Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): 

Spatial and temporal patterns in fish and benthic communities (2001-2007)

A cooperative investigation between NOAA and the University of Puerto Rico

NOAA Technical Memorandum NOS NCCOS 107
Mention of trade names or commercial products does not constitute endorsement or recommendation for their use 

by the United States government.

ACKNOWLEDGEMENTS:

This report was made possible through the participation of the individuals recognized above. Their efforts with data 

collection, field support, and report production are highly appreciated. Specific recognition goes to Sarah Hile and 

Jamie Higgins for report layout and production, to Matt Kendall and Richard Appeldoorn for critically reviewing 

the report, to Alicia Clarke for copy editing the document, and to Michelle Sharer for the Spanish translation of 

the Executive Summary. Very special gratitude goes to our dedicated boat captain Angel “Capitan” Nazario and 

his assistant Joel “Joeito” Rivera. Additional logistic support during field missions came from Victor La Santa and 

Roxanna Noriega (West Divers), Milton Carlo (University of Puerto Rico, Isla Magueyes), and our friends at Hotel 

Villa Parguera.

Citation for the entire document:

Pittman, S.J., S.D. Hile, C.F.G. Jeffrey, R. Clark, K. Woody, B.D. Herlach, C. Caldow, M.E. Monaco, R. Appeldoorn. 

2010. Coral reef ecosystems of Reserva Natural La Parguera (Puerto Rico): Spatial and temporal patterns in fish 

and benthic communities (2001-2007). NOAA Technical Memorandum NOS NCCOS 107. Silver Spring, MD. 202 

pp.
Coral reef ecosystems of Reserva natural de La Parguera (Puerto Rico): 

Spatial and temporal patterns in fish and benthic communities (2001-2007)

Simon J Pittman1,2,3, Sarah D Hile1,3, Christopher FG Jeffrey1,3, Randy Clark1, Kimberly Woody1,

Brook D Herlach1,3, Chris Caldow1

, Mark E Monaco1, Richard Appeldoorn4

1

 NOAA/National Ocean Service/National Centers for Coastal Ocean Science/Center for Coastal Monitoring and 

Assessment/Biogeography Branch

2

 Marine Science Center, University of the Virgin Islands, St. Thomas, U.S. Virgin Islands

3

 Consolidated Safety Services, Inc., Fairfax, Virginia, under NOAA Contract No. DG133C07NC0616

4 Department of Marine Sciences, University of Puerto Rico, Mayagüez, Puerto Rico

Biogeography Branch

Center for Coastal Monitoring and Assessment (CCMA) 

NOAA/NOS/National Centers for Coastal Ocean Science

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

Silver Spring, MD 20910

NOAA Technical Memorandum NOS NCCOS 107

April 2010

United States Department of 

Commerce

National Oceanic and 

Atmospheric Administration

National Ocean Service

Gary Locke

Secretary

Jane Lubchenco

Administrator

Jack Dunnigan

Assistant Administrator

Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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iii

About this Document

The report provides a spatial and temporal characterization of the fish and benthic communities of 

southwestern Puerto Rico, primarily within the La Parguera Natural Reserve. The reserve is a multiuse area that spans the continental shelf from the extensive mangrove forests fringing the shoreline 

to the complex shelfedge coral reefs that support a diverse and productive fish community. The coral 

reef ecosystem of La Parguera supports a locally important fishery, as well as recreational activities 

such as boating, snorkeling and diving. The data and synthesis in this report are intended to provide 

essential baseline biological information to support future management decision making. The project 

is a component of NOAA’s Caribbean Coral Reef Ecosystem Monitoring (CREM) project of NOAA’s 

Coral Reef Conservation Program (CRCP) and was conducted through an ongoing multi-agency 

collaboration between NOAA’s Center for Coastal Monitoring and Assessment Biogeography Branch 

(CCMA-BB), the University of Puerto Rico and the Puerto Rico Government’s Department of Natural 

and Environmental Resources (DNER).

This Technical Memorandum is part of a series of reports that focus on providing a quantitative spatial 

and temporal characterization of living marine resources and benthic communities associated with 

marine protected areas in the U.S. Caribbean. This project complements the National Coral Reef 

Ecosystem Monitoring Program’s (NCREMP) Coral Reef Ecosystem Monitoring grants awarded to 

DNER by CRCP. This project was funded by NOAA’s CRCP and National Centers for Coastal Ocean 

Science’s CCMA.

Related projects include:

Caribbean Coral Reef Ecosystem Monitoring

http://ccmaserver.nos.noaa.gov/ecosystems/coralreef/reef_fish.html

Development of Reef Fish Monitoring Protocols to Support the National Park Service Inventory and Monitoring Program

http://ccmaserver.nos.noaa.gov/ecosystems/coralreef/fish_protocol.html

Coral bleaching and recovery observed at Buck Island, St. Croix, U.S. Virgin Islands, October and December, 2005

http://ccma.nos.noaa.gov/ecosystems/coralreef/coral_bleaching.html

National Coral Reef Ecosystem Montoring Program

http://ccma.nos.noaa.gov/ecosystems/coralreef/coral_grant.html

Benthic Habitat Mapping of Puerto Rico and the U.S. Virgin Islands

http://ccma.nos.noaa.gov/ecosystems/coralreef/usvi_pr_mapping.html

Seafloor Characterization of the U.S. Caribbean - R/V Nancy Foster Missions

http://ccma.nos.noaa.gov/products/biogeography/usvi_nps/overview.html

All photographs provided in this document were taken by NOAA/NOS/NCCOS/Center for Coastal 

Monitoring Assessment Biogeography Branch in Puerto Rico unless otherwise indicated.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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iv

Project Team

Richard Appeldoorn (UPR)

Ivonne Bejarano (UPR)

Chris Caldow (NCCOS CCMA)

John Christensen (NCCOS CCMA)

Randy Clark (NCCOS CCMA)

Kimberly Edwards (NCCOS CCMA)

Brook Herlach (NCCOS CCMA)

Jamie Higgins (NCCOS CCMA)

Sarah Hile (NCCOS CCMA)

Chris Jeffrey (NCCOS CCMA)

Olaf Jensen (NCCOS CCMA)

Matt Kendall (NCCOS CCMA)

Tom McGrath (NCCOS CCMA)

Charles Menza (NCCOS CCMA)

Wendy Morrison (NCCOS CCMA)

Mark Monaco (NCCOS CCMA)

Simon Pittman (NCCOS CCMA)

Stephanie Williams (UPR)

Kimberly Woody (NCCOS CCMA)

Angel “Capitan” Nazario (Boat Captain, Aquanata)

Joel “Joeito” Rivera (Field Assistant, Aquanata)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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v Executive Summary

Executive Summary

Since 1999, NOAA’s Center for Coastal Monitoring and Assessment, Biogeography Branch (CCMA-BB) 

has been working with federal and territorial partners to characterize monitor and assess the status of 

the marine environment in southwestern Puerto Rico. This effort is part of the broader NOAA Coral Reef 

Conservation Program’s (CRCP) National Coral Reef Ecosystem Monitoring Program (NCREMP). With 

support from CRCP’s NCREMP, CCMA conducts the “Caribbean Coral Reef Ecosystem Monitoring 

project” (CREM) with goals to: (1) spatially characterize and monitor the distribution, abundance and 

size of marine fauna associated with shallow water coral reef seascapes (mosaics of coral reefs, 

seagrasses, sand and mangroves); (2) relate this information to in situ fine-scale habitat data and the 

spatial distribution and diversity of habitat types using benthic habitat maps; (3) use this information 

to establish the knowledge base necessary for enacting management decisions in a spatial setting; 

(4) establish the efficacy of those management decisions; and (5) develop data collection and data 

management protocols. The monitoring effort of the La Parguera region in southwestern Puerto Rico 

was conducted through partnerships with the University of Puerto Rico (UPR) and the Puerto Rico 

Department of Natural and Environmental Resources (DNER). Project funding was primarily provided 

by NOAA CRCP and CCMA.

In recent decades, scientific and non-scientific observations have indicated that the structure and 

function of the coral reef ecosystem in the La Parguera region have been adversely impacted by a 

wide range of environmental stressors. The major stressors have included the mass Diadema die 

off in the early 1980s, a suite of hurricanes, overfishing, mass mortality of Acropora corals due to 

disease and several coral bleaching events, with the most severe mass bleaching episode in 2005. 

The area is also an important recreational resource supporting boating, snorkeling, diving and other 

water based activities. With so many potential threats to the marine ecosystem several activities are 

underway or have been implemented to manage the marine resources. These efforts have been 

supported by the CREM project by identifying marine fauna and their spatial distributions and temporal 

dynamics. This provides ecologically meaningful data to assess ecosystem condition, support decision 

making in spatial planning (including the evaluation of efficacy of current management strategies) and 

determine future information needs. The ultimate goal of the work is to better understand the coral reef 

ecosystems and to provide information toward protecting and enhancing coral reef ecosystems for 

the benefit of the system itself and to sustain the many goods and services that it offers society. This 

Technical Memorandum contains analysis of the first seven years of fish survey data (2001-2007) and 

associated characterization of the benthos. The primary objectives were to quantify changes in fish 

species and assemblage diversity, abundance, biomass and size structure and to provide spatially 

explicit information on the distribution of key species or groups of species and to compare community 

structure across the seascape including fringing mangroves, inner, middle, and outer reef areas, and 

open ocean shelf bank areas.

Methods:

For each biannual survey mission, sample sites were selected via a stratified random design using 

hard and soft bottom habitat types delineated in NOAA’s benthic habitat map (Menza et al., 2006). Fish 

were surveyed during daylight hours along 25 m long by 4 m wide belt-transects for a fixed duration 

of 15 minutes. All species observed were identified to the lowest possible taxonomic level and their 

abundance was counted and grouped by size class. To quantify benthic habitat, five 1 m2

 quadrats were 

randomly placed along the transects and used to examine the relatively fine-scale biotic and abiotic 

components of the seascape (e.g., coral cover, macroalgal cover, etc.). A total of 1,167 fish surveys 

(572 from hardbottom, 437 from softbottom and 158 from mangrove habitats) were used in this analysis.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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vi Executive SummaryThe raw data were summarized by habitat type to examine differences in richness, abundance and 

biomass in different habitat types and were mapped across the study region to examine across shelf 

patterns in community and species attributes. Changes occurring between years and by season 

over the duration of the study were summarized and tested to detect periods with significant change. 

The data and synthesis provides essential baseline biological data via: 1) a comprehensive spatial 

characterization of resources and habitat condition, and 2) an examination of increases and decreases 

in the abundance of fish species and components of the benthic habitat across the study area.

Major findings: 

Diversity hotspots

• Highest coral cover was observed along the shelf edge southward of Terrumote I and around El Palo 

Reef. Cover of Acropora palmata (4.1-10.4%) was highest around El Palo Reef. Highest cover of 

Montastraea annularis complex (25.1-30.3%) was highest south-west of Turrumote I.

• Hot spots of coral species richness (10-14 species per 100 m2

) and diversity (H = 1.7-2.5) occurred 

offshore but were scattered throughout hard bottom habitat types. 

• Highest rugosity (0.81-1) occurred between Margarita Reef and El Palo Reef and at Romero. 

• Highest species richness of 41 fish per 100 m2

 area was recorded near the eastern shelf edge over 

colonized pavement with sand channels.

• Hotspots of high fish species richness, high fish biomass and high herbivore abundance and richness 

co-occurred along the shelf edge and around the complex of patch reefs between Margarita Reef 

and El Palo Reef.

Benthic habitat

• Hardbottom habitat types were dominated by algae, with an average of 5% live stony coral cover 

across the study area

• Montastraea and Porites spp. were the most common stony corals, with M. annularis complex 

comprising 35% of total coral cover, and Porites astreoides, the most frequently observed species 

occurring at 62% of hardbottom sites

• Acropora species were rarely observed, but Acropora cervicornis exhibited a wider spatial distribution 

than A. palmata

• Temporal analysis revealed that live coral cover varied significantly among some sampling years, but 

overall live coral cover decreased over the sampling period (2001-2007)

• Mangrove prop roots provided a substrate for colonization by a wide diversity of epiphytic organism

Fish

• Across the La Parguera study area a total of 210 fish species were identified to species level, with at 

least another 14 fish identified to genera

• Highest fish densities were associated with mangroves, with the assemblages composed mostly of 

juveniles
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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vii Executive Summary

• Twenty-five of the 30 most abundant fish species in mangroves were also observed over coral reefs 

indicating a high level of multi-habitat use, but with fish body length markedly smaller in mangroves 

than on coral reefs. This can be indicative of size dependent ontogenetic habitat shifts, particularly 

for grunts (Haemulidae), snapper (Lutjanidae), parrotfish (Scaridae) and barracuda (Sphyraenidae)

• Based on body size, mangroves appeared to function as an intermediate habitat type for some 

grunts and snappers, with smallest fish associated with seagrasses, larger fish in mangroves and 

the largest mean length recorded for fish on coral reefs

• The small-bodied parrotfish species, striped parrotfish (Scarus iseri) and redband parrotfish 

(Sparisoma aurofrenatum) were the most commonly occurring fish across the seascapes at La 

Parguera

• Although rainbow parrotfish (Scarus guacamaia) is thought to have a high dependence on mangroves 

and coral reefs, only two individuals were observed between 2001 and 2007 in the La Parguera region 

confirming the rarity of the species in the U.S. Caribbean and its status as vulnerable according to 

the IUCN red list of threatened species

• Sightings of many large-bodied fish species targeted by the fishery have declined substantially 

over the past 25 years based on comparison between 1980-1981 and 2001-2007. These include 

rainbow parrotfish, midnight parrotfish (Scarus coelestinus), Nassau grouper (Epinephelus striatus), 

tiger grouper (Mycteroperca tigris), rock hind (Epinephelus adscensionis), red hind (Epinephelus 

guttatus), coney (Cephalopholis fulva), graysby (Cephalopholis cruentata), yellowtail snapper 

(Ocyurus chrysurus), lane snapper (Lutjanus synagris), mahogany snapper (Lutjanus mahogoni), 

dog snapper (Lutjanus jocu) and queen triggerfish (Balistes vetula)

• The largest snappers seen in the study area were markedly smaller than the maximum known size for 

the species, particularly schoolmaster (Lutjanus apodus), O. chrysurus and gray snapper (Lutjanus 

griseus)

• From a total of 1,167 surveys (572 from hardbottom) over seven years, no M. tigris or yellowfin 

(Mycteroperca venenosa) were observed and only two E. striatus; two black grouper (Mycteroperca 

bonaci); two E. adscensionis and 43 E. guttatus were observed

• Small-bodied groupers including C. cruentata (n=246) and coney C. fulva (n=81) were significantly 

more abundant than large-bodied groupers

• None of the grouper species observed had attained the maximum known size for their species, with 

maximum length for Cephalopholis species in the La Parguera region estimated at 30 cm fork length 

(FL) compared with a maximum known for C. fulva of 41 cm total length (TL) and 43 cm TL for C. 

cruentata. The largest E. guttatus was approximately 50% of the maximum recorded size

• Grunt (Haemulidae) abundance and biomass was highest for mangroves in close proximity to coral 

reefs and seagrass beds. All common species of grunt showed a strong across shelf size-dependent 

distribution, with the majority of juveniles in lagoonal nearshore areas and adults in deeper mid- and 

outer-shelf zones. Juveniles and adults did also co-occur at several sites across the shelf indicating 

some flexibility in the strategy for ontogenetic segregation

• Five sharks (two species) and three stingrays were the only sharks and rays observed within transects 

at La Parguera between 2001 and 2007
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viii Executive Summary• Total fish biomass, herbivore biomass, grouper abundance, parrotfish and wrasse abundance were 

significantly higher in 2007 than in 2001

• Grouper biomass in 2007 was significantly lower than in 2001

• Only C. fulva and S. aurofrenatum increased in abundance consecutively for more than three years 

between 2001 and 2007

• Total snapper density and L. apodus density decreased from 2002 to 2005.

• The most striking inter-annual difference occurred between 2003 and 2004, whereby 65 metrics 

(approximately 80% of all metrics) decreased, with five decreasingly significantly; followed by 70% 

of metrics increasing the following year (2004-2005). A specific cause was not identified

• 65% of the 20 most abundant fish exhibited higher density in summer than in winter. Density of 

Thalassoma bifasciatum (bluehead wrasse) and Chromis cyanea (blue chromis) was more than 50% 

higher in summer. However, grunts, Haemulon aurolineatum (tomtate) and Haemulon flavolineatum

(french grunt), and S. iseri were more abundant in winter

• Mean biomass for grouper, snapper, parrotfish and grunts was lower in the winter than summer

• Planktivores and piscivores were more abundant in summer than winter

Recommendations:

Management plans are urgently required for the La Parguera Natural Reserve to effectively balance 

human resource usage with conservation objectives. Survey data from this study indicates that the coral 

reef ecosystem within and around the La Parguera Natural Reserve is impacted by multiple stressors, 

with low live coral cover, high macroalgal abundance and a depleted population of large-bodied species 

resulting in shifts in species dominance. 

The data collected through the NCREMP monitoring program and synthesized in this report together 

with related publications, CRES and UPR studies have made the La Parguera Natural Reserve one of 

the best studied regions in the Caribbean. 

These scientific survey data and associated map products provide a template of environmental data 

to support a process of marine zoning that could include conservation areas and ‘no-take’ reserves for 

selected areas, while maintaining access to other areas for human activities. In this report, diversity 

hotspots and distributions of key species have been identified and mapped and the population status of 

harvested species has been quantified, providing the prerequisites for the development of successful 

spatial management strategies. The shelfedge environment and the complex coral reef ecosystems 

between Margarita Reef and El Palo Reef support highest fish species diversity and high abundance 

for many species and should receive special management attention. Historical comparison, however, 

revealed that many of the large-bodied fish species that are targeted by the fishery have markedly 

declined in abundance since 1980. The very low abundance of large-bodied groupers and parrotfish is 

likely to have a major impact to ecosystem functioning and should receive priority attention. The shift 

towards declining live coral cover and increasing algal abundance over much of the region is usually 

associated with a degrading coral reef ecosystem. Mangroves are an important habitat type in this 

region and function as part of an interconnected mosaic of habitats for many fish species. Efforts to 

protect and restore mangroves, particularly those that are in close proximity to seagrasses and coral 

reefs will be beneficial to diversity and productivity. Restoration targets can be set for rebuilding depleted 

populations of large-bodied fish and for creating suitable water quality conditions to help restore benthic 
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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habitat structure and coral reef architecture. The Natural Reserve is a unique coral reef ecosystem in 

Puerto Rico, due to its relatively sheltered position on a wide and shallow section of the insular shelf 

of southern Puerto Rico, but like many such systems it is vulnerable to over-use and influenced by a 

rapidly changing local, regional and global environment. Comprehensive and strategic management is 

now urgently needed to restore ecological integrity to the La Parguera Natural Reserve and to ensure 

the long-term sustainability of the coral reef ecosystem for current and future generations.

References

Menza, C., J. Ault, J. Beets, C. Bohnsack, C. Caldow, J. Christensen, A. Friedlander, C. Jeffrey, M. 

Kendall, J. Luo, M.E. Monaco, S. Smith, and K. Woody. 2006. A guide to monitoring reef fish in the 

National Park Service’s South Florida/Caribbean Network. NOAA Technical Memorandum NOS NCCOS 

39. Silver Spring, MD. 166 pp. http://ccma.nos.noaa.gov/news/feature/FishMonitoring.html.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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x Executive SummaryResumen

Desde el 1999 el programa de Bio-geografía del Centro para Monitoreo y Evaluación Costera (CCMABB por sus siglas en inglés) de la Administración Nacional de Océanos y Atmosfera (NOAA por sus 

siglas en inglés) ha colaborado con agencias federales y territoriales para caracterizar, monitorear 

y evaluar la condición del ambiente marino del suroeste de Puerto Rico. Este esfuerzo es parte del 

Programa de Conservación de Arrecifes de Coral (CRCP por sus siglas en inglés) bajo el Programa 

Nacional de Monitoreo de Ecosistemas de Arrecifes de Coral (NCREMP por sus siglas en inglés) de 

NOAA. Con el apoyo del NCREMP del CRCP el CCMA lleva a cabo el proyecto titulado “Monitoreo de 

Ecosistemas de Arrecifes de Coral del Caribe” (CREM por sus siglas en inglés) cuyas metas incluyen: 

(1) caracterizar espacialmente y monitorear la distribución, abundancia y tamaño de la fauna asociada 

con los paisajes de arrecifes de coral (mosaico compuesto de arrecifes de coral, hierbas marinas, 

arenales y manglares); (2) relacionar esta información con datos de hábitat a pequeña escala y la 

distribución y diversidad de tipos de hábitat utilizando mapas de hábitat bénticos; (3) utilizar esta 

información para establecer una base de conocimiento sobre la cual formular medidas de manejo 

con un marco espacial; (4) evaluar la efectividad de medidas de manejo; y (5) desarrollar protocolos 

par la colección y distribución de datos de monitoreo. Los esfuerzos de monitoreo de la región de La 

Parguera, en el suroeste de Puerto Rico se llevaron a cabo en colaboración con la Universidad de 

Puerto Rico (UPR) y el Departamento de Recursos Naturales y Ambientales (DRNA) de Puerto Rico. 

La subvención de este proyecto proviene principalmente de NOAA CRCP y CCMA.

En tiempos recientes, observaciones científicas y no-científicas han indicado que la estructura y 

funcionamiento del ecosistema de arrecifes de coral de la región de La Parguera se han afectado por 

una gama de impactos ambientales. Algunos de estos impactos incluyen la mortandad masiva del 

erizo negro (Diadema) a principios de los 1980’s, varios huracanes, la sobre-explotación pesquera, 

mortandades significativas del coral Acropora por enfermedades y eventos de blanqueamientos, el 

mas severo de estos en el 2005. El área es también un importante recurso recreativo donde se llevan 

a cabo actividades relacionadas con el mar incluyendo la navegación, careteo o ‘snorkeling’ y buceo 

entre otras. Ante la variedad de amenazas potenciales al ecosistema marino se han implementado 

actividades para manejar el recurso marino. Estos esfuerzos han recibido el apoyo del proyecto CREM 

al identificar la dinámica temporal y la distribución espacial de la fauna marina. Este estudio provee 

datos ecológicos que sirven para evaluar la condición del ecosistema, apoyan la toma de decisiones en 

planificación espacial (incluyendo la evaluación de la efectividad de estrategias de manejo) y permite 

identificar las necesidades de información futuras. La meta fundamental de este trabajo es lograr 

entender el ecosistema de arrecifes de coral y proveer información para proteger y mejorar el mismo 

para el beneficio del ecosistema al igual que mantener los servicios y bienes que este provee a la 

sociedad. 

Este memorando técnico contiene un análisis de los primeros siete años (2001-2007) de censos 

de peces y caracterización béntica. El objetivo principal fue cuantificar cambios en la diversidad, 

abundancia, biomasa y estructura de tamaños de los peces y proveer información espacial explícita de 

la distribución de especies claves y grupos específicos. Esta información sirve a su vez para comparar 

la estructura de la comunidad a lo largo del paisaje marino compuesto de múltiples hábitats, incluyendo 

los manglares de franja, arrecifes en distintas posiciones de la plataforma y áreas de plataforma 

oceánica.

Métodos:

Para cada misión de muestreo semi-anual se seleccionaron sitios mediante un diseño aleatorio 

estratificado entre hábitats de fondos duros y blandos (arenosos) según el mapa de hábitat béntico de 

NOAA (Menza et al., 2006). Los censos de peces se llevaron a cabo durante el día en transectos de 
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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xi Executive Summary

banda de 25 por 4 metros durante 15 minutos. Todas las especies observadas se identificaron al nivel 

taxonómico mas bajo posible y las abundancias se cuantificaron en clases de tamaños. Para estimar 

la composición del hábitat béntico, cinco cuadratas de un metro cuadrado se colocaron al azar en 

el transecto y se cuantificaron los componentes bióticos y abióticos (por ejemplo cobertura de coral, 

algas, etc.). Para este análisis se utilizaron los datos de 1,167 censos de peces, de los cuales 572 

provienen de fondos duros y 595 de fondos blandos.

Los datos se resumieron por tipo de hábitat para examinar diferencias en la riqueza, abundancia y 

biomasa de peces en distintos hábitats bénticos y se mapearon a lo largo del área de estudio para 

examinar patrones en la distribución de los atributos de comunidades y especies. Se resumieron cambios 

entre años y por temporadas durante la duración del estudio para identificar mudanzas significativos 

entre periodos de muestreo. La síntesis de estos datos provee una base biológica mediante: 1) la 

caracterización espacial comprensiva de los recursos y la condición del hábitat y 2) los patrones de 

cambios en abundancias de peces y los componentes bénticos de hábitats en el área de estudio.

Resultados sobresalientes:

Centros de diversidad

• La cobertura de coral más alta se observo a lo largo del veril al sur de Cayo Turrumote I y en áreas 

circundantes al Cayo El Palo. La cobertura de Acropora palmata (4.1-10.4%) fue mayor en áreas de 

Cayo El Palo mientras la cobertura del complejo de Montastraea annularis (25.1-30.3%) fue mayor 

al suroeste de Cayo Turrumote I

• Puntos de alta riqueza de especies (10-14 especies por 100m2) y diversidad (H = 1.7-2.5) de corales 

se observaron lejos de la costa en distintas áreas de hábitats de fondos duros

• La rugosidad mayor (0.81-1) se observó en un área entre el Cayo Margarita y Cayo El Palo al igual 

que en Cayo Romero

• Se detectaron áreas donde la riqueza de especies de peces fue más alta (41 especies por 100m2) 

cerca del veril en hábitat de pavimento colonizado con canales de arena

• Puntos con alta riqueza de especies y alta biomasa de peces al igual que alta riqueza de especies 

y abundancia de peces herbívoros coincidieron a lo largo del veril y alrededor del grupo de arrecifes 

de parche localizados entre el Cayo Margarita y Cayo El Palo

Hábitat béntico

• Los hábitats duros tuvieron una cobertura béntica dominada por algas con un promedio de 5% de 

cobertura de coral vivo a lo largo del área de estudio

• Los hábitats duros tuvieron una cobertura béntica dominada por algas con un promedio de 5% de 

cobertura de coral vivo a lo largo del área de estudio

• Los corales más comunes pertenecen a los géneros Montastraea y Porites, donde M. annularis

compuso el 35% de la cobertura total y Porites astreoides se observó mas frecuentemente en 62% 

de los sitios de muestreo de hábitats duros

• Ejemplares del genero Acropora fueron observados muy pocas veces y entre estos Acropora 

cervicornis tuvo una distribución espacial más amplia que A. palmata
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xii Executive Summary• El análisis temporal revelo que la cobertura de coral vivo varió significativamente entre algunos 

años, pero en general declinó a lo largo del muestreo (2001-2007)

• Las raíces de mangles sumergidas proveen un sustrato béntico para la colonización de especies 

epífitas

Peces

• A lo largo del área de estudio en La Parguera se identificaron peces de 210 especies perteneciente 

a 14 géneros

• Las densidades más altas se encontraron en los manglares donde los juveniles dominaron el 

conjunto de peces 

• Veinticinco de las 30 especies más abundantes en los manglares también se observaron en los 

arrecifes de coral lo cual indica el uso de múltiples tipos de hábitats por estas especies. Además las 

tallas de peces en los manglares fueron menores a los arrecifes. Esto indica que están ocurriendo 

cambios ontogenéticos (a lo largo de su ciclo de vida) en los tipos de hábitat para las familias roncos 

(Haemulidae), pargos (Lutjanidae), loros (Scaridae) y picúas (Sphyraenidae)

• Basado en las tallas de los peces, los manglares aparentan tener una función de hábitat intermedio 

para algunos roncos y pargos, donde las tallas menores están asociadas a las hierbas marinas, las 

tallas intermedias en los manglares y el tamaño promedio mayor se encuentra en los arrecifes de 

coral

• Las especies más comunes a lo largo de todo el paisaje marino de La Parguera fueron los loros 

pequeños Scarus iseri y Sparisoma aurofrenatum

• Aunque se ha demostrado una alta dependencia de loro guacamayo (Scarus guacamaia) hacia los 

manglares y los arrecifes de coral, solamente se observaron dos individuos en La Parguera entre el 

2001 y 2007 lo cual revalida que la especie se considere escasa en el Caribe de Estados Unidos y 

vulnerable según la lista de especies amenazadas de la Unión Internacional para la Conservación 

de la Naturaleza (UICN)

• El avistamiento de varias especies de importancia comercial y de gran tamaño ha disminuido 

sustancialmente durante los pasados 25 años (basado en comparaciones entre 1980-81 y 2001-07) 

incluyendo a las siguientes especies: Scarus guacamaia, Scarus coeruleus, Epinephelus striatus, 

Mycteroperca tigris, Epinephelus adscencionis, Epinephelus guttatus, Cephalopholis cruentata, 

Cephalopholis fulva, Ocyurus chrysurus, Lutjanus synagris, Lutjanus mahogoni, Lutjanus jocu y

Balistes vetula

• Los pargos más grandes observados en La Parguera fueron de tallas menores al tamaño máximo 

reportado para estas especies, particularmente para Lutjanus apodus, O. chrysurus y Lutjanus 

griseus

• De la totalidad de 1,167 censos (572 en fondos duros) a lo largo de siete años no se observó 

ningún ejemplar de Mycteroperca tigris ni Mycteroperca venenosa y solamente se observaron 

dos ejemplares de E. striatus, Mycteroperca bonaci y E. adscensionis mientras se cuantificaron 

únicamente 43 Epinephelus guttatus

• Los meros de menor tamaño, incluyendo C. cruentata (n=246) y C. fulva (n=81) fueron 

significativamente mas abundantes que los meros de gran tamaño
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• Ninguna de las especies de meros observadas presentó el tamaño máximo reportado para su 

especie. Por ejemplo el tamaño máximo observado para especies de Cephalopholis en La Parguera 

se estimó en 30 cm (largo horquilla) aunque la talla máxima reportada para C. fulva es de 41 cm 

(largo total) y para C. cruentata es de 43 cm (largo total). La talla mayor observada en La Parguera 

para E. guttatus fue aproximadamente 50% del tamaño máximo reportado en la literatura científica

• La abundancia y biomasa de roncos (haemulidos) fue más alta en los manglares cercanos a arrecifes 

de coral y hierbas marinas. Todas las especies comunes de roncos demostraron un cambio en la 

distribución de tamaños que aumentó en sitios cercanos al veril, por ejemplo la mayoría de los 

juveniles se encontraron en lagunas cercanas a la costa y los adultos en áreas mas profundas de 

zonas intermedias y lejanas de la costa. Los juveniles y adultos coincidieron en varios lugares a lo 

largo de la plataforma lo cual indica alguna flexibilidad en su segregación ontogenética

• Solamente se observaron cinco tiburones (de dos especies) y tres rayas en La Parguera entre el 

2001 y 2007

• La biomasa total y la biomasa de herbívoros, al igual que la abundancia de meros, loros y lábridos 

fueron significativamente mayores en 2007 que durante el 2001

• La biomasa de meros fue significativamente menor el 2007 que durante el 2001

• Se observó un aumento consecutivo de más de tres años (entre 2001 y 2007) en la abundancia de 

C. fulva y S. aurofrenatum

• La densidad total de pargos y la densidad de L. apodus declinó entre 2002 y 2005

• La diferencia entre años más sobresaliente ocurrió entre 2003 y 2004 donde declinaron 65 métricas 

(aproximadamente 80% de todas las métricas), de las cuales cinco declinaron significativamente; 

seguido por un aumento de 70% de las métricas en el año siguiente (2004-2005) 

• Se observaron densidades más altas en verano que invierno para 65% de las 20 especies más 

abundantes. Por ejemplo la densidad de Thalassoma bifasciatum y Chromis cyanea fue 50% más 

alta durante el verano. Mientras tanto los roncos Haemulon aurolineatum y Haemulon flavolineatum

al igual que el loro S. iseri tuvieron mayores abundancias en temporada de invierno.

• La biomasa promedio para meros, pargos, loros y roncos fue menor en temporada de invierno en 

comparación con verano

• Los planctívoros y picívoros fueron más abundantes durante temporada de verano en comparación 

con invierno

Recomendaciones:

Para alcanzar un balance efectivo entre los usos del recurso marino por los humanos y los objetivos 

de conservación se requieren urgentemente unos planes de manejo para la Reserva Natural de La 

Parguera. Los datos del presente estudio indican que el ecosistema de arrecifes de coral dentro y 

alrededor de la Reserva Natural de La Parguera esta afectada por múltiples amenazas causando una 

baja cobertura de coral, mayor abundancia de algas y escasez de peces de gran tamaño lo cual resulta 

en una mudanza de las especies dominantes.
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Los datos colectados por el programa de monitoreo de N-CREMP y la síntesis de este informe al igual 

que otras publicaciones, e investigaciones de CRES y la UPR han establecido la Reserva Natural de 

La Parguera como una de las regiones más estudiadas del Caribe. Los resultados científicos al igual 

que los mapas proveen un borrador para comenzar un proceso de zonificación marina que puede 

incluir áreas de conservación, reservas marinas de no pesca, y garantizar el acceso a lugares para 

actividades recreativas. Los lugares de mayor diversidad y distribución de especies o grupos claves 

han sido identificados y cartografiados. Además se ha evaluado la condición de las poblaciones de 

especies de importancia comercial lo cual provee un pre-requisito para el desarrollo de estrategias 

de manejo y planificación espacial. El ambiente del veril y el ecosistema de arrecife de coral entre los 

cayos La Margarita y El Palo parecen tener mayor diversidad y abundancia para varias especies de 

peces lo cual merece atención especial por los manejadores. En una comparación histórica se destaca 

que la mayoría de los peces de importancia comercial de gran tamaño han declinado marcadamente 

en abundancia desde los años 80. Las abundancias bajas de los meros y loros de gran tamaño puede 

tener impactos a la función del ecosistema y deben recibir atención prioritaria. 

