" of the coral to the fishes may decrease as discussed earlier.

Under these conditions, we expect the following changes in behavior to occur:

1. The amount of agonistic encounters (chases), will increase as fish try to find more food in

their neighbor’s territories.

2. Territory sizes will change. Fish try to enlarge their territories to gain access to more food.

This may be possible for larger, dominant pairs at the expense of smaller, more submissive

pairs whose territories may become reduced in size. Subordinate fish with small, marginal

territories may be forced to leave. (These possibilities still need to be verified experimentally.) Thus, as the reef changes, territory sizes are expected to change as well.

3. Feeding rates will change. At first, as the nutrient food value of the corals decreases, feeding

rates will increase as the fish must feed on more coral polyps to gain the same amount of

nutrients and energy for metabolism, growth and reproduction. Later, as chasing behavior

increases, there may be less time for undisturbed feeding and feeding rates may decline.

Thus there is a dynamic relationship between the food value of the corals and the chasing,

territorial and feeding behavior of the fish.

Clearly, the more that is known about each of these behavioral steps, the greater the

sensitivity and predictive use of the monitoring method will be. Nevertheless, each suceeding

step of the method provides additional useful information.
32

A clear need exists for biological surveys of benthic, coral, and fish communities to

collect the raw data that can be converted into some form of condition indices. Ideally these

indices would be indicative of levels of stress or change before the “point of no return” so that

appropriate research can be initiated to provide information to the management community

responsible for mitigation strategies. There have been numerous calls (see Crosby et al., 1996;

Eichbaum et al., 1996) for a network of “index” sites for long-term monitoring and assessment at the national, regional, and global levels. The current dearth of individuals with high

level technical expertise, not to mention funding resources, precludes the implementation of

such a comprehensive network of coral reef monitoring sites (especially in remote regions)

without the significant involvement of adequately trained, local-level participants. Many nongovernmental groups have been at the forefront of various local coral reef monitoring efforts.

These grassroots efforts should be applauded and encouraged. However, long-term national

and globally coordinated coral reef monitoring programs are essential to manage, archive,

translate, and transfer data to scientists, managers, and other interest groups. NOAA is developing a nationally coordinated coral reef monitoring program to be implemented in 1996 and

actively pursuing partnership efforts with other agencies (such as the National Park Service

and the Environmental Protection Agency) and volunteer interest groups (such as American

Oceans, The Nature Conservancy, REEF, and Reefkeeper). The key to successful monitoring

programs, whether professions or volunteer based, is good data management.

From Data Collection to Long-term Data Management

It is very important that the data observations collected in the field on the underwater

analysis of results, it would not be prudent to advocate remedial action. The United Nations

Conference on Environment and Development (UNCED) in Rio De Janeiro in 1992 and

Agenda 21 brought worldwide attention to the need for monitoring and protection of coral reef

ecosystems. Globally, scientists are now working together and with other groups to promote

monitoring of coral reefs. For example, a Global Task Team on Climate Change and the

International Oceanographic Commission (IOC) have also published recommendations on

coral reef monitoring procedures and are organizing a global network for monitoring coral

reefs. A major activity in support of the International Coral Reef Initiative is the Global Coral

Reef Monitoring Network. This activity will involve coral reef researchers around the world

in coordinated monitoring of selected coral reefs.
33

data sheets be transferred as soon as possible to long-term data storage. The preferred method

is for all data observations to be entered onto computerized data spread sheets. At the very

least the field observations should be transferred to hard copy paper data spread sheets. The

advantage to storing your data on a computer format is that data manipulations, transfers of

data to central data bases such as those discussed above, and statistical analyses are much

easier than with hard copy paper storage. Whether one uses IBM based or Macintosh based

computers, there are numerous spreadsheet and statistical analyses packages available. The

pros and cons of these computer software packages are numerous and often subjective based

on individual preferences.

The following procedures should be followed to maintain quality control of your data files:

• Ensure that column headings in your data spread sheets are clearly and consistently labeled.

• Keep accurate descriptions of any abbreviations used in the actual data set and what units (feet,

meters, minutes, hours, etc.) the data are expressed as.

• Keep accurate and full descriptions of the how, who, when, and where each datum was collected.

