University of Rhode Island

DigitalCommons@URI

Open Access Master's Theses

2018

THE IMPACT OF FISHERIES MANAGEMENT ON CORAL REEF

FISH COMMUNITIES AND LIFE-HISTORY CHARACTERISTICS
Diky Suganda
University of Rhode Island, superdiky.fisheries@gmail.com

Follow this and additional works at: https://digitalcommons.uri.edu/theses

Terms of Use
All rights reserved under copyright.

Recommended Citation
Suganda, Diky, "THE IMPACT OF FISHERIES MANAGEMENT ON CORAL REEF FISH COMMUNITIES AND
LIFE-HISTORY CHARACTERISTICS" (2018). Open Access Master's Theses. Paper 1288.
https://digitalcommons.uri.edu/theses/1288

This Thesis is brought to you for free and open access by DigitalCommons@URI. It has been accepted for inclusion
in Open Access Master's Theses by an authorized administrator of DigitalCommons@URI. For more information,
please contact digitalcommons-group@uri.edu.
 THE IMPACT OF FISHERIES MANAGEMENT ON CORAL

REEF FISH COMMUNITIES AND LIFE-HISTORY

CHARACTERISTICS

BY

DIKY SUGANDA

A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE

REQUIREMENTS FOR THE DEGREE OF

MASTER OF SCIENCE

IN

BIOLOGICAL AND ENVIRONMENTAL SCIENCES

UNIVERSITY OF RHODE ISLAND

2018
 MASTER OF SCIENCE THESIS

OF

DIKY SUGANDA

APPROVED:

Thesis Committee:

Major Professor Austin Humphries

Kathleen Castro

Jeremy Collie

Nasser H. Zawia
DEAN OF THE GRADUATE SCHOOL

UNIVERSITY OF RHODE ISLAND

2018
 ABSTRACT

Indonesia’s coral reefs support over half of all small-scale fishers globally and are thus

an important economic and sociocultural resource. These coral reef fisheries, however,

are threatened from a variety of stressors including overexploitation. No-take fisheries

closures are thought to be a suitable management strategy in Indonesia that provide a

refuge for fish species with different ecological and life-history characteristics. While

examining such indicators is increasingly important to determine management efficacy,

few have done so in Indonesia. I investigate community ecological and life-history

responses to no-take fisheries closures using abundance data from 2009-2015 in three

regions across Indonesia’s Sunda Banda Seascape. Overall, fish biomass was 30%

greater in no-take closures than fished reefs. The only functional groups to respond to

management were corallivores and detritivores, and fished reefs had greater biomasses

of these groups. No-take closures had fish communities with greater maximum lengths,

longer life spans, slower growth rates, and higher mean trophic levels. Surprisingly,

total fish biomass was not a good predictor for life history values or functional group

biomass. These results indicate that non-target fish species may thrive in fished reefs

where predators with slow life histories are reduced. Also, no-take closures in

Indonesia’s Sunda Banda Seascape are facilitating recovery of life-history

characteristics and fish biomass but these impacts are not uniformly distributed across

functional groups. My findings are of value to current ecosystem-based management

objectives attempting to achieve broader conservation goals of maintaining ecological

sustainability.
 ACKNOWLEDGMENTS

The completion of this thesis would not have been possible without the guidance

and support of many people. First, thank you Dr. Austin Humphries for the opportunity

to learn about many different coral reef fish species, for his guidance and patience while

working with me. My sincere gratitude to my committee members Prof. Jeremy Collie

and Dr. Kathleen Castro for their advice.

I would like to thank my Indonesian friends and my lab mates in Rhode Island, for

their companionship during my study. I would like to acknowledge the sources of my

financial support: Bappenas (Pak Gellwynn) and Coremap DGCF MMAF in Jakarta.

Last but not least, I would like to thank my lovely wife (Asriyani), my beautiful daughter

(Aisha) and also my big family in Indonesia for their constant, continuing support so

that my thesis came together.

iii
 PREFACE

This thesis is written in manuscript format and adheres to the guidelines of the

Graduate School of the University of Rhode Island. The manuscript is formatted in

accordance with the guidelines set forth for publication in Fisheries Research.

iv
 TABLE OF CONTENTS

ABSTRACT ................................................................................................................... ii

ACKNOWLEDGMENTS ............................................................................................ iii

PREFACE ..................................................................................................................... iv

TABLE OF CONTENTS ............................................................................................... v

LIST OF TABLES ........................................................................................................ vi

LIST OF FIGURES ..................................................................................................... vii

MANUSCRIPT-1 ........................................................................................................... 1

1. Introduction .............................................................................................................. 3

2. Materials and methods ............................................................................................. 6

2.1. Field methods .................................................................................................... 6
2.2 Data analyses ..................................................................................................... 7
2.3. Statistical analyses ............................................................................................. 9
3. Results ...................................................................................................................... 9
3.1. Total fish biomass............................................................................................... 9
3.2. Functional group biomass................................................................................. 10
3.3. Life-history characteristics .............................................................................. 11

4. Discussion ............................................................................................................. 12

5. Conclusions ........................................................................................................... 16

6. References ............................................................................................................. 17

v
 LIST OF TABLES

TABLE PAGE

Table 1. Mean biomass (kg/ha) of reef fish communities on fished reefs and no-take

closures in three Sunda Banda Seascape regions ........................................................ 26

Table 2. Mean biomass (kg/ha) of reef fish families in fished reefs and no-take

closures of the Sunda Banda Seascape........................................................................ 27

Table 3. Statistical results testing the impact of region, management, and their

interaction on biomass of reef fish .............................................................................. 28

Table 4. Mean values (kg/ha) and results from individual t-tests of mean functional

group biomass from fished reefs versus no-take closures in the Sunda Banda Seascape

..................................................................................................................................... 29

Table 5. Functional group classifications and life history values of the 25 most

abundant reef fish in the Sunda Banda Seascape ........................................................ 30

Table 6. Mean values and results from individual t-tests of mean life-history

characteristics from fished reefs versus no-take closures in the Sunda Banda Seascape

..................................................................................................................................... 31

vi
 LIST OF FIGURES

FIGURE PAGE

Fig. 1. Study sites within Indonesia’s Sunda Banda Seascape across the Wakatobi (a),

Alor (b), and Koon-Kei (c) regions ............................................................................. 32

Fig. 2. Relationship of mean (and standard error) functional group fish biomass

(kg/ha) between fished reefs and no-take closures. Different letters above bars indicate

statistical differences between management based on individual t-tests ................... 33