El cambio en la composición béntica hacia mayor cobertura de algas en vez de corales en la mayoría de 

la región esta asociado a una degradación del ecosistema de arrecife de coral. Los manglares son un 

hábitat importante en esta región y tiene una función en el mosaico de hábitats que están entrelazados 

por el uso de varias especies. Los esfuerzos por proteger y restaurar los manglares, particularmente 

aquellos que se encuentran cercanos a las hierbas y los arrecifes serán de mayor beneficio para la 

diversidad y la productividad. Se deben establecer metas para recuperar las poblaciones de peces 

de gran tamaño y crear condiciones optimas de calidad de agua para ayudar a restaurar el hábitat 

béntico y la arquitectura del arrecife de coral. La Reserva Natural de La Parguera es un ecosistema 

de arrecifes único en Puerto Rico por su posición relativamente protegida en la plataforma insular de 

Puerto Rico, pero como muchos otros sistemas es vulnerable al mal uso y es influenciado por cambios 

ambientales regionales y globales. Actualmente se destaca la necesidad urgente de unas estrategias 

de manejo comprensivo para restaurar la integridad ecológica de la Reserva Natural de La Parguera 

y asegurar la sustentabilidad a largo plazo del ecosistema de arrecifes de coral para generaciones 

actuales y venideras.

Referencias

Menza, C., J. Ault, J. Beets, C. Bohnsack, C. Caldow, J. Christensen, A. Friedlander, C. Jeffrey, M. 

Kendall, J. Luo, M.E. Monaco, S. Smith, and K. Woody. 2006. A guide to monitoring reef fish in the 

National Park Service’s South Florida/Caribbean Network. NOAA Technical Memorandum NOS NCCOS 

39. Silver Spring, MD. 166 pp. http://ccma.nos.noaa.gov/news/feature/FishMonitoring.html.
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Contents

i Executive Summary

ii Table of Contents

iii List of Tables and Figures

1 Introduction ............................................................................................................................................ 1

1.1. Introduction to the La Parguera study region...................................................................................... 1

1.2. Ecosystem change and the impact of multiple stressors .................................................................... 4

1.3. Benthic habitat mapping in southwest Puerto Rico............................................................................. 8

1.4. References........................................................................................................................................ 10

2 Benthic Composition ............................................................................................................................... 13

2.1 Introduction ....................................................................................................................................... 13

 2.2 Methods .......................................................................................................................................... 13

2.2.1 Survey Data ............................................................................................................................... 13

2.2.2 Analytical Methods..................................................................................................................... 14

2.3 Results .......................................................................................................................................... 17

2.3.1 Benthic habitat composition....................................................................................................... 17

 2.3.2 Spatial distribution patterns in benthic cover ............................................................................. 20

2.3.3 Temporal patterns in benthic composition (2001-2007)............................................................. 29

 2.3.4 Abundance and distribution of macroinvertebrates.................................................................... 36

 2.3.5 Marine debris ............................................................................................................................. 38

2.3.6 Summary of results.................................................................................................................... 39

 2.4 Discussion......................................................................................................................................... 40

 2.4.1 Colonized hardbottom habitats: benthic characterization and spatial patterns.......................... 40

 2.4.2 Softbottom habitats: benthic characterization and spatial patterns............................................ 42

2.4.3 Benthic characterization of mangrove habitats.......................................................................... 43

 2.4.4 Temporal trends in benthic composition on hardbottom habitats............................................... 44

 2.4.5 Marine debris ............................................................................................................................. 44

2.4.6 Macroinvertebrates .................................................................................................................... 45

2.5 References........................................................................................................................................ 49

3 Fish Communities, Groups and Species ............................................................................................... 55

3.1 Introduction ....................................................................................................................................... 55

 3.2 Methods .......................................................................................................................................... 56

3.2.1 Survey Data .............................................................................................................................. 56

3.2.2 Data analysis ............................................................................................................................. 57

3.3 Results .......................................................................................................................................... 59 

3.3.1 Spatial distribution patterns and species-habitat associations................................................... 59 

 3.3.2 Taxonomic groups...................................................................................................................... 68

 3.4 Inter-annual trend in fish metrics (2001-2007) ................................................................................ 132

3.4.1 Fish community metrics ........................................................................................................... 134 

 3.4.2 Taxonomic groups.................................................................................................................... 135

 3.5 Seasonal patterns in fish metrics (2004-2007)................................................................................ 144 

3.5.1 Community ......................................................................................................................... 144 

 3.5.2 Taxonomic groups.................................................................................................................... 147 

3.6 Historical comparison of species occurrence between 1980-1981 and 2001-2007........................ 152

3.7 Summary of results ......................................................................................................................... 154

3.7.1 Fish assemblage composition.................................................................................................. 154 

3.7.2 Multi-habitat use....................................................................................................................... 154

3.7.3 Spatial patterns of fish diversity, biomass and abundance ...................................................... 154
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3.7.4 Habitat and ontogenetic space use patterns............................................................................ 155

3.7.5 Temporal patterns in fish abundance and biomass.................................................................. 156

3.8 Discussion....................................................................................................................................... 156

3.8.1 Fish community composition and mapped habitat types ......................................................... 156

3.8.2 Multi-habitat utilization ............................................................................................................. 158

3.8.3 Spatial patterns of fish diversity, biomass and abundance ...................................................... 159

3.8.4 Habitat and ontogenetic space use patterns............................................................................ 159

3.8.5 Size structure and shifts in predators for vulnerable species................................................... 161

3.8.6 Temporal patterns in fish abundance and biomass.................................................................. 162

3.9 References...................................................................................................................................... 163

Appendix A .................................................................................................................................................... 167

Appendix B .................................................................................................................................................... 168

Appendix C .................................................................................................................................................... 173

Appendix D .................................................................................................................................................... 174
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List of Tables

Table 2.1. Abiotic and biological variables measured................................................................................. 13

Table 2.2. Number of hardbottom benthic habitat sites surveyed............................................................... 14

Table 2.3. The number of benthic habitat sites surveyed by mapped habitat type..................................... 15

Table 2.4. Summary statistics for hard coral species found in surveys ...................................................... 22

Table 2.5. Summary statistics for coral species found in hardbottom sites by habitat type........................ 23

Table 2.6. Results of non-parametric ordered comparisons to determine trends in percent live cover of 

the five most abundant coral genera in Winter 2001 through summer 2007. ............................ 29

Table 2.7. Results of non-parametric ANOVA to determine significant differences in percent cover of live 

coral among sampling periods within hardbottom habitat type.................................................. 29

Table 2.8. Results of non-parametric ordered comparisons to determine trends in percent live cover of 

coral within habitat types............................................................................................................ 30

Table 2.9. Results of non-parametric ANOVA to determine significant differences in percent live cover of 

coral genera among sampling periods within hardbottom habitat type...................................... 31

Table 2.10. Results of non-parametric ordered comparisons to determine trends in percent live cover of 

coral genera within habitat types ............................................................................................... 31

Table 2.11. Results of non-parametric ordered comparisons to determine trends in percent cover of algal 

types .......................................................................................................................................... 33

Table 2.12. Results of non-parametric ANOVA (Wilcoxon test) to determine significant differences in 

percent live cover of algal types among sampling periods within hardbottom habitat type ....... 33

Table 2.13. Multiple non-parametric pair-wise comparisons of mean percent algae cover.......................... 34

Table 2.14. Number of total surveys where conch (Eustrombus gigas) were observed............................... 36

Table 2.15. Abundance of long-spined sea urchins (Diadema antillarum) observed within transects.......... 38

Table 2.16. Abundance of Caribbean spiny lobster (Panulirus argus) observed within transects................ 38

Table 3.1. ANOSIM R values measuring fish assemblage similarities between samples grouped by 

benthic habitat TYPE ................................................................................................................. 59

Table 3.2. ANOSIM R values measuring fish assemblage similarities between samples grouped by 

benthic habitat MODIFIER......................................................................................................... 60

Table 3.3. Percentage occurrence, range of major habitat types used and mean of fork length by 

habitat type for the 20 most abundant fish species/groups observed using mangroves ........... 62

Table 3.4. Summary data on selected species from key fish families showing maximum size observed

in the study region and maximum known size for the species found......................................... 68

Table 3.5. Density and biomass for selected fish species and families (2001-2007) ............................... 132

Table 3.6. Inter-annual metrics with significant differences (p<0.05) in 2001 and 2007 comparisons. .... 134

Table 3.7. Inter-annual metrics with significant differences (p<0.05) in 2002 and 2007 comparisons. .... 134

Table 3.8. Density and biomass for community metrics (2001-2007)....................................................... 134

Table 3.9. Density and biomass for selected grouper species (2001-2007)............................................. 135

Table 3.10. Density and biomass for selected snapper species (2001-2007) ........................................... 136

Table 3.11. Density and biomass for selected jack species (2001-2007)................................................... 137

Table 3.12. Density and biomass for selected parrotfish species (2001-2007) .......................................... 138

Table 3.13. Density and biomass for selected grunt species (2001-2007)................................................. 140

Table 3.14. Density and biomass for selected surgeonfish species (2001-2007)....................................... 141

Table 3.15. Density and biomass for additional selected families (2001-2007).......................................... 142

Table 3.16. Density and biomass for additional selected species (2001-2007).......................................... 143

Table 3.17. Winter and summer total and mean density for the 20 most abundant fish species................ 144

Table A1. Finfish landings as a proportion of the total finfish landings reported for the U.S. Caribbean

in 1980. Listed are the most commonly landed species and species groups.......................... 167

Table B1. Fish species list and summary data on occurrence, abundance and biomass (2001-2007) ... 168

Table C1. Summary information on selected species from five key fish families showing maximum size

observed compared with maximum known size for the species.............................................. 173
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List of Figures

Figure 1.1. La Parguera study region showing the boundary of the Natural Reserve and the bathymetry

from airborne laser altimetry light and detection data (LiDAR) collected in 2006. ..........................2

Figure 1.2. Modeled precipitation based on rainfall data from 108 National Weather Service stations. ...........2

Figure 1.3. A 3-dimensional visualization of the LiDAR bathymetry for the La Parguera region.......................3

Figure 1.4. Aerial photography from 1936, 1963 and 2005 showing the growth of urban development in the 

La Parguera area and modifications to vegetation in the watersheds, coastline and islands.........4

Figure 1.5. A) Aerial photograph showing sediment runoff in the 1980s; B) The depth distribution of the

terrestrial sediment percentage and terrestrial mass accumulation rate.........................................5

Figure 1.6. A) Instantaneous sea surface temperature during one of the days that coincided with the 2005 

mass bleaching event recorded in the U.S. Caribbean. B) A space-time chart showing the SST

across the region between 1985 and 2006; C) A space-time chart showing degree heating 

weeks; D) Legend for space-time charts.........................................................................................6

Figure 1.7. Examples of perceived causes of fishing decline............................................................................7

Figure 1.8. NOAA’s benthic habitat map showing four levels of the hierarchical classification scheme: 

1) ZONE; 2) HABITAT; 3) TYPE and 4) MODIFIER. .......................................................................8

Figure 1.9. Surface rugosity calculated from the LiDAR bathymetry.................................................................9

Figure 2.1. NOAA diver recording benthic habitat composition.......................................................................13

Figure 2.2. Photo of marine debris ..................................................................................................................13

Figure 2.3. A selection of habitat types designated in the hierarchical classification scheme of NOAA’s 

benthic habitat map.......................................................................................................................14

Figure 2.4. Mean percent cover for key benthic components on hardbottom sites.........................................17

Figure 2.5. Mean percent cover for key components of benthic community on hardbottom habitat types .....17

Figure 2.6. Aerial mean percent cover of coral genera found across hardbottom sites ..................................18

Figure 2.7. Mean percent cover of coral genera by hardbottom habitat type..................................................18

Figure 2.8. Mean percent cover of key benthic components on submerged aquatic vegetation sites ............19

Figure 2.9. Mean percent cover of seagrass species observed on SAV sites ................................................19

Figure 2.10. Mean percent cover of key benthic components on unconsolidated sediment sites ....................19

Figure 2.11. Mean percent cover of seagrass species observed on unconsolidated sediment sites................20

Figure 2.12. Mean percent cover of key benthic components on benthic substrates at mangrove sites ..........20

Figure 2.13. Maps of the spatial and interpolated distributions for: live percent coral cover, benthic rugosity,

coral species richness and coral species diversity........................................................................21

Figure 2.14. Maps of the spatial and interpolated distributions for live percent cover for coral species: 

Montastraea annularis complex and Porites astreoides ...............................................................24

Figure 2.15. Maps of the spatial and interpolated distributions for live percent cover for coral species: 

Montastraea cavernosa, Agaricia species and Siderastrea siderea .............................................25

Figure 2.16. Maps of the spatial and interpolated distributions for live percent cover for coral species:

Acropora cervicornis and Acropora palmata .................................................................................26

Figure 2.17. Maps of the spatial and interpolated distributions for percent cover for: macroalgae, turf algae, 

soft coral cover and sponges. .......................................................................................................27

Figure 2.18. Maps of the spatial and interpolated distributions for percent cover for: seagrass, Thalassia

testudinum and Syringodium filiforme ...........................................................................................28

Figure 2.19. Seasonal and inter-annual patterns in mean percent live coral cover of on hardbottom habitats

over a seven year period...............................................................................................................29

Figure 2.20. Seasonal and inter-annual patterns in mean percent live coral cover on two hardbottom habitat

types over a seven year period .....................................................................................................30

Figure 2.21. Seasonal and inter-annual patterns in mean percent live cover of Montastraea spp. on 

hardbottom habitats ......................................................................................................................30

Figure 2.22. Seasonal and inter-annual patterns in mean percent live cover of three of the five most

abundant coral genera: a) Montastraea spp., b) Porites spp. and c) Diploria spp., on 

hardbottom habitat types ..............................................................................................................32

Figure 2.23. Seasonal and inter-annual patterns in mean percent live cover of three of the five most 

abundantcoral genera: a) Siderastrea spp. and b) Agaricia spp., on hardbottom habitat types ...33

Figure 2.24. Seasonal and inter-annual patterns in mean percent cover of algal categories: a) macroalgae,

b) turf algae, c) cyanobacteria/filamentous algae and d) crustose coralline algae on hardbottom

habitat types..................................................................................................................................35
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Figure 2.25. Distribution map of queen conch (Eustrombus gigas) ..................................................................36

Figure 2.26. Distribution maps of a) immature and b) mature queen conch (Eustrombus gigas).....................37

Figure 2.27. Abundance of a) immature and b) mature queen conch (Eustrombus gigas)...............................37

Figure 3.1. NOAA trained observer recording fish census data along the belt transect..................................56

Figure 3.2. NOAA’s benthic habitat map showing the major classes of habitat type.......................................57

Figure 3.3. Multi-dimensional scaling plot of between site similarity in fish assemblage composition. ...........59

Figure 3.4. Maps of the interpolated and spatial distributions for: total fish abundance, total fish biomass

and total species richness.............................................................................................................61

Figure 3.5. Comparison of mean values by habitat type for: total fish density, total fish biomass and

total species richness....................................................................................................................62

Figure 3.6. Maps of the interpolated and spatial distributions for herbivore: abundance, biomass and 

species richness............................................................................................................................63

Figure 3.7. Comparison of mean values by habitat type for: herbivore: density, biomass and species 

richness.........................................................................................................................................64

Figure 3.8. Comparison of mean values by habitat type for: piscivore: density, biomass and species

richness.........................................................................................................................................64

Figure 3.9. Maps of the interpolated and spatial distributions for piscivore: abundance, biomass and

species richness............................................................................................................................65

Figure 3.10. Comparison of mean values by habitat type for planktivore: density, biomass and species

richness.........................................................................................................................................66

Figure 3.11. Maps of the interpolated and spatial distributions for planktivore: abundance, biomass

and species richness.....................................................................................................................67

Figure 3.12. Comparison of mean density and biomass by habitat type for select grouper (Serranidae).........66

Figure 3.13. Maps of the interpolated and spatial distributions for select large-bodied grouper (Serranidae):

abundance and biomass. ..............................................................................................................69

Figure 3.14. Size frequency histogram for select grouper (Serranidae); graysby (C. cruentata), 

coney (C. fulva) and red hind (E. guttatus). ..................................................................................69

Figure 3.15. Maps of the interpolated and spatial distributions for graysby (C. cruentata): abundance

and biomass..................................................................................................................................70

Figure 3.16. Comparison of mean density and biomass by substrate type for graysby (C. cruentata).............70

Figure 3.17. Mean density for juvenile/subadult and adult by mapped habitat type for graysby

(C. cruentata) ................................................................................................................................71

Figure 3.18. Maps of the interpolated and spatial distributions for coney (C. fulva): abundance 

and biomass..................................................................................................................................72

Figure 3.19. Comparison of mean density and biomass by substrate type for coney (C. fulva). ......................72

Figure 3.20. Mean density for juvenile/subadult and adult by mapped habitat type for coney (C. fulva). .........73

Figure 3.21. Maps of the interpolated and spatial distributions for red hind (E. guttatus): abundance

and biomass..................................................................................................................................74

Figure 3.22. Comparison of mean density and biomass by substrate type for red hind (E. guttatus)...............74

Figure 3.23. Mean density for juvenile/subadult and adult by mapped habitat type for red hind

(E. guttatus)...................................................................................................................................75

Figure 3.24. Maps of the interpolated and spatial distributions for snapper (Lutjanidae): abundance

and biomass..................................................................................................................................76

Figure 3.25. Comparison of mean density and biomass by habitat type for snapper species (Lutjanidae). .....76

Figure 3.26. Size frequency histogram for select snapper (Lutjanidae): schoolmaster (L. apodus), gray

snapper (L. griseus), mahogany snapper (L. mahogoni),lane snapper (L. synagris), and 

yellowtail snapper (O. chrysurus)..................................................................................................77

Figure 3.27. Maps of the interpolated and spatial distributions for schoolmaster (L. apodus): abundance

and biomass..................................................................................................................................78

Figure 3.28. Comparison of mean density and biomass by substrate type for schoolmaster (L. apodus). ......78

Figure 3.29. Mean density for juvenile/subadult and adult by mapped habitat type for schoolmaster 

(L. apodus). ...................................................................................................................................79

Figure 3.30. Maps of the interpolated and spatial distributions for gray snapper (L. griseus): abundance

and biomass..................................................................................................................................80

Figure 3.31. Comparison of mean density and biomass by substrate type for gray snapper (L. griseus) ........80
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Figure 3.32. Mean density for juvenile/subadult and adult by mapped habitat type for gray snapper 

(L. griseus). ...................................................................................................................................81

Figure 3.33. Maps of the interpolated and spatial distributions for mahogany snapper (L. mahogoni):

abundance and biomass. ..............................................................................................................82

Figure 3.34. Comparison of mean density and biomass by substrate type for mahogany snapper

(L. mahogoni). ..............................................................................................................................82

Figure 3.35. Mean density for juvenile/subadult and adult by mapped habitat type for mahogany snapper

(L. mahogoni). ..............................................................................................................................83

Figure 3.36. Maps of the interpolated and spatial distributions for lane snapper (L. synagris): abundance

and biomass..................................................................................................................................84

Figure 3.37. Comparison of mean density and biomass by substrate type for lane snapper (L. synagris).......84

Figure 3.38. Mean density for juvenile/subadult and adult by mapped habitat type for lane snapper

(L. synagris). .................................................................................................................................85

Figure 3.39. Maps of the interpolated and spatial distributions for yellowtail snapper (O. chysurus):

abundance and biomass. ..............................................................................................................86

Figure 3.40. Comparison of mean density and biomass by substrate type for yellowtail snapper

(O. chysurus).................................................................................................................................86

Figure 3.41. Mean density for juvenile/subadult and adult by mapped habitat type for yellowtail snapper

(O. chysurus).................................................................................................................................87

Figure 3.42. Maps of the interpolated and spatial distributions for parrotfish (Scaridae): abundance and

biomass.........................................................................................................................................88

Figure 3.43. Comparison of mean density and biomass by habitat type for parrotfish species (Scaridae).......88

Figure 3.44. Size frequency histogram for select parrotfish (Scaridae): striped parrotfish (S. iseri), princess 

parrotfish (S. taeniopterus), redband parrotfish (S. aurofrenatum), yellowtail parrotfish

(S. rubripinne) and stoplight parrotfish (S. viride)..........................................................................89

Figure 3.45. Maps of the interpolated and spatial distributions for striped parrotfish (S. iseri): abundance

and biomass..................................................................................................................................90

Figure 3.46. Comparison of mean density and biomass by substrate type for striped parrotfish (S. iseri) .......90

Figure 3.47. Mean density for juvenile/subadult and adult by mapped habitat type for striped parrotfish

(S. iseri).........................................................................................................................................91

Figure 3.48. Maps of the interpolated and spatial distributions for princess parrotfish (S. taeniopterus):

abundance and biomass. ..............................................................................................................92

Figure 3.49. Comparison of mean density and biomass by substrate type for princess parrotfish 

(S. taeniopterus). ..........................................................................................................................92

Figure 3.50. Mean density for juvenile/subadult and adult by mapped habitat type for princess parrotfish

(S. taeniopterus)............................................................................................................................93

Figure 3.51. Maps of the interpolated and spatial distributions for redband parrotfish (S. aurofrenatum):

abundance and biomass ...............................................................................................................94

Figure 3.52. Comparison of mean density and biomass by substrate type in the southwest Puerto Rico

study area for redband parrotfish (S. aurofrenatum) ....................................................................94

Figure 3.53. Mean density for juvenile/subadult and adult by mapped habitat type for redband parrotfish

(S. aurofrenatum)..........................................................................................................................95

Figure 3.54. Maps of the interpolated and spatial distributions for bucktooth parrotfish (S. radians):

abundance and biomass ...............................................................................................................96

Figure 3.55. Comparison of mean density and biomass by substrate type for bucktooth parrotfish

(S. radians)....................................................................................................................................96

Figure 3.56. Comparison of mean density and biomass by substrate type for yellowtail parrotfish 

(S. rubripinne) ...............................................................................................................................97

Figure 3.57. Maps of the interpolated and spatial distributions for yellowtail parrotfish (S. rubripinne):

abundance and biomass ...............................................................................................................97

Figure 3.58. Maps of the interpolated and spatial distributions for stoplight parrotfish (S. viride): abundance 

and biomass..................................................................................................................................98

Figure 3.59. Comparison of mean density and biomass by substrate type for stoplight parrotfish (S. viride)...98

Figure 3.60. Mean density for juvenile/subadult and adult by mapped habitat type for stoplight parrotfish

(S. viride).......................................................................................................................................99
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Figure 3.61. Maps of the interpolated and spatial distributions for grunt (Haemulidae): abundance and

biomass.......................................................................................................................................100

Figure 3.62. Comparison of mean density and biomass by habitat type for grunt species (Haemulidae) ......101

Figure 3.63. Size frequency histogram for select grunts (Haemulidae): tomtate (H. aurolineatum), French

grunt (H. flavolineatum), white grunt (H. plumierii) and bluestriped grunt (H. sciurus)................101

Figure 3.64. Maps of the interpolated and spatial distributions for tomtate (H. aurolineatum): abundance

and biomass................................................................................................................................102

Figure 3.65. Comparison of mean density and biomass by substrate type for tomtate (H. aurolineatum)......102

Figure 3.66. Mean density for juvenile/subadult and adult by mapped habitat type for tomtate

(H. aurolineatum) ........................................................................................................................103

Figure 3.67. Maps of the interpolated and spatial distributions for French grunt (H. flavolineatum):

abundance and biomass. ............................................................................................................104

Figure 3.68. Comparison of mean density and biomass by substrate type for French grunt 

(H. flavolineatum)........................................................................................................................104

Figure 3.69. Mean density for juvenile/subadult and adult by mapped habitat type for French grunt

(H. flavolineatum). .......................................................................................................................105

Figure 3.70. Maps of the interpolated and spatial distributions for white grunt (H. plumierii): abundance

and biomass................................................................................................................................106

Figure 3.71. Comparison of mean density and biomass by substrate type in the southwest Puerto Rico

study area for white grunt (H. plumierii). ....................................................................................106

Figure 3.72. Mean density for juvenile/subadult and adult by mapped habitat type for white grunt

(H. plumierii)................................................................................................................................107

Figure 3.73. Maps of the interpolated and spatial distributions for bluestriped grunt (H. sciurus):

abundance and biomass .............................................................................................................108

Figure 3.74. Comparison of mean density and biomass by substrate type for bluestriped grunt

(H. sciurus)..................................................................................................................................108

Figure 3.75. Mean density for juvenile/subadult and adult by mapped habitat type for bluestriped grunt

(H. sciurus)..................................................................................................................................109

Figure 3.76. Maps of the interpolated and spatial distributions for surgeonfish (Acanthuridae): abundance

and biomass................................................................................................................................ 110

Figure 3.77. Comparison of mean density and biomass by habitat type for surgeonfish (Acanthuridae) ....... 110

Figure 3.78. Size frequency histogram for surgeonfish (Acanthuridae): ocean surgeon (A. bahianus), 

doctorfish (A. chirurgus) and blue tang (A. coeruleus)................................................................ 111

Figure 3.79. Maps of the interpolated and spatial distributions for ocean surgeonfish (A. bahianus):

abundance and biomass ............................................................................................................. 112

Figure 3.80. Comparison of mean density and biomass by substrate type for ocean surgeonfish

(A. bahianus). ............................................................................................................................. 112

Figure 3.81. Maps of the interpolated and spatial distributions for doctorfish (A. chirurgus): abundance

and biomass................................................................................................................................ 113

Figure 3.82. Comparison of mean density and biomass by substrate type for doctorfish (A. chirurgus) ........ 113

Figure 3.83. Maps of the interpolated and spatial distributions for blue tang (A. coeruleus): abundance

and biomass................................................................................................................................ 114

Figure 3.84. Comparison of mean density and biomass by substrate type for blue tang (A. coeruleus) ........ 114

Figure 3.85. Size frequency histogram for goatfish (Mullidae): yellow goatfish (M. martinicus) and spotted 

goatfish (P. maculatus)................................................................................................................ 115

Figure 3.86. Maps of the interpolated and spatial distributions for yellow goatfish (M. martinicus):

abundance and biomass. ............................................................................................................ 116

Figure 3.87. Comparison of mean density and biomass by substrate type for yellow goatfish

(M. martinicus) ............................................................................................................................ 116

Figure 3.88. Maps of the interpolated and spatial distributions for spotted goatfish (P. maculatus):

abundance and biomass. ............................................................................................................ 117

Figure 3.89. Comparison of mean density and biomass by substrate type for spotted goatfish

(P. maculatus). ............................................................................................................................ 117

Figure 3.90. Maps of the interpolated and spatial distributions for jack (Carangidae): abundance and

biomass....................................................................................................................................... 118

Figure 3.91. Comparison of mean density and biomass by habitat type for jack (Carangidae)...................... 118
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Figure 3.92. Comparison of mean density and biomass by substrate type for blue runner (C. crysos).......... 119

Figure 3.93. Comparison of mean density and biomass by substrate type for bar jack (C. ruber) ................. 119

Figure 3.94. Maps of the interpolated and spatial distributions for yellowtail damselfish (M. chrysurus)

abundance and biomass. ............................................................................................................120

Figure 3.95. Comparison of mean density and biomass by substrate type for yellowtail damselfish

(M. chrysurus) .............................................................................................................................120

Figure 3.96. Maps of the interpolated and spatial distributions for dusky damselfish (S. adustus)

abundance and biomass .............................................................................................................121

Figure 3.97. Comparison of mean density and biomass by substrate type for dusky damselfish

(S. adustus).................................................................................................................................121

Figure 3.98. Maps of the interpolated and spatial distributions for beaugregory (S. leucostictus)

abundance and biomass .............................................................................................................122

Figure 3.99. Comparison of mean density and biomass by substrate type for beaugregory 

(S. leucostictus)...........................................................................................................................122

Figure 3.100. Maps of the interpolated and spatial distributions for bicolor damselfish (S. partitus)

 abundance and biomass ............................................................................................................123

Figure 3.101. Comparison of mean density and biomass by substrate type for bicolor damselfish

 (S. partitus).................................................................................................................................123

Figure 3.102. Maps of the interpolated and spatial distributions for threespot damselfish (S. planifrons)

 abundance and biomass ............................................................................................................124

Figure 3.103. Comparison of mean density and biomass by substrate type for threespot damselfish

 (S. planifrons).............................................................................................................................122

Figure 3.104. Maps of the interpolated and spatial distributions for cocoa damselfish (S. variabilis)

 abundance and biomass ............................................................................................................125

Figure 3.105. Comparison of mean density and biomass by substrate type cocoa damselfish

 (S. variabilis) ..............................................................................................................................125

Figure 3.106. Maps of the interpolated and spatial distributions for wrasse (Labridae) abundance and

 biomass......................................................................................................................................126

Figure 3.107. Comparison of mean density and biomass by substrate type for wrasse (Labridae).................126

Figure 3.108. Maps of the interpolated and spatial distributions for porgy (Sparidae) abundance and

 biomass......................................................................................................................................127

Figure 3.109. Comparison of mean density and biomass by substrate type for porgy (Labridae)...................127

Figure 3.110. Map of the spatial distributions for sharks and rays: abundance and biomass..........................128

Figure 3.111. Comparison of mean density and biomass by substrate type for sharks and rays ....................128

Figure 3.112. Maps of the interpolated and spatial distributions for queen triggerfish (B. vetula) abundance

 and biomass...............................................................................................................................129

Figure 3.113. Comparison of mean density and biomass by substrate type queen triggerfish (B. vetula) ......129

Figure 3.114. Maps of the interpolated and spatial distributions for great barracuda (S. barracuda)

 abundance and biomass ............................................................................................................130

Figure 3.115. Comparison of mean density and biomass by substrate type for great barracuda

 (S. barracuda). ..........................................................................................................................130

Figure 3.116. Size frequency histogram for queen triggerfish (B. vetula) and great barracuda

 (S. barracuda) ............................................................................................................................131

Figure 3.118. Seasonal change in mean herbivore: density, biomass and richness........................................145

Figure 3.119. Seasonal change in mean piscivore: density, biomass and richness.........................................146

Figure 3.120. Seasonal change in mean planktivore: density, biomass and richness. ....................................147

Figure 3.121. Seasonal change in mean fish biomass for : groupers, snappers, parrotfish and grunts ..........148

Figure 3.122. Seasonal change in mean grouper biomass for: graysby (C. cruentata), coney (C. fulva) and

red hind (E. guttatus)..................................................................................................................148

Figure 3.123. Seasonal change in mean snapper biomass for: schoolmaster (L. apodus), gray (L. griseus)

and yellowtail snapper (O. chrysurus)........................................................................................149

Figure 3.124. Seasonal change in mean parrotfish biomass for: striped (S. iseri), princess 

(S. taeniopterus), redband (S. aurofrenatum) and stoplight parrotfish (S. viride) ......................150

Figure 3.125. Seasonal change in mean grunt biomass for: French (H. flavolineatum), white (H. plumierii)

and bluestriped grunt (H. sciurus). .............................................................................................151
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Figure 3.126. Seasonal changes in mean goatfish biomass for: yellow (M. martinicus) and spotted goatfish

(P. maculatus).............................................................................................................................151

Figure 3.127. Seasonal changes in mean biomass for: queen triggerfish (B. vetula) and great barracuda

(S. barracuda) ............................................................................................................................152

Figure 3.128. A spearfisherman with a rainbow parrotfish (Scarus guacamaia) caught in Puerto Rico...........152

Figure E1. Spatial predictive map of fish species richness across the seascapes of La Parguera..............177
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1 1 - Introduction

1. Introduction

Since 1999, NOAA’s Center for Coastal Monitoring and 

Assessment Biogeography Branch (CCMA-BB) has been 

working with federal and territorial partners to characterize, 

monitor and assess the status of the marine environment 

in southwestern Puerto Rico. This effort is part of the 

broader NOAA Coral Reef Conservation Program’s 

(CRCP) National Coral Reef Ecosystem Monitoring 

Program (NCREMP). With support from CRCP’s NCREMP, 

CCMA conducts the “Caribbean Coral Reef Ecosystem 

Monitoring project” (CREM) with goals to: (1) spatially 

characterize and monitor the distribution, abundance, and 

size of marine fauna associated with shallow water coral 

reef seascapes (mosaics of coral reefs, seagrasses, sand 

and mangroves); (2) relate this information to in situ finescale habitat data and the spatial distribution and diversity 

of habitat types using benthic habitat maps; (3) use this 

information to establish the knowledge base necessary 

for enacting management decisions in a spatial setting; 

(4) establish the efficacy of those management decisions; 

and (5) develop data collection and data management 

protocols. Since 2002, CREM surveys have contributed 

to the Coral Reef Ecosystem Studies (CRES) program in 

the La Parguera region. CRES was a five-year research 

program funded through NOAA’s Center for Sponsored Coastal Ocean Research and coordinated by 

the Department of Marine Sciences of University of Puerto Rico (UPR) to define and understand causes 

and effects of reef degradation, and provide managers with information and tools to aid in reversing 

the degradation of U.S. Caribbean coral reef ecosystems. The monitoring effort for the La Parguera 

region was conducted through partnerships with UPR and the Puerto Rico Department of Natural and 

Environmental Resources (DNER). The intention of this report is to provide a comprehensive spatial 

and temporal characterization to support management decision making and baseline data to facilitate 

the development of an effective adaptive management plan for the Reserva Natural La Parguera.

1.1. Introduction to the La Parguera study region

The coral reef ecosystem of the La Parguera region of southwestern Puerto Rico is a complex spatial 

mosaic of habitat types dominated by coral reefs, seagrasses, macroalgal beds, unconsolidated 

sediments and mangroves. The broad shelf and coastal embayment at La Parguera provides a sheltered 

shallow-water environment that has facilitated the development of a diverse and productive seascape, 

with important ecological, economic and cultural value. The mangrove forests of the La Parguera region 

are some of the most extensive in Puerto Rico and have been protected as insular forests since 1943 

(Aguiler-Perera et al., 2006). In 1979, the La Parguera region (327 km2

) was designated as a Natural 

Reserve (NR), known as the Reserva Natural La Parguera, becoming the second marine protected 

area to be established (after Aguirre in 1918) in Puerto Rico. The La Parguera NR is managed by 

DNER Bureau of Coastal Reserves and Refuges (BCRR) as a multiple use zone and currently has no 

restrictions on fishing and no formal management plan. In the 1980s, La Parguera was proposed as a 

potential multi-use marine sanctuary under the National Marine Sanctuaries Act, but the proposal was 

abandoned due to insufficient local agreement (Valdés-Pizzini, 1990; Aguiler-Perera et al., 2006). In 

addition, a small “no-take” marine reserve (7.6 km2

) has been proposed for Turrumote, south of Corral 

(Figure 1.1).