• Routinely check and double check (it is best if you have someone else double check your data for

you) to ensure that the data you have entered in the long-term data file accurately reflects the data

observations in the field.

• Maintain at least one complete set of “backup” data files on floppy disks and/or other hard drives,

and update these backups periodically.

Please Note: While it is often not practical to maintain the full and complete descriptions as part of

the actual data files, always make sure to reference where the complete “meta-data” files exist.

Examples of data spread sheets that can be used with the Indicator Species Monitoring

approach described in this manual are in Figures 8-11. These data sheets were created using

the Microsoft Excel program. Figure 8 may be used as a format to record the observations

from the fish transect method. [“Trip” refers to the sequential number of field trips in this

study. “Site” refers to one of three sites being examined as part of this study. “PI” refers to the

individual who collected this data. “Station” refers to one of a number of different specific
34

transects conducted by a particular PI at a given site during a specific trip. “Date” is the

specific calendar point in time that the data was collected. “C. multi” refers to the number of

C. multicinctus observed by the PI on this particular transect line. “C. multi juv” refers to

juvenile C. multicinctus. The next five column headings are abbreviations for other species of

butterfly fishes observed on the transect line. The final column is a summation of all butterfly

fishes observed.]

Figure 9 may be used as a format to record the observations from the coral transect

method. [The first five column headings are the same as in Figure 8. The next seven column

headings are abbreviations for different coral species found in this study. “Unknown” refers to

any coral species that can not be identified to species or genus. “Noncoral” refers to an observation point on the transect line that has neither living nor dead coral. “Deadcoral” is self

explanatory. “∑ counts” refers to the total number of observation points along the transect

line. “∑ live coral” refers to the total number of live coral occurring at observation points

along the transect line. “% coverage” refers to the percent of observation points along the

transect line that had living coral.]

Figure 10 may be used as a format to record the observations from the butterfly fish

chasing method. The first five column headings are the same as in Figure 8. “Time” data are

presented sequentially in each of the 10 minute time periods in which they occurred (i.e., 2

refers to the second 10 minute time period), and for the total 50 minute period of observation.

The next 12 columns represent different fish species that the focal pair of fished had agonistic

encounters with, broken down as whether the focal pair was the aggressor (agg) or submissive

(sub) to the species noted in the column heading. The sum of both aggressive and submissive

encounters with the species noted in the column heading are given under the “∑” headings.

The “agg?”, “sub?’ and “∑?” headings refer to encounters the focal pair had with a species that

the data collector could not identify. The “∑agg” represents the sum of all encounters that the

focal pair had with other fished in which the focal pair was the aggressor. “∑sub” is likewise

for all encounters in which the focal pair was submissive, and “∑∑” represents the sum of all

encounters that the focal pair had with all other fishes. “Terr sqM” refers to the territory size

in square meters, of the focal pair, “hour” refers to the time of day (rounded to the nearest 15

minutes) that the observations began, and “fish cm” refers to the size estimate for one individual of the focal pair. (As an example - during the sixth trip of this study, Javier Mendez

made an observation at station 1 of site 2 on June 3, 1995 that during the first 10 minute period
35

of observing his focal pair of fish, an individual in his focal pair made two aggressive chases

against a C. multicinctus. In the fifth 10 minute period of this total 50 minute observation

period, Javier noted that an individual in his focal pair again made an aggressive chase against

a C. multicinctus. No other encounters occurred with his focal pair during this 50 minute

observation period. The territory size of the focal pair that he observed during this period was

53.6 m2, and the size of one individual in his focal pair was 8.0 cm.)

Figure 11 may be used as a format to record the observations from the butterflyfish

feeding method. The first five column headings are the same as in Figure 8. “Min” refers to

each of the 10 minute time periods (i.e., 2 refers to the second 10 minute time period) during

the total 30 minute period of observation. The next seven columns represent the number of

bites made on different species of coral that an individual of the focal pair of fish fed upon.

“Unknown” refers to bites that were made on an unknown species of coral, and “noncoral”

refers to bites made upon a substrate other than living coral. “∑coral” refers to the sum of all

bites on coral during the given time period, and “∑bites” refers to all bites regardless of

substrate. “Hour” refers to the time of day (to the nearest 15 minutes) that the observation

period began.