Fig. 3. Relationship between log biomass and functional group biomass by

management type (blue = fished reefs; orange = no-take closures) ............................ 35

Fig. 4. Boxplot of median and quartile range of community-level mean maximum

length (cm), life span (yrs), growth rate (cm/yr), and trophic level for fished reefs and

no-take closures. Letters above boxplots indicate statistical differences between

management based on individual t-tests .................................................................... 39

Fig. 5. Relationship between reef fish biomass (log) and functional group by

management (blue = fished reefs; orange = no-take closures).................................... 40

vii
 Manuscript-1

Prepared for submission to Fisheries Research

The Impact of Fisheries Management on Coral Reef Fish Communities and Life-

History Characteristics

Diky Suganda1, Austin Humphries1,2

1
Department of Fisheries, Animal and Veterinary Sciences, University of Rhode
Island, Kingston, RI, 02881 USA
2
Graduate School of Oceanography, University of Rhode Island, Narragansett, RI,
02882 USA

Corresponding Author: Diky Suganda

Department of Fisheries, Animal and Veterinary
Sciences
University of Rhode Island
Kingston, RI 02881
Tel: 401-932-3665
Email: superdiky@my.uri.edu

1
Abstract

Indonesia’s coral reefs support over half of all small-scale fishers globally and are thus

an important economic and sociocultural resource. These coral reef fisheries, however,

are threatened from a variety of stressors including overexploitation. No-take fisheries

closures are thought to be a suitable management strategy in Indonesia that provide a

refuge for fish species with different ecological and life-history characteristics. While

examining such indicators is increasingly important to determine management efficacy,

few have done so in Indonesia. We investigate community ecological and life-history

responses to no-take fisheries closures using abundance data from 2009-2015 in three

regions across Indonesia’s Sunda Banda Seascape. Overall, fish biomass was 30%

greater in no-take closures than fished reefs. The only functional groups to respond

significantly to management were corallivores and detritivores, and fished reefs had

greater biomasses of these groups. No-take closures had fish communities with greater

maximum lengths, longer life spans, slower growth rates, and higher mean trophic

levels. Surprisingly, total fish biomass was not a good predictor for life history values

or functional group biomass, as none of these indicators responded to fish recovery. Our

results indicate that non-target fish species may thrive in fished reefs where predators

with slow life histories may be reduced. Also, no-take closures in Indonesia’s Sunda

Banda Seascape are facilitating recovery of life-history characteristics and fish biomass

but these impacts are not uniformly distributed across functional groups. These findings

are of value to current ecosystem-based management objectives attempting to achieve

broader conservation goals of maintaining ecological sustainability.

2
1. Introduction

Coral reefs are an extremely important ecosystem because of biological, ecological,

and socioeconomic reasons. They are one of the most biodiverse and productive

ecosystems in the world, matching rainforests in their richness of life (Birkeland 1997).

They constitute less than 0.1% of the ocean floor (Spalding et al. 2001) but support

greater than 30% of all known fish species (Sorokin 2013). Coral reefs are critical for

both economic security and food security because they sustain the lives of more than

130 million coastal communities and supply global markets, providing ecosystem

services to coastal communities that include fisheries, aquaculture, tourism, coastal

protection, and transport (Moberg & Folke 1999). Coral reefs, however, are sensitive to

a myriad of natural and anthropogenic stressors, primarily through invasive species,

climate change, pollution, and destructive fishing practices (Edinger 1998; McClanahan

2007, Wenger 2016). In particular, the impact of overfishing has been shown to severely

damage coral reef fish community structure and functioning (Jennings & Kaiser 1998;

Roberts 1995). These changes come as a result of a reduction in fish abundance, shifts

in body size distribution, diversity, and biomass (Jennings et al. 1999; Graham et al.

2007; Worm et al. 2009; McClanahan & Humphries 2012).

Indonesian coral reefs cover 32,000 km2 or about 18% of the world’s coral reef area,

which is the second largest in the world after the Great Barrier Reef in Australia (Nontji

2002). Indonesia’s Sunda Banda Seascape (SBS) has been designated as the second

most important marine ecological region in Indonesia in terms of its biodiversity (Wang

et al. 2015). The SBS is 151 million hectares that stretches from Bali to Nusa Tenggara

and Maluku Tenggara, as well as the southern and eastern sides of Sulawesi. The SBS

3
provides habitat for 76% of known coral species globally, and over 3000 fish species

(Veron et al. 2009). These coral reefs, however, are at constant risk of degradation due

to natural and anthropogenic stressors. A recent study by the Indonesian government

suggests that less than 7% of coral reefs in the SBS are in ‘excellent’ condition (76-100

percent of live coral cover), and 35% are functionally obsolete (Giyanto et al. 2017).

Fisheries management and harvest tools, such as gear restrictions, periodically

harvested closures, and Marine Protected Areas (MPA) have been implemented to

protect coral reef fisheries from decline and collapse (Friedlander 2015). MPAs, for

example, have been widely promoted as strategy for an ecosystem approach to fisheries

management (Hastings & Botsford 1999; Claudet et al. 2006), potentially benefiting

both fisheries and conservation objectives (Claudet et al. 2010, Rodrigues et al. 2004).

Evidence supporting these claims, however, is limited; managing for coral reef fisheries

production along with biodiversity conservation remains as a major challenge

(Balmford et al. 2005). While such MPAs provide a refuge to a variety of fish species

(Di Lorenzo et al. 2016), they may not provide protection to fish with different life-

history and ecological characteristics. Furthermore, the efficacy of MPAs to increase

fish biomass and diversity has been proven to be dictated by the ability of management

to enforce regulations (Gill et al. 2017). This presents particular problems for Indonesia

where MPAs are often touted as a panacea for management solutions but capacity to

enforce them is low.

Ecosystem functionality highlights feeding functional groups and multiple

processes involved in the transfer of energy and matter over time and space (Reiss et al.

2009). Trophic guilds (i.e., feeding functional groups) have thus become important tools

4
in providing a better understanding of the structure and functioning of fish assemblages

(Mathieson et al. 2000). Feeding functional group affects distribution, abundance, and

demographics of the fish community. These processes can be dynamic, changing

population productivity and trophic structure at ecological or evolutionary time scales

(Brodeur et al. 2017). Viewing ecosystems through this functional group lens is

particularly well-suited for coral reef systems that are incredibly diverse. It is important

in coral reefs to enable description of community-level feeding patterns and determine

how habitat is influenced by fish functional groups and which species or taxa are critical

in maintaining an ecosystem (Edwards et al. 2014).