La Parguera, Puerto Rico

Diver collecting benthic habitat composition data
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2 1 - Introduction

Figure 1.1. La Parguera study region in southwest Puerto Rico showing the boundary of the 

Natural Reserve and the bathymetry from airborne laser altimetry light and detection data (LiDAR) 

collected in 2006. Several prominent coral reefs, islands and mangrove cays are labeled.

Figure 1.2. Modeled precipitation based on rainfall data from 108 National Weather Service stations. Lowest annual precipitation 

is recorded for the La Parguera region with 770 mm yr-1. Source: http://www.climatesource.com/pr/fact_sheets/prppt_xl.jpg
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3 1 - Introduction

The well-developed coral reef ecosystem at La Parguera exists in an area with relatively low human 

population size and coastal development, an absence of local rivers and the lowest rainfall in Puerto 

Rico (Figure 1.2). 

Marine geological studies have revealed that the extensive coral 

reefs along the 8-10 km insular shelf developed over the past 

10,000 years and were originally dominated by branching corals 

(Acropora palmata) in deeper shelf edge waters (Hubbard et al., 

2008). Then in the Holocene period, reef development tracked 

sea level rise shoreward to create the distinct across shelf 

spatial arrangement of coral reef ecosystems observed today. 

This process led Hubbard et al. (2008) to classify coral reefs 

across the shelf at La Parguera into inner shelf reefs (some of 

which support mangrove stands), mid-shelf reefs and shelf edge 

reefs. The classification of inner, mid and outer shelf zones was 

adopted in this report to describe the spatial patterns in fish distribution across the shelf. Figure 1.3 

shows examples of the complex structural features that exist across the shelf at La Parguera including 

collections of patch reefs, mangrove islands and shelfedge ridges and valleys.

Mangrove islands of La Parguera, PR

1

1

2

3

2

3

Figure 1.3. A 3-dimensional visualization of the LiDAR bathymetry for the La Parguera region showing (1) the ridges and valleys along the 

shelf edge; (2) the highly heterogeneous patch reefs that exist around El Palo reef, and (3) the distinctive mangrove cays that separate the 

lagoonal areas from the mid shelf zone.
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4 1 - IntroductionTides in the study area are diurnal and of low amplitude (<25 cm). The circulation of water is generally 

westward (Ojeda, 2002), with velocities of 10-15 cm/s (Hensley et al., 1994). Internal waves breaking 

on the shelf edge create nutrient pulses and periodically reduce water temperature. Offshore winds can 

cause an eastward subsurface current on the La Parguera shelf (Tyler and Sanderson, 1996).

1.2 Ecosystem change and the impact of multiple stressors

Until the past few decades, La Parguera coral reef ecosystems 

had been subjected to relatively few land-based stressors, but 

increasing deforestation of the coastal limestone hills, coastal 

resort development and housing have been linked to increased 

sedimentation and nutrient input to the system (Warne et al., 

2005; Ryan et al., 2008). A time series of aerial photographs 

from 1936, 1963 and 2005 showing the La Parguera town and 

surrounding hills reveals an increase in urban development, 

roads and modification to the coastline, watershed and coastal 

vegetation (Figure 1.4). 

Multiple interacting stressors including sedimentation, 

nutrient runoff, elevated sea water temperature and 

fishing are changing the structure and function of 

coral reef ecosystems of La Parguera (Garcia-Sais et 

al., 2005, 2008; Ballantine et al., 2008). Deterioration 

in water quality due largely to increased nutrients and 

turbidity as a result of land-based sources has been 

reported as a primary threat to nearshore coral reef 

ecosystems in South West Puerto Rico (Garcia-Sais 

et al., 2005, 2008). Bush (1991) estimated that 90 

percent of the river sediment discharged at the coast 

is transported to the shelf edge and slope within a 

few months due to dominant across-shelf processes. 

Storm events can also redistribute sediment. 

Suspended sediment yields during the passage of 

Hurricane Georges (September 20-25, 1998) were 

highest in the watersheds on the south and southwest 

coast (Rio Portugues) yielding 7,800 tonnes/km2

and Rio Jacquas yielding 3,700 tonnes/km2

 (Larsen 

and Webb, 2009). Sediment sampling (Figure 1.5) 

has demonstrated an overall doubling of terrestrial 

material in the marine sediments of backreef areas 

of La Parguera over the last century (Ryan et al., 

2008). In southwest Puerto Rico, turbidity is a strong 

positive linear predictor of live coral cover along the 

shelf (Bejarano-Rodríguez, 2006) and has resulted in 

a compression of coral depth zonation, accompanied 

by changes in the relative abundance of coral species, 

which is directly related to individual species tolerance 

to sediment stress (Acevedo et al.,1989). Significant 

environmental changes have been observed, but 

relatively little is known about the interaction between 

ocean circulation, water clarity and the resilience of 

biotic communities across the shelf. 

Coastal development along mangrove lined coastline at La 

Parguera

Figure 1.4. Aerial photography from 1936, 1963 and 2005 

showing the growth of urban development in the La Parguera 

area and modifications to vegetation in the watersheds, 

coastline and islands. Source: http://www.log.furg.br/WEBens/

morelock/clnpho.htm#par.
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5 1 - Introduction

Figure 1.5. A) Aerial photograph showing sediment runoff in the 1980s along the developed shoreline at La Parguera. Source: http://

www.log.furg.br/WEBens/morelock/clnpho.htm#par. B) The depth distribution of the terrestrial sediment percentage and terrestrial mass 

accumulation rate in a sediment core collected from a back-reef setting near Corral Reef, La Parguera. A gradual increase in both variables 

is observed with depth and time, with an overall doubling of terrestrial material over the last century. Source: Ryan et al., 2008.

In addition, disease and bleaching have led to substantial deterioration of coral reefs in the past two 

decades. White plague II and Caribbean yellow band diseases (YBD) are the most widespread and 

damaging diseases affecting scleractinian corals in Puerto Rico. 

White plague was first reported from La Parguera in 1995 (Bruckner 

and Bruckner, 1997) and since 1999 has been reported with 

increasing frequency from both shallow and deep coral reefs across 

the shelf at La Parguera. Disease, particularly white band, has 

caused a decline in A. palmata since it was first reported in the early 

1980s (Davis et al., 1986). For more details on monitoring programs 

and results for the region see data in Garcia-Sais et al. (2008). Mass 

bleaching events have occurred periodically in the Caribbean, when 

anomalously high sea surface temperatures (SST) have persisted 

for extended periods of time, usually in late summer and fall. At 

least seven mass bleaching events have been reported in Puerto 

Rico and the U.S. Virgin Islands (USVI) since 1985, four of which 

occurred between 1998 and 2006 (Figure 1.6). Assessment of the 

2005 bleaching event in southwest and west Puerto Rico revealed 

that 65% of 4,000 corals examined exhibited bleaching, with highest 

occurrence of bleaching found in La Parguera (42% fully bleached, 

31% partially bleached; Garcia-Sais et al., 2008). In Puerto Rico, 

the combined effect of the 2005 bleaching event and subsequent 

disease, led to a failure of sexual reproduction in Acropora spp. and 

Montastraea spp. in 2006 (Ballantine et al., 2008).

Bleached Siderastrea sidera colony

Bleached Montastraea sp. colony

A B
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6 1 - Introduction

Figure 1.6. A) Shows instantaneous sea 

surface temperature (SST) during one 

of the days that coincided with the 2005 

mass bleaching event recorded in the U.S. 

Caribbean (Clark et al., 2009). The SST 

data were recorded by the Advanced Very 

High Resolution Radiometer (AVHRR). 

B) A space-time chart showing the SST 

across the region between 1985 and 

2006. Wider orange and red horizontal 

bands indicate greater persistence of high 

SST and alignment with bleaching records 

show that bleaching events typically occur 

where high water temperatures persist for 

weeks or months. C) A space-time chart 

showing degree heating weeks (DHW), 

a measure of cumulative thermal stress. 

For example, if the current temperature 

is above the maximum expected 

summertime temperature for a period 

of two weeks, the site would receive a 

rating of 2 DHWs. D) Legend for spacetime charts. Source: SST data processed 

by Varis Ransi of NOAA CCMA Remote 

Sensing Group.

A

B C D
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7 1 - Introduction

Reduction in the abundance of many of the key predators and herbivores on coral reef ecosystems 

through deterioration of habitat, disease and extraction by the fishery is likely to have a major influence 

on ecosystem integrity and ecological processes, but the overall ecological consequences for coral 

reef ecosystem structure and function in southwest Puerto Rico are not known. The integrity of 

coral reef ecosystems and the sustainability of commercially exploited fish species are key resource 

management concerns. Substantial reductions in commercial fish catches have been recorded for coral 

reef associated fishes over the past two decades (Matos-Caraballo, 2004; Garcia-Sais et al., 2005). 

Landings of demersal reef fishes peaked in 1979 at 2,400 metric tonnes (mt) then declined to 394 mt in 

1988 (Appeldoorn and Myers, 1993). Spiny lobster (Panulirus argus) landings followed a similar trend 

(Bohnsack et al., 1991). In addition, the fishery has shown many of the other classic signs of overfishing 

which include shifts to catching smaller sized individuals and recruitment failures (Matos-Caraballo, 

2004). This has occurred even though the number of active commercial fishers has decreased from 

1,758 in 1996 to 1,163 in 2002. Surveys with fishers reveal that the insular shelf is the most heavily 

fished area, with coral reef associated fish being the primary targets. For instance, 85% of the fishers 

target resources on the continental shelf and 94% target “reef fishes”. Fishing pressure on the southern 

shelf increased from 70% in 1996 to 83% in 2002, but the fishing on the shelf edge decreased in 

the same period. A decrease in net use and traps has occurred and an increase in “hook-and-line” 

between 1996 and 2002 and most fishers now use a wide range of gears to diversify their catch. In 

2002, the La Parguera region had the highest number of fishers on the south coast with a total of 26 full 

time and 37 part time utilizing 47 fishing vessels (Matos-Caraballo, 2004). Seventy-four percent of the 

fishers from the La Parguera region believe that the fishery has worsened over time with the perceived 

causes being overfishing (31%), pollution (13%) and habitat destruction (11%; Figure 1.7). The shallow 

water reef fish fishery in Puerto Rico primarily targets grunts (Haemulidae), groupers (large-bodied 

Serranidae), goatfish (Mullidae), parrotfish (Scaridae) and snappers (Lutjanidae; Caribbean Fishery 

Management Council [CFMC], 1985; see Appendix A). Triggerfish, squirrelfish, hogfish, porgies and 

trunkfish combined represent approximately 15% of the total catch (Appendix A).

a b

c d

Figure 1.7. Examples of perceived causes of fishing decline according to the majority of fishers from the La Parguera region. a) fishing 

pressure; b) noise/recreation; c) habitat degregation, and d) habitat destruction. 
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8 1 - IntroductionA recent numerical evaluation of the sustainability of a wide range of important fishery species in 

Puerto Rico indicated that the majority of species are fished at unsustainable levels (Ault et al., 2008). 

Of the 25 fish species assessed, 16 were below the spawning potential ratio of 30%, implying that the 

stocks are no longer sustainable at current exploitation levels. Therefore, the Ault et al. (2008) study 

using length-based assessment corroborates local fishers knowledge. At time of writing, only three 

species (Nassau grouper, Epinephelus striatus, goliath grouper, Epinephelus itajara, and queen conch, 

Strombus gigas) were designated as overfished by NOAA’s National Marine Fisheries Service (NMFS), 

but under the Sustainable Fisheries Act, National Standard 1 Amendments, the yellowtail snapper 

(Ocyurus chrysurus) and several large-bodied grouper including misty (Epinephelus mystacinus), red 

(Epinephelus morio), yellowedge (Epinephelus flavolimbatus), tiger (Mycteroperca tigris) and yellowfin 

(Mycteroperca venenosa) groupers should also be designated as overfished, while several species of 

parrotfish, triggerfishes and boxfishes would be considered “at risk” (Appeldoorn, 2008).

1.3. Benthic habitat mapping 

in southwest Puerto Rico

CCMA BB initiated benthic 

mapping activities in 1998 and 

developed a benthic habitat map 

with a 1 acre (4,000 m2) minimum 

mapping unit (MMU) that used a 

hierarchical classification scheme 

of 21 distinct benthic habitat types 

within eight geomorphological 

zones (Figure 1.8). Habitat classes 

and boundaries were delineated 

based on visual interpretation of 

aerial photography and extensive 

ground-truthing (Kendall et al., 

2001). The classification scheme 

included ZONE (geomorphological 

zone: lagoon, bank, etc.); 

HABITAT (broad substratum 

type: uncolonized sediments, 

submerged vegetation, colonized 

hardbottom, etc.); TYPE 

(mangrove, seagrass, linear reef, 

etc.); MODIFIERS (percentage 

cover of aquatic vegetation: patchy/

continuous or <30% seagrass, 

etc.); and DESCRIPTORS (a 

combined description of HABITAT, 

TYPE and MODIFIER). The zone/

habitat approach to classifying 

benthic structure was developed 

in consultation with CFMC, Dr. 

Kenyon Lindeman and CCMA 

BB to support multiple needs 

in resource management and 

ecosystem-based science. A 

relatively large “UNKNOWN” area 

remained unclassified due to poor Figure 1.8. NOAA’s benthic habitat map showing four levels of the hierarchical classification 

scheme: 1) ZONE; 2) HABITAT; 3) TYPE and 4) MODIFIER. Source: Kendall et al., 2001.

N a)

b)

c)

d)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

9 1 - Introduction

light penetration in the water column. In 2005, spectral analysis of Landsat Thematic Mapper data 

(CCMA-BB) and locally available side-scan sonar data (Triana, 2002) was used to classify the deeper 

water “unknown” area to provide a complete coverage for the La Parguera study site. The benthic 

habitat map includes coral reef ecosystems to a depth of approximately 35 m and therefore does not 

provide benthic information for the large proportion of the La Parguera NR that exists in deeper (>100 

to >1,500 m) shelf edge waters extending to the 9 mile NR boundary.

In 2006, hydrographic Light Detection and Ranging (LiDAR) data were collected to map bathymetry in 

southwestern Puerto Rico using the ADS Mk II Airborne System (Figure 1.9). The 900 Hertz (1,064 nm) 

A Nd: Yag laser acquired 4x4 m spot spacing and 200% seabed coverage. In total, 265 square nautical 

miles of LiDAR were collected between -20 m (topographic) up to 50 m (depth). This data provided 

a bathymetric surface and reflectivity 

surface. More details on the methods 

can be found via online metadata files 

(http://ccma.nos.noaa.gov/products/

biogeography/lidar_pr/welcome.html).

To highlight the changes in surface 

complexity across the region we 

measured surface rugosity as the ratio 

between the horizontal surface and the 

actual convoluted 3-dimensional surface 

bathymetry. Areas with high change 

in surface complexity occured at reef 

and shelf edges and where clusters of 

patch reefs existed and spur and groove 

formations. Relatively high topographic 

complexity (orange/red in Figure 1.9) 

was found along the shelf edge, the 

fringing coral reefs of the backreef zone 

(innershelf reefs) and linear midshelf 

reefs, the highly heterogeneous patch 

reefs at El Palo and north of Margarita 

Reef (Figure 1.9) and the large expanse of 

colonized pavement with sand channels 

extending east-west parallel to the shelf 

edge. This study and earlier analyses 

have revealed that measures of surface 

complexity are useful and cost-effective 

predictors of the distribution of fish 

species, fish diversity and biomass and 

coral diversity and abundance (Pittman 

et al., 2009).

Figure 1.9. Surface rugosity calculated from the LiDAR bathymetry to show 

the pattern of topographic variability across the study region. A distinctive 

area with high rugosity created through the clustering of linear and patch reefs 

exists between El Palo and Margarita Reefs. The La Parguera Natural Reserve 

encompasses almost all of the shallow water high complexity coral reef in the 

region.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

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stress. Palaios 4(1): 92-100.

Appeldoorn, R.S. 2008. Transforming reef fisheries management: application of an ecosystem-based approach 

in the USA Caribbean. Environ. Conserv. 35: 232-241.

Aguilar-Pererra, A., M. Scharer, M. Valdes-Pizzini. 2006. Marine protected areas in Puerto Rico: Historical and 

current perspectives. 49(12): 961-975.

Appeldoorn, R.S. and S. Meyers. 1993. Part 2. Puerto Rico and Hispaniola. pp. 99-158. In: Marine fishery 

resources of the Antilles: Lesser Antilles, Puerto Rico and Hispaniola, Jamaica, Cuba. FAO Fisheries Technical 

Paper No. 326. Rome, FAO. 325 pp.

Ault, J.S., S.G. Smith, J. Luo, M.E. Monaco, and R.S. Appeldoorn. 2008. Length-Based Assessment of 

Sustainability Benchmarks for Coral Reef Fishes in Puerto Rico. Environ. Conserv. 35: 221-231.

Ballantine, D.L., R.S. Appeldoorn, P. Yoshioka, E. Weil, R. Armstrong, J.R. Garcia, E. Otero, F. Pagan, C. 

Sherman, E.A. Hernandez-Delgado, A. Bruckner, and C. Lilyestrom. 2008. Biology and Ecology of Puerto Rican 

Coral Reefs. pp. 375-406. In: B. Riegl and R.E. Dodge (eds.). Coral Reefs of the USA. Coral Reefs of the World, 

Volume 1. Springer. 806 pp.

Bejarano-Rodríguez, I. 2006. Relationships between reef fish communities, water and habitat quality on coral 

reefs. M.S. Thesis. University Puerto Rico at Mayagüez. Mayagüez, PR. 51 pp.

Bohnsack, J., S. Meyers, R. Appledoorn, J. Beets, D. Matos-Caraballo, and Y. Sadovy. 1991. Stock Assessment 

of Spiny Lobster, Panulirus argus, in the U.S. Caribbean. NMFS SEFSC-Miami Laboratory Contribution No. MIA9C/91-49. Miami, FL. 50 pp.

Bruckner, A.W. and R.J. Bruckner. 1997. Outbreak of coral disease in Puerto Rico. Coral Reefs 16: 260.

Bush, D.M. 1991. Storm sedimentation on the northern shelf of Puerto Rico. Doctoral dissertation. Department 

of Geology, Duke University. Durham, NC. 

Caribbean Fishery Management Council (CFMC). 1985. Fishery Management Plan, Final Environmental Impact 

Statement, and Draft Regulatory Impact Review, for the Shallow-Water Reef fish fishery of Puerto Rico and 

the U.S. Virgin Islands. Caribbean Fishery Management Council. San Juan, Puerto Rico. 178 pp. http://www.

caribbeanfmc.com/SCANNED%20FMPS/REEF%20FISH/RF%20FMP.pdf.

Clark, R., C. Jeffrey, K. Woody, Z. Hillis-Starr, and M. Monaco. 2009. Spatial and temporal patterns of coral 

bleaching around Buck Island Reef National Monument, St. Croix, U.S. Virgin Islands. Bull. Mar. Sci. 84(2): 167-

182.

Davis, M., E. Gladfelter, H. Lund, and M. Anderson. 1986. Geographic range and research plan for monitoring 

white band disease. Virgin Islands Biosphere Reserve Research Report 6. Virgin Islands Resource Management 

Cooperative. St. John, U.S. Virgin Islands. 28 pp.

García-Sais, J.R., R. Appeldoorn, A. Bruckner, C. Caldow, J.D. Christensen, C. Lilyestrom, M.E. Monaco, J. 

Sabater, E. Williams, and E. Diaz. 2005. The state of coral reef ecosystems of the commonwealth of Puerto Rico. 

pp. 91-134. In: J.E. Waddell (ed.). The state of coral reef ecosystems of the United States and Pacific Freely 

Associated States: 2005. NOAA Technical Memorandum NOS NCCOS 11. Silver Spring, MD. 522 pp.

Garcia-Sais J., R. Appeldoorn,T. Battista, L. Bauer, A. Bruckner, C. Caldow, L. Carrubba, J. Corredor, E. Diaz, 

C. Lilyestrom, G. Garcia-Moliner, E. Hernandez-Delgado, C. Menza, J. Morell, A. Pait, J. Sabater, E. Weil, E. 

Williams, and S. Willimas. 2008. The state of coral reef ecosystems of the commonwealth of Puerto Rico. pp. 

75-116. In: J.E. Waddell and A. Clarke (eds.). The state of coral reef ecosystems of the United States and Pacific 
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11 1 - Introduction

Freely Associated States: 2008. NOAA Technical Memorandum NOS NCCOS 73. Silver Spring, MD. 569 pp.

Hensley, D.A., R.S. Appeldoorn, D.Y. Shapiro, M. Ray, and R.G. Turingan. 1994. Egg dispersal in a Caribbean 

coral reef fish Thalassoma bifasciatum. I. Dispersal over the reef platform. Bull. Mar. Sci. 54(1): 256-270.

Hubbard, D.K., R.B. Burke, I.P. Gill, W.R. Ramirez, and C. Sherman. 2008, Coral-Reef Geology: Puerto Rico and 

the U.S. Virgin Islnds. pp. 263-302. In: B. Riegl and R.E. Dodge (eds.). Coral Reefs of the USA. Coral Reefs of 

the World, Volume 1. Springer. 806 pp.

Kendall, M.S., M.E. Monaco, K.R. Buja, J.D. Christensen, C.R. Kruer, M. Finkbeiner, and R.A. Warner. 2001. 

Methods used to map the benthic habitats of Puerto Rico. NOAA Technical Memorandum NOS NCCOS CCMA 

152 (on-line). http://ccma.nos.noaa.gov/products/biogeography/benthic/welcome.html.

Larsen, M.C. and R.M.T. Webb. 2009. Potential Effects of Runoff, Fluvial Sediment, and Nutrient Discharges on 

the Coral Reefs of Puerto Rico. J. Coast. Res. 25(1): 189-208.

Matos-Caraballo, D. 2004. Comprehensive Census of the Marine Fishery of Puerto Rico, 2002. Commerical 

Fisheries Statistics Program, Fisheries Research Laboratory, Puerto Rico Department of Natural and 

Environmental Resources. Mayagüez, PR. 85 pp.

Ojeda, E. 2002. Description of larval development of the red hind (Epinephelus guttatus) and the spatio-temporal 

distributions of ichthyoplankton during a red hind spawning aggregation of La Parguera, Puerto Rico. University 

of Puerto Rico, Mayaguez. 190 p.

Pittman, S.J., B. Costa, and T. Battista. 2009. Using Lidar bathymetry and boosted regression trees to predict the 

diversity and abundance of fish and corals. J. Coast. Res. Spec. Issue 53(1): 27-38.

Ryan, K.E., J.P. Walsh, D.R. Corbett, and A. Winter. 2008. A record of recent change in terrestrial sedimentation 

in a coral-reef environment, La Parguera: A response to coastal development? Mar. Pollut. Bull. 56(6): 1177-

1183.

Triana, P. 2002. Mapping benthic habitats on the southwest of Puerto Rico as determined by side scan sonar. 

PhD thesis. Department of Marine Science, University of Puerto Rico.

Tyler, R.H. and B.G. Sanderson.1996. Wind-driven pressure and flow around an island. Cont. Shelf Res. 16(4): 

469-488.

Valdes-Pizzini, M. 1990. Fishermen associations in Puerto Rico: praxis and discourse in the politics of fishing. 

Hum. Org. 49: 164–73.

Warne, A.G., R.M.T. Webb, and M.C. Larsen. 2005. Water, Sediment, and Nutrient Discharge Characteristics 

of Rivers in Puerto Rico, and their Potential Influence on Coral Reefs: U.S. Geological Survey Scientific 

Investigations Report 2005-5206. U.S. Geological Survey. Reston, VA. 58 p.
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13 2 - Benthic Composition

Chapter 2. Benthic Composition 

2.1. Introduction

Coral reef ecosystems in Puerto Rico comprise a system of connected mangrove forests, seagrass beds, 

unconsolidated sediments and coral reefs and hardbottom substrates. These habitats provide shelter 

and sustenance to fishes and invertebrates that form the basis of important fisheries in the region. A major 

goal of CCMA’s Caribbean Coral Reef Ecosystem Monitoring (CREM) project is to characterize benthic 

composition and correlate such information to spatial and 

temporal patterns in the distribution of fish and invertebrate 

populations. Benthic characterizations provide the basis 

for identifying species-habitat relationships, increasing 

understanding of spatial patterns in the distributions of 

habitats, and illustrating important and crucial linkages 

for the successful management of coral reef fisheries and 

other important resources. This chapter provides baseline 

estimates of benthic substrate composition of hardbottom, 

seagrass, macroalgae, mangrove and unconsolidated 

habitats in La Parguera, Puerto Rico as defined by Kendall 

et al. (2001). More specifically, data are presented to 

characterize the types, distributions and percent cover of 

benthic flora and fauna within mapped substrates.

2.2. Methods

2.2.1. Survey Data

Underwater visual surveys were conducted for benthic 

composition data collection within the La Parguera study 

area in southwest Puerto Rico. At each site, benthic data 

were collected from five 1 m2 quadrats randomly placed 

along a 25 m belt transect used for fish census (Figure 2.1). 

For detailed methods, see Appendix D.

Table 2.1 provides a list of measured benthic composition 

variables and types of measurements collected. In 

addition to abiotic and biotic benthic cover, data were 

collected on queen conch (Eustrombus gigas), longspined sea urchin (Diadema antillarum), Caribbean spiny 

lobster (Panulirus argus) and marine debris. Collection of 

abundance and distribution data on these selected macro 

invertebrates began in 2004, with their abundance being 

recorded only if individual(s) were observed within the 4 x 

25 m transect (Appendix B). Marine debris data collection 

began in January 2007 

(Figure 2.2). Between 

2001 and 2004, 

benthic habitats were 

categorized according 

to the habitat types 

defined by (Kendall et 

al., 2001; Figure 2.3) 

for sampling allocation 

Figure 2.1. NOAA diver recording benthic habitat 

composition within the randomly placed 1 m2

 quadrat 

along the belt transect.

Benthic Biota

Measurements

Cover 

(%)

Height 

(cm)

Abund. 

(#)

Abiotic

Hardbottom X X

Sand X

Rubble X

Fine sediment X

Rugosity

Water depth

Biotic

Corals (by species) X

Macroalgae X X

Seagrass (by species) X X

Gorgonians

Sea rods, whips and plumes X X X

Sea fans X X X

Encrusting form X

Sponges

Barrel, tubes, rope, vase morph. X X X

Encrusting morphology X

Other benthic macrofauna

Anemonies and hydroids X X

Tunicates and zoanthids X

Macro-invertebrates

Queen conch (by sexual maturity) X

Spiny lobster X

Long-spined urchin X

Mangroves

Prop roots X

Prop roots colonized by algae X

Prop roots colonized by sponges X

Prop roots colonized by other X

Table 2.1. Abiotic and biological variables measured to 

characterize benthic assemblages along fish transects in 

the southwest Puerto Rico study region.

Figure 2.2. Marine debris.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

14 2 - Benthic Composition Figure 2.3. A selection of habitat types designated in the hierarchical classification scheme of NOAA’s benthic habitat map (Kendall et al., 

2001) for the U.S. Caribbean (clockwise from left to right): colonized pavement, patch reef, scattered coral and/or rock, linear reef, seagrass 

and sand. 

and design. Between 2004 and 2007 however, habitats were re-categorized into three habitat types 

(hard, soft and mangrove) for sampling and data collection.

2.2.2. Analytical Methods

In situ data on the cover of benthic biota were summarized from 1,167 surveys (approximately 5,835 

quadrats) during 2001 to 2007 (Tables 2.2 and 2.3). Surveyed sites were located within four broad 

thematic habitat types in the study area: colonized coral reef and hardbottom areas (hereafter hardbottom 

habitats; n=572), seagrass and algal communities (hereafter submerged aquatic vegetation or SAV; 

n=272), unconsolidated sediments (sand and mud habitats; n=91) and mangroves (n=158). An additional 

74 sites that were sampled were mapped as unknown habitat (Table 2.2). These sites had no biotic 

cover and were excluded from subsequent analyses. Note that during any single mission, the number 

of surveys conducted within hardbottom types varied and was relatively low for the least abundant 

habitat types (colonized bedrock and reef rubble) but high for more abundant habitats such as colonized 

Table 2.2. Number of hardbottom benthic habitat sites surveyed in the southwest Puerto Rico study area by mission and mapped habitat 

type from 2001-2007. Mapped habitat categories are from Kendall et al. (2001).

Sample period

Colonized 

Pavement

Col. pav. 

w/ sand

channels

Linear 

reef

Patch 

reef

Scattered 

coral/rock 

in sand

Bedrock/

reef rubble

Sand/ 

mud

Seagrass/ 

macroalgae Unknown* Mangrove Total

2001 Winter 1 2 2 3 1 1 11 1 7 29

2001 Spring 2 3 2 2 1 8 13 7 38

2001 Fall 2 4 1 3 3 16 6 35

2002 Winter 4 5 2 1 5 28 3 9 57

2002 Summer 3 3 7 3 3 4 12 1 9 45

2003 Winter 5 7 6 6 5 8 22 1 15 75

2003 Spring 4 13 9 5 11 12 22 1 13 90

2003 Fall 5 13 6 7 12 8 26 13 90

2004 Spring 11 14 4 7 6 9 26 13 90

2004 Summer 10 16 4 5 10 11 20 13 89

2005 Winter 7 18 7 4 7 1 14 19 13 90

2005 Summer 10 23 12 2 6 2 10 15 80

2006 Winter 12 23 8 8 1 11 16 10 89

2006 Summer 14 22 8 2 6 1 2 14 11 10 90

2007 Winter 14 22 9 2 12 10 11 10 90

2007 Summer 10 25 10 5 1 3 12 14 10 90

Total 110 212 100 53 94 3 91 272 74 158 1,167

*Unknown refers to benthic substrates that were unmappable by Kendall et al. (2001) because of turbid conditions.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

15 2 - Benthic Composition

pavement and patch reefs (Table 

2.3). Hence observed differences 

in benthic composition or the 

lack thereof among hardbottom 

habitat types should be 

interpreted with caution, given 

that mean estimates of metrics 

from least abundant habitats 

were more variable compared 

with estimates from more 

abundant habitats. 

Many benthic variables were 

measured during the surveys, 

but data analyses for this report focused primarily on describing broad-scale spatial patterns and 

temporal trends in the area abundance (percent cover) of the sessile biotic components as described 

in Table 2.1. Specifically, data were analyzed to examine the following: 1) benthic habitat composition 

of broad thematic habitat types and more resolved hardbottom habitat types; 2) broad-scale seascape 

patterns in cover of live coral, macro algae and seagrasses; and 3) temporal trends in live scleractinian 

(hard) coral and algal cover.

2.2.2.1. Characterizing spatial distributions of benthic biotic components among habitats

Estimates of percent cover (mean ± standard error [SE]) of selected benthic biota were calculated 

for each site. Sites were used as independent sample units and were considered replicates within 

survey missions and habitat types. Multiple quadrat measurements (percent cover) for biota within 

each transect were averaged using the equation: 

Σ(Qi - n) / n

where Qi = quadrat i, and n is the total number of quadrats. Average site values were then used to 

calculate means and SE of measured variables per 100 m2

 for each habitat type. Standard errors of 

means represent variability among sites rather than variability among quadrats within a site. Differences 

in the cover of benthic biota among habitat types were determined by using a series of One-Way nonparametric ANOVA (Wilcoxon) tests to identify significant differences among habitat types (Zar, 1999). 

When significant differences were found, non-parametric multiple pair-wise comparisons were used to 

determine the pairs of habitat types that were significantly different (Zar, 1999).

2.2.2.2. Characterizing seascape spatial patterns

Site values of benthic community metrics averaged from quadrat data were interpolated and mapped 

using inverse distance weighting (IDW) to guide interpretation of spatial patterns within the broader 

seascape of the study region. IDW is a method of interpolation that estimates values by averaging the 

values of sample data points in a predetermined neighborhood. In this case, the interpolation used a 

minimum of 10 neighbors. Only hardbottom sites were used to map typical hardbottom features such as 

coral, gorgonian, and sponge percent cover. Similarly, only softbottom sites were used to map seagrass 

percent cover. Some caution should be taken with resulting mapped patterns especially in areas of low 

point abundance and at the edges of the study area.

2.2.2.3. Characterizing temporal trends in live coral and algal cover

Characterization of temporal trends in benthic composition metrics were based on data collected from 

2001 to 2007. Survey missions were categorized as occurring during winter or spring (December 

through June), summer (August), or fall (September through October). Average site values were then 

Table 2.3. The number of benthic habitat sites surveyed by mapped habitat type for the 

southwestern Puerto Rico study region from 2001-2007. Mapped habitat categories are 

from Kendall et al. (2001).

Mapped habitat types

Mapped 

Area (km2

)

% Mapped 

Area

# sites 

surveyed

Area 

surveyed 

(km2)

% area 

surveyed

Colonized Bedrock 212.76 0.2% 2 0.2 0.09

Colonized Pavement 18,505.23 16.5% 110 11 0.06

Col. Pavement w/ Sand Channels 32,058.04 28.6% 212 21.2 0.07

Linear Reef 11,306.15 10.1% 100 10 0.09

Patch Reefs (Aggregate & Individual) 2,738.33 2.4% 53 5.3 0.19

Reef Rubble 270.98 0.2% 1 0.1 0.04

Scattered Coral/Rock in sand 10,345.36 9.2% 94 9.4 0.09

Unconsolidated sediments 6,620.71 5.9% 91 9.1 0.14

Macroalgae 5,160.26 4.6% 40 4 0.08

Seagrass 24,692.53 22.1% 234 23.4 0.09

Total 111,910.3624 100% 937 93.7 0.08
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

16 2 - Benthic Composition used to calculate means and SE of measured variables per 100 m2

 for each survey mission. Standard 

errors of means represent variability among sites rather than variability among quadrats within a site. 

Temporal trends in mean estimates of live coral and algal cover were determined by using two statistical 

approaches. First, a series of One-Way non-parametric ANOVA (Wilcoxon) tests were done to determine 

if significant differences occurred among the sampling periods, and non-parametric multiple pair-wise 

comparisons were used to identify pairs of sampling periods that were significantly different. Second, 

overall temporal trends in mean estimates of live coral and algal cover were determined by using the 

nonparametric Jonckheere Test (JT) for Ordered Alternatives to examine whether or not significant 

change occurred in percent coral cover and algae between 2001 and 2007. The JT statistical procedure 

assumed that there were no differences in coral and algae cover among sampling periods and tested 

against a postulated sequential increase or decrease in those benthic metrics across sampling periods. 