If one desires, conventional statistical methods may be used to analyze changes over

time and differences between sites (see Sokal and Rohlf, 1981 and Mendenhall, 1971 for

complete overview of statistical analyses of data sets). In the ecological sciences, a probability level >0.05 is generally used for rejection of null hypotheses. In laymen’s terms this means

that in order to make the conclusion that observations between two or more times or sites are

“significantly” different, analyses must indicate that there is a 95% probability that there is a

difference. Just as there are numerous data spread sheet software packages available commercially for both IBM and Macintosh computers, there are also a number of easy to use statistical

software packages that one can employ to do a range of elementary descriptive and more

sophisticated statistical analyses of your data sets. Any of a number of paired-comparison

tests can be used to detect trends of directional changes in measurements, an Electivity Index

(Gore, 1984) is used to compare feeding preferences to coral abundance, and a number of

different parametric and non-parametric approaches can be used to determine differences

between two or more sites or events in time and over time. Please note: It is highly advisable

to consult with an individual who is experienced with statistical analyses and discuss the

questions that you are trying to answer in order to design the appropriate data formats and

conduct the appropriate statistical analyses.
36

TRIP SITE P I STATION DATE C. multi C. multi juv. C orn. C. unimac C. kleinii C. trifas C. miliaris total fish

5 1 JM 1 3/15/95 16 0 0 10 6 1 0 33

5 2 JM 1 3/15/95 14 6 0 0 0 0 02 05 3 JM 1 3/18/95 8 0 0 7 5 4 02 45 3 JM 2 3/18/95 6 0 2 7 2 2 01 95 1 ESR 1 3 / 1 5 / 9 5 8 0 0 4 4 6 02 25 2 ESR 1 3 / 1 5 / 9 5 2 0 0 0 0 0 0 02 05 3 ESR 1 3 / 1 8 / 9 5 1 2 0 0 8 3 2 12 65 3 ESR 2 3 / 1 8 / 9 5 8 0 1 17 12 8 0 465 1 MPC 1 3 / 1 5 / 9 5 1 6 0 4 6 0 2 02 85 2 MPC 1 3 / 1 5 / 9 5 1 8 0 0 0 3 0 02 15 3 MPC 1 3 / 1 8 / 9 5 2 0 0 4 1 4 01 15 3 MPC 2 3 / 1 8 / 9 5 4 0 0 6 2 1 11 46 1 JM 1 6/3/95 10 1 1 3 1 2 01 86 2 JM 1 6/3/95 16 7 0 0 0 0 02 36 1 JM 2 6/4/95 8 1 1 9 0 2 02 16 3 JM 1 6/6/95 10 0 0 25 14 0 0 497 2 JM 1 11/17/95 10 2 4 3 1 0 0 03 87 1 JM 1 11/18/95 13 8 2 9 0 2 03 4

Figure 8. Fish Transect Data
37

TRIP SITE P I STATION DATE P.compressa P.lobata M.patula M.verrucosa Poc.meand Pav.varians P.deur unknown noncoral deadcoral ∑counts ∑livecoral %coverage

5 1 ESR 1 3/15/95 39 1 3 5 3 1 0 0 8 0 60 52 86.675 2 ESR 1 3/15/95 32 2 2 1 2 0 0 0 21 0 60 39 65.005 3 ESR 1 3/18/95 2 1 28 2 7 1 1 0 0 0 0 60 60 100.00

5 3 ESR 2 3/18/95 1 5 31 2 1 1 1 0 0 0 0 60 60 100.00

5 1 JM 1 3/15/95 21 9 16 6 2 2 0 0 0 0 56 56 100.00

5 2 JM 1 3/15/95 21 10 12 5 6 0 0 0 0 6 60 54 90.005 3 JM 1 3/18/95 3 5 20 2 3 4 3 0 0 0 2 60 58 96.675 3 JM 2 3/18/95 7 11 26 1 1 1 0 0 0 1 3 60 56 93.335 1 MPC 1 3 / 1 5 / 9 5 29 2 5 4 0 1 0 0 2 11 54 41 75.935 2 MPC 1 3 / 1 5 / 9 5 25 6 4 2 1 1 0 0 3 17 59 39 66.105 3 MPC 1 3 / 1 8 / 9 5 4 12 20 1 1 2 0 1 0 3 2 55 50 90.915 3 MPC 2 3 / 1 8 / 9 5 10 10 24 6 2 0 0 0 2 4 58 52 89.666 1 JM 1 6/3/95 24 2 7 7 7 0 0 0 0 13 60 47 78.336 1 JM 2 6/4/95 14 7 25 5 1 2 0 0 0 6 60 54 90.006 2 JM 1 6/3/95 28 15 9 2 3 0 0 0 0 3 60 57 95.006 3 JM 1 6/6/95 2 6 26 2 3 2 1 0 0 0 0 60 60 100.00