Patterns of recovery are controlled by ecological organization principles such as

competition and predator-prey interactions (McClanahan et al. 2007). For example,

species with life-history characteristics that support fast growth and short generation

times can maintain populations and production under high fishing mortality, but they

are expected to experience genetic change under new conditions and associated

selection pressures (McClanahan & Graham 2015). The opposite is true with fishes

characterized by slow growth rates and low natural mortality and protection from fishing

can take a long time to recover such life histories (McClanahan & Humphries 2012).

These trade-offs are poorly understood in diverse coral reef fish communities. Thus, in

order for fisheries management to be successful at maintaining not only fisheries-

specific goods and services from coral reefs but also the portfolio of characteristics

necessary to support resilience and tourism, such life-history characteristics should be

examined (Taylor et al. 2014).

5
 Classical fisheries management relies on life-history characteristics such as growth,

reproduction, and longevity as the basis for key indicators of productivity and maximum

sustainable yield (Walters & Martell 2004). In coral reef systems that lack data, less

attention has been given to life-history characteristics which are also important in

evaluations of the increasing frequency and magnitude of anthropogenic and climate

stressors (McClanahan & Humphries 2012). This study is intended to provide a greater

understanding of how fisheries management impacts fish communities by comparing

ecological indicators between fished reefs and no-take fisheries closures. Specifically,

we investigated the efficacy of no-take closures in protecting or recovering coral reef

fish assemblages and traits, thus bolstering ecosystem functioning. Utilizing a large

dataset of coral reef fish abundances, we evaluated changes in biomass, functional

groups, as well as key life-history characteristics of coral reef fish communities of the

Sunda Banda Seascape in Indonesia. We hypothesized that reef fish communities would

shift towards larger, slower growing fish in no-take closures, and community biomass

would be higher in these closures, particularly for piscivores and herbivores that are

target species.

2. Materials and Methods

2.1. Field Methods

Abundance data for fish were collected by the non-governmental organization

World Wildlife Fund (WWF) within the three regions in the Sunda Banda Seascape

(SBS; Fig. 1). Regions studied included Wakatobi, Alor Timur, Koon-Kei, and data

were collected between 2009 and 2015. Individual sampling sites were classified as

6
either being a fished reef, where fishing activities were open to everyone, or no-take

closures where fishing activities were prohibited and enforced. If enforcement

capabilities of local authorities were unknown or in question, the data were not used in

analyses.

At sampling sites, species, size, and number of fish found in 5 x 50 meters belt

transects were recorded and tabulated using underwater visual census (UVC; English et

al. 1994; Samoilys & Carlos 2000). Three to five replicates were carried out for each

site. In order to minimize any biases during UVC, divers were trained prior to the field

sampling on species identification and all trained divers swam slowly (approximately

30 minutes per transect) with an average depth of 10 meters. Coral boulders within

transects were circumnavigated in order to count and identify reef fish hidden from

view. Fish smaller than 10 cm were excluded to reduce errors in density comparisons.

Large transect widths and lengths were selected to reduce inconsistency between

transects, to include species at low densities, and to minimize flight distance effects

observed in certain species (Bellwood & Alcala 1988).

2.2. Data analyses

Estimated fish size and count data were converted into biomass using species-

specific length-weight relationships (Letourneur et al. 1998; Froese & Pauly 2012;

McClanahan et al. 2015) where:

W = aLb

a and b are growth coefficients retrieved from FishBase and L is total length (cm). We

then converted each 250 m2 transect to kilograms per hectare (kg/ha) to be consistent

with the literature.

7
 We determined each individual’s functional group at the genus-level based on diet

and foraging strategies (Wilson et al. 2008; Froese & Pauly 2012). We obtained these

classifications from FishBase and categorized each individual as one of the followings:

corallivore, detritivore, grazer, invertivore, piscivore, planktivore, and scraper. Grazers

and scrapers are both types of herbivores but were not lumped together for our analyses

because they have been shown to have differential effects on reef functioning and

structure (Humphries et al. 2015).

Life history characteristics of individual species included in this study were

maximum length, growth rate, and life span. We retrieved species-level values for these

using FishBase (www.fishbase.org). Non-existing life-history parameters of certain

species were estimated from studies of the closest genus or family level. Maximum

length represents the greatest size ever reported for that particular species. Intrinsic

rowth rate is an estimate of the growth of individuals in size or length for a given time

period as a fraction of the population at the beginning of a time period. Life span is the

approximate maximum age that fish of a given population would reach. Trophic level

of a species is a unitless metric which represents its position in the food chain and it is

estimated based on diet composition. For example, a high trophic level value such as 4

may represent a piscivorous species (e.g., apex predators such as sharks), and

conversely, a low trophic level value may represent an herbivorous fish species.

Community-level weighted averages for life histories and trophic level (resulting in

‘mean trophic level’) were calculated for each year (k) as:

8
where Yi is the biomass of species i (per transect), and LH is the life-history

characteristic of interest (McClanahan & Humphries 2012).

2.3. Statistical analyses

We used a linear mixed effects model to determine if region, management, or their

interaction influenced fish biomass. To account for site-level differences within each

region (n = 3), site was included in the model as an orthogonal nested random effect.

Biomass was not normally distributed (determined using a Kolmogorov-Smirnov test),

and thus a log-transformation was necessary. We used an alpha value of 0.05 for

statistical significance. We used individual t-tests to compare functional group biomass

of fishes between fished reefs and no-take closures for corallivores, detritivores, grazers,

invertivores, piscivores, planktivores, and scrapers. We also used individual t-tests to

compare life-history characteristics of maximum length, growth rate, life span, or

trophic level between management zones. Last, we used linear regression to explore

relationships between the recovery of fish biomass on reefs and functional group

biomass as well as individual life-history characteristics.

3. Results

The dataset ended up consisting of 193 unique sites: Wakatobi contained the most

sites (n = 126), followed by Alor (n = 47), and Koon-Kei (n = 20; Fig. 1). A total of

9,899 individuals were included, belonging to 293 species, 77 genera, and 24 families.

3.1. Total fish biomass

Mean biomass of reef fish in the no-take closures was 30% greater than biomass on

fished reefs (Table 1). The difference was the most prevalent in Alor, where biomass in

9
the no-take closures was 86% greater than that in the fished reefs. Similarly, although

smaller, there was 14% increase of reef fish biomass in Wakatabi’s no-take closures

compared to its fished reef areas. On the contrary, Koon-Kei acted differently, where

reef fish biomass in no-take closure (401 kg/ha) was found to be 5% lower than the

biomass in fished reef (424 kg/ha).