The JT test also assumed that estimates of means were derived from random independent samples and 

that estimates of variance were homogeneous across sampling periods and the frequency distribution 

of data was similar among periods. To calculate the test statistic, the k(k-1)/2 Mann-Whitney U counts 

were derived with the following equation (Wolfe and Hollander, 1999):

Where ø(a,b)=1 if a < b, or 0 if otherwise. The Jonckheere test Statistic (J) was then calculated as the sum 

of these U counts and was compared against a significance threshold (Jα = 0.05) that was dependent on 

the number of sampling periods and number of sites surveyed within each sampling period (Wolfe and 

Hollander 1999). If Jmetric > Jα

 = 0.05 , we concluded that estimates of mean cover were not equal across 

sampling periods, and that there was an overall sequential increase or decrease in mean estimates 

during the study period. Additionally, residual plots of coral and algae percent cover were examined to 

determine if variance estimates were homogenous and comparable over time.

Montastraea annularis complex and gorgonians
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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17 2 - Benthic Composition

2.3. Results

2.3.1. Benthic habitat composition

Estimates of percent cover (mean ± SE) of selected benthic organisms are reported for four habitat 

types observed in the study area: hardbottom habitats, SAV communities, sand and mud habitats, and 

mangroves. Data are presented at these broader thematic habitat resolutions because the number of 

surveyed sites varied among the more resolved habitat types and was extremely low for rare habitats 

such as bedrock and reef rubble (Table 2.3). Comparisons presented here are intended to characterize 

broad differences in benthic composition among the four habitat types.

2.3.1.1. Characterization of colonized hardbottom habitat types

Pavement habitat was the most spatially extensive habitat type (45.1% of the study area) and was 

therefore the most intensively surveyed, followed by seagrass (22.1%) and linear reefs (10.1%; Table 

2.2). The remainder of the area surveyed comprised of scattered coral and rock in sand, unconsolidated 

sediments, macro algae, patch reefs, reef rubble and bedrock (Table 2.2); Hardbottom habitats combined 

formed a larger proportion (67.4%) of the study region than did soft bottom areas.

Overall, mean benthic cover on 

hardbottom habitats was 60.5 ± 1.3%. 

Generally, hardbottom habitat types 

were dominated by algae (27.0 ± 

1.0% "turf-like" algae [turf algae], 14.0 

± 0.7% macroalgae, and 1.3 ± 0.3% 

crustose coralline algae [CCA]; Figure 

2.4). The next most abundant benthic 

group was the gorgonians (soft corals), 

which had an average percent cover 

of 7.4 ± 0.4% (Figure 2.4). Mean live 

scleractinian coral cover averaged 

5.3 ± 0.3% across the study area. 

Cyanobacteria and filamentous algae 

were grouped as a single component 

and had a mean cover of 1.4 ± 0.2%. 

Other benthic organisms observed on 

hardbottom habitats included sponges 

(2.3 ± 0.1%), seagrasses (1.7 ± 0.3%) 

and hydroids such as fire coral (0.3 ± 

0.1%; Figure 2.4).

Benthic composition was variable 

among the hardbottom habitat types 

surveyed (Figure 2.5). The mean 

percent cover of live scleractinian 

coral was highest on linear reef (6.8 ± 

0.7%, n=100), followed by colonized 

pavement (6.1 ± 0.4%), patch reef (4.8 

± 0.7%), but was lowest on reef rubble 

(1.2 ± 0%, n=1; Figure 2.5). Fire corals 

were the most commonly recorded 

hydroid species on pavement, linear 

reef, and patch reef habitat types, 

with highest percent cover occurring 

0

10

20

30

Mean cover (%)

Figure 2.4. Mean (± SE) percent cover for key benthic components on hardbottom 

sites (n=572) in the southwest Puerto Rico study region from 2001-2007. 

CCA=crustose coralline algae; CB and FA=cyanobacteria and filamentous algae). 

0

20

40

60

Bedrock Pavement Linear Reef Patch Reef Reef Rubble Scattered 

Coral/Rock

Mean cover (%)

Turf algae Macroalgae

Corals Soft corals

CCA FA and CB

Hydroids Sponges

Figure 2.5. Vertical bars represent mean (± SE) percent cover for Key components 

of benthic community on hardbottom habitat types (n=572) in the southwest Puerto 

Rico study region from 2001-2007. 
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

18 2 - Benthic Composition on linear reef habitats (1.1 ± 0.6%). The percent cover of 

sponges and fire corals were similar among the habitat 

types surveyed (Figure 2.5).

Live scleractinian coral cover included at 

least 24 coral genera, but only nine with a 

mean cover greater than 0.01% (Figure 

2.6). The two coral genera with the 

highest mean cover were Montastraea

spp. (2.3 ± 1.9%) and Porites spp. (0.8 ± 

0.1%). Three genera (Siderastrea spp., 

Diploria spp. and Agaricia spp.) had the 

next highest mean cover (0.4 ± 0.0%) 

on hardbottom throughout the study 

region. Acropora corals had a mean 

cover of 0.2 ± 0.1%, (Figures 2.6 and 

2.7). Boulder brain coral (Colphophyllia 

natans) was not as commonly 

observed and had a mean cover of 

0.2 ± 0.1%). Other genera observed 

on hardbottom habitats were rare 

and included Cladocora, Dendrogyra, 

Dichocoenia, Eusmilia, Favia, 

Helioceris, Isophyllastrea, Isophyllia, 

Madracis, Manicina, Mycetophyllia, 

Oculina, Mussa, Scolymia, Solenastrea

and Stephanocoenia. These corals had 

a combined mean cover of 0.6 ± 0.1% 

(Figure 2.6).

The distribution of coral genera 

among hardbottom habitats reflected 

the overall spatial patterns of coral 

distribution observed on all hardbottom 

habitats combined. Montastraea

and Porites species were the most 

ubiquitous corals, with the highest 

mean coral cover in the most abundant 

hardbottom habitats: pavement, linear 

reef and patch reefs habitats (Figure 

2.6). Additionally, Montastraea and 

Porites were the genera with the most 

coral cover for all hardbottom habitats 

except for colonized bedrock, where 

only two sites were surveyed. Acroporid 

coral cover generally was low, but was 

highest on linear reef habitat (Figure 

2.7). Other coral genera such as 

Diploria, Siderastrea and Colpophyllia

appeared to have similarly low cover 

in both abundant and rare hardbottom habitats (Figure 2.7). Interestingly, several genera were not 

Diploria labyrintheformes (left) and Montastraea annularis complex (right)

0

1

2

3

Mean cover (%)

Figure 2.6. Aerial mean (± SE) percent cover of coral genera found across 

hardbottom sites in the southwest Puerto Rico study region from 2001-2007. 

Asterisks (*) indicates the other genus: Claocora, Dendrogyra, Dichocoenia, 

Eusmilia, Favia, Heloceris, Isophyllastrea, Isophyllia, Madracis, Manicina, 

Mycetophyllia, Oculina, Musa, Scolymia, Solenastrea and Stephanocoenia.

0

1

2

3

4

Pavement Linear Reef Patch Reef Scattered 

Coral / Rock

Bedrock

Mean cover (%)

Montastraea

Porites

Agaricia

Siderastrea

Diploria

Acropora

Colpophyllia

Meandrina

Other

(Abundant habitat) (Rare habitat)

Figure 2.7. Mean (± SE) percent cover of coral genera by hardbottom habitat type 

in the southwest Puerto Rico study region from 2001-2007.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

19 2 - Benthic Composition

observed on colonized bedrock habitats, 

but this pattern may have reflected the 

low number of colonized bedrock sites 

surveyed (n=2).

2.3.1.2. Characterization of submerged 

aquatic vegetation (SAV) habitat types

The benthic organisms observed in SAV 

habitats had total cover of 48.4 ± 2%. 

Seagrasses had the highest mean cover 

(28.1 ± 1.8%) in habitats classified as SAV 

followed by macroalgae (12.4 ± 0.9%; 

Figure 2.8). Four seagrass species were 

observed, of which Thalassia testudinum

(turtle grass) had the most cover (21.2 ± 

1.7%; Figure 2.9). Some turf algae (4.2 ± 

0.8% cover) was observed in SAV habitat, 

most likely colonizing fragments of hard 

substrates that are commonly observed 

in seagrass beds. Cyanobacteria and 

unidentified filamentous algae (2.3 ± 

0.6%) were also observed colonizing 

seagrasses, macroalgae, and patches 

of hardbottom substrates encountered 

in SAV habitats. Other organisms 

found inhabiting SAV habitats included 

sponges, gorgonians, corals, CCA algae 

and hydroids such as fire corals. These 

organisms were rare with mean estimates 

of cover less than 0.4 ± 0.1%. 

2.3.1.3. Characterization of 

unconsolidated sediment types

Overall, the total benthic cover on 

unconsolidated sediments was low (18.5 

± 2.6%). Most of the cover observed on 

this habitat type was turf algae (6 ± 1.6%), 

followed by seagrass (5.7 ± 1.4%), and 

macroalgae (4.6 ± 0.7%; Figure 2.10). 

Hard and soft corals, sponges, crustose 

coralline algae, cyanobacteria and 

filamentous algae were also observed, 

but their mean cover was less than 0.7 

± 0.3%. These organisms were often 

encountered on small patches of hard 

substrate that often occurred within 

unconsolidated sediment habitats. Unlike 

seagrass habitats in which T. testudinum

dominated, Syringodium filiformeA 

0

8

16

24

32

Mean cover (%)

Figure 2.8. Mean (± SE) percent cover of key benthic components on submerged 

aquatic vegetation (SAV) sites (n=272) in the study region from 2001-2007. 

CCA=crustose coralline algae; CB and FA=cyanobacteria and filamentous 

algae.

0

12

24

Thalassia 

testudinum

Syringodium 

filiforme

Halodule 

wrightii

Halophila 

decipiens

Mean cover (%)

Figure 2.9. Mean (± SE) percent cover of seagrass species observed on SAV 

sites (n=272) in the southwest Puerto Rico study region from 2001-2007.A

0

2

4

6

8

Mean cover (%)

Figure 2.10. Mean (± SE) percent cover of key benthic components on 

unconsolidated sediment sites (n=91) in the study region from 2001-2007. CCA= 

crustose coralline algae; CB and FA=cyanobacteria and filamentous algae.

A The taxonomic name for Halodule wrightii has been recently changed to Halodule beaudettei and Syringodium filiforme has recently 

changed to Syringodium isoetifolium.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

20 2 - Benthic Composition (manatee grass) was the most dominant 

of the seagrass species in unconsolidated 

sediments habitats, with a mean cover of 

3.1 ± 1.0% (Figure 2.11).

2.3.1.4. Characterization of benthic 

substrates in mangrove habitats

Benthic substrates within mangrove 

habitats were colonized by benthic 

organisms and had an overall mean 

benthic cover of 54.9 ± 2.7%. At many 

sites, mangrove prop roots provided a 

major benthic substrate for colonization 

by epiphytic organisms. Macroalgae 

had the highest benthic cover (14.5 ± 

1.5%) and was the most dominant subaquatic benthic organism identified on 

mangrove prop roots and abiotic benthic 

substrates (Figure 2.12). One commonly 

encountered feature of benthic substrates 

in mangrove habitats was a thick layer of 

detritus, which had a mean cover of 7.8 

± 1.2% (Figure 2.12). Other organisms 

found colonizing mangrove habitats 

included sponges, hard and soft corals, 

CCA and hydroids such as fire corals, turf 

algae, seagrasses, cyanobacteria and 

filamentous algae. 

2.3.2. Spatial distribution patterns in 

benthic cover

The following sections describe broad 

spatial patterns in metrics for live 

corals, algae, gorgonians, sponges 

and seagrasses that were derived 

from interpolations of percent cover data. Maps of interpolated distributions are useful in that they 

help elucidate broad-scale patterns (e.g., the degree of patchiness and location of hotspots) in the 

seascape that are not discernible from point-data. Interpolations for corals and rugosity were confined 

to hardbottom areas leaving softbottom areas as a white space in the interpolated maps. Several 

distinct spatial patterns were observed and are described below.

2.3.2.1. Spatial patterns in live coral cover 

Interpolated live coral data reveal patchy areas of higher (10-15%) 

live coral cover along the nearshore fringing reefs in the center 

of the study area (Figure 2.13a). More robust areas of high coral 

cover were located in the western portion of the study area with 

cover ranging from 10-37%. Two smaller patches of high live coral 

cover were evident on the bank shelf with cover ranging from 10-

52%. These areas were associated with increased benthic rugosity 

(Figure 2.13b) and also featured higher coral species richness 

0

2

4

6

Thalassia 

testudinum

Syringodium 

filiforme

Halodule 

wrightii

Halophila 

decipiens

Mean cover (%)

Figure 2.11. Mean (± SE) percent cover of seagrass species observed on 

unconsolidated sediment sites (n=91) in the southwest Puerto Rico study region 

from 2001-2007.

0

6

12

18

Macroalgae Detritus Seagrasses Prop Roots Sponges Soft Coral

Mean cover (%)

Figure 2.12. Mean (± SE) percent cover of key benthic components on benthic 

substrates at mangrove sites (n=158) in the southwest Puerto Rico study region 

from 2001-2007.

Diploria strigosa
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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21 2 - Benthic Composition

Figure 2.13. Maps of the interpolated (left map) and spatial (right map) distributions for: (a) live percent coral cover, (b) benthic rugosity, 

(c) coral species richness and (d) coral species diversity.

Interpolation

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25.01 - 51.98

Live coral cover (%)

< 0.10

0.11 - 0.2

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0.66 - 1

Index of rugosity

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Coral cover (%)

0 0.01 - 5 5.1 - 10 10.1 - 20 20.1 - 52

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0.01 - 0.2 0.21 - 0.4 0.41 - 0.6 0.61 - 0.8 0.81 - 1

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Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

22 2 - Benthic Composition 

(Figure 2.13c). These areas of higher benthic 

rugosity and live coral cover contained six coral 

species on average with maximum richness (10-14 

species) typically observed on the bank shelf. Coral 

species diversity (Figure 2.13d) appeared greatest 

on the bank shelf with values ranging from 1.4-

2.5 per 100 m2. In general, diversity declined with 

decreasing depth (shallower) and rugosity. 

The broad study area exhibited overall low live coral 

cover (5%) on hardbottom with scattered patches 

of high cover and species richness. Live coral 

cover was dominated by five species (Montastraea 

annularis complex, Porites astreoides, Montastraea 

cavernosa, Agaricia spp. and Siderastrea siderea) 

that accounted for over 60% of the total coral cover 

in the study area (Table 2.4). These species also 

varied in percent cover and frequency of occurrence 

among hardbottom habitat types (Table 2.5). 

Species

Measurements

Frequency

occurance

% cover

Mean ± SE

Acropora cervicornis 0.11 0.18 (0.06)

Acropora palmata 0.01 0.04 (0.02)

Agaricia agaricites 0.13 0.06 (0.01)

Agaricia fragilis 0.00 0.00 (0.00)

Agaricia lamarcki 0.01 0.01 (0.00)

Agaricia spp. 0.38 0.31 (0.03)

Cladocora arbuscula 0.00 0.00 (0.00)

Colpophyllia natans 0.16 0.18 (0.05)

Dendrogyra cylindricus 0.02 0.02 (0.01)

Dichocoenia stokesii 0.17 0.03 (0.00)

Diploria clivosa 0.15 0.12 (0.02)

Diploria labyrnithiformis 0.12 0.06 (0.01)

Diploria spp. 0.03 0.03 (0.01)

Diploria strigosa 0.28 0.18 (0.03)

Eusmilia fastigiata 0.02 0.00 (0.00)

Favia fragum 0.05 0.01 (0.00)

Helioceris cucullata 0.02 0.03 (0.01)

Isophyllastrea rigida 0.00 0.00 (0.00)

Isophyllia sinuosus 0.01 0.00 (0.00)

Madracis mirabilis 0.01 0.00 (0.00)

Madracis mirabilis 0.01 0.00 (0.00)

Madracis spp. 0.04 0.01 (0.00)

Manicina areolata 0.04 0.01 (0.00)

Meandrina meandrites 0.22 0.07 (0.01)

Millepora alcicornis 0.24 0.07 (0.01)

Millepora complanata 0.04 0.01 (0.00)

Millepora spp. 0.20 0.22 (0.11)

Montastraea annularis complex 0.49 1.89 (0.17)

Montastraea cavernosa 0.47 0.43 (0.04)

Mussa angulosa 0.01 0.00 (0.00)

Mycetophyllia aliciae 0.00 0.00 (0.00)

Mycetophyllia daanana 0.00 0.00 (0.00)

Mycetophyllia ferox 0.03 0.01 (0.00)

Mycetophyllia lamarckiana 0.01 0.00 (0.00)

Mycetophyllia reesi 0.00 0.00 (0.00)

Mycetophyllia spp. 0.03 0.01 (0.00)

Oculina diffusa 0.01 0.01 (0.01)

Porites asteroides 0.62 0.56 (0.04)

Porites branneri 0.01 0.01 (0.01)

Porites colonensis 0.01 0.01 (0.00)

Porites porites 0.32 0.20 (0.04)

Porites spp. 0.01 0.03 (0.03)

Scleractinia 0.05 0.36 (0.12)

Scolymia cubensis 0.00 0.00 (0.00)

Scolymia spp. 0.04 0.01 (0.00)

Siderastrea radians 0.31 0.16 (0.02)

Siderastrea siderea 0.42 0.21 (0.02)

Siderastrea spp. 0.05 0.03 (0.01)

Solenastrae spp. 0.02 0.00 (0.00)

Stephanocoenia intercepta 0.16 0.03 (0.01)

Table 2.4. Summary statistics for hard coral species found in 

surveys in the southwest Puerto Rico study area from 2001-2007.

Porites astreoides

Agaricia lamarckii

Montastrea cavernosa 
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

23 2 - Benthic Composition

Species

Col. Bedrock

(n=2)

Col. Pavement

(n=322)

Linear Reef

(n=100)

Patch Reef

(n=53)

Reef Rubble

(n=1)

Scat. Coral/Rock in 

Sand (n=94)

Freq. 

occur

% cover

Mean ± SE

Freq. 

occur

% cover

Mean ± SE

Freq. 

occur

% cover

Mean ± SE

Freq. 

occur

% cover

Mean ± SE

Freq. 

occur

% cover

Mean ± SE

Freq. 

occur

% cover

Mean ± SE

A. cervicornis 0.14 0.16 (0.04) 0.09 0.42 (0.29) 0.08 0.14 (0.08) 0.03 0.05 (0.04)

A. palmata 0.01 0.04 (0.03) 0.04 0.11 (0.07)

A. agaricites 0.50 0.03 (0.03) 0.16 0.07 (0.02) 0.12 0.05 (0.02) 0.15 0.06 (0.03) 0.02 0.03 (0.02)

A. fragilis 0.01 <0.01 (<0.01)

A. lamarcki 0.02 0.02 (0.01) 0.01 <0.01 (<0.01)

Agaricia spp. 0.44 0.37 (0.05) 0.41 0.30 (0.09) 0.32 0.37 (0.15) 1.00 0.24 0.17 0.11 (0.04)

C. arbuscula 0.01 <0.01 (<0.01)

C. natans 0.17 0.16 (0.04) 0.17 0.38 (0.26) 0.15 0.18 (0.11) 0.09 0.06 (0.03)

D. cylindricus 0.02 0.03 (0.02) 0.01 0.01 (0.01) 0.01 0.01 (0.01)

D. stokesii 0.50 0.05 (0.05) 0.21 0.04 (0.01) 0.17 0.04 (0.01) 0.09 0.01 (<0.01) 0.07 0.01 (<0.01)

D. clivosa 0.50 0.75 (0.75) 0.16 0.10 (0.02) 0.18 0.22 (0.07) 0.21 0.19 (0.07) 0.03 0.01 (0.01)

D. labyrinthiformis 0.13 0.07 (0.02) 0.17 0.06 (0.02) 0.13 0.09 (0.04) 0.03 0.02 (0.01)

Diploria spp. 0.02 0.03 (0.01) 0.05 0.04 (0.02) 0.02 <0.01 (<0.01)

D. strigosa 0.31 0.14 (0.02) 0.39 0.43 (0.15) 0.23 0.17 (0.07) 1.00 0.30 0.09 0.06 (0.03)

E. fastigiata 0.02 <0.01 (<0.01) 0.02 <0.01 (<0.01) 0.01 0.00 (<0.01)

F. fragum 0.03 0.01 (<0.01) 0.08 0.02 (0.01) 0.15 0.04 (0.02)

H. cucullata 0.03 0.05 (0.02) 0.02 0.03 (0.02)

I. rigida <0.01 <0.01 (<0.01) 0.01 0.01 (0.01)

I. sinuosus 0.01 <0.01 (<0.01) 0.02 <0.01 (<0.01)

M. decactis 0.03 0.01 (<0.01) 0.04 <0.01 (<0.01) 0.01 <0.01 (<0.01)

M. mirabilis 0.01 <0.01 (<0.01) 0.01 <0.01 (<0.01) 0.02 0.01 (0.01)

Madracis spp. 0.05 0.01 (<0.01) 0.05 0.01 (<0.01) 0.03 <0.01 (<0.01)

M. areolata 0.04 0.01 (<0.01) 0.06 0.01 (0.01) 0.02 <0.01 (<0.01) 0.02 0.01 (<0.01)

M. meandrites 0.29 0.10 (0.02) 0.18 0.04 (0.01) 0.09 0.03 (0.02) 0.13 0.04 (0.02)

M. alcicornis 0.26 0.05 (0.01) 0.30 0.12 (0.03) 0.30 0.21 (0.12) 0.11 0.02 (0.01)

M. complanata 0.03 0.01 (<0.01) 0.10 0.03 (0.01) 0.04 0.00 (<0.01) 0.02 <0.01 (<0.01)

Millepora spp. 0.19 0.07 (0.02) 0.35 0.92 (0.60) 0.21 0.16 (0.11) 0.09 0.02 (0.01)

M. annularis cmplx 0.50 0.35 (0.35) 0.57 2.40 (0.26) 0.49 1.74 (0.38) 0.53 1.54 (0.38) 1.00 0.20 0.20 0.56 (0.26)

M. cavernosa 0.56 0.47 (0.05) 0.45 0.62 (0.15) 0.43 0.33 (0.10) 0.20 0.19 (0.06)

M. angulosa 0.01 <0.01 (<0.01) 0.01 <0.01 (<0.01)

M. aliciae <0.01 <0.01 (<0.01)

M. danaana <0.01 <0.01 (<0.01)

M. ferox 0.04 0.02 (0.01) 0.02 0.01 (<0.01) 0.02 0.03 (0.03)

M. lamarckiana 0.01 <0.01 (<0.01) 0.01 0.01 (0.01)

M. reesi 0.01 <0.01 (<0.01)

Mycetophyllia spp. 0.04 0.01 (<0.01) 0.02 0.01 (0.01) 0.04 0.01 (0.00)

O. diffusa 0.01 0.01 (0.01) 0.01 0.01 (0.01)

P. asteroides 1.00 0.10 (<0.01) 0.67 0.62 (0.05) 0.77 0.75 (0.08) 0.66 0.56 (0.11) 1.00 0.10 0.24 0.15 (0.05)

P. branneri 0.01 <0.01 (<0.01) 0.02 0.04 (0.04) 0.01 <0.01 (<0.01)

P. colonensis <0.01 0.01 (0.01) 0.02 0.01 (0.01) 0.01 0.01 (0.01)

P. porites 0.50 0.20 (0.20) 0.34 0.21 (0.07) 0.34 0.25 (0.08) 0.42 0.20 (0.05) 1.00 0.14 0.14 0.07 (0.03)

Porites spp. 0.01 0.05 (0.05) 0.02 0.01 (<0.01)

Scleractinia 0.04 0.36 (0.18) 0.11 0.68 (0.32) 0.09 0.44 (0.26) 0.02 0.02 (0.02)

S. cubensis 0.02 0.01 (0.01)

Scolymia spp. 0.05 0.01 (<0.01) 0.02 0.00 (<0.01) 0.04 <0.01 (<0.01) 0.04 0.01 (<0.01)

S. radians 0.50 0.65 (0.65) 0.34 0.17 (0.03) 0.40 0.26 (0.06) 0.26 0.14 (0.05) 1.00 0.22 0.15 0.07 (0.03)

S. siderea 0.50 0.25 (0.03) 0.46 0.23 (0.04) 0.38 0.23 (0.07) 0.18 0.07 (0.02)

Siderastrea spp. 0.50 0.20 (0.20) 0.05 0.02 (0.01) 0.05 0.03 (0.01) 0.06 0.02 (0.01) 0.05 0.05 (0.03)

Solenastraea spp. 0.02 <0.01 (<0.01) 0.01 0.00 (<0.01) 0.02 0.01 (0.01) 0.02 <0.01 (<0.01)

S. intercepta 0.50 0.10 (0.10) 0.18 0.04 (0.01) 0.20 0.04 (0.01) 0.09 0.02 (0.02) 0.10 0.02 (0.01)

Table 2.5. Summary statistics for coral species found in hardbottom sites by habitat type in the southwest Puerto Rico study area from 

2001-2007.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

24 2 - Benthic Composition 

M. annularis complex was the most dominant coral taxa (35% of total cover; 45% frequency of 

occurrence) with areas of highest cover (5-25%) located in the central and western portions of the study 

area (Figure 2.14a). Highest coral cover (>25%) occurred on the bank shelf in relatively deeper waters 

(>15 m). P. astreoides was the most frequently observed coral taxa (62%) with extensive presence 

within the study area but limited in terms of cover (Figure 2.14b). Overall, cover was low (0.1-1%) 

with localized patches of high cover (5-9%) associated with increased rugosity on the bank shelf. M. 

cavernosa (Figure 2.15a) was observed in nearly 50% of surveys (Table 2.4) and exhibited low cover 

throughout the study area. 

Isolated patches of high cover (4-12%) were observed along 

the nearshore fringing reefs and moderate cover (1-4%) was 

patchily distributed throughout the study area. Colonies of 

Agaricia spp. were common (38%), but generally low in cover 

throughout the study area (Figure 2.15b). Highest cover (3-

8%) was generally observed at moderate depths (6-18 m) and 

high rugosity. S. siderea was commonly observed (42%) but 

cover was low throughout the study area (Figure 2.15c). Small 

isolated patches of cover >1% were widely dispersed but not 

spatially associated with depth or rugosity. 

Figure 2.14. Maps of the interpolated (left map) and spatial (right map) distributions for live percent cover for coral species: (a) Montastraea 

annularis complex and (b) Porites astreoides.

Interpolation

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Interpolation

0 1 km

3 a)

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0.2 - 5

5.1 - 15

15.1 - 25

25.1 - 30.3

M. annularis complex cover (%)

0 1 km

Survey Sites N=572

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Interpolation

0 1 km

3 b)

< 0.10

0.1 - 1

1.1 - 3

3.1 - 5

5.1 - 9

P. astreoides cover (%)

M. annularis

0 0.1 - 5 5.1 - 15 15.1 - 25 25.1 - 30.3

0 1 km

complex % cover

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M. annularis

0 0.1 - 5 5.1 - 15 15.1 - 25 25.1 - 30.3

Survey Sites N=572

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Interpolation

P. asteroides

0 0.1- 1 1.1 - 3 3.1 - 5 5.1 - 9

0 1 km

% cover

0 1 km

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0 0.1- 1 1.1 - 3 3.1 - 5 5.1 - 9

Survey Sites N=572

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Acropora cervicornis
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

25 2 - Benthic Composition

Threatened Acropora species were rare or uncommon 

(Acropora cervicornis 11%, Acropora palmata 1%) with patchy 

distribution throughout the study area (Figure 2.16). Colonies 

were generally observed at depths <10 m and both exhibited 

highest cover in relatively close proximity in the western portion 

of the study area. A. cervicornis (Figure 2.16a) was more widely 

distributed than Acropora palmata, (Figure 2.16b) but both were 

low overall. It should be noted, our sampling approach was not 

optimized for detecting Acropora species.

a)

< 0.1

0.1 - 1

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M. cavernosa cover (%)

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Agaricia species cover (%)

0 1 km

Survey Sites N=572

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S. siderea cover (%)

Figure 2.15. Maps of the interpolated (left map) and spatial (right map) distributions for live percent cover for coral species: (a) Montastraea 

cavernosa, (b) Agaricia species and (c) Siderastrea siderea.

Interpolation

M. cavernosa

0 0.1 - 1 1.1 - 4 4.1 - 8 8.1 - 12

0 1 km

% cover

0 1 km

M. cavernosa

0 0.1 - 1 1.1 - 4 4.1 - 8 8.1 - 12

Survey Sites N=572

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Interpolation

Agaricia spp.

0 0.01 - 1 1.1 - 3 3.1 - 5 5.1 - 8

0 1 km

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0 1 km

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0 0.01 - 1 1.1 - 3 3.1 - 5 5.1 - 8

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0 01 - 1 11 - 2 21 - 3 31 - 53

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0 1 km

S. siderastrea

0 0.1 - 1 1.1 - 2 2.1 - 3 3.1 - 5.3

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Acropora palmata
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

26 2 - Benthic Composition 

2.3.2.2. Spatial patterns in macroalgae cover

Macroalgae (Figure 2.17a) cover was similar on both hard and soft bottom habitats (mean=12-14%) 

with considerable variability in the shallow waters close to the mangrove shoreline and the edge of 

the bank shelf. Macroalgal cover was highest (60-90%) along the western edge of the study area 

exhibited by broad shallow (<4 m) sand flats. Significant macroalgal cover was also observed in and 

adjacent to the mangroves that line the shore and some offshore islands. Turf algae were a significant 

component to the hardbottom benthic community. Mean cover (30%) was nearly twice that of macroalgal 

cover and combined accounted for 40-45% of the hardbottom cover. Turf algae (Figure 2.17b) were 

widely distributed across the hardbottom seascape and 

higher cover was spatially associated with high rugosity. 

Soft corals (gorgonians and sea fans) were common 

hardbottom features (Figure 2.17c), in particular on patch 

reefs and colonized pavement. Cover ranged from 0-58% 

and, in general, declined from the shore to the edge of 

the bank shelf. High soft coral cover was associated with 

high rugosity. Sponge cover ranged from 0-30% and 

was variable on all hardbottom types (Figure 2.17d). The 

general pattern indicated higher cover at greater depth 

and higher rugosity. Sponge cover appeared to be greater 

in the central and eastern portions of the study area.

Figure 2.16. Maps of the interpolated (left map) and spatial (right map) distributions for live percent cover for coral species: (a) Acropora 

cervicornis and (c) Acropora palmata.

Interpolation

0 1 km

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Interpolation

0 1 km

3 a)

< 0.1

0.1 - 1

1.1 - 5

5.1 - 9

9.1 - 27.2

A. cervicornis cover (%)

0 1 km

Survey Sites N=572

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Interpolation

0 1 km

3 b)

< 0.1

0.1 - 1

1.1 - 2

1.1 - 4

4.1 - 10.4

A. palmata cover (%)

A. cervicornis

0 0.1 - 1 1.1 - 5 5.1 - 9 9.1 - 27.2

0 1 km

% cover

0 1 km

A. cervicornis

0 0.1 - 1 1.1 - 5 5.1 - 9 9.1 - 27.2

Survey Sites N=572

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Interpolation

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0 0.1 - 1 1.1 - 2 2.1 - 4 4.1 - 10.4

0 1 km

% cover

0 1 km

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0 0.1 - 1 1.1 - 2 2.1 - 4 4.1 - 10.4

Survey Sites N=572

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Macroalgae
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

27 2 - Benthic Composition

Figure 2.17. Maps of the interpolated (left map) and spatial (right map) distributions for: (a) macroalgae, (b) turf algae, (c) soft coral cover 

and (d) sponges.

Interpolation

0 1 km

Survey Sites N=1093

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Interpolation

0 1 km

3 a)

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5.1 - 15

15.1 - 25

25.1 - 40

40.1 - 89.7

Macroalgal cover (%)

0 1 km

Survey Sites N=572

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Interpolation

0 1 km

3 b)

< 10

10.1 - 25

25.1 - 50

50.1 - 75

75.1 - 93.8

Turf algae cover (%)

0 1 km

Survey Sites N=572

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Interpolation

0 1 km

3 c)

< 5

5.1 - 10

10.1 - 15

15.1 - 20

20.1 - 58.4

Soft coral cover (%)

0 1 km

Survey Sites N=572

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Interpolation

0 1 km

3 d)

< 3

3.1 - 5

5.1 - 10

10.1 - 20

20.1 - 30

Sponge cover (%)

Macroalgal cover (%)

0 0.1 - 10 10.1 - 25 25.1 - 40 40.1 - 90

0 1 km

3

0 1 km

Macroalgal cover (%)

0 0.1 - 10 10.1 - 25 25.1 - 40 40.1 - 90

Survey Sites N=1093

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Interpolation

Turf algae

0 0.1 - 25 25.1 - 50 50.1 - 75 75.1 - 93.8

0 1 km

% cover

0 1 km

Turf algae

0 0.1 - 25 25.1 - 50 50.1 - 75 75.1 - 93.8

Survey Sites N=572

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Interpolation

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Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

28 2 - Benthic Composition 2.3.2.3. Spatial patterns in seagrass cover

Seagrasses were common on the softbottom habitats in the study area 

(Figure 2.18a). Seagrass cover was most extensive close to shore and 

declined toward the shelf break. Large continuous areas of seagrass 

are present nearshore, adjacent to mangroves and inside the fringing 

reefs. In this zone, seagrass was predominantly T. testudinum that was 

abundant to depths of approximately 16 m (Figure 2.18b). T. testudinum

cover ranged from 0-96% and occurred primarily in the central and 

western portion of the study area. S. filiforme (Figure 2.18c), was 

uncommon (<10% cover) at depths <10 m, but surpassed Thalassia

with increasing depth where cover ranged between 30-85%.

Thalassia testudinum

0 1 km Survey Sites N=521

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Figure 2.18. Maps of the interpolated (left map) and spatial (right map) distributions for: (a) seagrass, (b) Thalassia testudinum and (c) 

Syringodium filiforme.