6 1 MPC 1 6 / 3 / 9 5 26 2 19 5 0 2 0 0 0 6 60 54 90.006 1 MPC 2 6 / 4 / 9 5 26 3 19 7 1 1 0 0 0 3 60 57 95.006 2 MPC 1 6 / 3 / 9 5 23 4 10 4 3 2 0 1 0 13 60 46 76.676 3 MPC 1 6 / 6 / 9 5 9 12 17 1 0 4 1 0 0 3 3 59 53 89.837 1 JM 1 11/18/95 19 8 23 3 4 0 0 1 0 2 60 57 95.00

7 2 JM 1 11/17/95 20 7 11 5 6 5 0 0 0 6 60 54 90.00

7 1 MPC 1 1 1 / 1 8 / 9 5 27 0 18 6 1 0 0 0 0 8 60 52 86.677 2 MPC 1 1 1 / 1 7 / 9 5 22 8 8 2 3 1 0 0 0 15 59 44 74.58

Figure 9. Coral Transect Data
38

TRIP SITE P I STATION DATE TIME aggCm subCm ∑Cm aggCo subCo ∑Co aggCmil subCmil ∑Cmil aggPj subPj ∑ P j aggCs subCs ∑Cs aggCt subCt ∑Ct aggCh subCh ∑Ch aggAn subAn ∑An agg? sub? ∑ ? ∑agg ∑sub ∑∑ terr sqM hour fish cm

6 1 JM 1 6/3/95 1 0 0 0 0 0 0 0 0 0 0 4 4 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 4 4 82.5 1000 9.0

6 1 JM 1 6/3/95 2 0 0 0 0 0 0 0 0 0 0 3 3 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 3 3 82.5 1000 9.0

6 1 JM 1 6/3/95 3 0 0 0 0 0 0 0 0 0 0 3 3 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 3 3 82.5 1000 9.0

6 1 JM 1 6/3/95 4 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 2 2 82.5 1000 9.0

6 1 JM 1 6/3/95 5 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 2 2 82.5 1000 9.0

6 1 JM 1 6/3/95 50 0 0 0 0 0 0 0 0 0 0 14 14 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 14 14 82.5 1000 9.0

6 1 JM 2 6/4/95 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 1 1 142.8 1000 10.0

6 1 JM 2 6/4/95 2 0 0 0 0 0 0 0 0 0 0 3 3 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 3 3 142.8 1000 10.0

6 1 JM 2 6/4/95 3 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 1 1 142.8 1000 10.0

6 1 JM 2 6/4/95 4 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 2 0 2 0 0 0 0 0 0 0 00 2 1 3 142.8 1000 10.0

6 1 JM 2 6/4/95 5 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 1 1 142.8 1000 10.0

6 1 JM 2 6/4/95 50 0 0 0 0 0 0 0 0 0 0 7 7 0 0 0 2 0 2 0 0 0 0 0 0 0 00 2 7 9 142.8 1000 10.0

6 2 JM 1 6/3/95 1 2 0 2 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 2 1 3 53.6 1300 8.0

6 2 JM 1 6/3/95 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 53.6 1300 8.0

6 2 JM 1 6/3/95 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 53.6 1300 8.0

6 2 JM 1 6/3/95 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 53.6 1300 8.0

6 2 JM 1 6/3/95 5 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 0 1 53.6 1300 8.0

6 2 JM 1 6/3/95 50 3 0 3 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 3 1 4 53.6 1300 8.0

6 2 JM 2 6/5/95 1 1 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 1 2 48.9 1030 6.5

6 2 JM 2 6/5/95 2 1 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 1 2 48.9 1030 6.5

6 2 JM 2 6/5/95 3 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 0 1 48.9 1030 6.5

6 2 JM 2 6/5/95 4 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 1 1 48.9 1030 6.5