Based on their high abundance, Acanthuridae was the most dominant family in the

Sunda Banda Seascape, followed by Scaridae, and Lutjanidae (Table 2). All of these

families had greater than 200 kg/ha on both fished reefs and no-take closures. After

these top 3 families, there were numerous less abundant families with biomass levels

below 100 kg/ha, such as: Haemulidae, Serranidae, Lethrinidae, Siganidae,

Holocentridae. The families of Pomacentridae, Tetradontidae, and Zanclidae were

found only in fished reefs but not in no-take closures.

There was a significant effect of management on the overall biomass of fish in the

Sunda Banda Seascape (p = 0.023), whereas region had no significant effect (Table 3).

Also, there was no significant interaction between region and management for biomass.

3.2. Functional group biomass

Based on total functional group biomass, no-take closures were dominated by

grazers (1067 kg/ha), piscivores (495 kg/ha), and scrapers (454 kg/ha; Fig. 2). There

were very few corallivores (7 kg/ha) or detritivores (37 kg/ha) in no-take closures,

whereas their existence was abundant in fished reefs. Comparisons of individual

functional groups revealed that corallivore and detritivore were significantly different

between management zones (p < 0.05), with both being higher in fished reefs (Table 4).

The biomass of grazers, invertivores, piscivores, planktivores, and scrapers found in

10
fished reefs were comparable to that in no-take closures (p > 0.05). As fish biomass

increased across the sites, functional group biomass did not respond predictably (Fig.

3). No linear regressions had an R2 value greater than 0.15 and none were statistically

significant. Although, there were trends of an inverse relationship between total biomass

and functional group biomass for corallivores, detritivores, invertivores, and

planktivores.

3.3. Life-history characteristics

We found that Bolbometopon muricatum, Naso hexacanthus and Lutjanus bohar

had the highest maximum length values of all species in the dataset: 130 cm, 100 cm

and 90 cm, respectively (Table 5). Life spans of reef fish were highly varied, ranging

from the shortest of 2 years (Cephalopholis urodeta) to the longest-lived species of 31.8

years (Lutjanus bohar). On the contrary, these two fish were found as the fastest and

slowest growth rate species, respectively. Meanwhile, feeding functional groups of

dominant reef fish were grazer, piscivore, invertivore, and scraper. The species with the

highest trophic level (TL) found in the study sites was Cephalopholis argus, belonging

to piscivore group with TL value of 4.9; all piscivores had trophic level values above

3.9. Herbivores were represented as either grazers or scrapers and they all had trophic

levels between 2 and 3. Planktivores and invertivores occupied trophic levels between

those of the herbivores and piscivores, around 3. There were only two detritivores in the

top 25 species and they have a low trophic level of 2, similar to that of some herbivores.

Management had a significant effect on each life-history characteristic (Fig. 4).

Individual t-tests showed that no-take closures had fish communities with greater

maximum lengths, lower growth rates, greater life spans, and higher mean trophic levels

11
(Table 6). The maximum length for fished reefs was 47.2 cm compared to 55 cm in no-

take closures. The mean trophic levels were both below 3, however, the no-take closures

were greater at 2.83 versus 2.69 in the fished reefs. Life span was 11.2 years in the

closures and only 9.3 years in the fished reefs.

Linear regressions indicated that while there were increasing trends of maximum

length and life span of reef fish in both management areas as total biomass increased,

these were not significant or very predictive (R2 < 0.1; Fig. 5). Trophic level showed no

sign of increase or decrease with fish biomass on reefs, regardless of management. In

addition, growth rates decreased slightly along the fish biomass gradient.

4. Discussion

Our primary hypotheses in this study were supported but with some important

nuances. No-take fisheries closures in Indonesia’s Sunda Banda Seascape were an

effective management strategy to increase total fish biomass, but this was spread rather

indiscriminately across functional groups. In other words, target groups such as

piscivores and scrapers showed no statistically significant differences between fished

and protected reefs. Community life histories changed as expected towards larger,

slower growing and long-lived fauna in the no-take closures. Our results agree with life-

history theory and a number of previous studies that measure fish abundance and

biomass responses to management (e.g., Halpern & Warner 2002; Rakitin & Kramer

1996; Russ & Alcala 1996; Watson et al. 1996; Jennings & Kaiser 1998; and

McClanahan & Humphries 2012). What is less clear is why functional groups did not

12
respond as strongly to management and why our indicators were not strongly correlated

on a continuous gradient with biomass.

One of our sites, Koon-Kei, had fish biomass on fished reefs that was slightly higher

than the biomass in no-take closures. In this case, level of protection did not necessarily

associate with an increase in biomass. However, McClanahan et al. (2008) and Claudet

et al. (2008) found that size and age of no-take closures can also influence fish

communities. The no-take closures in Koon-Kei were established in 2012, or three years

before the sampling was conducted in 2015. Some studies suggested that fish density

and species richness increasing after three years of protection (Halpern & Warner 2002,

Russ et al. 2005). We also note that fishing is not the lone factor responsible to fish

biomass but there are other aspects such as habitat and environmental parameters which

create sub-regions within the SBS (Rochet & Trenkel 2003). For instance, Wang et al.

(2015) delineated the SBS based on environmental conditions such as temperature,

currents, and primary productivity, and Koon-Kei was in a different class than Wakatobi

and Alor. Furthermore, Tegner (1993) and Armstrong et al. (1993) indicated that no-

take closures performed poorly when they were located in unfavorable habitats or

contained an insufficient portion of critical habitats. A number of these factors may help

explain why this region did not show the same trends as the other, but we do not have

sufficient data to tease apart these mechanisms.

The coral reef fish communities in the SBS were multispecies and spanned a broad

trophic spectrum from detritivore to apex predators (piscivore). We found that

Acanthuridae, Scaridae, Lutjanidae and Haemulidae were the dominant families in both

management regimes. This result, along with the fact that the associated functional

13
groups also had similar compositions in both management regimes, was somewhat

surprising. However, non-selective fishing has been shown to have an equal impact

across all families and functional groups for fisheries in Kenya (McClanahan 1995).