Interpolation

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0 0.1 - 20 20.1 - 40 40.1 - 60 60.1 - 96.5

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Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

29 2 - Benthic Composition

2.3.3. Temporal patterns in benthic composition (2001-2007)

Temporal trends in the percent cover of live coral and algae were based on multiple pair-wise 

comparisons among sampling periods. A common approach to determining temporal trends in coral 

reef communities is the use of permanent sites and stations that are revisited periodically to examine 

successive changes in selected metrics. The data used in this study instead were collected using a 

stratified sampling design to randomly select sites that were never revisited. Thus, observed temporal 

trends reflect changes in average conditions within a habitat type between sampling periods rather than 

changes at specific sites. An underlying premise is that randomly selected sites are representative of 

the habitat strata from which they were selected.

2.3.3.1. Temporal patterns in live 

coral cover

Examination of differences in mean 

live coral cover among sampling 

periods revealed interesting patterns, 

which indicated that there has been a 

general decline in overall coral cover 

(Figures 2.19). Except for February 

2002, mean estimates among sites 

during earlier missions (January 2001 

through June 2006) were greater than 

those of later missions (December 

2002 through August 2007). The 

higher variance in coral cover 

observed during earlier missions may 

have resulted from smaller sample 

size (n<19) during earlier missions 

compared with later missions (n>29). 

However, examination of residual 

plots of percent coral cover ordered 

from Winter 2001 to Summer 2007 indicates that the 

frequency distribution and variance estimates of coral 

cover were similar across sampling periods and that 

the statistical populations can be compared across 

years. Further statistical analysis revealed that there 

was a consistent overall decline in live coral cover from 

Winter 2001 through Summer 2007, and the decline 

was significant for the five most abundant genera: 

Montastraea, Porites, Agaricia, and Siderastrea (Figure 

2.19, Table 2.6). 

When coral cover data were analyzed 

separately for different habitat types, 

statistically significant differences among 

years were detected. Live coral cover varied 

significantly among sampling periods within two 

hardbottom habitat types: colonized pavement 

and colonized pavement with sand channels 

habitats (Table 2.7). Non-parametric multiple 

comparisons of mean live coral cover among 

sampling periods within colonized pavement 

Table 2.7. Results of non-parametric ANOVA (Wilcoxon test) to 

determine significant differences in percent cover of live coral among 

sampling periods within each in hardbottom habitat in the southwest 

Puerto Rico study region.

0

5

10

15

Mean live coral cover (%)

Figure 2.19.Seasonal and inter-annual patterns in mean (± SE) percent live coral 

cover of on hardbottom habitats over a seven year period in the southwest Puerto 

Rico study region.

Hardbottom habitat types N X2 DF P

Colonized Pavement 110 29.7 14 0.008*

Col. Pavement with Sand Channels 212 33.3 15 0.004*

Linear Reef 100 17.0 15 0.317

Patch Reef (Aggregated) 27 11.6 9 0.237

Patch Reef (Individual) 26 8.1 10 0.621

Scattered Coral/Rock in Sand 94 12.3 15 0.654

Hard genus J Z P N

All hard corals 113,946.50 16.9 < 0.0001 572

Montastraea 95,339.90 9.0 < 0.0001 572

Diploria 85,418.50 4.8 < 0.0001 572

Porites 90,439.00 6.7 < 0.0001 572

Siderastrea 94,496.50 8.6 < 0.0001 572

Agaricia 81,357.50 2.8 0.0029 572

Table 2.6. Results of non-parametric ordered comparisons 

(JT test) to determine trends in percent live cover of the five 

most abundant coral genera in SW Puerto Rico from Winter 

2001 through Summer 2007. N = 572.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

30 2 - Benthic Composition 

with sand channels revealed 

that mean live coral cover was 

significantly higher during September 

2003 compared with January 2006 

and August 2007 (p<0.05, Figure 

2.20). Non-parametric multiple pairwise comparisons of mean live coral 

in colonized pavement habitats did 

not reveal which sampling periods 

were statistically different, although 

a non-parametric One-Way ANOVA 

indicated statistically significant 

variation among sampling periods 

(Table 2.7; Figure 2.20). In terms of 

overall trends, there was a decreasing 

trend in hard coral cover from Winter 

2001 through Summer 2007 in 

five habitats (colonized pavement, 

colonized pavement with Sand 

channels, Linear reef, Patch reef, and 

scattered coral/rock in unconsolidated 

sediment (Table 2.8). 

Non-parametric Wilcoxon tests 

revealed that mean live cover of 

M. annularis complex also varied 

significantly among sampling periods 

(x2

=39.0, p=0.0006). The temporal 

trend in mean estimates suggested 

that live Montastraea cover may have 

decreased through time (Figure 2.21). 

Non-parametric multiple pair-wise 

comparisons revealed that observed 

live cover of M. annularis complex 

was significantly higher during later 

sampling periods (September 2003 

through August 2007) compared 

with June 2002 (p<0.05; Figure 

2.21). However, there was an overall 

significant declining trend in cover of 

M. annularis complex during the study 

period, and the decline was strongest 

from winter 2003 through summer 

2007 (J=95,339, P<0.0001, N=572).

Further analysis revealed disparate 

results for significant differences 

among sampling periods in live coral 

cover of the five most common coral 

genera on hardbottom habitat types 

surveyed in La Parguera (Table 2.9). 

0

8

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24

32

Mean coral cover (%)

Colonized Pavement

Colonized Pavement with Sand Channels

Figure 2.20. Seasonal and inter-annual patterns in mean (± SE) percent live coral 

cover on two hardbottom habitat types over a seven year period in the southwest 

Puerto Rico study region. Means with similarly shaded arrows were not significantly 

different from each other (p<0.05).

0

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9

Montastraea live cover (%)

Figure 2.21. Seasonal and inter-annual patterns in mean (± SE) percent live cover of 

Montastraea spp. on hardbottom habitats over a seven year period in the southwest 

Puerto Rico study region. Means with similarly shaded arrows were not significantly 

different from each other (p<0.05).

Table 2.8. Results of non-parametric ordered comparisons (JT test) to determine 

trends in percent live cover of coral within habitat types in southwest Puerto Rico 

from Winter 2001 through summer 2007.

Habitat type J Z P N

Colonized pavement 4,321.50 8.2 < 0.0001 110

Colonized pavement with Sand channels 18,429.50 15.9 < 0.0001 212

Linear reef 4,340.50 12.2 < 0.0001 100

Patch reef (aggregated) 292.50 5.6 < 0.0001 27

Patch reef (individual) 279.50 5.9 < 0.0001 26

Scattered Coral / rock in unconsolidated 

sediment 2,461.50 3.2 0.0 94
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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31 2 - Benthic Composition

All five genera (Montastraea, Porites, 

Siderastrea, Diploria and Agaricia) 

showed significant differences among 

sampling periods within at least one 

hardbottom habitat (p<0.05; Table 2.10). 

Significant differences in mean live cover 

of these coral genera among sampling 

periods were detected for pavement and 

linear reef habitats, which were the more 

common habitats in La Parguera, and also 

were the ones most intensely surveyed 

(Table 2.10). 

Hard bottom Habitat Montastraea Porites Siderastrea Diploria Agaricia

Colonized Pavement 0.009 0.064 0.193 0.001 0.070

Col. Pav. w/ Sand Channels 0.098 0.003 0.027 0.837 0.001

Linear Reef 0.174 0.051 0.037 0.110 0.009

Patch Reef (Aggregated) 0.409 0.400 0.141 0.511 0.555

Patch Reef (Individual) 0.271 0.249 0.133 0.571 0.162

Scattered Coral/Rock in Sand 0.747 0.972 0.470 0.231 0.849

Table 2.9. Results of non-parametric ANOVA (Wilcoxon test) to determine 

significant differences in percent live cover of coral genera among sampling 

periods within each in hardbottom habitat in the southwest Puerto Rico study 

region.

A) Montastraea

Habitat type J Z P N

Colonized pavement 3,573.50 4.4 < 0.0001 110

Colonized pavement with Sand channels 14,141.50 7.6 < 0.0001 212

Linear reef 3,426.50 6.9 < 0.0001 100

Patch reef (aggregated) 256.00 4.1 < 0.0001 27

Patch reef (individual) 244.00 4.4 < 0.0001 26

Scattered Coral / rock in unconsolidated sediment 2,089.00 0.6 0.271 94

B) Diploria

Habitat type J Z P N

Colonized pavement 2,421.50 -2.0 0.022 110

Colonized pavement with Sand channels 12,650.00 4.8 < 0.0001 212

Linear reef 3,130.50 5.1 < 0.0001 100

Patch reef (aggregated) 198.00 2.0 < 0.0001 27

Patch reef (individual) 223.50 3.5 0.0002 26

Scattered Coral / rock in unconsolidated sediment 2,046.00 0.3 0.367 94

C) Porites

Habitat type J Z P N

Colonized pavement 3,034.00 1.5 0.0636 110

Colonized pavement with Sand channels 13,330.00 6.0 < 0.0001 212

Linear reef 3,525.00 7.3 < 0.0001 100

Patch reef (aggregated) 251.00 3.9 < 0.0001 27

Patch reef (individual) 243.50 4.4 < 0.0001 26

Scattered Coral / rock in unconsolidated sediment 2,147.00 1.1 0.1368 94

D) Siderastrea

Habitat type J Z P N

Colonized pavement 3,313.50 3.0 0.0012 110

Colonized pavement with Sand channels 14,924.00 9.1 < 0.0001 212

Linear reef 3,381.00 6.5 < 0.0001 100

Patch reef (aggregated) 199.50 1.8 0.0398 27

Patch reef (individual) 221.50 3.4 0.0003 26

Scattered Coral / rock in unconsolidated sediment 2,101.00 0.7 0.2447 94

E) Agaricia

Habitat type J Z P N

Colonized pavement 3,123.00 2.0 0.0202 110

Colonized pavement with Sand channels 12,094.50 3.6 0.0001 212

Linear reef 2,786.50 3.0 0.0012 100

Patch reef (aggregated) 187.00 1.3 0.105 27

Patch reef (individual) 196.50 2.4 0.0079 26

Scattered Coral / rock in unconsolidated sediment 2,026.50 0.1 0.4524 94

Table 2.10. A-E. Results of non-parametric ordered comparisons (JT test) to determine trends in percent 

live cover of coral genera within habitat types in southwest Puerto Rico from Winter 2001 - Summer 2007.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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32 2 - Benthic Composition 

Non-parametric multiple pair-wise 

comparisons between sampling 

periods revealed significant 

differences in mean live cover for 

the most abundant coral genera 

within a few hardbottom habitat 

types. Temporal variability in mean 

live cover of Montastraea spp. was 

significant for colonized pavement 

(p=0.009), and appeared to be 

lower during later sampling periods 

compared with earlier ones. Mean 

live cover of Montastraea spp. 

was 19.3 ± 2.4%, in October 2001 

(n=2) but over time gradually 

decreased to 2.7 ± 1.0 (n=15) by 

August 2007 (Figure 2.22a). On 

colonized pavement with sand 

channel hardbottom habitats, 

mean live cover of Porites spp. was 

4.1 ± 0.2% in May 2001 (n=3) but 

gradually decreased significantly 

to 0.4 ± 0.1% (n=23) by January 

2006 (Figure 2.22b). Likewise, 

mean live cover of Diploria spp. in 

colonized pavement habitat was 

1.6 ± 0.8% (n=3) in June 2002 but 

was significantly lower (p<0.05) in 

January 2006 (0.1 ± 0.1%, n=12). 

During January 2007 and August 

2007, live cover of Diploria spp. 

was essentially 0% (Figure 2.22c).

On both linear reef and colonized 

pavement with sand channel 

hardbottom habitats, mean live 

cover of Siderastrea spp. varied 

significantly among sampling 

periods, although it appeared not to 

increase or decrease over time. Yet 

multiple comparison tests detected 

significant pair-wise differences 

among sampling periods only for 

linear reef habitats, where mean 

live cover of Siderastrea spp. was higher in August 2006 and August 2007 compared with June 2002 

(p<0.05; Figures 2.23a). Similarly, mean live cover of Agaricia spp. on linear reef habitats was essentially 

0% in June 2006 (n=7) but was significantly higher (0.7 ± 0.3%, n=9) by May 2003 (Figure 2.23b). 

Significant pair-wise differences in mean live cover of Agaricia spp. on colonized pavement with sand 

channel habitats were not detected among sampling period (Figure 2.23b). Interestingly, significant 

declining trends in percent cover were observed for all five genera (Montastraea, Porites, Siderastrea, 

Diploria and Agaricia) in most habitats (Table 2.10 A-E).

0

12

24

Montastraea cover (%)

Colonized Pavement

0

6

12

Porites cover (%)

Colonized Pavement with Sand Channels

0

2

4

Diploria cover (%)

Colonized Pavement

c)

b)

a)

Figure 2.22. Seasonal and inter-annual patterns in mean (± SE) percent live cover of three of the five 

most abundant coral genera: a) Montastraea spp., b) Porites spp. and c) Diploria spp., on hardbottom 

habitat types over a seven year period. Arrows indicate non-parametric pair-wise comparisons 

where significant differences among sampling periods were found. Means with similarly shaded 

arrows were not significantly different from each other (p< 0.05).
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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33 2 - Benthic Composition

a) 4

Colonized Pavement with Sand Channels

)

Linear Reef

%( ervco ea

2

rasteridS

0

b) 4

Colonized Pavement with Sand Channels

Linear Reef

)%( ervco2 iciaarAg

0

Figure 2.23. Seasonal and inter-annual patterns in mean (± SE) percent live cover of 

two of the five most abundant coral genera: a) Siderastrea spp. and b) Agaricia spp., on 

hardbottom habitat types over a seven year period. Arrows indicate non-parametric pairwise comparisons where significant differences among sampling periods were found. 

Means with similarly shaded arrows were not significantly different from each other (p< 

0.05).

2.3.3.2. Temporal patterns in algal cover

Mean algal cover varied significantly among sampling periods (Table 2.11). Non-parametric ordered 

comparisons indicated that there was a significant declining trend in cover of various types of algae 

(Table 2.12); whereas, non-parametric multiple comparison tests revealed several pair-wise differences 

in mean algal cover among years (Table 2.13a-d). Mean cover of macroalgae followed a sinusoidal 

pattern with highs of 31.9 ± 7.9% in January 2001, 17.4 ± 2.1% in March 2004, and 21.7 ± 2.8% in 

August 2008 (Figure 2.24a). Mean cover of turf algae exhibited a similar pattern, with higher peaks in 

mean cover occurring during February and December 2002 and lower peaks in cover during January 

and August 2005 (Figures 2.24b). Temporal pattern in mean cover of cyanobacteria/filamentous algae 

and CCA suggest that episodic blooms alternated with periods of low cover of those algal types between 

January 2001 and August 2007 (Figures 2.24c and 2.24d). 

Table 2.12. Results of non-parametric ANOVA (Wilcoxon test) to 

determine significant differences in percent live cover of algal types 

among sampling periods within each in hardbottom habitat in the 

southwest Puerto Rico study region.

Table 2.11. Results of non-parametric ordered comparisons (JT 

test) to determine trends in percent cover of algal types from 

Winter 2001 through summer 2007 in southwest Puerto Rico 

study region. N = 422.

Algal Group J Z P N

All algae 78,954.5 26.41 < 0.0001 422

Turf algae 66,370.5 17.70 < 0.0001 422

Crustose coralline algae 47,279.0 5.06 < 0.0001 422

Filamentous algae 52,848.0 8.76 < 0.0001 422

Macro algae 67,059.0 18.16 < 0.0001 422

Algae types X2 DF P

All algal types 56.8 15 0.000001

Macroalgae 60.8 15 0.000000

Turf algae 87.5 15 0.000000

Cyanobacteria / Filamentous algae 132.7 15 0.000000

Crustose coralline algae 90.6 15 0.000000
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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34 2 - Benthic Composition Table 2.13. Multiple non-parametric pair-wise comparisons of mean percent algae cover for: a) macroalgae, b) turf algae, c) cyanobacteria/

filamentous algae and d) crustose coralline algae among sampling periods (2001-2007) in the southwest Puerto Rico study region. Green 

boxes with dots indicate significant pair-wise differences (p<0.05).

Win-01 Spr-01 Fall-01 Win-02 Spr-02 Win-03 Spr-03 Fall-03 Spr-04 Sum-04 Win-05 Sum-05 Win-06 Sum-06 Win-07 Sum-07

Win-01

Spr-01

Fall-01

Win-02

Spr-02 ●

Win-03 ● ● ●

Spr-03

Fall-03

Spr-04 ●

Sum-04

Win-05

Sum-05 ●

Win-06 ●

Sum-06 ●

Win-07 ●

a) Macroalgae Sum-07

Win-01 Spr-01 Fall-01 Win-02 Spr-02 Win-03 Spr-03 Fall-03 Spr-04 Sum-04 Win-05 Sum-05 Win-06 Sum-06 Win-07 Sum-07

Win-01 ● ● ● ●

Spr-01 ●

Fall-01 ●

Win-02 ● ●

Spr-02 ●

Win-03 ● ● ● ● ●

Spr-03 ●

Fall-03 ●

Spr-04 ● ●

Sum-04

Win-05

Sum-05

Win-06 ●

Sum-06 ●

Win-07 ●

b) Turf algae Sum-07

Win-01 Spr-01 Fall-01 Win-02 Spr-02 Win-03 Spr-03 Fall-03 Spr-04 Sum-04 Win-05 Sum-05 Win-06 Sum-06 Win-07 Sum-07

Win-01 ● ● ●

Spr-01 ● ●

Fall-01 ● ● ● ●

Win-02 ● ●

Spr-02 ● ● ●

Win-03 ● ● ●

Spr-03 ● ● ●

Fall-03 ● ● ●

Spr-04 ●

Sum-04 ●

Win-05 ●

Sum-05 ●

Win-06 ●

Sum-06 ●

Win-07 ●

c) Cyanobacteria / filamentous algae Sum-07

Win-01 Spr-01 Fall-01 Win-02 Spr-02 Win-03 Spr-03 Fall-03 Spr-04 Sum-04 Win-05 Sum-05 Win-06 Sum-06 Win-07 Sum-07

Win-01

Spr-01 ● ●

Fall-01

Win-02 ● ● ● ●

Spr-02 ● ●

Win-03

Spr-03 ● ● ●

Fall-03 ●

Spr-04 ● ● ●

Sum-04 ● ● ●

Win-05

Sum-05 ● ●

Win-06 ●

Sum-06

Win-07 ●

d) Crustose coralline algae Sum-07
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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35 2 - Benthic Composition

0

25

50

Mean cover (%) 

Macroalgae

a)

0

30

60

Mean cover (%) 

Turf algae

0

4

8

Mean cover (%) 

cyanobacteria / filamentous algae

0

3

6

Mean cover (%) 

CCA

c)

b)

d)

Figure 2.24. Seasonal and inter-annual patterns in mean (± SE) percent cover of algal 

categories: a) macroalgae, b) turf algae, c) cyanobacteria/filamentous algae and d) 

crustose coralline algae on hardbottom habitat types over a seven year period. Results 

of non-parametric comparisons of means among sampling periods are shown in Tables 

2.13a-d.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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36 2 - Benthic Composition 2.3.4. Abundance and distribution of macroinvertebrates

2.3.4.1. Queen conch (Eustrombus gigas)

A total of 45 individual queen conch (Eustrombus gigas; 34 

immature and 11 mature) were recorded from 35 of 618 survey 

sites in the La Parguera study area since conch data collection 

began in August 2004 (Table 2.14). Seventy six percent of all 

conch observed were immature, and these occurred at 26 of the 

35 sites (Table 2.14). Mature conch were recorded at 10 of the 

35 sites, comprising approximately 24% of all conch recorded 

(Table 2.14). Immature and mature conch were found across the 

shelf, with no clear inshore-offshore pattern of spatial or habitat 

segregation, although immature and mature conch were present 

together at only one site (Figure 2.25). In contrast to other regions of the U.S. Caribbean, extensive 

areas of seagrass did not appear to support markedly higher occurrence of conch than other habitat 

types. 

Queen conch (Eustrombus gigas)

Year Habitat type

Number of 

sites where 

conch were 

observed

Number of 

immature 

conch

Number 

of mature 

conch

Total number 

of conch

Total

number 

of sites 

surveyed

Hard 3 2 1 3

2004 Soft 5 8 0 8 89

Mangrove 1 1 0 1

Hard 1 1 0 1

2005 Soft 2 4 0 4 170

Mangrove 1 1 0 1

Hard 4 4 1 5

2006 Soft 11 10 5 15 179

Mangrove 0 0 0 0

Hard 4 2 2 4

2007 Soft 3 1 2 3 180

Mangrove 0 0 0 0

Table 2.14. Number of total surveys, surveys where conch were observed and individuals 

counted during 2004-2007. Conch totals per year represent two missions per year, 

January and August, except in 2004: conch data collection began in August 2004.

Eustrombus gigas

Submerged

vegetation

Unconsolidated

sediment

Hard bottom

Mangrove

Substrate type

Figure 2.25. Distribution map of queen conch (Eustrombus gigas) in the southwest Puerto Rico study 

area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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37 2 - Benthic Composition

E. gigas was most abundant 

in 2006 (n=20) and least 

abundant in 2005 when 

no adults were recorded 

(n=6; Table 2.14). In almost 

all years abundance was 

higher in softbottom habitat 

than hardbottom and lowest 

in mangroves. Highest 

density of mature conch was 

2 individuals per 100 m2

 of 

softbottom (Figure 2.26b). 

The greatest number (n=3) 

of conch observed at any 

one site occurred twice: at 

an unconsolidated sediment 

site during the January 2005 

field mission and again at 

an unconsolidated sediment 

site during the August 2006 

field mission (Figure 2.26a). 

Only immature E. gigas were 

recorded during mangrove 

surveys in 2004 and 2005 

(Figure 2.27); no mature 

E. gigas were recorded 

in mangroves. Immature 

E. gigas were also more 

likely to be encountered 

over hardbottom than 

were mature conch (Figure 

2.27a). 

Immature Eustrombus gigas

Submerged

vegetation

Unconsolidated

sediment

Hard bottom

Mangrove

Substrate type

Submerged

vegetation

Unconsolidated

sediment

Hard bottom

Mangrove

Substrate type

Mature Eustrombus gigas

a)

b)

Figure 2.26. Distribution maps of a) immature and b) mature queen conch (Eustrombus gigas) in 

the southwest Puerto Rico study area.

Figure 2.27. Abundance of a) immature and b) mature queen conch (Eustrombus gigas) in the southwest Puerto Rico study area.

0

5

10

2004 2005 2006 2007

Immature E. gigas abundance (100 m2)

Hard

Soft

Mangrove

a)

0

5

10

2004 2005 2006 2007

Mature E. gigas abundance (100 m2)

Hard

Soft

Mangrove

b)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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38 2 - Benthic Composition 2.3.4.2. Long-spined sea urchin (Diadema antillarum)

Long-spined sea urchins (Diadema antillarum) have been included in habitat surveys since 2006 and 

have been observed primarily at hardbottom sites (Table 2.15). Across the study area, D. antillarum were 

observed at relatively few sites (n=12) out of 314 surveys, but were clustered in large numbers. A total 

of 192 individuals from 12 surveys 

sites were recorded between 2006 

and 2007 (Table 2.15). The highest 

abundance at a single site was 75 

individuals observed along a linear 

reef site. The one softbottom site 

surveyed with D. antillarum present 

was a seagrass habitat surveyed 

in August 2006, with 30 individuals 

recorded, also the second highest 

density for D. antillarum at any one 

site (Table 2.15).

2.3.4.3. Caribbean spiny lobster (Panulirus argus)

A total of 27 Caribbean spiny lobster 

(Panulirus argus) were observed 

at 14 of 469 (approximately 3.0%) 

sites from 2005-2007. Lobster were 

observed on all three substrate 

types in 2005, on hard and 

mangrove sites in 2006, and only on 

hardbottom habitats in 2007 (Table 

2.16). The highest abundance at 

one survey was seven individuals 

recorded at a mangrove survey site 

in August 2006 (Table 2.16).

2.3.5. Marine debris

Collection of marine debris data began in January 2007. One piece of cloth was recorded in January 

2007, and four pieces of debris were seen in August 2007. In addition to the cloth, debris found included 

a champagne bottle, a plastic cylinder, a PVC rectangular 

frame and a glass bottle. The area of all debris equaled 

1,014 cm2

 with a total affected area of 1,054 cm2. Debris 

were colonized by a combination of cyanobacteria, turf 

algae, sponges, crabs and macroalgae. The sites where 

debris was recorded ranged in average depth from 1 to 

12 meters. Habitat types included hardbottom, scattered 

coral and rock in sand, and mangrove. The largest amount 

of debris was seen at the mangrove site on the north side 

of Magueyes, where a large PVC rectangular frame and a 

plastic cylinder were found. A champagne bottle was seen 

on the north side of Media Luna in the sand of a hardbottom 

site; a glass bottle was north of Corral on a hardbottom site 

with some sand, and the cloth was found toward the shelf 

edge on hardbottom.

Table 2.15. Abundance of long-spined sea urchins (Diadema antillarum) observed 

within transects by year in the southwest Puerto Rico study area.

Year Habitat 

type

Sites 

surveyed

Sites 

observed

Total 

individuals

2006 Hard 91 7 64

2006 Soft 41 1 30

2006 Mangrove 20 0 0

2007 Hard 100 4 98

2007 Soft 42 0 0

2007 Mangrove 20 0 0

Total 314 12 192

Year Habitat 

type

Sites 

surveyed

Sites 

observed

Total 

individuals

2005 Hard 90 2 2

2005 Soft 52 1 2

2005 Mangrove 13 2 6

2006 Hard 91 4 5

2006 Soft 41 0 0

2006 Mangrove 20 1 7

2007 Hard 100 4 5

2007 Soft 42 0 0

2007 Mangrove 20 0 0

Total 469 14 27

Table 2.16. Abundance of Caribbean spiny lobster (Panulirus argus) observed within 

transects by year in the southwest Puerto Rico study area.

Marine debris in the mangroves of La Parguera.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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39 2 - Benthic Composition

2.3.6. Summary of results

• Colonized pavement habitat was the most common habitat type occurring in La Parguera.

• Hardbottom habitat types were dominated by algae, followed by gorgonians and stony corals.

• The cover of live stony coral averaged about 5% on hardbottom throughout the study area.

• Linear reefs represented 10% of the study area, but had the highest percent cover of live coral 

(approximately 7%).

• Live stony coral cover comprised of at least 24 genera, but only nine had a mean cover greater than 

0.01.

• Montastraea and Porites spp. were the most ubiquitous coral genera among habitat types and had 

the highest mean cover among observed genera. M. annularis complex had the most coverage 

(35% of total coral cover) whereas P. asteroides was the most frequently observed species (62% 

occurrence).

• On average, T. testudinum was the most abundant seagrass species in SAV habitats whereas S. 

filiforme had the most coverage in unconsolidated sediments such as sand or mud.

• Mangrove prop roots provide a major benthic substrate for colonization by epiphytic organism, with 

macroalgae having the highest coverage of benthic organisms observed thereon.

• Live coral cover was patchy in spatial distribution, and interpolations revealed that live coral cover 

and taxa richness correlated positively with substrate rugosity.

• Acropora species were rarely observed, most likely due to the sampling approach used in this study; 

A. cervicornis had a wider spatial distribution compared with A. palmata. 

• Highest coral cover was observed along the shelf edge southward of Terrumote I and around El Palo. 

Cover of A. palmata (4.1-10.4 %) was highest around El Palo, whereas highest cover of M. annularis

complex (25.1-30.3%) wast south-west of Turrumote I.

• Hot spots of coral species richness (10-14 species per 100 m2) and diversity (H = 1.7-2.5) occurred 

offshore, but were scattered throughout hard bottom habitat types. 

• Highest rugosity (0.8-1) occurred between Margarita Reef and El Palo and at Romero.

• Temporal analysis revealed that live coral cover varied significantly among sampling years. Overall, 

there was a trend of decreasing coral cover over time.

• Mean algal cover varied significantly over time, with different seasonal patterns being observed for 

various algal types.

• Densities of E. gigas were relatively low, with 76% of observed conch being immature and below the 

legal size class for the fishery.

• D. antillarum were only observed at a few sites, but occurred 

in large numbers when they were encountered.

• P. argus were infrequently observed; with the highest number 

of individuals were observed in mangrove during 2006. 

• The largest piece of debris was seen on the northern side 

of Magueyes, where a large PVC rectangular frame and 

a plastic cylinder were found. Debris were colonized by a 

combination of algae, cyanobacteria, sponges, crabs and 

shrimp. Shrimp
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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40 2 - Benthic Composition 2.4. Discussion

2.4.1. Colonized hardbottom habitats: benthic characterization and spatial patterns

Marine benthic maps of nearshore environments have become an important tool for conservation and 

management of biological resources. Due to cost restraints, such maps often are produced at low 

resolutions (i.e., large scales) that typically do not capture the full spectrum of spatial variation in the 

distribution and composition of benthic resources. A good example of this is the minimum mapping unit 

(MMU) of 0.4 ha (1 acre or approximately 4,047 m2) used to develop benthic maps of La Parguera, 

Puerto Rico and which resulted in the identification of 26 distinct benthic habitat types (Kendall et al., 

2001). However, some softbottom polygons (i.e., habitat types) included unidentified benthic features 

such as sand halos and patch reefs that were smaller than the MMU. These unidentified features are 

known to influence the spatial distribution and occurrence of marine fauna at multiple scales (Parrish, 

1989; Kendall et al., 2003; Chittaro, 2004). Likewise, spatial patterns in benthic composition can be 

influenced by marine fauna such as fishes and invertebrates at spatial scales more resolved than a 

MMU of 0.4 ha (Helfman, 1978, 1982; Meyer et al., 1983; Pittman et al., 2004; Burkepile and Hay, 2008). 

By quantitatively characterizing temporal and sub-meter spatial variation in benthic composition and 

physical attributes of mapped polygons, this study provided additional information for use in elucidating 

species-habitat relationships, understanding spatial patterns in the distribution of marine fauna, and 

identifying faunal effects on benthic composition. 

Although the composition of benthic substrates varied spatially within and among habitat types, 

some general spatial patterns in occurrence and cover of benthic organisms were observed. For 

example, turf algae – defined as a multispecific assemblage of small filamentous algae – was the most 

extensively occurring benthic organism group within all hardbottom 

habitat types, followed by macroalgae and a low occurrence 

of CCA. A widely accepted hypothesis is that the abundance of 

algae on coral reefs typically is controlled by herbivory (Steneck 

and Dethier, 1994). Under high rates of herbivory, coral reefs and 

hardbottom substrates generally are characterized by low-biomass 

algal assemblages dominated by turf, with low cover of CCA, and 

macroalgae (Steneck and Dethier, 1994). In contrast, under low 

rates of herbivory, macroalgae forms the dominant algal cover type, 

followed by turf algae, and few crustose coralline algae (Steneck 

and Dethier, 1994). 

The extensive coverage of turf algae observed during this study suggests that rates of herbivory on 

reefs and hardbottom areas in la Parguera may be high. This seems counter-intuitive, given the widely 

accepted belief that a Caribbean-wide decline in the abundance of D. antillarum along with serial overfishing have reduced herbivory and have resulted in a phase-shift from coral-dominated to macroalgaldominated reefs in the Caribbean (Lessios et al., 1984; Gardner et al., 2003; Hughes, 1994). During this 

study, the abundance of long-spined sea urchins was relatively low, but small-bodied herbivorous fishes 

were numerically abundant and dominated fish assemblages across the 

shelf (Fish chapter, this report). Idjadi et al. (2006) implicated structural 

refugia from long-lived colonies of M. annularis complex in a phase-shift 

reversal from macroalgae-dominated back to coral-dominated reefs 

in Jamaica. Bechtel et al. (2006) also implicated a resurgence of D. 

antillarum and an abundance of other echinoids in an observed decline 

in macroalgal cover and a corresponding increase in scleractinian coral 

cover on a reef in Jamaica. Based on our data from La Parguera, it 

is possible that rates of herbivory from relatively abundant fishes and 

other observed echinoids may have provided enough herbivory to keep 

the cover of macroalgae relatively low while turf algae flourished. 

Dictyota sp. surrounding Dichocoenia stokesi

M. annularis complex
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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41 2 - Benthic Composition

The high cover of turf algae on reef and hardbottom areas in La 

Parguera also may be due to relatively quick growth rates that provide 

a competitive advantage to this algal group over slower-growing coral 

and crustose coralline algae. CCA play a crucial role in coral reef ecology 

by contributing calcium carbonate to reef structure and by facilitating 

the settlement and colonization of scleractinian corals (Steneck and 

Dethier, 1994). Turf algae trap and stabilize unconsolidated sediments, 

but also rapidly overgrow and kill underlying coralline algae and 

coral colonies through encroachment (Steneck and Dethier, 1994). 

In La Parguera, the coverage of CCA was very low, indicating that 

overgrowth of CCA and corals may have occurred, and that very little 

suitable substrates exist for newly settling coral colonies.

Another general pattern in benthic composition observed during this study was the low average cover 

of live scleractinian coral (approximately 5-7%) on reef and hardbottom areas and the virtual absence 

of Acropora spp. thickets on forereefs (0.5-6 m depth) in La Parguera. Such low coral live cover is 

now typical of most reefs in the Caribbean and has resulted from the synergistic effect of natural 

and anthropogenic factors operating over the past three decades (Gardener et al., 2003). Acroporid 

populations were formally very abundant on reefs near La Parguera (Weil et al., 2002). Since the 

late 1970s however, successive disease outbreaks, periodic hurricanes, bleaching events, predators, 

and increased anthropogenic activities have contributed to the demise of acroporid and other coral 

populations in La Parguera (Weil et al., 2002). Following the major outbreak of white band disease, several 

tropical cyclones decimated shallow-water acroporid populations in La 

Parguera between 1979 and 1998 and thereby removed a major source 

of structural complexity from those coral reefs (reviewed in GarciaSais et al., 2005, 2008; Weil et al., 2002). No major hurricane has hit 

Puerto Rico since 1998, but per this study, reefs at depths less than 18 

m in La Parguera remain depauperate of acroporid corals and hence 

also their important function of providing high structural complexity for 

fish and other organisms. Most recently, the major Caribbean-wide 

bleaching event in 2005 resulted in additional coral mortality, such that 

coral cover declined by 40-60% (Garcia-Sais et al., 2008). 