6 2 JM 2 6/5/95 5 1 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 1 2 48.9 1030 6.5

6 2 JM 2 6/5/95 50 4 0 4 0 0 0 0 0 0 0 4 4 0 0 0 0 0 0 0 0 0 0 0 0 0 00 4 4 8 48.9 1030 6.5

6 3 JM 1 6/6/95 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 133.0 930 10.06 3 JM 1 6/6/95 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 133.0 930 10.06 3 JM 1 6/6/95 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 133.0 930 10.06 3 JM 1 6/6/95 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 133.0 930 10.06 3 JM 1 6/6/95 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 133.0 930 10.06 3 JM 1 6/6/95 50 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 133.0 930 10.06 3 JM 2 6/7/95 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 122.3 930 10.06 3 JM 2 6/7/95 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 122.3 930 10.06 3 JM 2 6/7/95 3 0 0 0 0 0 0 0 0 0 0 3 3 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 3 3 122.3 930 10.06 3 JM 2 6/7/95 4 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 2 2 122.3 930 10.06 3 JM 2 6/7/95 5 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 11 0 2 2 122.3 930 10.06 3 JM 2 6/7/95 50 0 0 0 0 0 0 0 0 0 0 6 6 0 0 0 0 0 0 0 0 0 0 0 0 0 11 0 7 7 122.3 930 10.06 1 MPC 1 6 / 3 / 9 5 1 5 0 5 0 0 0 0 0 0 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 00 6 1 7 59.5 1000 10.0

6 1 MPC 1 6 / 3 / 9 5 2 5 0 5 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 5 2 7 59.5 1000 10.0

6 1 MPC 1 6 / 3 / 9 5 3 4 0 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 4 0 4 59.5 1000 10.0

6 1 MPC 1 6 / 3 / 9 5 4 2 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 2 0 2 59.5 1000 10.0

6 1 MPC 1 6 / 3 / 9 5 5 4 0 4 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 4 1 5 59.5 1000 10.0

6 1 MPC 1 6 / 3 / 9 5 5 0 2 0 0 2 0 0 0 0 0 0 0 0 4 4 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 21 4 25 59.5 1000 10.0

6 1 MPC 2 6 / 4 / 9 5 1 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 2 2 66.9 1000 8.0

6 1 MPC 2 6 / 4 / 9 5 2 1 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 11 1 2 3 66.9 1000 8.0

6 1 MPC 2 6 / 4 / 9 5 3 2 0 2 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 2 1 3 66.9 1000 8.0

6 1 MPC 2 6 / 4 / 9 5 4 2 2 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 2 2 4 66.9 1000 8.0

6 1 MPC 2 6 / 4 / 9 5 5 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 2 2 66.9 1000 8.0

6 1 MPC 2 6 / 4 / 9 5 5 0 5 3 8 0 0 0 0 0 0 0 5 5 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 5 9 14 66.9 1000 8.0

6 2 MPC 1 6 / 3 / 9 5 1 2 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 2 0 2 46.3 1300 7.0

6 2 MPC 1 6 / 3 / 9 5 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 46.3 1300 7.0

6 2 MPC 1 6 / 3 / 9 5 3 2 0 2 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 2 2 4 46.3 1300 7.0

6 2 MPC 1 6 / 3 / 9 5 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 46.3 1300 7.0

6 2 MPC 1 6 / 3 / 9 5 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 46.3 1300 7.0

6 2 MPC 1 6 / 3 / 9 5 5 0 4 0 4 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 4 2 6 46.3 1300 7.0

6 2 MPC 2 6 / 5 / 9 5 1 5 0 5 0 0 0 0 0 0 1 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 6 1 7 40.7 1000 8.0

6 2 MPC 2 6 / 5 / 9 5 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 40.7 1000 8.0

6 2 MPC 2 6 / 5 / 9 5 3 5 1 6 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 00 5 3 8 40.7 1000 8.0

6 2 MPC 2 6 / 5 / 9 5 4 3 0 3 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 3 1 4 40.7 1000 8.0

6 2 MPC 2 6 / 5 / 9 5 5 1 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 1 2 40.7 1000 8.0

6 2 MPC 2 6 / 5 / 9 5 5 0 1 4 1 1 5 0 0 0 0 0 0 1 5 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 15 6 21 40.7 1000 8.0