Our study also revealed that lower trophic level fish were more abundant in fished reef

area, suggesting that these fish is non-targeted by the fishers or their predators have been

removed (e.g., Botsford 1997; Hall 1999). Chaetodontidae are also highly dependent on

coral cover, so habitat quality and quantity may be interacting to influence these findings

(Coker et al. 2014).

Our study revealed that all life-history characteristics showed a predictable and

significant response to management. No-take closures contained fish communities that

had greater maximum lengths, slower growth rates, and longer life spans. The result

conformed with our central hypothesis and previous studies (e.g., McClanahan et al.

2014). Trophic level was also greater in the no-take closures, which has been shown to

be highly variable and even lower because of a dominance of herbivores that are slow

to recover (McClanahan & Humphries 2012).

Growth rate is a plastic trait which can vary in response to environmental conditions

such as food availability (Overholtz 1989; Lorenzen & Enberg 2002) and water

temperature (Jorgensen 1992; Brander 1995; Daufresne et al. 2009). Our study showed

that growth rate was slower in no-take closure compared to fished reef, indicating that

no-take closure provided a better environment for bigger reef fish to grow and to live

longer. In contrast, higher growth rates in fished reefs indicate that fish can mature

faster, turning over faster and providing increases in fisheries production that is resilient

to high fishing pressure.

14
 Most fisheries preferentially target larger species and individuals (Jennings &

Polunin 1997). However, we also noted that coral reef fisheries in Indonesia are

characterized by artisanal fishers where the fishers do not necessarily only target

specific species and sizes to meet their daily consumption. It is important to protect

large, old, and mature fish from fishing as these have greater reproductive potential than

the younger ones (Birkeland & Dayton 2005). Our findings revealed that fish found in

no-take closure had higher life span and maximum length compared to that in fished

reef, suggesting that a more mature and larger fish are abundant and the management

can be successful for bolstering recruitment in surrounding areas.

Our study also showed that lower trophic level species were almost non-existent in

no-take closure. This result indicated that protected areas favored higher trophic level

species. We also noted that piscivory is considered as the main process of energy

transfer in coral reef ecosystems (Parish et al. 1986; Hixon 1991). Therefore, this result

indicates that piscivory may be contributing to a rapid decrease of these prey species

(low trophic level) in the no-take closure area and future analyses should investigate

size-spectra as another indicator for management efficacy. On the flip side of this, we

found a very high proportion of lower trophic level species such as corallivores and

detritivores on fished reefs. These functional groups were fast growing species and have

relatively low contribution to fisheries yield in the SBS.

We recognize our findings have some limitations to consider. For instance, using

global databases such as FishBase in this study may introduce bias in the analysis of

fish life-history characteristics. This is because those data are not geographically

specific to Indonesia and some of the values were obtained from studies in the Caribbean

15
or when species were not available, the next closest relative. However, due to limited

data available in Indonesia for the myriad of coral reef fish species, we have to assume

that the data from those previous studies are better than any other alternative at this time.

In addition to this is the issue of covariates driving fish community composition outside

of fishing pressure. We previously discussed things like habitat quality and quantity, as

well as environmental characteristics, but enforcement has been shown to be the

strongest predictor of fish biomass on a global scale (Gill et al. 2017). We addressed

this issue by removing any data from no-take closures where it was not clear how well

it was managed by communicating with WWF and the field team that collected the data.

We have no way of knowing these values for our dataset and therefore have to treat all

no-take closures as equal even though we acknowledge the bias this may introduce.

5. Conclusions

This study demonstrated that fishing activity affected the biomass and community

level life-history characteristics of reef fish in Sunda Banda Seascape (SBS) of

Indonesia. Our findings corroborate other study by Davis & Dodrill (1989) that no-take

closure produced positive results for some species and functional groups, but not others.

Although community biomass did not increase in all three regions with no-take closures

or across all functional groups, closures did provide protection for larger, slower

growing high trophic level species. Non-target fish species may thrive in fished reefs

where predators with slow life histories may be reduced. Future research should look to

incorporate things like habitat, enforcement and environmental conditions to test

whether these have greater impacts on fish community characteristics than fishing.

16
 To safeguard sustainable coral reef fisheries in the SBS, we suggest no-take closures

should continue to established but to also look for alternative strategies that may provide

complimentary benefits for functional groups and fish life histories. Such adaptive

management strategies might include things like a gear restriction or temporal closures

in heavily fished areas. These strategies may improve coral reef fish resilience to

interacting effects of fishing and climate.

6. References

Armstrong, D. A., Wainwright, T. C., Jensen, G. C., Dinnel, P. A., and Andersen, H. B.

Taking refuge from by-catch issues: red king crab (Paralithodes camtschaticus)

and trawl fisheries in the Eastern Bering Sea. Canadian Journal of Fisheries and

Aquatic Sciences, no. 50 (1993): 1993–2000.

Balmford, Andrew, Peter Crane, Andy Dobson, Rhys E. Green, and Georgina M. Mace.

"The 2010 challenge: data availability, information needs and extraterrestrial

insights." Philosophical Transactions of the Royal Society B: Biological

Sciences 360, no. 1454 (2005): 221-228.

Bellwood, D. R., and A. C. Alcala. "The effect of a minimum length specification on

visual estimates of density and biomass of coral reef fishes." Coral reefs 7, no.

1 (1988): 23-27.

Birkeland, Charles. Life and death of coral reefs. Springer Science & Business Media,

1997.

Birkeland, Charles, and Paul K. Dayton. "The importance in fishery management of

leaving the big ones." Trends in Ecology & Evolution 20, no. 7 (2005): 356-358.

17
Botsford, Louis W., Juan Carlos Castilla, and Charles H. Peterson. "The management

of fisheries and marine ecosystems." Science 277, no. 5325 (1997): 509-515.

Brander, K. M. "The effect of temperature on growth of Atlantic cod (Gadus morhua

L.)." ICES Journal of Marine Science 52, no. 1 (1995): 1-10.

Brodeur, Richard D., Brian E. Smith, Richard S. McBride, Ron Heintz, and Ed Farley.

"New perspectives on the feeding ecology and trophic dynamics of

fishes." Environmental Biology of Fishes 100, no. 4 (2017): 293-297.

Claudet, Joachim, Dominique Pelletier, J-Y. Jouvenel, F. Bachet, and R. Galzin.

"Assessing the effects of marine protected area (MPA) on a reef fish assemblage

in a northwestern Mediterranean marine reserve: Identifying community-based

indicators." Biological conservation 130, no. 3 (2006): 349-369.