Interpolations of this study’s synoptic estimates of live coral cover summed across species revealed 

a few areas of relatively high cover that could be considered hot spots of live coral (see Figure 2.13a, 

pg. 21). These hotspots may be refuge areas where demographic processes have resulted in coral 

populations that are resilient to multiple synergistic stressors. If so, corals at these locations are more 

likely to persist longer in the future than corals at other locations. Additionally, the locations of such 

hotspots corresponded with areas of relatively high rugosity, coral species richness, and diversity (Figures 

2.13b, c, pg. 21). Protection of such hotspots may benefit ecosystem conservation. Interestingly, the 

five most dominant species in terms of coral cover in La Parguera were two frame building species (M. 

cavernosa and M. annularis complex) and three more weedy species 

(P. astreoides, Agaricia spp. and S. siderea). Frame building corals are 

important in that they provide structural complexity and are also major 

contributors to reef growth and persistence, whereas weedy species 

provide very little complexity and contribute relatively little to reef growth 

(Hoegh-Guldberg et al., 2007). If these hotspots are to be selected for 

increased management and protection from anthropogenic stressors, 

further work is needed to understand the physical and oceanographic 

properties that correlate with their enhanced ecological features.

Crustose coralline algae

Diseased S. sidera

S. radians
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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42 2 - Benthic Composition An interesting feature of reef and hardbottom areas in La 

Parguera, is the preponderance of gorgonians (sea rods, 

sea whips sea plumes, and sea fans) at sites throughout 

the shelf. Also known as soft corals, gorgonians were 

commonly observed intermingled with other sessile 

benthic organisms, and often dominated colonized 

pavement and patch reef habitats in terms of cover. At 

some colonized pavement sites for example, the canopy 

created by the high density and cover of gorgonians 

virtually obscured the sea floor such that other sessile 

benthic organisms were not visible. Percent cover of soft 

corals also correlated positively with rugosity, probably 

because through niche partitioning, more rugose areas 

provide a wider variety of benthic substrate for settlement 

and colonization by soft corals than less rugose areas (MacArthur and Levins, 1964). Although they 

contribute minimally to calcium carbonate accretion and reef growth, soft corals represent important 

ecosystem components on the La Parguera shelf in that they provide important habitat for fishes and 

feeding sea turtles (Gratwicke et al., 2006; Blumenthal et al., 2009). Additionally, soft corals can affect 

the spatial distributions of other sessile benthic organisms because they are superior competitors for 

space, and they are able to secrete allelo-chemicals that deter growth of other benthic organisms 

(Fenical, 1987; Harvell and Fenical, 1989).

2.4.2. Softbottom habitats: benthic characterization and spatial patterns

Analysis of the benthic maps used in this study showed that softbottom habitats comprised approximately 

33% of the study area (22.1% seagrass, 4.6% macroalgae and 5.9% unconsolidated sediments (Table 

2.3, pg. 15). As shown by the spatial interpolations of synoptic estimates from this study, softbottom 

areas in La Parguera exhibited a zonation pattern typical of 

Caribbean shallow-water ecosystems; seagrass percent cover 

was highest near the shore but decreased toward deeper 

offshore areas. Similarly, the spatial distributions of the two 

most commonly occurring seagrass species also were zoned. 

T. testudinum dominated shallow water near shore areas down 

to a depth of 16 m, whereas S. filiforme dominated deeper 

areas offshore. Such zonation patterns result generally from 

decreasing nearshore-to-offshore gradients of nutrients and light 

penetration. Sponges and native coral species (e.g., Dichocoenia 

stokesii) also were observed frequently in seagrass and macro 

algae habitats. Calcareous macroalgae (e.g., Halimeda spp., 

Udotea spp., and Penicillus spp.) were commonly encountered 

on soft bottom habitats, but their percent cover was low relative 

to those of seagrass and more foliose algae such as Lobophora, 

Dictyota, and Padina spp. Nevertheless calcareous algae are 

ecologically important to coral reef communities because their 

skeletal remains (e.g., Halimeda spp.) are a major component 

of carbonate sediments occurring within coral reef ecosystems 

(Hubbard et al., 1981; Drew, 1983).

Interestingly, benthic organisms typical of coral reefs and hardbottom substrates (e.g., turf algae, CCA, 

scleractinian corals and gorgonians) occasionally were encountered within areas mapped as softbottom. 

The atypical occurrence of these reef-associated organisms within softbottom polygons was likely an 

artifact of differences between the scale at which map polygons were delineated and the scale at which 

S. filiforme

Dense gorgonian habitat

T. testudinum
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43 2 - Benthic Composition

benthic data were collected. An MMU of 0.4 ha did not allow delineation of reef and hardbottom patches 

less than 0.4 ha that were encountered within areas mapped as softbottom. Thus, our fine-scale 1 m2

quadrat benthic surveys on these hardbottom patches that occurred within softbottom areas provide 

additional data that may be crucial for understanding observed relationships between faunal species 

and their mapped habitats. 

Several studies have shown that softbottom habitats are ecologically important components of coral reef 

ecosystems. For example, reef fishes are known to migrate from reef and hard bottom areas, forage on 

adjacent non coral reef habitats (sand, seagrasses, algal plains), and they represent a trophic pathway 

of energy transfer among habitats (McFarland et al., 1979; Meyer et al., 1983). Furthermore, several 

landscape analyses have correlated various seagrass metrics with increased probability of juvenile grunt 

occurrence on reef and hardbottom areas in St. Croix (Kendall et al., 2003), higher sighting frequencies 

of groupers on hardbottom habitats in the Florida Keys (Jeffrey, 2004), and increased fish abundance 

and species richness in mangrove communities in Puerto Rico (Pittman et al., 2007a). Several other 

studies have demonstrated that both vegetated and non-vegetated softbottom areas are known to 

provide habitat and food for several coral reef fishery species, endangered and threatened species, 

and many other marine organisms (Parrish, 1989; Nagelkerken et al., 2000; Dahlgren and Marr, 2004; 

Adams et al., 2006). Fine-scale benthic characterizations, such as those conducted during this study, 

should provide additional information to further explain these faunal species-habitat relationships.

2.4.3. Benthic characterization of mangrove habitats

In La Parguera, mangroves are very distinctive intertidal and nearshore habitat features which are dominated by red mangrove 

(Rhizopora mangle). Extensive mangrove stands occur along the 

shoreline as well as form islands in the back reef lagoon areas of 

the La Parguera coral reef ecosystem, where they support abundant 

fish populations (Christensen et al., 2003; Pittman et al., 2007b). The 

tidal range in La Parguera is less than 0.5 m, thus most prop roots 

seaward of the mangrove community are continuously submerged. 

Our benthic characterizations of mangrove habitats found extensive 

cover of benthic organisms, particularly algae, sponges and hydroids, 

on both the seafloor and as epiphytes on submerged prop roots. The 

high mean cover (>50%) of vegetation observed on benthic substrates 

indicates that the La Parguera mangroves are highly productive 

systems which may be providing enough food and nutrients to support 

resident fish and invertebrate populations. Additionally, many sites had 

thick deposits of detritus, suggesting that mangroves were performing 

their function of entrapping sediments. 

The benthic characterizations of mangrove habitats reported in this 

study have provided information that could be used as covariates to 

explain spatial variance in fish populations among mangroves and 

other benthic habitats. Further research is necessary however to 

identify additional characteristics of mangrove communities that may 

vary spatially and may be influencing benthic composition and faunal 

distributions. For example, mangroves are major producers of detritus 

that may be contributing to offshore productivity (Odum and Heald, 

1972, 1975). However, rates of productivity and generation of detritus 

in mangroves will vary based on location, oceanographic properties, 

composition of submerged vegetation, and frequency of tidal flushing.

Tunicates on mangrove prop root

Sponges on mangrove prop root

 Mussels on mangrove prop root
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44 2 - Benthic Composition 2.4.4. Temporal trends in benthic composition on hardbottom habitats

Temporal analysis of data on percent cover revealed a general decrease in live coral in La Parguera, 

particularly on pavement habitats between fall of 2003 and summer of 2007. Our observations of a 

temporal decline in live coral cover at La Parguera are consistent with those of other recent studies 

from Puerto Rico and the USVI. Temporal declines ranging from 40-50% in live coral were reported 

at several sites in Puerto Rico including reefs off Isla Desecheo, Mayguez, Guanica and Ponce, with 

most of the loss occurring after the 2005 bleaching event (Garcia-Sais et al., 2008). At Buck Island, St. 

Croix, mean estimates of live coral cover on reefs were lowest in 2006 after four years of observations 

(Pittman et al., 2008; Clark et al., 2009). Similarly, in St. John, average live coral cover declined from 

21.4% in 2005 to 8% by October 2007 (Miller at al., 2009). 

Much of the reported loss in live coral occurred in a few species, 

namely M. annularis complex, C. natans and Agaricia agaricites

(St. John), M. annularis complex (Puerto Rico), and M. annularis

complex and Agaricia spp. (Buck Island, St. Croix). After the 

drastic decline in acroporid corals, Montastraea remained one of 

the most abundant coral species in La Parguera (per this study) 

and in other areas of the U.S. Caribbean (Garcia-Sais et al., 

2008; Rothenberger et al. 2008; Miller et al., 2009). Given their 

dominance and key ecological roles as reef-building species, the 

recent declines in the cover of Montastraea species represent a 

severe degradation to already fragile reef ecosystems. 

Such coral loss and reef degradation have ultimate ecological and economic consequences. 

However, there is still a lack of understanding about the ecosystem properties that confer resilience 

and sustainable ecological function to coral reefs (Done, 1992). Consequently, further research is 

needed to identify areas within near-shore ecosystems with physical and ecological properties that 

correlate well with enhanced ecosystem resilience to multiple stressors. Identification of such areas 

can help managers design and manage protected areas to promote ecosystem conservation. Our 

characterizations of benthic composition and descriptions of spatial patterns at La Parguera provide 

a foundation for identifying locations with enhanced ecological properties that may be resistant and 

resilient to manageable anthropogenic stressors such as over-fishing, land-based sources of pollution, 

and habitat destruction.

2.4.5. Marine debris

Marine debris, as defined by NOAA’s Marine Debris program, 

is “any persistent solid material that is manufactured 

or processed and directly or indirectly, intentionally or 

unintentionally, disposed of or abandoned into the marine 

environment or the Great Lakes” (http://marinedebris.noaa.

gov/info/welcome.html, accessed 26 Jan 2010). There are 

several types of marine debris from a variety of sources 

that have multiple impacts on the marine environment. 

Marine debris is composed mainly of plastics, glass, metal, 

rubber, derelict fishing gear and derelict vessels. The types 

of marine debris commonly seen in the waters around La 

Parguera include plastics, glass, rubber and derelict fishing 

gear. 

Marine debris originates from ocean-based and land-based sources including cargo ships, fishing and 

other vessels, littering, dumping, poor waste management, storm water discharge and extreme natural 

events such as hurricanes and floods. The effects of marine debris are difficult to measure but affect 

Colpophyllia natans

Derelict trap
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the aesthetics of shorelines and coastal environments world-wide. Debris can also cause damage by 

scouring, breaking or smothering important and fragile marine habitats and entangling wildlife such 

as sea turtles, whales and fish. Ingestion of marine debris can prove fatal to animals by leading to 

loss of nutrition, internal injury or blockage and starvation. Marine debris can transport organisms 

from native habitats to non-native ones, serving as the vector for potentially catastrophic alien species 

introductions. Marine debris can be dangerous to safe navigation of vessels and has in the past caused 

vessel damage and loss of navigation abilities of large NOAA research vessels. 

The debris data collected in La Parguera from 2006-2007 revealed only four locations of debris. It is 

important to recall that only debris on the 25 x 4 m transect was recorded, explaining the low number of 

sites with debris. Several (unquantified) pieces of debris were seen outside transects. 

The marine debris around La Parguera may have a detrimental economic impact due to degradation 

of the marine habitat that tourism and human health depend upon. Boat damage, water pollution from 

storm runoff, and health issues are all potential effects. Further studies into the amount, distribution 

and affects of marine debris around the study area will provide useful information for future action and 

protection of this marine environment.

2.4.6. Macroinvertebrates

2.4.6.1. Queen conch (E. gigas)

Queen conch are ecologically important components of faunal assemblages that occur in Caribbean 

coastal ecosystems. They are important herbivores that feed on micro- and macro-algae (Stoner ,1997), 

become prey for several marine organisms such as reptiles, fishes, and crustaceans (Randall, 1964), 

and their populations support commercial fisheries that were valued as much as U.S. $30 million in 

1992 (Appeldoorn and Rodriguez, 1994). However, Caribbean-wide declines in annual queen conch 

landings – most likely from overfishing and habitat degradation – resulted in the species being listed as 

commercially threatened and being protected under Appendix II of CITES in 1992 (Wells et al., 1985; 

Appeldoorn, 1994). Protection under Appendix II means that management of queen conch stocks and 

monitoring of exports are necessary to prevent extinction of the species.

Reported commercial landings of queen conch from Puerto Rico decreased from 402,510 pounds 

in 1983 to 90,947 pounds in 1992 (CFMC, 1996). Although reported conch landings subsequently 

increased in 1993 to 164,612 pounds and to as much as 281,378 pounds by 2002, the Puerto Rican 

queen conch fishery has not rebounded to 1983 levels (SEDAR, 2007). It is likely that annual landings 

of queen conch from La Parguera mirrored trends in the reported conch landings for Puerto Rico. 

Queen conch is a major fishery in La Parguera. Conch meat (locally known as "carucho") is an easily 

accessible source of protein for locals because extensive shallow seagrass and algal beds typically 

inhabited by conch are in close proximity to coastal villages and towns. Additionally, conch meat is 

a staple on the menu of many local restaurants (pers. obs). Conch assemblages in La Parguera are 

fished both by local artisanal (recreational) fishers for home consumption (pers. obs) and by commercial 

fishers from Puerto Rico’s west coast (Appeldoorn, 1992). Landings from west coast fishers accounted 

for a substantial proportion of conch landings reported for Puerto Rico (Appeldoorn, 1992; SEDAR, 

2007). It is unclear what proportion of conch landings from La Parguera are taken by artisanal fishers 

because most of their catch is unreported, but artisanal catch was estimated to be 35% of commercial 

landings in 1986 (CFMC, 2005; Matos-Caraballo, 2004; Diaz, 2007). If this estimate is realistic, then it 

is reasonable assume that both artisanal and commercial fishing over the past few decades has had 

negative demographic consequences on conch assemblages in La Parguera.

Concern that historical over-fishing may have been decimating queen conch populations resulted in 

the implementation of federal and territorial regulations during the past 16 years to reduce the harvest 

of conch from U.S. Caribbean territories (CFMC, 1996). Implemented regulations applicable to Puerto 
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46 2 - Benthic Composition Rico include: (1) the requirement of a territorial fishing license and permit since 1994 to catch conch; 

(2) prohibiting the use of hookahs as fishing gear within Puerto Rico’s Exclusive Economic Zone (EEZ) 

since 1997 and within territorial waters since 2004; (3) a legal size limit of 23 cm (9 inches) or greater 

in total shell length and lip thickness greater than 3 mm (1/8th of an inch) the EEZ since 1997 and for 

territorial waters since 2004; (4) maximum daily quotas of 150 queen conch per commercial fisher or 

450 per commercial fishing vessel and three queen conch per recreational fisher or 12 per recreational 

vessel from the EEZ since 1996 and from territorial waters since 2004; (5) complete closure of the 

queen conch fishery within Puerto Rico’s EEZ since 1997; and (6) an annual seasonal closure from July 

1st through September 30th within the EEZ since 1996 and within Puerto Rico’s territorial waters since 

1997 (CFMC, 1996; SEDAR, 2007). Interestingly, there was a seven-year delay in the implementation 

of daily quotas and gear restrictions in territorial waters compared with federal (EEZ) waters.

The federal and territorial regulations for queen conch were implemented to reduce harvest, with the 

expectation that if fishing mortality on conch assemblages was reduced, then in situ abundance and 

occurrence of conch would increase over time. The data collected by this study suggest that regulations 

have not been effective in protecting queen conch assemblages at La Parguera. The densities and 

occurrence of queen conch observed in La Parguera region were relatively low. This study surveyed 

6.18 ha and observed queen conch at only 6% of the 618 surveys region between August 2004 and 

August 2007. Average queen conch density was 0.073 individuals per 100 m2

, with highest density 

being two adult mature individuals per 100 m2

. Additionally, the maximum number of conch observed 

during a sampling mission was 12 individuals in August 2006. Not only was observed conch abundance 

very low in La Parguera, 76% of conch encountered were immature (i.e., their shells had not yet 

developed a lip) and were below the legal size class for the fishery.

Data from a previous study that surveyed 81 randomly selected sites in La Parguera observed similarly 

low densities of queen conch between May 1985 and April 1986. Torres Rosado (1987) visually surveyed 

40.81 ha of bottom substrate and observed 331 queen conch, with a density of 0.08 conch per 100 m2. 

Of the 331 queen conch observed, 227 (68%) were immature and had shells without lips. The similarity 

in queen conch densities observed during 1985-1986 and those observed during this study period 

(2001-2007), suggests that queen conch stocks were 

already depressed in 1985, and have not rebounded 

since then. Stock assessment studies indicate that 

queen conch requires minimum population densities of 

56 individuals per ha for successful reproduction and 

recruitment (Stoner and Ray-Culp, 2000), although 

observations in the Florida Keys have also revealed 

that successful mating occurred when densities were 

200 individuals per ha or higher (Bob Glazer, pers. 

comm). Density of queen conch was only eight conch 

per ha during 1985-86 (Torres Rosado, 1987) and only 

seven conch per ha during 2001-2007 (this study). 

It is very likely that densities of queen conch in La 

Parguera now are too low for successful reproduction 

and recruitment, which may be preventing queen 

conch abundance and occurrence from increasing.

It is possible too that the territorial and federal fishery regulations have reduced fishing mortality on 

queen conch, but that the demographic effects of this reduction are yet to be seen. The sites surveyed 

during this study were all within territorial waters, where daily quotas and gear restrictions have 

been in place for only six years; this may not have been sufficient time for queen conch abundance 

and occurrence to increase. Also, we have no comparable fishery-independent survey data on the 

E. gigas
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47 2 - Benthic Composition

abundance and occurrence of queen conch from the EEZ, where fishery regulations have been in effect 

for a longer time. According to local fishers, queen conch populations have rebounded within the EEZ. 

At recent stock assessment workshops (http://www.sefsc.noaa.gov/sedar/), west coast Puerto Rican 

fishers insist that at La Parguera, most conch occur off-shore toward the shelf edge in deeper waters 

at 30 m rather than at shallower depths where visual surveys for conch were conducted (Diaz, 2007). 

Benthic substrates that occur at depths greater than 30 m however, were beyond the sampling domain 

of this study. 

2.4.6.2. Long-spined sea urchin (D. antillarum) 

The long-spined sea urchin is another important component of faunal assemblages found in coral reef 

ecosystem. It is considered a major herbivore that controls macroalgae abundance on Caribbean coral 

reefs (Lessios et al., 1984; Lessios, 1998). Prior to the massive Caribbean-wide die-off of Diadema in 

the late 1980s, urchins were present throughout the wider Caribbean in most habitats including coral 

reefs, seagrass beds, rocky shores, softbottom and mangrove; they were abundant in shallow areas 

down to 15 m, with some found as deep as 40 m (Randall et al., 1964; Sammarco, 1972; Weil et al., 

1984). The Caribbean-wide mass mortality of D. antillarum was estimated to be greater than 93% 

(Lessios et al., 1984; Lessios, 1988), and this had catastrophic effects on reef health (Knowlton, 2001). 

With the disappearance of these keystone herbivores from the reefs, many Caribbean reefs became 

dominated by macroalgae, and the recruitment of scleractinian corals was inhibited (Edmunds and 

Carpenter, 2001). Diseased urchins were first reported in Puerto Rico in January 1984 near Laurel reef 

in La Parguera (Vicente and Goenaga, 1984).

A total of 192 long spined sea urchins (D. 

antillarum) were recorded on only 12 of 314 

(3.8%) sites surveyed, and had an overall density 

0.53 per 100 m2

. These observations suggest 

that the abundance of long-spined sea urchins 

on reefs at La Parguera is low, but the estimates 

being reported here may not precisely reflect in 

situ abundance. Sea-urchins can be very patchy 

in their distributions, and it may be that survey 

effort (i.e., the number of sites surveyed) was 

inadequate to characterize population abundance 

at La Parguera. For example, one of the twelve 

sites surveyed contained 75 of the 192 individuals 

recorded. In addition, long-spined sea urchins 

were enumerated as part of generalized surveys 

that were intended to characterize benthic 

composition but were not optimized to search and identify urchins only. For example, in extremely 

rugose and complex reef habitats, long-spined urchins could be located in deep crevices, holes or 

behind overhangs, where they would be undetected unless exhaustive searches were made to locate 

them. 

Nevertheless, the occurrence of D. antillarum on La Parguera reefs is an indication of potential population 

recovery, given the recent mass mortality of the species. La Parguera has higher densities of urchins 

than other localities in the Caribbean, with large numbers of reproductive adults, abundant juveniles 

and no signs of disease or unhealthy individuals (Weil et al., 2005). Weil et al. (2005) also reported that 

the distribution of urchins across the La Parguera region is patchy, but suggested that recovery of local 

populations may be occurring. The sampling design used in this study and the data generated would be 

useful in detecting long-term increases in the sea urchin density, if such changes were to occur. More 

targeted surveys that monitor urchin population recovery may be required. 

D. antillarum
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48 2 - Benthic Composition 2.4.6.3. Caribbean spiny lobster (P. argus) 

The stratified random sampling design utilized by this study also provided limited opportunities to 

conduct spiny lobster surveys and determine their broad-scale distribution in the La Parguera region. 

Lobsters were rarely encountered during surveys and were observed only at approximatrely 3.0% 

of 3 sites visited between 2005 and 2006. However, sightings of spiny lobsters during surveys were 

opportunistic rather than deliberate, and exhaustive searches of overhangs and crevices were not 

conducted to determine lobster presence because of time dive time limitation. Thus, it is quite likely 

that the sightings reported from this study underestimated the frequency of lobster encounters in La 

Parguera. 

Like queen conch however, spiny lobster is a treasured local delicacy in La Parguera, and local 

populations are targeted by both commercial and artisanal fishers. Stock assessments conducted by 

over the past two decades have indicated that the spiny lobster fishery in Puerto Rico has shown signs 

of overfishing; and landings, catch rates, and relative abundance have declined significantly since 

the fishery began in 1969 (Morris et al., 2004). Analysis of data on spiny lobster landings from Puerto 

Rico indicates that the fishery is concentrated around the southwestern shelf, which includes the La 

Parguera region (Valle-Esquivel, 2005). 

Fishing pressure on spiny lobster is controlled under federal regulations 

implemented by the CFMC since 1985 and by territorial regulations 

implemented by DNER since 1936 (CFMC, 1985). Regulations include 

prohibiting the harvest of females with eggs and individuals measuring less 

than 9 cm (3.5 inches) in carapace length; barring the use of chemicals, 

explosives, poisons, drugs, spears, and hooks or similar devices to 

harvest lobsters; requiring the use of traps with self-destruct panels; and 

limiting entry into the fishery to only fishers with a permit (CFMC, 1985). It 

is doubtful whether these regulations have been effective however, given 

that between 1985 and 1989, undersized lobsters accounted for 40% of 

the total lobster catch from Puerto Rico (Bohnsack et al., 1991; Morris et 

al., 2004). In summary, better enforcement of existing regulations may be 

needed to improve lobster abundance over time, and a dedicated lobstermonitoring program will be needed to document long-term changes in 

lobster populations in La Parguera.

P. argus
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2.5. References

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Puerto Rico and the U.S. Virgin Islands. pp. 95-105. In: M.H. Goodwin, S.M. Kau, and G.T. Waugh (eds.). 

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Islands.

Appeldoorn, R.S. 1994. Queen conch management and research: status, needs, and priorities. pp. 301-320. 

In: R.S. Appeldoorn and B. Rodriguez (eds.). Queen Conch biology, fisheries, and mariculture. Fundación, 

Científíca. Los Roques, Caracas Venezuela. 356 pp.

Appeldoorn, R.S. and B. Rodriguez, (eds.). 1994. Queen Conch biology, fisheries, and mariculture. Fundación, 

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Bechtel, J.D., P. Gayle, and L. Kaufman. 2006. The return of Diadema antillarum to Discovery Bay: patterns of 

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Okinawa, Japan. 1950 pp. 

Blumenthal, J.M., T.J. Austin, C.D.L. Bell, J.B. Bothwell, A.C. Broderick, G. Ebanks-Petrie, J.A. Gibb, K.E. Luke, 

J.R. Olynik, M.F. Orr, J.L. Solomon, and B.J. Godley. 2009. Ecology of Hawksbill Turtles, Eretmochelys imbricata, 

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Chapter 3. Fish Communities, Groups and Species 

3.1. Introduction

Previous studies of fish distributions and diversity across the 

shelf at La Parguera have either been limited to selected 

coral reef locations or specific habitat types and have been 

brief snapshots of communities from discrete periods within 

a single year or two. Such data do not provide sufficient 

spatial information to develop comprehensive management 

plans across the complex multiple-habitat seascapes that 

characterize the La Parguera Natural Reserve. In addition, to 

determine changes over time, sampling must be conducted 

on a regular basis across the region. In response, the current 

Coral Reef Ecosystem Monitoring (CREM) project was 

designed to provide information on fish community structure 

and change across all components of the seascape and to 

examine inter-annual and seasonal changes and multi-year 

trends for fish species and assemblage biomass, abundance and diversity.

The CREM data were first utilized in 2003 to examine fish communities and their distribution across the 

shelf at La Parguera (Christensen et al., 2003). The study examined data on fish species distributions 

and size classes from multiple habitat types across multiple zones (lagoonal, etc.) and found that the 

type of habitat (i.e., mangrove, seagrass, hardbottom) was a better predictor of the spatial distribution 

of fish, particularly snappers (Lutjanidae) and grunts (Haemulidae), than the location across the shelf. 

For several fish species, distinct body size dependent distribution patterns were considered to be 

indicative of ontogenetic shifts in habitat use, with smallest individuals more abundant in seagrasses 

and mangroves and larger adults more abundant at coral reef sites. In similar studies, Aguilar-Perera 

and Appeldoorn (2007, 2008) also highlighted the importance of connectivity between mangroves, 

seagrasses and coral reefs, particularly for the many species of fish that undertake multiple habitat 

shifts in their life-cycle. In addition, CREM data were utilized to examine the ecological relevance of a 

mosaic of habitat types to fish (Pittman et al., 2007a) demonstrating that fish diversity and abundance in 

mangroves is influenced by the marine habitat types adjacent to mangroves. For instance, mangroves 

and seagrasses that coexisted in close proximity (≤100 m) supported higher fish species richness and 

abundance than mangroves with adjacent unvegetated sediments. For some species, such as yellowtail 

snapper (Ocyurus chrysurus), the close proximity of both seagrasses and coral reefs explained the 

observed distribution patterns.

Other studies have found that the topographic complexity of the surrounding coral reef ecosystem at 

La Parguera is also important in explaining patterns of fish species diversity and abundance across 

the shelf. Work by Kimmel (1985) conducted in 1980 and 1981 provides a rare example of a spatially 

comprehensive fish-habitat study of fish communities from 21 discrete biotopes across the insular shelf at 

La Parguera. These data provide an important historical reference point with which to compare changes 

in fish species occurrence. Kimmel’s work also highlighted the importance of integrating topographic 

complexity into the definition of habitat types and the potential for predicting fish assemblages from 

mapped habitat types. Recent studies in Puerto Rico using NOAA CCMA Biogeography Branch (CCMABB) fish survey data demonstrated the importance of topographic complexity for predicting fish species 

richness and other key metrics (Pittman et al., 2007b; Pittman et al., 2009) in the La Parguera region 

and neighboring USVI. Spatial predictive modeling using regression trees and geographic information 

system (GIS) was used to predict and map fish species richness across the entire seascape at La 

Parguera, with greater than 70% map accuracy (Appendix E).

Balloonfish (Diodon holocanthus) in turtle grass. 
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56 3 - Fish CommunitiesWhile spatial patterns in species distributions are becoming better known, comparatively little is 

known about the temporal change that may be occurring in both the size and abundance of fish 

and macroinvertebrates. This report provides both a spatial and temporal characterization of fish 

community metrics, fish species (with emphasis on fishery species), macroinvertebrates and benthic 

habitat composition using underwater survey data collected during a seven year period (2001-2007). 

This section of the report focuses on the spatial distribution of species and community metrics (i.e., 

composition, species richness, biomass, abundance of assemblages) and the temporal patterns (2001-

2007) across mosaics of habitat types in the study area at La Parguera, southwest Puerto Rico. The 

intention is to provide a spatial and temporal characterization for the area and does not therefore 

establish relationships between environmental structure and fish distributions, which will be the focus of 

subsequent publications. Fish communities are highly heterogeneous in time and space and can also 

function as indicators of ecosystem integrity and health. Examination of fish community composition 

and fish species distributions provides important baseline information for ecological studies, as well 

as, critical information to support resource management decision making with regard to understanding 

essential fish habitat, identifying where species of concern are located, identifying diversity and 

productivity hotspots, prioritizing activities in marine protection, mapping environmental sensitivity, 

designing restoration strategies and monitoring programs.

3.2. Methods

3.2.1. Survey Data

Fish surveys were conducted along a 25 m long by 

4 m wide belt transect (100 m2

) using a fixed survey 

duration of 15 minutes (Menza et al., 2006; Figure 3.1). 

The number of individuals per species is recorded in 5 

cm size class increments up to 35 cm using the visual 

estimation of fork length. Individuals greater than 35 cm 

are recorded as an estimate of the actual fork length to 

the nearest centimeter. A benthic habitat map was used 

to develop and implement a stratified-random sampling 

design based on two strata: hard and softbottom habitat 

types to minimize variance in population estimates and 

maximize the power to detect changes (Figure 3.2). A 

total of 1,167 fish surveys (572 from hardbottom and 

595 from softbottom habitats, including mangroves) 

were used in this analysis. For a detailed description of 

CCMA-BB’s fish census survey methods see Appendix

D. 

Figure 3.1. NOAA trained observer recording fish census 

data along the belt transect.
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Figure 3.2. NOAA’s benthic habitat map showing the major classes of habitat type across the study region of Reserva Natural La Parguera. 

Survey sites were allocated at spatially random locations distributed within the strata of hardbottom and softbottom benthic classes between 

2001 and 2007.

3.2.2. Data analysis

3.2.2.1. Benthic composition and fish community composition

Differences and similarities in the species composition of communities between samples (often referred 

to as assemblage or community structure) were examined using a species-abundance by site data 

matrix. Infrequently observed fish that were not identified to species level were removed. The matrix 

was fourth-root transformed to ensure that rare and intermediate abundance species, in addition to the 

highly abundant species, played a significant role in determining patterns in community composition. 

The data was then used to construct a matrix of the percentage similarity in community composition 

between all pairs of sites using the Bray-Curtis Coefficient, 

S’jk = 1 - 

Sn Xij i=1 - Xik

Sn Xij + Xik i=1

where xij is the abundance of the ith species in the jth sample and where there are n species overall.

This algorithm is considered a robust estimator of ecological distance and has had widespread usage 

in ecology particularly for comparison of biological data on community structure (Faith et al., 1987). 

Its robustness is in part due to its exclusion of double zeros, that is, if two samples are missing the 

same species, they will not be regarded as similar based on the same absentees (Legendre and 

Legendre, 1998). This similarity coefficient reduces the comparison between all pairs of samples to 

single numerical values that are arranged in a secondary matrix from which pattern is examined. 
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58 3 - Fish CommunitiesAn ANOSIM test, a multivariate version of Analysis of Variance (ANOVA; Primer v6; Clarke and Warwick, 

1994), with 999 permutations was used to test for significant differences in fish assemblage composition 

between mapped classes at multiple thematic resolutions including: 1) HABITAT Structure - soft, hard 

and mangrove; 2) all habitat TYPES (see Table 3.1), and 3) modifiers for seagrass (e.g., classes of 

percentage cover). The R value is a better relative indicator of the amount of dissimilarity between 

groups than the significance test and is thus given greater emphasis here. The R is usually interpreted 

as the pairs of fish assemblage composition being: R<0.25 = barely separable; R>0.5 = overlapping, 

but clearly different and R>0.75 = well separated. 

For a visual examination of patterns of between site similarity a two-dimensional non-metric dimensional 

scaling plot (nMDS) was constructed. This information determines whether benthic map classes 

and thematic levels are delineated in an ecologically meaningful way for fish. For instance: Do fish 

communities respond to the structural differences perceived by the map maker (i.e., geomorphological 

and biological features)? The multivariate analyses also provide an assessment of the ability of the 

benthic habitat map to predict patterns of fish assemblage composition. The similarities/dissimilarities 

should not be interpreted as a measure of connectivity between habitat types as has been suggested 

by Chittaro et al. (2005), although similarity between neighboring habitat types may result from interhabitat movements and resource utilization that would need to be validated by direct observations of 

space use patterns.

3.2.2.2. Species habitat association

Species-habitat associations were determined by overlaying fish survey points (start of the 25 m 

transect) on the NOAA benthic habitat map and linking to the class of habitat type at the point location.

3.2.2.3. Inter-annual Patterns

Inter-annual patterns were examined by comparing means using ANOVA and SigmaPlot© (SAS 

Institute, 2006) in a wide range of community metrics and individual species data amongst years. Data 

were tabulated and where means decreased significantly from one year to the next then a red arrow 

was assigned and if increased significantly then a green arrow was assigned. Consecutive years of 

significant decline or increase where denoted with double arrows. 

3.2.2.4. Seasonal Patterns

Seasonal patterns across years (2004-2007) were examined by grouping fish data into winter (DecemberMarch) and summer field seasons (August) and then plotting means (± SE) using bar charts.
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3.3. Results

3.3.1. Spatial distribution patterns and species-habitat associations

3.3.1.1. Fish assemblages

Overall a total of 210 species were 

identified to the species level, with at least 

another 14 fishes identified to genera. For 

the complete list of species size details and 

summary information refer to Appendix 

B. At the coarsest thematic resolution 

within the hierarchical classification 

scheme, using mapped classes of 

hardbottom, softbottom and mangroves, 

fish assemblages were significantly 

different (ANOSIM R=0.54, p<0.01). 