6 3 MPC 1 6 / 6 / 9 5 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 117.4 930 10.06 3 MPC 1 6 / 6 / 9 5 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 117.4 930 10.06 3 MPC 1 6 / 6 / 9 5 3 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 1 1 117.4 930 10.06 3 MPC 1 6 / 6 / 9 5 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 117.4 930 10.06 3 MPC 1 6 / 6 / 9 5 5 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 0 1 117.4 930 10.06 3 MPC 1 6 / 6 / 9 5 5 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 00 1 1 2 117.4 930 10.0

Figure 10. Fish Chase Data
39

TRIP SITE P I STATION DATE M I N P.compressa P.lobata M.patula M.verrucosa Poc.meand Pav.varians P.deur unknown noncoral ∑coral ∑bites hour

5 3 JM 1 3/18/95 1 1 4 146 28 1 2 0 0 0 182 182 1215

5 3 JM 1 3/18/95 2 0 4 108 11 0 0 0 0 0 123 123 1215

5 3 JM 1 3/18/95 3 0 4 192 16 0 0 0 1 0 213 213 1215

5 3 JM 2 3/19/95 1 3 0 86 34 1 5 0 0 0 129 129 8305 3 JM 2 3/19/95 2 6 4 46 36 5 0 0 0 0 97 97 830

5 3 JM 2 3/19/95 3 1 4 99 47 3 0 0 0 0 154 154 8305 1 MPC 1 3 / 1 6 / 9 5 1 21 4 31 11 11 5 0 0 0 83 83 1030

5 1 MPC 1 3 / 1 6 / 9 5 2 19 3 45 18 5 3 0 0 0 93 93 1030

5 1 MPC 1 3 / 1 6 / 9 5 3 17 0 59 12 3 1 0 0 0 92 92 1030

5 1 MPC 2 3 / 1 6 / 9 5 1 8 0 46 13 28 0 0 0 0 95 95 1500

5 1 MPC 2 3 / 1 6 / 9 5 2 12 16 38 26 15 2 0 0 1 109 110 15005 1 MPC 2 3 / 1 6 / 9 5 3 12 0 31 9 19 0 0 0 1 71 72 1500

5 2 MPC 1 3 / 1 7 / 9 5 1 49 3 23 4 14 1 0 0 2 94 96 1030

5 2 MPC 1 3 / 1 7 / 9 5 2 23 10 29 5 17 2 0 0 1 86 87 1030

5 2 MPC 1 3 / 1 7 / 9 5 3 30 9 45 6 11 0 0 0 1 101 102 1030

5 2 MPC 2 3 / 1 7 / 9 5 1 37 14 52 10 14 7 0 0 4 134 138 1430

5 2 MPC 2 3 / 1 7 / 9 5 2 36 2 34 4 8 0 0 0 8 84 92 1430

5 2 MPC 2 3 / 1 7 / 9 5 3 40 2 25 5 18 0 0 0 5 90 95 1430

5 3 MPC 1 3 / 1 8 / 9 5 1 12 38 33 18 3 0 0 0 0 104 104 1200

5 3 MPC 1 3 / 1 8 / 9 5 2 14 41 5 16 0 0 0 0 0 76 76 1200

5 3 MPC 1 3 / 1 8 / 9 5 3 8 54 6 23 3 0 0 0 0 94 94 1200

5 3 MPC 2 3 / 1 9 / 9 5 1 13 40 25 39 0 2 0 0 0 119 119 930

5 3 MPC 2 3 / 1 9 / 9 5 2 15 61 1 38 2 0 0 0 0 117 117 9305 3 MPC 2 3 / 1 9 / 9 5 3 1 41 27 29 8 8 0 0 0 114 114 930

Figure 11. Fish Feeding Data
40

APPENDIX 1

Possible Indicator Species of Butterflyfishes from Different Indo-Pacific Regions

The pictures on pages 40 and 41 are examples of possible indicator species of

butterflyfishes from different Indo-Pacific regions. The sizes indicated below the pictures

represent the total length of mature fish from a number of resources.