Claudet, Joachim, Craig W. Osenberg, Lisandro Benedetti‐Cecchi, Paolo Domenici,

José‐Antonio García‐Charton, Ángel Pérez‐Ruzafa, Fabio Badalamenti et al.

"Marine reserves: size and age do matter." Ecology letters 11, no. 5 (2008): 481-

489.

Claudet, Joachim, C. W. Osenberg, Paolo Domenici, Fabio Badalamenti, Marco

Milazzo, Jesús Manuel Falcón, Iacopo Bertocci et al. "Marine reserves: fish life

history and ecological traits matter." Ecological applications 20, no. 3 (2010):

830-839.

Coker, Darren J., Shaun K. Wilson, and Morgan S. Pratchett. "Importance of live coral

habitat for reef fishes." Reviews in Fish Biology and Fisheries 24, no. 1 (2014):

89-126.

18
Daufresne, Martin, Kathrin Lengfellner, and Ulrich Sommer. "Global warming benefits

the small in aquatic ecosystems." Proceedings of the National Academy of

Sciences 106, no. 31 (2009): 12788-12793.

Davis, Gary E., and Jon W. Dodrill. "Recreational fishery and population dynamics of

spiny lobsters, Panulirus argus, in Florida Bay, Everglades National Park, 1977–

1980." Bulletin of Marine Science 44, no. 1 (1989): 78-88.

Di Lorenzo, Manfredi, Joachim Claudet, and Paolo Guidetti. "Spillover from marine

protected areas to adjacent fisheries has an ecological and a fishery

component." Journal for Nature Conservation 32 (2016): 62-66.

Edinger, Evan N., Jamaluddin Jompa, Gino V. Limmon, Wisnu Widjatmoko, and

Michael J. Risk. "Reef degradation and coral biodiversity in Indonesia: effects

of land-based pollution, destructive fishing practices and changes over

time." Marine Pollution Bulletin 36, no. 8 (1998): 617-630.

Edwards C.B., Friedlander A.M., Green A.G., Hardt M.J., Sala E., Sweatman H.P.,

Williams I.D., Zgliczynski B., Sandin S.A. and Smith J.E. Global assessment of

the status of coral reef herbivorous fishes: evidence for fishing effects.

Proceedings of the Royal Society Biological Sciences no. 281(2014): 20131835.

English, Susan S., Clive CR Wilkinson, and Valonna V. Baker. Survey manual for

tropical marine resources. Australian Institute of Marine Science, 1997.

Friedlander, Alan M. "21 A perspective on the management of coral reef

fisheries." Ecology of fishes on coral reefs (2015): 208.

19
Froese, Rainer, and Daniel Pauly. "Fishbase (www database)." World Wide Web

Electronic Publications. Available at: http://www. fishbase. org (accessed June

2012) (2012).

Gill, David A., Michael B. Mascia, Gabby N. Ahmadia, Louise Glew, Sarah E. Lester,

Megan Barnes, Ian Craigie et al. "Capacity shortfalls hinder the performance of

marine protected areas globally." Nature 543, no. 7647 (2017): 665.

Giyanto et al. Status Terumbu Karang Indonesia. COREMAP CTI. Jakarta: Puslit

Oseanografi – LIPI, 2017.

Graham, Nicholas AJ, Shaun K. Wilson, Simon Jennings, Nicholas VC Polunin, J. A.

N. Robinson, Jude P. Bijoux, and Tim M. Daw. "Lag effects in the impacts of

mass coral bleaching on coral reef fish, fisheries, and

ecosystems." Conservation biology 21, no. 5 (2007): 1291-1300.

Hall, Stephen J. The effects of fishing on marine ecosystems and communities.

Blackewel Science Ltd, 1999.

Halpern, Benjamin S., and Robert R. Warner. "Marine reserves have rapid and lasting

effects." Ecology letters 5, no. 3 (2002): 361-366.

Hastings, Alan, and Louis W. Botsford. "Equivalence in yield from marine reserves and

traditional fisheries management." Science 284, no. 5419 (1999): 1537-1538.

Hixon, M. "Tropical and temperate reef fishes: comparison of community

structures." The ecology of fishes on coral reefs (1991): 509-563.

Humphries, Austin T., Christopher D. McQuaid, and Tim R. McClanahan. "Context-

Dependent Diversity-Effects of Seaweed Consumption on Coral Reefs in

Kenya." PloS one10, no. 12 (2015): e0144204.

20
Jennings, S., and N. V. C. Polunin. "Impacts of predator depletion by fishing on the

biomass and diversity of non-target reef fish communities." Coral reefs 16, no.

2 (1997): 71-82.

Jennings, Simon, and Michel J. Kaiser. "The effects of fishing on marine ecosystems."

In Advances in marine biology, vol. 34, pp. 201-352. Academic Press, 1998.

Jennings, Simon, Simon Greenstreet, and John Reynolds. "Structural change in an

exploited fish community: a consequence of differential fishing effects on

species with contrasting life histories." Journal of Animal Ecology 68, no. 3

(1999): 617-627.

Jørgensen, Terje. "Long-term changes in growth of North-east Arctic cod (Gadus

morhua) and some environmental influences." ICES Journal of Marine

Science 49, no. 3 (1992): 263-278.

Letourneur, Y., Michel Kulbicki, and P. Labrosse. "Length-weight relationship of fishes

from coral reefs and lagoons of New Caledonia: an update." Naga, the ICLARM

Quarterly 21, no. 4 (1998): 39-46.

Lorenzen, Kai, and Katja Enberg. "Density-dependent growth as a key mechanism in

the regulation of fish populations: evidence from among-population

comparisons." Proceedings of the Royal Society of London B: Biological

Sciences 269, no. 1486 (2002): 49-54.

Mathieson, S., André Cattrijsse, M. J. Costa, Pilar Drake, M. Elliott, J. Gardner, and J.

Marchand. "Fish assemblages of European tidal marshes: a comparison based

on species, families and functional guilds." Marine Ecology Progress Series 204

(2000): 225-242.

21
McClanahan, T. R. "A coral reef ecosystem-fisheries model: impacts of fishing intensity

and catch selection on reef structure and processes." Ecological Modelling 80,

no. 1 (1995): 1-19.

McClanahan, Tim R., Nicholas AJ Graham, Jacqulyn M. Calnan, and M. Aaron

MacNeil. "Toward pristine biomass: reef fish recovery in coral reef marine

protected areas in Kenya." Ecological applications 17, no. 4 (2007): 1055-1067.