Greatest difference in fish assemblage 

composition was found for hardbottom 

versus mangroves (ANOSIM R=0.83, 

p<0.01), which were very distinct from 

one another. In contrast, softbottom and 

hardbottom fish assemblages exhibited 

overlap (ANOSIM R=0.32, p<0.01; Figure 

3.3). 

At the more detailed thematic level of habitat TYPE (all hard and soft types included), fish assemblages 

were less distinct from one another (ANOSIM R=0.39, p<0.01).

At the level of habitat TYPE, fish assemblages within mangroves 

were significantly well separated from all hardbottom and softbottom 

habitat types, with most overlap existing between fish assemblages 

of seagrasses and mangroves (Table 3.1). Fish assemblages 

associated with patch reefs were least distinct and showed high 

overlaps with all soft- and hardbottom habitat classes, but were 

more similar to other colonized hardbottom habitat types than to 

macroalgae and sand. Individual patch reefs and aggregated patch 

reefs were barely separable although a statistically significant 

difference was calculated. Fish assemblage composition of 

macroalgae and sand and macroalgae and seagrass habitat types 

were not significantly different and exhibited very high overlap. Fish assemblages associated with 

linear reefs, colonized pavement with sand channels, colonized pavement and the two classes of patch 

Figure 3.3. Multi-dimensional scaling plot (nMDS) showing between site 

similarity in fish assemblages composition. Points are individual survey sites 

that have been assigned a benthic habitat class based on geographical location 

within a mapped class of the NOAA benthic habitat map. Stress <0.2 indicates 

an adequate two-dimensional representation of the data.

Jenkinsia sp. and schoolmaster (Lutjanus apodus)

Table 3.1. ANOSIM R values measuring fish assemblage similarities between samples grouped by benthic habitat TYPE. Uncommon 

habitat types were not shown. R values in bold are statistically significant (p=<0.01). Dark green= well separated, medium green= some 

overlap, light green= barely separable.

Mapped habitat type Patch Reef 

(Ind)

Patch Reef 

(Agg)

Linear

Reef

Colonized 

Pavement

Col. Pav. w/ 

Sand Chan. Seagrass Macroalgae Unconsol.Sediment

Patch Reef (Individual)

Patch Reef (Aggregate) 0.18

Linear Reef 0.05 -0.06

Colonized Pavement -0.03 -0.08 0.07

Col. Pavement w/ Sand Channels 0.13 0.12 0.12 0.11

Seagrass 0.16 0.10 0.29 0.25 0.43

Macroalgae 0.26 0.34 0.74 0.49 0.78 0.09

Unconsolidated Sediments 0.24 0.21 0.56 0.40 0.71 0.28 0.02

Mangrove 0.89 0.86 0.89 0.85 0.90 0.47 0.94 0.82
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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60 3 - Fish Communitiesreef were barely separable (Table 3.1). This result was confirmed when only hardbottom types were 

analyzed for pairwise differences (R=0.16), with the fish assemblages associated with the biologically 

and topographically complex habitat types showing no significant pairwise differences. Significant 

differences in fish assemblages only occurred between the most complex and the least complex habitat 

types (i.e. linear reef versus scattered coral/rock with sand). Highest dissimilarity in fish assemblages of 

hardbottom types occurred between individual patch reefs and scattered coral/rock in sand (80.1% mean 

dissimilarity). The higher abundance (>50% higher) of redband parrotfish (Sparisoma aurofrenatum) and 

striped parrotfish (Scarus iseri), and foureye butterflyfish (Chaetodon capistratus) on patch reefs versus 

scattered coral/rock explained the highest proportion of the dissimilarity. Most species were in higher 

abundance on patch reefs, except for very small-bodied benthic fish such as orangespotted goby (Nes 

longus), goldspot goby (Gnatholepis thompsoni) and bridled goby (Coryphopterus glaucofraenum), 

which were more abundant on scattered coral/rock and sandy substrata.

When pairwise differences were examined at 

the MODIFIER level for classes of seagrass 

cover and sand statistically significant 

difference was detected (Global R=0.54, 

p<0.01). Fish assemblages associated with 

continuous seagrass cover (>90%) were 

most different than other levels of cover 

including the sand habitat type (Table 3.2). 

Fish assemblages of continuous seagrass, 

however, did show some overlap with fish 

assemblages associated with sparser 

seagrass cover, but was most dissimilar to 

patchy seagrass (50-70%) and sand.

Fish assemblages of the sparser 30-50% seagrass cover were not significantly different than assemblages 

associated with 10-30% cover and sand habitat types (Table 3.2). High variability in the response was 

evident, with assemblages of 50-70% seagrass being significantly different to 30-50% seagrass and 10-

30 % seagrass, although some overlap in assemblage composition was evident. These data indicate 

that thresholds in seagrass 

cover may occur at the >90% 

level of cover, since fish 

assemblage composition was 

most different between this 

level of cover and almost all 

other classes exhibiting lower 

seagrass cover. Therefore, 

the presence of seagrasses 

with >90% cover makes a 

significant difference to the 

fish community highlighting 

the fact that not all seagrass 

beds provide equal function 

and spatial heterogeneity 

in benthic structure is very 

important for many fish.

Table 3.2. ANOSIM R values measuring fish assemblage similarities 

between samples grouped by benthic habitat MODIFIER (i.e., % seagrass 

cover). Sand was included to examine differences in fish assemblages 

between classes of patchy seagrass and sand. R values in bold are 

statistically significant (p<0.01). Medium green=some overlap, light 

green=barely separable.

Mapped habitat 

type modifier

Continuous

>90%

Patchy

70-90%

Patchy

50-70%

Patchy

30-50%

Patchy

10-30%

Continuous >90%

Patchy 70-90% 0.28

Patchy 50-70% 0.52 0.17

Patchy 30-50% 0.18 0.27 0.58

Patchy 10-30% 0.36 0.34 0.47 0.01

Sand (Uncon. sed.) 0.46 0.05 0.04 0.11 0.13

Baitfish in mangroves
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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61 3 - Fish Communities

3.3.1.2. Fish community metrics

Highest fish abundance was observed in two zones: (i) the mangroves and some coral reefs in the 

nearshore lagoonal zone, as well as offshore mangrove islands within 3 km of the coastline; and (ii) 

colonized hardbottom in deeper waters (>10 m) on the shelf approximately 7-10 km from the coastline 

(Figure 3.4a). Lowest fish abundance was associated with unvegetated sediments (i.e., sand and 

muddy sand) across the study area (Figure 3.5a). The majority of fish in mangroves were juveniles 

of multi-habitat species including fish species that also utilize coral reefs (Table 3.3). Large schools 

(>1,000) of small-bodied planktivorous fish (Atherinidae, Clupeidae, Engraulidae) were observed at 

many (n=9) mangrove sites. Greatest concentrations of high fish biomass were found over colonized 

hardbottom at the most offshore portion of the study area and also at several nearshore sites including 

colonized hardbottom immediately east of Margarita Reef. Other high biomass sites were near the 

interface between colonized hardbottom and softbottom habitat types (Figure 3.4b). 

Overall, mangroves supported intermediate levels of fish biomass and unvegetated sediments supported 

lowest mean fish biomass (Figure 3.4b). Similarly, highest fish species density (number of species 

Total fish abundance (100 m2)

<10

11 - 50

51 - 200

>200

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Total fish biomass (g/100 m2)

<1000

1001 - 5000

5001 - 10000

>10000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish species richness (100m2)

0 >30

1 - 5

6 - 10

11 - 20

21 - 30

c)

Figure 3.4. Maps of the interpolated (left map) and spatial (right map) distributions for: (a) total fish abundance, (b) total fish biomass and 

(c) total species richness.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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62 3 - Fish Communities

0

750

1500

Mean number of individuals (#/100 m2)

Habitat type

0

3

6

Mean biomass (kg/100 m2)

Habitat type

0

11

22

Mean number of fish species (#/100 m2)

Habitat type

a)

c)

b)

Figure 3.5. Comparison of mean (+ SE) values by habitat type in the southwest Puerto Rico study area for: (a) total fish density, (b) total 

fish biomass and (c) total species richness.

per 100 m2) was observed over 

colonized hardbottom along 

the shelf edge and along the 

mangrove fringe closest to 

colonized hardbottom areas 

(Figure 3.4c). The majority of 

hardbottom sites supported 

fish species density of 11 or 

more fish per 100 m2

, with 

several sites in deeper water 

areas along the shelf edge and 

around El Palo and Margarita 

Reef supporting more than 30 

fish species per 100 m2. Lowest 

mean fish density was recorded 

over unvegetated sediments 

across the study area (Figure 

3.5a).

Herbivorous fish species 

richness was highly 

heterogeneous across 

hardbottom habitat types, with 

Species/Family

%

occur

Habitats

used

Mean (±SD) Fork Length, cm

Mangrove Seagrass Unvegetated Coral reef

Jenkinsia spp. 51.9 MSUC 3.1 (0.6) 3.0 (0.0) 3.0 (0.0) 3.0 (0.0)

Atherinomorus spp. 14.2 M 3.0 (0.0) - - - - - -

Clupeidae 14.2 MS 3.2 (1.0) 3.0 (0.0) - - - -

Haemulon flavolineatum 76.4 MSC 6.4 (3.3) 4.5 (2.6) - - 13.4 (3.9)

Haemulon sciurus 77.2 MSUC 9.9 (5.0) 4.1 (2.7) 12.5 (0.0) 16.5 (7.5)

Engraulidae 2.4 M 3.0 (0.0) - - - - - -

Lutjanus apodus 100 MSC 12.3 (6.4) 10.0 (3.5) - - 17.4 (5.8)

Stegastes leucostictus 85.8 MSUC 5.9 (2.4) 4.5 (2.1) 4.8 (2.3) 5.8 (2.8)

Scarus iseri 35.4 MSUC 5.6 (3.2) 4.4 (2.1) 7.4 (6.0) 10.9 (6.0)

Abudefduf saxatilis 59.1 MC 5.8 (2.8) - - - - 8.8 (4.3)

Eucinostomus melanopterus 52.8 MUC 5.3 (2.8) - - 9.2 (2.9) 7.8 (6.7)

Lutjanus griseus 45.7 MSC 16.7 (8.8) 12.5 (7.1) - - 21.9 (8.7)

Gerres cinereus 52.7 MSUC 9.7 (5.2) 10.0 (3.5) 17.5 (0.0) 19.0 (5.2)

Sparisoma radians 35.4 MSUC 6.4 (3.9) 5.1 (2.8) 4.6 (2.8) 4.1 (2.1)

Haemulon spp. 20.5 MSUC 4.4 (4.1) 3.4 (1.2) 3.0 (0.0) 4.9 (4.7)

Chaetodon capistratus 58.3 MSUC 5.0 (2.4) 3.7 (1.6) 5.7 (2.5) 8.0 (2.9)

Coryphopterus glaucofraenum 14.7 MSUC 3.4 (1.3) 3.2 (0.9) 3.3 (1.1) 3.3 (1.8)

Sphyraena barracuda 65.4 MUC 18.6 (13.8) - - 46.5 (61.5) 75.9 (40.9)

Acanthurus chirurgus 27.6 MSUC 7.3 (3.6) 6.4 (5.0) 11.1 (4.0) 14.9 (5.7)

Haemulon aurolineatum 5.5 MSUC 3.0 (0.0) 3.0 (0.0) 6.8 (5.4) 13.3 (6.3)

Table 3.3. Percentage occurrence, range of major habitat types used and mean (± Standard 

Deviation) of fork length by habitat type for the 20 most abundant fish species/groups 

observed using mangroves in southwest Puerto Rico from 2001-2006. Habitat types used 

are indicated by M-mangroves, S-seagrasses, U-unvegetated sediments and C-coral reefs.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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63 3 - Fish Communities

highest mean species richness along the shelf edge 

and around the El Palo and Margarita Reef region 

(Figure 3.6c). Although mean herbivore richness 

was relatively high in mangroves too (Figure 3.6c), 

this was elevated at offshore mangroves rather than 

the inshore fringing mangroves. Mean herbivorous 

fish species richness was more than 50% lower at 

seagrass/macroalgal sites than colonized hardbottom 

sites and lowest at unvegetated sediment sites 

(Figure 3.7c).

Mean herbivore fish biomass was markedly higher over colonized hardbottom than any other major 

habitat type (Figure 3.7b) and highest abundance was observed along the outer shelf sites and around 

the topographically complex collection of patch reefs at El Palo (Figure 3.6b). High herbivore fish 

Herbivorous surgeonfish (Acanthurus species).

a)

Herbivore fish abundance (100m2)

0

0.1 - 10

10.1 - 50

50.1 - 100

>100

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Herbivore fish biomass (g/100m2)

0

0.1 - 1000

1000.1 - 5000

5000.1 - 10000

>10000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

c)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Herbivore species richness (100m2)

0

1 - 5

6 - 10

11 - 16

Figure 3.6. Maps of the interpolated (left map) and spatial (right map) distributions for herbivore: (a) abundance, (b) biomass and 

(c) species richness.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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64 3 - Fish Communities

abundance was also observed across colonized hardbottom and seagrass habitat types in inshore 

lagoonal environments (Figure 3.6a and Figure 3.7a). 

Mean herbivorous fish species richness (Figures 3.7c) was considerably higher than piscivorous fish 

species richness and no distinct areas of high piscivorous fish diversity were observed in the La Parguera 

study area. Mean piscivorous fish species richness was highest in mangroves and lowest in macroalgae/

seagrass habitat (Figure 3.8c). Only 0.01% of all samples had more than five piscivorous fish species. 

Planktivorous fish species richness (Figure 3.10c) was also considerable lower than herbivorous fish 

richness, although areas of high planktivorous fish richness were more spatially distinct, with between 

six and 10 species observed in fish assemblages over colonized hardbottom sites along the shelf edge 

(Figure 3.11c). Overall, however, a higher mean diversity of planktivorous fish was found utilizing the 

mangrove edge (Figure 3.11c). 

0

25

50

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean herbivore density (#/100 m2)

0

1

2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean herbivore biomass (kg/100 m2)

0

5

10

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean no. of herbivore species (#/100 m2)

Habitat type

a)

b)

c)

Figure 3.7. Comparison of mean (+ SE) values by habitat type in 

the southwest Puerto Rico study area for herbivore: (a) density, (b) 

biomass and (c) species richness.

Habitat type

Habitat type

0

12

24

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean piscivore density (#/100 m2)

0

1

2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean piscivore biomass (kg/100 m2)

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean no. of piscivore species (#/100 m2)

Habitat type

a)

b)

c)

Figure 3.8. Comparison of mean (+ SE) values by habitat type in 

the southwest Puerto Rico study area for piscivore: (a) density, (b) 

biomass and (c) species richness.

Habitat type

Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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65 3 - Fish Communities

In contrast, piscivorous fish were most abundant in 

the mangroves (Figure 3.9a), but with some isolated 

sites in both midshelf and outershelf zones exhibiting 

high abundance (Figure 3.8a). The piscivorous fish 

observed within mangroves were primarily juveniles 

of fish that are piscivorous as subadults and adults. 

Several sites over colonized hardbottom that exhibited 

high biomass were sightings of cartilaginous fish such 

as sharks and rays (Figure 3.8b and Figure 3.9b). 

Highest mean biomass of piscivorous fish was recorded 

for mangroves (Figure 3.9b). 

a)

Piscivore fish abundance (100m2)

0

1 - 10

11 - 50

51 - 100

>100

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Piscivore fish biomass (g/100m2)

0

0.1 - 1000

1000.1 - 5000

5000.1 - 10000

>10000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

c)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Piscivore species richness (100m2)

0

1 - 5

6 - 7

Figure 3.9. Maps of the interpolated (left map) and spatial (right map) distributions for piscivore: (a) abundance, (b) biomass and (c) species 

richness.

Piscivorous fish, cero mackrel (Scomberomorus regalis). 
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

66 3 - Fish CommunitiesIn the mangroves, planktivorous fish abundance was dominated by frequently observed schools of 

small-bodied silvery fish from families including Clupeidae, Atherinidae and Engraulidae (Figure 3.10a). 

These fish often swim in mixed species schools and could not be identified to species level using visual 

surveys. Mean biomass, however, was highest for colonized hardbottom habitat types particularly 

near the outershelf zone, where planktivorous fish included large numbers of creole wrasse (Clepticus 

parrae; Figure 3.11b).

0

600

1200

Colonized Macroalgae/ Unvegetated Mangrove

Mean planktivore density (#/100 m2)

a)

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mean no. of planktivore species (#/100 m

Mangrove

2)

c)

0

0.2

0.4

Colonized Macroalgae/ Unvegetated Mangrove

Mean planktivore biomass (kg/100 m2)

b)

Figure 3.10. Comparison of mean (+ SE) values by habitat type in 

the southwest Puerto Rico study area for planktivore: (a) density, (b) 

biomass and (c) species richness.

Hardbottom Seagrass Sediments

Habitat type

Habitat type

Hardbottom Seagrass Sediments

Habitat type

Planktivorous species, creole wrasse (Clepticus parrae).

Jenkinsia species in mangroves.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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67 3 - Fish Communities

a)

Planktivore fish abundance (100m2)

0

1 - 10

11 - 50

51 - 100

>100

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Planktivore fish biomass (g/100m2)

0

0.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

c)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Planktivore species richness (100m2)

0

1 - 5

6 - 10

Figure 3.11. Maps of the interpolated (left map) and spatial (right map) distributions for planktivore: (a) abundance, (b) biomass and (c) 

species richness.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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68 3 - Fish Communities3.3.2. Taxonomic groups

Select Large-body Groupers (Serranidae)

Abundance and biomass of the large-bodied serranids or grouper (species from the Subfamily 

Epinephelinae; Epinephelus spp., Cephalopholis spp. and Mycteroperca spp.) was highest (7 

individuals/100 m2 and 1.4 kg/100 m2

, respectively) along the deeper water (>10 m) topographically 

complex shelf edge habitat types (linear reef and colonized pavement with sand channels; Figures 

3.12a-b and 3.13a-b). Grouper were absent from mangroves and only found in very low abundance in 

macroalgal and seagrasses of nearshore and lagoonal zones (Figures 3.12a and 3.13a). For much of 

the colonized hardbottom across the study area, groupers were either absent or at very low densities 

(≤2/100 m2). Biomass was highest over colonized pavement on the mid and outershelf (Figure 3.11b).

The smallest juveniles of the three most 

common grouper species were absent for coney 

(Cephalopholis fulva) and red hind (Epinephelus 

guttatus) and were very rarely seen for graysby 

(Cephalopholis cruentata; Figure 3.14a). C. fulva

and E. guttatus were not observed until the 10-

15 cm size class (subadults). In contrast, a larger 

proportion of the total sightings of C. cruentata were 

large juveniles and subadults (Figure 3.12a). The 

most frequently seen size classes of C. fulva were 

the 15-25 cm (measured in fork length [FL]) and 

25-30 for E. guttatus (Figures 3.13 and 3.14b). No 

Cephalopholis species longer than 30 cm FL were 

recorded in the study area, although the maximum 

known size for C. fulva is 41 cm TL and for C. cruentata

is 42.6 cm TL (Table 3.4). The maximum size for E. 

guttatus in this study was 40 cm FL compared with 

a maximum for the species of 76 cm TL (Table 3.4). 

Species of the genus Mycteroperca (M. tigris; M. 

bonaci, M. venenosa, M. interstitialis) were entirely 

absent from the surveyed sites between 2001-2007, 

even though tiger grouper (Mycteroperca tigris) and 

yellowfin grouper (Mycteroperca venenosa) may once have spawned along the shelf edge (OjedaSerrano, 2007).

Habitat type Habitat type

Species

Approx. size 

class at first 

maturity*

Max. known 

size*, TL

Max. size

 observed in 

PR, FL

Balistes vetula 20-25 60 40

Cephalopholis cruentata 15-20 42.6 30

Cephalopholis fulva 15-20 41 30

Epinephelus guttatus 20-25 76 40

Haemulon aurolineatum 15-20 25 25

Haemulon flavolineatum 15-20 30 30

Haemulon plumierii 15-20 53 30

Haemulon sciurus 15-20 46 35

Lutjanus apodus 20-25 67.2 45

Lutjanus griseus 25-30 89 65

Lutjanus mahogoni 15-20 48 30

Lutjanus synagris 20-25 60 65

Ocyurus chrysurus 20-25 86.3 40

Mulloidichthys martinicus 15-20 39.4 35

Pseudupeneus maculatus 15-20 30 30

Scarus iseri 10-15 35 35

Scarus taeniopterus 10-15 35 35

Sparisoma aurofrenatum 10-15 28 35

Sparisoma rubripinne -- 47.8 40

Sparisoma viride 15-20 64 50

Sphyraena barracuda -- 200 150

Table 3.4. Summary data on selected species from key fish 

families showing maximum size observed in the study region 

and maximum known size for the species found from 1,167 

samples from 2001-2007. * Values from FishBase (http://www.

fishbase.org). TL=total length; FL= fork length. Refer to Appendix 

C for detailed information.

0

0.4

0.8

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean grouper density (#/100 m2)

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean grouper biomass (kg/100 m2)

a) b)

Figure 3.12. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for select grouper 

(Serranidae).
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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69 3 - Fish Communities

Figure 3.13. Maps of the interpolated (left map) and spatial (right map) distributions for select large-bodied grouper (Serranidae): (a) 

abundance and (b) biomass.

Grouper abundance (100 m2)

0

1 - 2

3 - 7

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Grouper biomass (g/100 m2)

0

0.1 - 1000

>1000

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

C. cruentata Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

C. fulva Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

E. guttatus Frequency (%)

Size class (cm)

Juveniles/subadults

a)

c)

b)

Figure 3.14. Size frequency histogram for select grouper (Serranidae) in the southwest Puerto Rico study area. (a) graysby (C. cruentata), 

(b) coney (C. fulva) and (c) red hind (E. guttatus).
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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70 3 - Fish CommunitiesGraysby (Cephalopholis cruentata)

C. cruentata abundance and biomass were highest over 

colonized hardbottom with higher occurrence on mid and 

outer shelf zones than the nearshore (Figures 3.15 and 

3.16). Densities were relatively low across the study area, 

but with several high rugosity areas of contiguous coral reefs 

on the outer shelf supporting densities of >2 C. cruentata

per 100 m2

 (Figure 3.15a). Abundance and biomass were 

comparatively low in seagrasses and unvegetated sandy 

areas (Figure 3.16).

Graysby (Cephalopholis cruentata) in the Flower Garden Banks.

C. cruentata abundance (100 m2)

0

1 - 2

>2

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

C. cruentata biomass (g/100 m2)

0

0.1 - 500

>500

Figure 3.15. Maps of the interpolated (left map) and spatial (right map) distributions for graysby (C. cruentata): (a) abundance and (b) 

biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. cruentata biomass (kg/100 m2)

0

0.3

0.6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. cruentata density (#/100 m2)

a) b)

Figure 3.16. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for graysby (C. 

cruentata).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

71 3 - Fish Communities

The majority of C. cruentata individuals observed were adults and subadults, with highest densities 

occurring in the topographically complex colonized pavement with sand channels, linear coral reefs 

and patch reefs habitat types (Figure 3.17). All life stages co-occurred in the same zones and habitat 

types across the shelf. No distinct inshore-offshore ontogenetic patterns were evident and juveniles 

exhibited no distinct preference for shallow inshore areas (Figure 3.17). C. cruentata were absent from 

mangroves and reef rubble and rarely seen over unvegetated sediments, macroalgae and seagrasses 

(Figure 3.17).

0

0.2

0.4

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean C. cruentata density (#/100 m2)

Mapped habitat type

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Juveniles/Subadults

Adults

0 0

20.7

10.8

16.0

15.0

9.1

0 0

2.4 2.4

16.4

1.1

4.3

3.8 2.1

7.4

10.6

0 0

20.7

13.5

Figure 3.17. Top: Mean density (+SE) for juvenile/subadult and adult by mapped habitat type for graysby (C. cruentata). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for graysby 

(C. cruentata) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

72 3 - Fish CommunitiesConey (Cephalopholis fulva)

C. fulva exhibited a very restricted distribution within 

the study area with abundance and biomass highest on 

colonized hardbottom along the shelf edge (Figures 3.18 

and 2.18). In contrast, in nearshore and mid-shelf zones 

only one C. fulva individual was observed (Figure 3.18). 

Seagrasses, mangroves and unvegetated soft sediments 

were not utilized during daylight hours (Figure 3.19). 

Coney (Cephalopholis fulva)

C. fulva abundance (100 m2)

0

1 - 2

>2

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

C. fulva biomass (g/100m2)

0

1 - 500

500.1 - 1000

>1000

b)

Figure 3.18. Maps of the interpolated (left map) and spatial (right map) distributions for coney (C. fulva): (a) abundance and (b) biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. fulva biomass (kg/100 m2)

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. fulva density (#/100 m2)

a) b)

Figure 3.19. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for coney (C. fulva). Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

73 3 - Fish Communities

Juvenile C. fulva have not been observed in this monitoring program and very few subadults were 

sighted (only subadults are shown in the juvenile/subadult size class; Figures 3.14b and 3.20). Subadults 

and adults almost exclusively occupy outer shelf linear reef, colonized pavements with sand channels, 

colonized pavement and scattered corals that exist at the southern outer edge of the study area (Figure 

3.20).

0

0.25

0.5

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean C. fulva density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0 0

6.3

1.4

6.1

18.0

3.0

0 0 0 0 0 0 0 0 0 0 0

4.3

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Figure 3.20. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for coney (C. fulva). Number above 

error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for coney (C. fulva) in the 

southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

74 3 - Fish CommunitiesRed hind (Epinephelus guttatus)

E. guttatus exhibited a very patchy spatial distribution, with 

sightings confined to colonized hardbottom habitat types 

predominantly on the mid and outer shelf zones (Figure 3.21a; 

3.22). Several individuals were also observed on coral reefs 

fronting the lagoonal zone. Highest biomass was estimated 

at sites along the topographically complex linear reefs and 

colonized pavement with sand channels habitat types at the 

shelf edge (Figure 3.21b). 

 Red hind (Epinephelus guttatus)

E. guttatus abundance (100 m2)

0

1 - 2

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

E. guttatus biomass (g/100m2)

0

1 - 500

500.1 - 1000

>1000

b)

Figure 3.21. Maps of the interpolated (left map) and spatial (right map) distributions for red hind (E. guttatus): (a) abundance and (b) 

biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean E. guttatus biomass (kg/100 m2)

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean E. guttatus density (#/100 m2)

a) b)

Figure 3.22. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for red hind (E. 

guttatus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

75 3 - Fish Communities

Juvenile E. guttatus have not been observed in this monitoring program (Figure 3.14c). Subadults and 

adults occurred predominantly in the mid and outer shelf zones, with highest densities of adults in the 

topographically complex patch reefs, colonized pavement with sand channels and linear reef habitat 

types (Figure 3.23). Subadults and adults showed no obvious geographical segregation. Five out of 11 

habitat types were utilized and E. guttatus was absent from all softbottom habitat types, reef rubble, 

colonized bedrock and mangroves (Figure 3.23).

0

0.1

0.2

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean E. guttatus density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0

2.7 2.7

0.5

8.5

3.0 4.0

0 0 0 0 0

12.7

0 0 0 0

1.1

2.1

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Subadult 10 - 20 cm

Adult >20

Figure 3.23. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for red hind (E. guttatus). Number 

above error bar indicates percent occurrence. Bottom: Spatial distribution of subadult and adult for red hind (E. guttatus) in 

the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

76 3 - Fish CommunitiesSnappers (Lutjanidae)

Lutjanids were widely distributed over the study region, but 

with abundance highest in nearshore areas, particularly 

fringing mangroves and mangrove cays (Figure 3.24a). 

Biomass was also highest in nearshore mangroves, although 

several hardbottom sites on the mid and outer shelf also 

exhibited high biomass represented by a few large-bodied 

individual snapper (Figure 3.24b). Lowest abundance and 

biomass were observed over soft bottomed habitat types 

(Figure 3.25).

L. apodus

Snapper abundance (100 m2)

0

1 - 10

11 - 50

51 - 100

>100

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Snapper biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Figure 3.24. Maps of the interpolated (left map) and spatial (right map) distributions for snapper (Lutjanidae): (a) abundance and (b) 

biomass.

0

10

20

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean snapper density (#/100 m2)

0

0.6

1.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean snapper biomass (kg/100 m2)

a) b)

Figure 3.25. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for snapper species 

(Lutjanidae).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

77 3 - Fish Communities

The smallest size classes of lutjanids were well represented, particularly for lane snapper (Lutjanus 

synagris) with approximately 35% of all individuals <5 cm (Figure 3.26). Relatively few of the smallest 

juveniles were observed for mahogany snapper (Lutjanus mahogoni), but this species exhibited the 

largest proportion of adult individuals compared with other lutjanids (Figure 3.26c). The largest adult 

schoolmaster (Lutjanus apodus) and L. synagris were no larger than 30 cm FL (Figures 3.26a and 

3.26d); the largest gray snapper (Lutjanus griseus) was 50 cm FL and the largest yellowtail snapper 

(Ocyurus chrysurus) was 40 cm FL (Figures 3.26b and 3.26e). The largest individuals observed in 

the study area were markedly smaller than the maximum known size for the species (Table 3.4). The 

largest L. apodus and L. synagris was less than 50% of the maximum size for the species. L. mahogoni

was 62% of the maximum size for the species (Table 3.4). Some snapper species were very rarely 

seen. Only two blackfin snapper (Lutjanus buccanella), one cubera snapper (Lutjanus cyanopterus) 

and 13 dog snapper (Lutjanus jocu) were observed between 2001 and 2007.

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

L. apodus Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

L. grisieus Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

L. mahogoni Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

L. synagris Frequency (%)

Size class (cm)

d) Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

O. chrysurus Frequency (%)

Size class (cm)

e) Juveniles/subadults

a)

c)

b)

Figure 3.26. Size frequency histogram for select snapper (Lutjanidae) in the southwest Puerto Rico study area. (a) schoolmaster (L. 

apodus), (b) gray snapper (L. griseus), (c) mahogany snapper (L. mahogoni), (d) lane snapper (L. synagris) and (e) yellowtail snapper (O. 

chrysurus).
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

78 3 - Fish CommunitiesSchoolmaster (Lutjanus apodus)

L. apodus were most frequently observed relatively close 

to shore in the lagoonal zone, with mean abundance and 

biomass markedly higher in mangroves than any other 

habitat type (Figures 3.27 and 3.28). High biomass was 

found at several sites on mangrove cays and at one site 

over colonized hardbottom in the outer shelf zone (Figure 

3.27).

L. apodus

a)

L. apodus abundance (100 m2)

0

1 - 10

10.1 - 25

25.1 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

L. apodus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.27. Maps of the interpolated (left map) and spatial (right map) distributions for schoolmaster (L. apodus): (a) abundance and (b) 

biomass.

0

0.4

0.8

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. apodus biomass (kg/100 m2)

0

7

14

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. apodus density (#/100 m2)

Habitat type

a) b)

Figure 3.28. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for schoolmaster 

(L. apodus). 

Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

79 3 - Fish Communities

L. apodus juveniles and adults were observed in eight of 11 habitat types, with highest densities for all 

life stages recorded for mangroves (Figure 3.29). The majority of juvenile L. apodus were sighted within 

mangroves, but with some juveniles also seen at sites from the mid and outer shelf zones. Although 

L. apodus was observed across all shelf zones, occurrence was lowest in the outer shelf zone (Figure 

3.29).

0

7

14

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean L. apodus density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 9.1 3.6 7.5 2.3 4.0 9.0 9.1

29.5

96.2

0 0 5.5 2.1 1.1 3.2 3.8 0 0 0 5.3 1.1

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 15 cm

Subadult 15 - 20 cm

Adult >20

Figure 3.29. Mean density (+SE) for juvenile/subadult and adult by mapped habitat type for schoolmaster (L. apodus). Number 

above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for schoolmaster (L. 

apodus) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

80 3 - Fish CommunitiesGray snapper (Lutjanus griseus)

L. griseus exhibited a relatively restricted distribution, with more 

than 95% of sightings occurring within mangroves, particularly 

along the fringe of the largest area of mangrove forest in 

the western portion of the study area (Figure 3.30). Highest 

biomass was also associated with mangroves (Figures 3.30b 

and 3.31b). L. griseus were very rarely observed on the mid 

and outer shelf. No individuals were observed over unvegetated 

sediments (Figure 3.31).

Gray snapper (Lutjanus griseus)

a)

L. griseus abundance (100 m2)

0

1 - 10

11 - 25

25.1 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

L. griseus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.30. Maps of the interpolated (left map) and spatial (right map) distributions for gray snapper (L. griseus): (a) abundance and (b) 

biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. griseus biomass (kg/100 m2)

0

3

6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. griseus density (#/100 m2)

a) b)

Figure 3.31. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for gray snapper 

(L. griseus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

81 3 - Fish Communities

L. griseus juveniles, subadults and adults coexisted both geographically and in the same habitat 

types across the study area (Figure 3.32). Most of the juveniles, subadults and adults were sighted in 

mangroves, but subadults and adults were also very occasionally seen over colonized hardbottom of 

the mid and outer shelf zones (Figure 3.32). L. griseus exhibited a more inshore distribution than did 

L. apodus (Figures 3.29 and 3.32). While juveniles/subadults were sighted in eight of 11 habitat types, 

adults exhibited a less general distribution being only observed in four of 11 habitat types (Figure 3.32).

0

3

6

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean L. griseus density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

50.0

0 1.8 0.1 0.5 0 1.0 2.0 1.8

9.6

48.7

0 0 0 0 0 0 2.1 1.7 0 0 1.1

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 15 cm

Subadult 15 - 20 cm

Adult >20

Figure 3.32. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for gray snapper (L. griseus). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for gray 

snapper (L. griseus) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

82 3 - Fish CommunitiesMahogany snapper (Lutjanus mahogoni)

L. mahogoni was infrequently sighted across the study area (Figure 

3.33a), with highest mean abundance recorded for mangroves 

and lowest for seagrasses (Figure 3.34a). Highest biomass was 

recorded for unvegetated sediments (Figure 3.34b) all of which 

were in close proximity to colonized hardbottom areas (Figure 

3.33b).