A convenient source for excellent pictures of all these species, excepting Chaetodon

multicinctus, which only occurs in Hawaii, is Robert Myers’ Micronesian Reef Fishes, or John

Randall, Gerald Allen and Roger Steene’s Fishes of the Great Barrier Reef and Coral Sea,

Gerald Allen and Roger Swainston's Reef Fishes of New Guinea. For Hawaii, see Randall’s

Guide to Hawaiian Reef Fishes and John Hoover's Hawaii's Fishes. See the literature cited

for complete references.

The species marked with an asterisk (*) are particularly promising indicator species.

* Chaetodon austriacus occurs only in the Red Sea. It is replaced by C. melapterus in the

Arabian Gulf.

* Chaetodon baronessa occurs from Cocos-Keeling Island in the Indian Ocean, east to Fiji,

north to Japan, and south to the Great Barrier Reef. It is replaced by C. triangulum in the rest

of the Indian Ocean.

* Chaetodon larvatus occurs in the Red Sea. It appears to be ecologically equivalent to C.

triangulum in the Indian Ocean and C. baronessa in the Pacific Ocean.

Chaetodon melannotus occurs from East Africa and the Red Sea, east to Samoa, north to

Japan, and south to the Great Barrier Reef. This species specializes in feeding on soft corals as

well as hard corals.

* Chaetodon melapterus occurs in the Arabian Gulf but is rare in the Red Sea. Ecologically it

is similar to C. austriacus in the Red Sea, and C. trifasciatus in the rest of the Indo-Pacific.

* Chaetodon meyeri occurs from East Africa across the Indian Ocean extending eastward to
41

the Line Islands. It occurs widely in Micronesia and extends south to the Great Barrier Reef

and north to Japan.

* Chaetodon multicinctus occurs throughout the Hawaiian Archipelago and Johnston Atoll.

* Chaetodon ornatissimus occurs from Sri Lanka in the Indian Ocean eastward to Hawaii,

Tahiti and Polynesia, north to Japan, and south to the Great Barrier Reef.

Chaetodon plebeius occurs from the Andaman Sea in the eastern Indian Ocean, extending

eastward to Fiji, north to Japan, and south to the Great Barrier Reef. Apparently it is rare or

does not occur in Micronesia and Polynesia.

Chaetodon quadrimaculatus occurs throughout Micronesia and eastward to Polynesia, Tahiti

and Hawaii, and north to Japan. This species specializes in feeding on corals of the genus

Pocillopora.

Chaetodon reticulatus occurs form the Great Barrier Reef in the west, northward to Japan, and

eastward through Micronesia to Polynesia, Tahiti, and northwest to Hawaii where it is relatively rare.

* Chaetodon triangulum occurs in the Indian Ocean. Ecologically it is similar to C. baronessa

in the Pacific Ocean and C. larvatus in the Red Sea.

* Chaetodon trifascialis occurs from East Africa and the Red Sea, eastward to Tahiti and

throughout Polynesia to Johnston Atoll and the Northwest Hawaiian Islands, but not the high

islands of Hawaii. It is widespread throughout Melanesia, and Micronesia, extends southward

to the Great Barrier Reef and northward to Japan. It specializes in feeding on corals of the

genus Acropora.

* Chaetodon trifasciatus occurs from East Africa eastward in Polynesia to Hawaii and Tahiti.

It is widespread in Melanesia and Micronesia. It extends north to Japan and south to the Great

Barrier Reef. It is replaced in the Red Sea by C. austriacus, and by C. melapterus in the

Persian Gulf.
42Ornate butterflyfish, Chaetodon ornatissimus, to 7.5 in. (190 mm) Oval butterflyfish, Chaetodon trifasciatus, to 6 in. (152 mm)

 Reticulated butterflyfish, Chaetodon reticulatus, to 6.3 in. (160 mm) Brown-barred/Multiband butterflyfish, Chaetodon multicinctus, to 3.7 in. (93 mm)
43

Triangle butterflyfish, Chaetodon baronessa, to 6 in. (152 mm)

Chevron butterflyfish, Chaetodon trifascialis, to 7 in. (178 mm)

Meyer's butterflyfish, Chaetodon meyeri, to 7 in. (178 mm)

Photographs by John E. Randall, BerniceP. Bishop Museum, Honolulu, Hawaii.Following the common and scientificname of each fish, the approximatelargest total length the species attains isgiven in inches and millimeters.
44

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