McClanahan, T. R., M. Ateweberhan, N. A. J. Graham, S. K. Wilson, C. Ruiz Sebastián,

Mireille MM Guillaume, and J. Henrich Bruggemann. "Western Indian Ocean

coral communities: bleaching responses and susceptibility to

extinction." Marine Ecology Progress Series 337 (2007): 1-13.

McClanahan, Tim R., Nicholas AJ Graham, Shaun K. Wilson, Yves Letourneur, and

Rebecca Fisher. "Effects of fisheries closure size, age, and history of compliance

on coral reef fish communities in the western Indian Ocean." Marine Ecology

Progress Series 396 (2009): 99-109.

McClanahan, T. R., and A. T. Humphries. "Differential and slow life-history responses

of fishes to coral reef closures." Marine Ecology Progress Series 469 (2012):

121-131.

McClanahan, T. R., and N. A. J. Graham. "Marine reserve recovery rates towards a

baseline are slower for reef fish community life histories than biomass." Proc.

R. Soc. B 282, no. 1821 (2015): 20151938.

McClanahan, T. R., N. A. J. Graham, M. A. MacNeil, and J. E. Cinner. "Biomass‐based

targets and the management of multispecies coral reef fisheries." Conservation

Biology 29, no. 2 (2015): 409-417.

22
Moberg, Fredrik, and Carl Folke. "Ecological goods and services of coral reef

ecosystems." Ecological economics 29, no. 2 (1999): 215-233.

Nontji, Anugerah. "Coral reefs of Indonesia: past, present and future." In Proceedings

of the Ninth International Coral Reef Symposium, Bali, 23-27 October 2000,

vol. 1, pp. 17-27. 2002.

Overholtz, W. J. "Density-dependent growth in the Northwest Atlantic stock of Atlantic

mackerel (Scomber scombrus)." J. Northw. Atl. Fish. Sci 9 (1989): 115-121.

Parrish, James D., James E. Norris, Michael W. Callahan, Janice K. Callahan, Eric J.

Magarifuji, and Robert E. Schroeder. "Piscivory in a coral reef fish community."

In Contemporary studies on fish feeding: the proceedings of GUTSHOP’84, pp.

285-298. Springer, Dordrecht, 1986.

Rakitin, Ana, and Donald L. Kramer. "Effect of a marine reserve on the distribution of

coral reef fishes in Barbados." Marine Ecology Progress Series (1996): 97-113.

Reiss, Julia, Jon R. Bridle, Jose M. Montoya, and Guy Woodward. "Emerging horizons

in biodiversity and ecosystem functioning research." Trends in ecology &

evolution 24, no. 9 (2009): 505-514.

Roberts, Callum M. "Effects of fishing on the ecosystem structure of coral

reefs." Conservation Biology 9, no. 5 (1995): 988-995.

Rochet, Marie-Joëlle, and Verena M. Trenkel. "Which community indicators can

measure the impact of fishing? A review and proposals." Canadian Journal of

Fisheries and Aquatic Sciences 60, no. 1 (2003): 86-99.

Rodrigues, Ana SL, H. Resit Akcakaya, Sandy J. Andelman, Mohamed I. Bakarr, Luigi

Boitani, Thomas M. Brooks, Janice S. Chanson et al. "Global gap analysis:

23
 priority regions for expanding the global protected-area network." AIBS

Bulletin54, no. 12 (2004): 1092-1100.

Russ, Garry R., and Angel C. Alcala. "Do marine reserves export adult fish biomass?

Evidence from Apo Island, central Philippines." Marine Ecology Progress

Series (1996): 1-9.

Samoilys, Melita A., and Gary Carlos. "Determining methods of underwater visual

census for estimating the abundance of coral reef fishes." Environmental

Biology of Fishes 57, no. 3 (2000): 289-304.

Spalding, Mark, Mark D. Spalding, Corinna Ravilious, and Edmund Peter Green. World

atlas of coral reefs. Univ of California Press, 2001.

Sorokin, Yuri I. Coral reef ecology. Vol. 102. Springer Science & Business Media,

2013.

Taylor, Brett M., Peter Houk, Garry R. Russ, and J. Howard Choat. "Life histories

predict vulnerability to overexploitation in parrotfishes." Coral Reefs 33, no. 4

(2014): 869-878.

Tegner, M. J. Southern California abalones: can stocks be rebuilt using marine refugia?

Canadian Journal of Fisheries and Aquatic Sciences no. 50. (1993): 2010–2018.

Veron, J. E. N., Lyndon M. Devantier, Emre Turak, Alison L. Green, Stuart

Kininmonth, Mary Stafford-Smith, and Nate Peterson. "Delineating the coral

triangle." Galaxea, Journal of Coral Reef Studies 11, no. 2 (2009): 91-100.

Walters, Carl J., and Steven JD Martell. Fisheries ecology and management. Princeton

University Press, 2004.

24
Wang, Mingshu, Gabby N. Ahmadia, Iliana Chollett, Charles Huang, Helen Fox, Anton

Wijonarno, and Marguerite Madden. "Delineating biophysical environments of

the Sunda Banda Seascape, Indonesia." International journal of environmental

research and public health 12, no. 2 (2015): 1069-1082.

Watson, Maggie, Dave Righton, Tim Austin, and Rupert Ormond. "The effects of

fishing on coral reef fish abundance and diversity." Journal of the Marine

Biological Association of the United Kingdom 76, no. 1 (1996): 229-233.

Wenger, Amelia S., David H. Williamson, Eduardo T. da Silva, Daniela M. Ceccarelli,

Nicola K. Browne, Caroline Petus, and Michelle J. Devlin. "Effects of reduced

water quality on coral reefs in and out of no‐take marine reserves." Conservation

Biology 30, no. 1 (2016): 142-153.

Wilson, Shaun K., Scott C. Burgess, Alistair J. Cheal, Mike Emslie, Rebecca Fisher, Ian

Miller, Nicholas VC Polunin, and Hugh Sweatman. "Habitat utilization by coral

reef fish: implications for specialists vs. generalists in a changing

environment." Journal of Animal Ecology 77, no. 2 (2008): 220-228.

Worm, Boris, Ray Hilborn, Julia K. Baum, Trevor A. Branch, Jeremy S. Collie,

Christopher Costello, Michael J. Fogarty et al. "Rebuilding global

fisheries." Science 325, no. 5940 (2009): 578-585.

25
Tables

Table 1

Mean biomass (kg/ha) of reef fish communities on fished reefs and no-take closures in
three Sunda Banda Seascape regions.

Region (Year of MPA Mean Biomass (kg/ha) Management

MPA area (ha)
established) Fished Reef No-take Closure Impact
Alor (2002) 21.85 356 664 +86.6%
Koon-Kei (2012) 150 424 401 -5.4%
Wakatobi (1996) 1,390,000 496 565 +13.9%
Total 435 564 +29.7%

26
Table 2

Mean biomass (kg/ha) of reef fish families in fished reefs and no-take closures of the
Sunda Banda Seascape.

Mean Biomass (kg/ha)

Family
Fished Reef No-take Closure
Acanthuridae 666.2 735.2
Scaridae 395.4 454.2
Lutjanidae 237.5 333.6
Haemulidae 62.4 67.3
Serranidae 62.4 98.2
Lethrinidae 49.8 44.1
Siganidae 44.8 37.3
Holocentridae 28.5 2.6
Chaetodontidae 27.7 7.1
Mullidae 24.7 5.4
Nemipteridae 17.0 7.2
Pomacanthidae 11.3 4.4
Ephippidae 10.5 2.1
Balistidae 10.1 0.5
Kyphosidae 8.0 5.4
Pomacentridae 8.0 0
Labridae 7.3 9.7
Tetraodontidae 0.4 0
Zanclidae 0.4 0
Pempheridae 0.4 1.3
Sphyraenidae 0.2 12.5

27
Table 3

Statistical results testing the impact of region, management, and their interaction on
biomass of reef fish.

Effect numDF denDF F-value p-value

Region 2 105 2.12 0.125
Management 1 105 5.55 0.023
Region * Management 2 105 0.91 0.406

28
Table 4

Mean values (kg/ha) and results from individual t-tests of mean functional group
biomass from fished reefs versus no-take closures in the Sunda Banda Seascape.

Functional Group Fished Reef No-take Closure df t-value p-value

Corallivore 25.33 6.97 186.9 3.43 < 0.001
Detritivore 104.89 37.31 189.9 3.30 < 0.01
Grazer 965.32 1,067.02 153.2 -0.58 0.468
Invertivore 75.88 77.96 144.3 -0.11 0.913
Piscivore 390.91 495.44 171.7 -1.28 0.204
Planktivore 94.21 75.59 110.4 0.09 0.924
Scraper 395.39 454.20 117.7 -0.76 0.448

29
Table 5

Functional group classifications and life history values of the 25 most dominant reef
fish in the Sunda Banda Seascape.

Growth
Max Life
Functional Rate Trophic
Family Species Length Span
Group (cm / Level
(cm) (year)
year)
Acanthuridae Acanthurus leucocheilus Grazer 45.00 10.60 0.27 2.00
Acanthuridae Acanthurus mata Grazer 50.00 12.90 0.22 2.50
Acanthuridae Acanthurus nigrofuscus Grazer 38.00 4.00 0.70 2.00
Acanthuridae Acanthurus olivaceus Grazer 35.00 2.70 1.06 2.30
Acanthuridae Acanthurus pyroferus Grazer 29.00 6.70 0.42 2.00
Acanthuridae Acanthurus spp. Grazer 40.72 7.42 0.49 2.21
Scaridae Bolbometopon muricatum Scraper 130.00 28.80 0.10 2.70
Serranidae Cephalopholis argus Piscivore 60.00 15.90 0.18 4.50
Serranidae Cephalopholis urodeta Piscivore 28.00 2.00 1.40 4.00
Acanthuridae Ctenochaetus binotatus Detritivore 22.00 4.50 0.63 2.00
Acanthuridae Ctenochaetus striatus Detritivore 26.00 3.50 0.80 2.00
Lutjanidae Lutjanus bohar Piscivore 90.00 31.80 0.09 4.30
Lutjanidae Lutjanus gibbus Piscivore 50.00 9.20 0.31 4.10
Lutjanidae Lutjanus kasmira Piscivore 40.00 13.60 0.21 3.90
Lutjanidae Macolor macularis Piscivore 60.00 11.50 0.25 4.00
Lutjanidae Macolor niger Piscivore 75.00 13.70 0.21 4.00
Lethrinidae Monotaxis grandoculis Invertivore 60.00 13.00 0.22 3.40
Acanthuridae Naso annulatus Grazer 100.00 13.70 0.21 2.10
Acanthuridae Naso hexacanthus Planktivore 75.00 13.00 0.22 3.10
Acanthuridae Naso lituratus Grazer 46.00 8.10 0.35 2.30
Acanthuridae Naso lopezi Planktivore 60.00 13.60 0.21 2.90
Acanthuridae Naso spp. Grazer 59.67 11.66 0.29 2.49
Haemulidae Plectorhinchus chaetodonoides Invertivore 72.00 17.90 0.16 3.80
Haemulidae Plectorhinchus lineatus Invertivore 72.00 9.60 0.30 3.90
Scaridae Scarus spp. Scraper 45.89 7.59 0.47 2.00

30
Table 6

Mean values and results from individual t-tests of mean life-history characteristics
from fished reefs versus no-take closures in the Sunda Banda Seascape.

Life-history Characteristic Fished Reef No-take Closure df t-value p-value

Max length (cm) 47.2 55.0 149.8 -5.06 > 0.001
Growth rate (cm/yr) 0.47 0.40 188.7 -5.11 > 0.001
Life span (yr) 9.3 11.2 164.8 -5.37 > 0.001
Trophic level 2.69 2.83 163.0 -2.94 > 0.01

31
Figures

Fig. 1. Study sites within Indonesia’s Sunda Banda Seascape across the Wakatobi (a),
Alor (b), and Koon-Kei (c) regions.

32
Fig. 2. Relationship of mean (and standard error) functional group fish biomass (kg/ha)
between fished reefs and no-take closures. Different letters above bars indicate
statistical differences between management based on individual t-tests.

a a

b
b

a a
a a

a a

a a

33
 a
a

34
Fig. 3. Relationship between log biomass and functional group biomass by management
type (blue = fished reefs; orange = no-take closures).

35
36
37
38
Fig. 4. Boxplot of median and quartile range of community-level mean maximum length
(cm), life span (yrs), growth rate (cm/yr), and trophic level for fished reefs and no-take
closures. Letters above boxplots indicate statistical differences between management
based on individual t-tests.

39
 a
b

a b

40
Fig. 5. Relationship between reef fish biomass (log) and functional group by
management (blue = fished reefs; orange = no-take closures).

41
42