Mahogany snapper (Lutjanus mahogoni) in the Bahamas

L. mahogoni abundance (100 m2)

0

0.1 - 7

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

L. mahogoni biomass (g/100 m2)

0

0.1 - 500

Figure 3.33. Maps of the interpolated (left map) and spatial (right map) distributions for mahogany snapper (L. mahogoni): (a) abundance 

and (b) biomass.

0

0.01

0.02

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. mahogoni biomass (kg/100 m2)

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. mahogoni density (#/100 m2)

a) b)

Figure 3.34. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for mahogany 

snapper (L. mahogoni). 

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

83 3 - Fish Communities

L. mahogoni were relatively uncommon in the study area. Juveniles/subadults were associated with 

five of 11 habitat types and adults with six of 11 habitat types (Figure 3.35). Juveniles of L. mahogoni 

were found only in the nearshore and lagoonal zone, with highest mean densities in mangroves and 

linear reefs (Figure 3.35). In contrast, subadults and adults were sighted most frequently over patch 

reefs, linear reefs and unconsolidated sediments on the mid and outer shelf zones (Figure 3.35). 

0

0.2

0.4

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean L. mahogoni density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0 0

0.9 0.5 0.5

4.0

5.0

0

0.6

5.1

0 0

1.8

0

2.2

0

0.4

0 0 0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Figure 3.35. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for mahogany snapper (L. 

mahogoni). Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult 

for mahogany snapper (L. mahogoni) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

84 3 - Fish CommunitiesLane snapper (Lutjanus synagris)

L. synagris was observed across all major habitat types 

and occurred in lagoonal, mid shelf and outer shelf zones 

(Figure 3.36a). Mean abundance and biomass were highest 

over unvegetated sediments in close proximity to coral 

reefs (Figure 3.36b). Mean abundance in mangroves and 

seagrass beds was higher than for colonized hardbottom, 

but the reverse was true for mean biomass (Figure 3.37).

Lane snapper (Lutjanus synagris) in Honduras. Photo: Les Wilk

L. synagris abundance (100 m2)

0

1 - 10

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

L. synagris biomass (g/100 m2)

0

0.1 - 500

Figure 3.36. Maps of the interpolated (left map) and spatial (right map) distributions for lane snapper (L. synagris): (a) abundance and (b) 

biomass.

0

0.01

0.02

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. synagris biomass (kg/100 m2)

0

0.15

0.3

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean L. synagris density (#/100 m2)

a) b)

Figure 3.37. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for lane snapper 

(L. synagris). 

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

85 3 - Fish Communities

Juvenile L. synagris were sighted in nearshore and lagoonal environments, as well as the mid-shelf 

zone, but were not observed on the outer shelf (Figure 3.38). Juveniles/subadults utilized a wide range 

of hard and softbottom habitat types, with highest densities in colonized bedrock, patch reefs and 

mangroves. Although juveniles were most frequently seen over colonized hardbottom, the sightings 

were always in close proximity to seagrasses. Adults were only sighted over colonized pavement with 

sand channels, colonized pavement and unconsolidated sediments mostly located farther offshore, 

near the boundary between extensive contiguous hardbottom and adjacent softbottom habitat types 

(Figure 3.38).

0

0.6

1.2

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean L. synagris density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

50.0

0 0.9 1.8 0.9 0.9 0 0

5.5

0

5.8

0

7.3

0

4.3

0 0

3.0

6.4

0 0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 15 cm

Subadult 15 - 20 cm

Adult >20

Figure 3.38. Mean density (+SE) for juvenile/subadult and adult by mapped habitat type for lane snapper (L. synagris). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for lane 

snapper (L. synagris) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

86 3 - Fish CommunitiesYellowtail snapper (Ocyurus chrysurus)

O. chrysurus is the most widely distributed and most 

abundant of the Lutjanidae in the La Parguera region, with 

highest mean abundance over colonized hardbottom then 

seagrasses and mangroves (Figures 3.39a and 3.40a). 

Highest mean biomass was also calculated for colonized 

hardbottom, with several high biomass sites occurring on 

the mid and outer shelf zones (Figure 3.39b). Lowest mean 

biomass was calculated for mangroves and seagrasses, 

while lowest mean abundance was for sites classified as 

unvegetated sediment (Figure 3.40b).

Yellowtail snapper (Ocyurus chrysurus)

O. chrysurus abundance (100 m2)

0

0.1 - 10

10.1 - 20

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

b)

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

O. chrysurus biomass (g/100m2)

0

1 - 500

500.1 - 1000

>1000

Figure 3.39. Maps of the interpolated (left map) and spatial (right map) distributions for yellowtail snapper (O. chysurus): (a) abundance 

and (b) biomass.

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean O. chrysurus biomass (kg/100 m2)

0

0.6

1.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean O. chrysurus density (#/100 m2)

a) b)

Figure 3.40. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for yellowtail

snapper (O. chysurus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

87 3 - Fish Communities

O. chrysurus was the most ubiquitous of all Lutjanidae species, with juveniles sighted in 10 of the 11 

habitat types and adults in eight of 11 habitat types (Figure 3.41). All life history stages coexisted across 

hardbottom habitat types (except reef rubble) of the study area. No geographical segregation was 

evident as juveniles were frequently sighted on the mid and outer shelf zones, however, juveniles utilized 

nearshore and lagoonal environments more frequently than did subadults and adults (Figure 3.41). In 

contrast to other common Lutjanidae species, adult O. chrysurus were not observed in mangroves 

(Figure 3.41). Juveniles and adults were observed at high occurrence over colonized pavement with 

sand channels, linear reefs and colonized pavement and juveniles were markedly more common over 

seagrasses than adults. High densities at colonized bedrock sites were due to relatively high abundance 

at only two sites (Figure 3.41).

0

9

18

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean O. chrysurus density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0

100.0

29.721.6

39.0

21.1 17.0

30.7 23.6

0 10.9 9.8 7.3 16.4 0.9 5.4 3.2

14.9 16.2 0 0 6.4

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 15 cm

Subadult 15 - 20 cm

Adult >20

Figure 3.41. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for yellowtail snapper (O. chysurus). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for yellowtail

snapper (O. chysurus) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

88 3 - Fish CommunitiesParrotfish (Scaridae)

Scaridae abundance follows closely the distribution of colonized 

hardbottom, although the abundance is highly heterogeneous, with highest 

abundance across the large expanse of the most topographically 

complex colonized hardbottom habitat in the mid and outer shelf zones 

including El Palo and along some of the inner shelf linear reefs (Figures 

3.42a and 3.43a). Very similar patterns were documented for Scaridae 

biomass, but with very low biomass for nearshore lagoonal habitat types 

such as fringing mangroves, sand and macroalgae and seagrasses 

(Figures 3.42b and 3.43b). This suggests that many of the Scaridae 

found in high abundance in lagoonal areas were small bodied individuals 

and juveniles. The largest Caribbean parrotfish were rare or absent from the study including rainbow 

Redband (Sparisoma aurofrenatum), striped 

(Scarus iseri), and stoplight (Scarus viride) 

parrotfish

a)

Parrotfish abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Parrotfish biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.42. Maps of the interpolated (left map) and spatial (right map) distributions for parrotfish (Scaridae): (a) abundance and (b) 

biomass.

0

12

24

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean parrotfish density (#/100 m2)

0

0.6

1.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean parrotfish biomass (kg/100 m2)

a) b)

Figure 3.43. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for parrotfish species 

(Scaridae).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

89 3 - Fish Communities

parrotfish (Scarus guacamaia), two individuals observed. Midnight parrotfish (Scarus coelestinus) and 

blue parrotfish (Scarus coeruleus) were absent during the 2001-2007 sampling period.

Approximately 40% of all striped parrotfish (Scarus iseri) were the smallest juveniles (<5 cm; Figure 

3.44a). In contrast, for princess parrotfish (Scarus taeniopterus) adults were more frequently seen 

than juveniles (Figure 3.44b). The largest S. iseri, S. taeniopterus and redband parrotfish (Sparisoma 

aurofrenatum) of more than 30 cm FL were very rarely seen (<3%; Figure 3.44). Yellowtail parrotfish 

(Sparisoma rubripinne) and stoplight parrotfish (Sparisoma viride), were the only species with individuals 

more than 35 cm (Figures 3.44d and 3.44e), albeit a very small proportion of the total (2 and 4%, 

respectively). The largest individual S. rubripinne in the study area was estimated at 40 cm FL (Figure 

3.44d) and the maximum recorded for the species is 47.8 cm TL (Table 3.4). The largest individual S. 

viride in the study area was estimated at 50 cm FL (Figure 3.44e) and the maximum recorded for the 

species is 64 cm TL (Table 3.4).

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

S. iseri Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

S. taeniopterus Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

S. aurofrenatum Frequency (%)

Size class (cm)

Juveniles/subadults

a)

c)

b)

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

Sp. rubipinne Frequency (%)

Size class (cm)

d) Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

S. viride Frequency (%)

Size class (cm)

e) Juveniles/subadults

Figure 3.44. Size frequency histogram for select parrotfish (Scaridae) in the southwest Puerto Rico study area. (a) striped parrotfish (S. 

iseri), (b) princess parrotfish (S. taeniopterus), (c) redband parrotfish (S. aurofrenatum), (d) yellowtail parrotfish (S. rubripinne) and (e) 

stoplight parrotfish (S. viride).
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

90 3 - Fish CommunitiesStriped parrotfish (Scarus iseri)

S. iseri is the most frequently observed parrotfish across 

the study area found in all major habitat types and 

distributed widely across lagoonal, mid-shelf and outer 

shelf zones (Figure 3.45). Abundance and biomass is 

higher on the western side of the study area, particularly 

over highly rugose colonized hardbottom areas (Figure 

3.45). Mean abundance was relatively high in colonized 

hardbottom, seagrasses and mangroves, but biomass 

was markedly higher in colonized hardbottom than any 

other habitat type (Figure 3.46).

Terminal (left) and juvenile/initial (right) phase S. iseri.

a)

S. iseri abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. iseri biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.45. Maps of the interpolated (left map) and spatial (right map) distributions for striped parrotfish (S. iseri): (a) abundance and (b) 

biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. iseri biomass (kg/100 m2)

0

5

10

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. iseri density (#/100 m2)

a) b)

Figure 3.46. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for striped 

parrotfish (S. iseri).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

91 3 - Fish Communities

S. iseri juveniles and adults co-occurred in all zones and in all abundant habitat types throughout the 

study area, although adults used inshore mangroves and seagrasses less frequently than did juveniles 

(Figure 3.47). Mean densities of juveniles and adults were higher in the colonized hardbottom habitat 

types than softbottom habitat types such as macroalgae, seagrasses and sand. Highest densities of 

adults were calculated for colonized pavement with sand channels, colonized pavements and patch 

reefs (Figure 3.47). Highest mean densities of juveniles were calculated for patch reefs, linear reefs 

and seagrasses. High densities at colonized bedrock sites were due to relatively high abundance at 

only two sites (Figure 3.47).

0

15

30

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean S. iseri density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

100.0

0

54.4

55.9

61.0

71.4

57.0

52.0

74.5

1.3

28.2

2.4

12.2

72.7

1.7 4.3 5.4

25.6

35.1

24.5

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <10 cm

Adult >10

Figure 3.47. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for striped parrotfish (S. iseri). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile and adult for striped parrotfish 

(S. iseri) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

92 3 - Fish CommunitiesPrincess parrotfish (Scarus taeniopterus)

S. taeniopterus has a more geographically restricted distribution 

than S. iseri, with it being almost absent from nearshore areas 

and highest abundance and biomass observed over colonized 

hardbottom along the outer shelf zone (Figure 

3.48). S. taeniopterus was more frequently 

observed over areas of highly contiguous 

colonized hardbottom. It was absent from 

mangroves and only found in very low 

abundance over seagrasses and unvegetated 

sediments (Figure 3.49).

Terminal (bottom) and juvenile/initial (top) phase princess parrotfish (Scarus taeniopterus)

a)

S. taeniopterus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. taeniopterus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.48. Maps of the interpolated (left map) and spatial (right map) distributions for princess parrotfish (S. taeniopterus): (a) abundance 

and (b) biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. taeniopterus biomass (kg/100 m2)

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. taeniopterus density (#/100 m2)

a) b)

Figure 3.49. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for princess 

parrotfish (S. taeniopterus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

93 3 - Fish Communities

Juvenile and adult S. taeniopterus co-occurred across colonized hardbottom habitat types of the mid 

and outer shelf zones, but in contrast to S. iseri were rarely encountered in nearshore and lagoonal 

environments (Figures 3.47 and 3.50). Highest densities of juveniles and adults were calculated for the 

most topographically complex areas of colonized pavement, colonized pavement with sand channels, 

linear reefs and patch reefs. Juveniles and adults were absent from mangroves and reef rubble and 

occurred only once in seagrass beds (Figure 3.50).

0

2

4

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean S. taeniopterus density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0

29.7

36.9

36.6

48.8

27.0

20.0

10.9

0 0

2.4 4.9

23.6

4.3 3.2

0.4 0.4

10.6

12.8

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <10 cm

Adult >10

Figure 3.50. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for princess parrotfish (S. 

taeniopterus). Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile and adult for 

princess parrotfish (S. taeniopterus) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

94 3 - Fish CommunitiesRedband parrotfish (Sparisoma aurofrenatum)

S. aurofrenatum exhibited a close affinity with 

colonized hardbottom, with more than 90% of all 

sightings occurring over colonized hardbottom 

habitat types (Figures 3.51 and 3.52). Abundance 

and biomass were generally highest in the mid to 

outer shelf zones (Figure 3.51).

Terminal (bottom) and initial (top) phase (S. aurofrenatum.)

a)

S. aurofrenatum abundance (100 m2)

0

1 - 10

11 - 25

>25

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. aurofrenatum biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.51. Maps of the interpolated (left map) and spatial (right map) distributions for redband parrotfish (S. aurofrenatum): (a) 

abundance and (b) biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. aurofrenatum biomass (kg/100 m2)

0

3

6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. aurofrenatum density (#/100 m2)

a) b)

Figure 3.52. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for redband 

parrotfish (S. aurofrenatum).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

95 3 - Fish Communities

Juvenile and adult S. aurofrenatum co-occurred throughout the study area and in all zones and in all 

habitat types except colonized bedrock (Figure 3.53). Density of juvenile and adult S. aurofrenatum

was markedly higher over colonized hardbottom habitat types than softbottom, particularly colonized 

hardbottom with sand channels, patch reefs, linear reefs and colonized pavement. Softbottom habitat 

types and mangroves supported relatively low densities of both juveniles and adults (Figure 3.53).

0

3

6

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean S. aurofrenatum density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0

63.1 68.5

83.6

83.1

81.0

67.0

67.3

0.6 7.1 7.3

2.4

78.2

4.3 6.5

1.3 4.7

30.9

34.0

100.0

0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <10 cm

Adult >10

Figure 3.53. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for redband parrotfish (S. 

aurofrenatum). Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile and adult for 

redband parrotfish (S. aurofrenatum) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

96 3 - Fish CommunitiesBucktooth parrotfish (Sparisoma radians)

S. radians is a widespread resident species in seagrass beds and is also 

found at the interface between seagrasses and mangroves and coral 

reefs and seagrasses (Figure 3.54). High abundance was recorded in 

the large continuous seagrass beds on the western side of the study 

area and on some surveys on the mid- and outer-shelf within patches 

of seagrasses surrounded by colonized hardbottom (Figure 3.55). 

Lowest mean abundance and biomass were recorded for colonized 

hardbottom and unvegetated sediments (Figure 3.55). Bucktooth parrotfish (Sparisoma radians)

a)

S. radians abundance (100 m2)

0

1 - 10

11 - 25

>25

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. radians biomass (g/100 m2)

0

0.1 - 370

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.54. Maps of the interpolated (left map) and spatial (right map) distributions for bucktooth parrotfish (S. radians): (a) abundance 

and (b) biomass.

0

0.01

0.02

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. radians biomass (kg/100 m2)

0

3

6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. radians density (#/100 m2)

a) b)

Figure 3.55. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for bucktooth 

parrotfish (S. radians).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

97 3 - Fish Communities

Yellowtail parrotfish (Sparisoma rubripinne)

S. rubripinne was sighted in all major habitat types, but was 

more frequently sighted in colonized hardbottom habitat types 

than any others (Figure 3.56). The majority of S. rubripinne

were from nearshore and mid-shelf zones (Figure 3.57). 

Terminal phase yellowtail parrotfish (Sparisoma rubripinne) in St. 

Croix, USVI.

0

0.02

0.04

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. rubripinne biomass (kg/100 m2)

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. rubripinne density (#/100 m2)

a) b)

Figure 3.56. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for yellowtail 

parrotfish (S. rubripinne).

Habitat type Habitat type

a)

S. rubripinne abundance (100 m2)

0

1 - 10

>10

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. rubripinne biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.57. Maps of the interpolated (left) and spatial (right) distributions for yellowtail parrotfish (S. rubripinne): (a) abundance and (b) 

biomass.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

98 3 - Fish CommunitiesStoplight parrotfish (Sparisoma viride)

S. viride was associated with all major habitat types, but with markedly higher 

mean abundance and mean biomass over colonized hardbottom (Figure 

3.59). Mid- and outer-shelf zones were used more than the nearshore and 

lagoonal zone (Figure 3.58). High biomass was recorded 

for the high rugosity coral reefs in the vicinity of El Palo 

and Margarita reefs and several of the nearshore reefs 

on the northeastern side of the study area (Figure 3.58b). 

Terminal (bottom; St. Croix) and initial (top; PR) phase S. viride

a)

S. viride abundance (100 m2)

0

1 - 10

>10

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. viride biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.58. Maps of the interpolated (left map) and spatial (right map) distributions for stoplight parrotfish (S. viride): (a) abundance and 

(b) biomass.

0

0.3

0.6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. viride biomass (kg/100 m2)

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. viride density (#/100 m2)

a) b)

Habitat type Habitat type Figure 3.59. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for stoplight 

parrotfish (S. viride).
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

99 3 - Fish Communities

Like other common parrotfish species, life stages of S. viride co-occurred across the study area showing 

no recognizable spatial segregation (Figure 3.60). Both juveniles and adults were found in all of the 

most abundant habitat types, with highest densities over colonized hardbottom including patch reefs, 

colonized pavement with sand channels, colonized pavement and linear reefs (Figure 3.60). Lowest 

densities were recorded for unconsolidated sediments, seagrasses and mangroves. Both juveniles and 

adults were recorded in low abundance from seagrasses and mangroves. Similar to other Scaridae 

species, presence across the study area was higher in the most topographically complex hardbottom 

areas (Figure 3.60).

0

2

4

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean S. viride density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

100.0

50.0 32.4

35.1

37.1

47.9

46.0

48.0

65.5

2.6 10.3

0 0

49.1

1.7 3.2 3.2 4.7

25.5

13.8

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Figure 3.60. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for stoplight parrotfish (S. viride). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for stoplight 

parrotfish (S. viride) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

100 3 - Fish CommunitiesGrunts (Haemulidae)

Haemulidae abundance was highest in the nearshore 

and midshelf mangroves, particularly around the 

mangrove islands in close proximity to coral reefs 

and seagrass beds and on shallow water patch 

reefs surrounded by seagrasses in the lagoonal and 

backreef zones (Figures 3.61a and 3.62a). Haemulidae 

abundance was also high in several locations over 

colonized pavement with sand channels in the outer 

shelf zone. Highest mean Haemulidae biomass was 

also found in mangroves (Figure 3.62b), with highest 

biomass calculated for several sites along the fringing 

mangroves and mangrove islands and associated 

backreef zones on the midshelf zone (Figure 3.61b). 

Several sites on colonized hardbottom sites (colonized 

pavement with sand channels and patch reefs) in close 

proximity to seagrasses and sand patches also exhibited high haemulidae biomass.

Assemblage of tomtates (Haemulon aurolineatum). 

a)

Grunt abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Grunt biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.61. Maps of the interpolated (left map) and spatial (right map) distributions for grunt (Haemulidae): (a) abundance and (b) biomass.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

101 3 - Fish Communities

0

25

50

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean grunt density (#/100 m2)

0

0.4

0.8

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean grunt biomass (kg/100 m2)

a) b)

Figure 3.62. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for grunt species 

(Haemulidae).

The largest proportion of haemulids in the study area were represented by individuals of the smallest 

size class (<5 cm FL; Figure 3.63). This is likely to be an underestimate since some observers classify 

small grunts as unknown haemulids due to the difficulty of identifying the juveniles to species level. 

Approximately 95% of Haemulon aurolineatum (tomtate) and Haemulon flavolineatum (French grunt)

were juveniles and subadults (Figures 3.63a and 3.63b) and approximately 85% of Haemulon sciurus 

(bluestriped grunt) and 80% of Haemulon plumierii (white grunt; Figures 3.63d and 3.63c). No individuals 

greater than 20 cm FL were recorded for H. flavolineatum; 25 cm FL for H. aurolineatum; 30 cm FL 

for H. sciurus and 30 cm FL for H. plumierii (Figure 3.63). None of the common haemulid individuals 

reached the maximum known for the species (Table 3.4). 

0

25

50

5-10 10-15 15-20 20-25 25-30 30-35 >35

H. aurolineatum Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

5-10 10-15 15-20 20-25 25-30 30-35 >35

H. flavolineatum Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

5-10 10-15 15-20 20-25 25-30 30-35 >35

H. plumierii Frequency (%)

Size class (cm)

Juveniles/subadults

a)

c)

b)

0

25

50

5-10 10-15 15-20 20-25 25-30 30-35 >35

H. sciurus Frequency (%)

Size class (cm)

d) Juveniles/subadults

Figure 3.63. Size frequency histogram for select grunts (Haemulidae) in the southwest Puerto Rico study area. (a) tomtate (H. aurolineatum), 

(b) French grunt (H. flavolineatum), (c) white grunt (H. plumierii) and (d) bluestriped grunt (H. sciurus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

102 3 - Fish CommunitiesTomtate (Haemulon aurolineatum)

H. aurolineatum utilized all major habitat types, with highest 

abundance recorded on nearshore patch reefs in close proximity to 

extensive softbottom habitat types (seagrasses and unvegetated 

sediments; Figure 3.64). Although the benthic habitat map 

indicates highest mean abundance and biomass over unvegetated 

sediments it is likely that some finer-scale unmapped hardbottom 

structure also occurred along these sand dominated sample sites 

(Figures 3.64 and 3.65). Abundance and biomass were lowest on 

colonized hardbottom areas of the outer shelf.

H. aurolineatum

a)

H. aurolineatum abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

H. aurolineatum biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.64. Maps of the interpolated (left map) and spatial (right map) distributions for tomtate (H. aurolineatum): (a) abundance and (b) 

biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. aurolineatum biomass (kg/100 

m2)

Habitat type

0

6

12

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. aurolineatum density (#/100 m2)

a) b)

Figure 3.65. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for tomtate (H. 

aurolineatum).

Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

103 3 - Fish Communities

H. aurolineatum juveniles, subadults and adults showed a less distinct pattern of spatial segregation 

across the shelf compared to other Haemulid species, although similarly, few adults were found amongst 

mangroves and in shallow nearshore vegetated habitat types and few juveniles were found on the outer 

shelf (Figure 3.66). Although high geographical overlap across lifestages was observed, highest mean 

densities of juveniles/subadults were recorded over scattered coral in sand, macroalgal beds, patch 

reefs and linear reefs, seagrasses and mangroves of the mid and nearshore zones, while highest 

densities of adults were recorded over scattered coral and patch reefs in mid- and outer-shelf zones 

(Figure 3.66).

0

4

8

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean H. aurolineatum density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0 10.8 3.6

6.1

2.8 1.0

8.0 5.5

0

5.8

2.4

7.3

12.7

6.5

0.4 1.1

4.7

18.1

8.5

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Figure 3.66. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for tomtate (H. aurolineatum). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for tomtate

(H. aurolineatum) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

104 3 - Fish CommunitiesFrench grunt (Haemulon flavolineatum)

H. flavolineatum were widely distributed throughout the 

study area, with highest mean abundance and biomass 

in mangroves, particularly the mangrove cays around Isla 

Magueyes and Enrique reefs (Figure 3.67). Mean biomass 

was higher in colonized hardbottom than seagrasses and 

unvegetated sediments (Figure 3.68b).

 French grunt (Haemulon flavolineatum)

a)

H. flavolineatum abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

H. flavolineatum biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.67. Maps of the interpolated (left map) and spatial (right map) distributions for French grunt (H. flavolineatum): (a) abundance and 

(b) biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. flavolineatum biomass (kg/100 m2)

Habitat type

0

15

30

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. flavolineatum density (#/100 m2)

a) b)

Figure 3.68. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for French grunt

(H. flavolineatum).

Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

105 3 - Fish Communities

H. flavolineatum juveniles were distributed primarily in fringing mangroves and mangrove cays, but were 

also observed co-occurring with adults and subadults at several hardbottom sites on the mid and outer 

shelf (Figure 3.69). Subadult, and particularly adults, occurred in lower densities than juveniles overall 

and exhibited a distinctly more offshore distribution, with highest densities on patch reefs, colonized 

pavement with sand channels and linear coral reefs. Subadults, and occasionally adults, were also 

observed using inshore habitat types (Figure 3.69).

0

15

30

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean H. flavolineatum density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0 24.3 7.2 27.3

16.0 22.0

23.0

0.6 16.4

71.8

2.4 0 4.3

6.5

0.4 2.2

5.1

16.0

0 0 4.3

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Figure 3.69. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for French grunt (H. flavolineatum). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for French 

grunt (H. flavolineatum) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

106 3 - Fish CommunitiesWhite grunt (Haemulon plumierii)

H. plumierii were widely distributed in the nearshore lagoonal zone 

and across all major habitat types (Figure 3.70). Mean density 

was highest in neashore seagrasses and mangroves and mean 

biomass was highest in colonized hardbottom habitat types (Figures 

3.70 and 3.71). Several high biomass sites existed over colonized 

hardbottom areas along the mid to outer shelf zone.

White grunt (Haemulon plumierii)

a)

H. plumierii abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

H. plumierii biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.70. Maps of the interpolated (left map) and spatial (right map) distributions for white grunt (H. plumierii): (a) abundance and (b) 

biomass.

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. plumierii biomass (kg/100 m2)

0

1.5

3

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. plumierii density (#/100 m2)

a) b)

Figure 3.71. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for white grunt

(H. plumierii).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

107 3 - Fish Communities

H. plumierii juveniles were more frequently distributed in mangroves and seagrasses in lagoonal and 

nearshore zones (Figure 3.72), but like H. flavolineatum, juvenile H. plumierii were also observed in 

mid and outer shelf zones usually on or very near to seagrasses (Figures 3.69 and 3.72). Subadults 

and adults also inhabited nearshore habitat types, but were more frequently observed over colonized 

hardbottom sites on the mid and outer shelf zones (Figure 3.72). Highest densities of juveniles/subadults 

were recorded for macroalgal and seagrass habitat types and highest densities for adults were recorded 

for patch reefs and colonized pavement with sand channels.

0

2

4

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean H. plumierii density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0 0 7.2

12.6 7.5 10.8

5.0

8.0

7.3

0

14.1

2.4

14.6

12.7

1.1 3.2

0

18.4

8.5

9.6

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Figure 3.72. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for white grunt (H. plumierii). Number 

above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for white grunt (H. 

plumierii) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

108 3 - Fish CommunitiesBluestriped grunt (Haemulon sciurus)

Although H. sciurus were observed in all major habitat types, 

both the abundance and biomass of H. sciurus was highest 

in the mangroves of the nearshore lagoonal zone and around 

many of the island cays, particularly on the western side of the 

study area (Figure 3.73). Lowest abundance was recorded for 

unvegetated sediments and lowest biomass for seagrasses 

and macroalgal beds (Figure 3.74).

Bluestriped grunt (Haemulon sciurus)

a)

H. sciurus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

H. sciurus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.73. Maps of the interpolated (left map) and spatial (right map) distributions for bluestriped grunt (H. sciurus): (a) abundance and 

(b) biomass.

0

0.3

0.6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. sciurus biomass (kg/100 m2)

0

7

14

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean H. sciurus density (#/100 m2)

a) b)

Figure 3.74. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for bluestriped 

grunt (H. sciurus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

109 3 - Fish Communities

H. sciurus juveniles were most frequently sighted in mangroves (fringing and cays), but also utilized 

several other habitat types including seagrasses, macroalgal beds, scattered coral in sand, patch 

reefs, linear reefs and colonized bedrock (Figure 3.75). In contrast to other haemulids, subadult H. 

sciurus were more frequently seen in nearshore and lagoonal habitat types than on the mid and outer 

shelf suggesting an offshore ontogenetic shift at a later stage than appears evident for other common 

haemulids (Figure 3.75). Subadults appeared to co-occur more often with juveniles (very occasionally 

with adults) and were frequently observed using mangroves, but adults were more frequently sighted 

on mid and outer shelf zones (Figure 3.75). Highest densities of all life stages were recorded for 

mangroves.

0

7

14

Colonized 

bedrock

Colonized 

pavement

Col. pav. 

sand chan.

Linear 

reef

MacroalgaeMangrove Patch 

reef

Reef 

rubble

Scat coral/ 

rock sand

Seagrass Uncon. 

sediment

Mean H. sciurus density (#/100 m2)

Mapped habitat type

Juveniles/Subadults

Adults

0

50.0

0 2.7 4.2 10.3 3.0 14.0

7.2 25.6

75.0

0

4.9

11.0 0.9 1.1 3.2

8.1

3.2

8.5

0 0

Benthic HABITAT

Coral reef and 

colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Fish size classes

Juvenile <5 - 10 cm

Subadult 10 - 15 cm

Adult >15

Figure 3.75. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for bluestriped grunt (H. sciurus). 

Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for 

bluestriped grunt (H. sciurus) in the southwest Puerto Rico study area.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

110 3 - Fish CommunitiesSurgeonfish (Acanthuridae)

Acanthurids, a major family of herbivorous fish, 

were markedly more abundant across colonized 

hardbottom habitat types, with abundance and 

biomass hotspots at several locations around El 

Palo and Margarita Reefs and across colonized 

hardbottom of the outer shelf (Figures 3.76 and 

3.77). Lowest abundance and biomass were 

observed in the shallow lagoonal habitat types 

including mangroves closest to the shoreline 

(Figure 3.76). Assemblage of blue tangs (Acanthurus coeruleus).

a)

Surgeonfish abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Surgeonfish biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.76. Maps of the interpolated (left map) and spatial (right map) distributions for surgeonfish (Acanthuridae): (a) abundance and (b) 

biomass.

0

3

6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean surgeonfish density (#/100 m2)

0

0.3

0.6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean surgeonfish biomass (kg/100 m2)

a) b)

Figure 3.77. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for surgeonfish 

(Acanthuridae).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

111 3 - Fish Communities

Less than 35% of the surgeonfish population sampled in the La Parguera study area were classified as 

juveniles and subadults (≤10 cm; Figure 3.78). The majority of Acanthurus bahianus (ocean surgeon) 

and Acanthurus coeruleus (blue tang) were between 10 and 20 cm FL (Figures 3.78a and 3.78c) and 

between 10 and 25 cm for Acanthurus chirurgus (doctorfish; Figure 3.78b). Less than 5% of surgeonfish 

were larger than 25 cm FL. No individuals were seen above 25 cm FL for A. bahianus, 30 cm FL for 

A. coeruleus and 35 cm FL for A. chirurgus and the maximum sizes known for these species were not 

attained in the study area (Table 3.4).

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

A. bahianus Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

A. chirurgus Frequency (%)

Size class (cm)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

A. coeruleusFrequency (%)

Size class (cm)

Juveniles/subadults

a)

c)

b)

Figure 3.78. Size frequency histogram for surgeonfish (Acanthuridae) in the southwest Puerto Rico study area. (a) ocean surgeon (A. 

bahianus), (b) doctorfish (A. chirurgus) and (c) blue tang (A. coeruleus).
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

112 3 - Fish CommunitiesOcean surgeonfish (Acanthurus bahianus)

The most frequently occurring of the surgeonfish species, A. 

bahianus, exhibited highest abundance and biomass over 

colonized hardbottom habitat types of the mid and outer shelf 

zones (Figures 3.79 and 3.80). Lowest mean abundance and 

biomass were observed in mangroves and over unvegetated 

sediments (Figure 3.80), particularly in the very nearshore 

zones (Figure 3.79). 

Ocean surgeonfish (Acanthurus bahianus; top) and spotted 

goatfish (Pseudupeneus maculatus).

a)

A. bahianus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

A. bahianus biomass (g/100 m2)

0

0.1 - 100

100.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.79. Maps of the interpolated (left map) and spatial (right map) distributions for ocean surgeonfish (A. bahianus): (a) abundance 

and (b) biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean A. bahianus biomass (kg/100 m2)

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean A. bahianus density (#/100 m2)

a) b)

Figure 3.80. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for ocean 

surgeonfish (A. bahianus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

113 3 - Fish Communities

Doctorfish (Acanthurus chirurgus)

A. chirurgus is the least widely distributed of Acanthuridae 

species, with lowest abundance and biomass. An area of high 

abundance and biomass occured in close proximity to the coral 

reefs in the El Palo and Margarita Reef areas, as well as several 

locations along the linear reef on the outer shelf (Figure 3.81). In 

contrast to other Acanthuridae species, a large proportion of the 

total abundance is observed within mangroves and seagrasses, 

although biomass was relatively low (Figure 3.82).

 Doctorfish (Acanthurus chirurgus)

a)

A. chirurgus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

A. chirurgus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.81. Maps of the interpolated (left map) and spatial (right map) distributions for doctorfish (A. chirurgus): (a) abundance and (b) 

biomass.

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean A. chirurgus biomass (kg/100 m2)

0

0.5

1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean A. chirurgus density (#/100 m2)

a) b)

Figure 3.82. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for doctorfish (A. 

chirurgus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

114 3 - Fish CommunitiesBlue tang (Acanthurus coeruleus)

A. coeruleus distribution closely follows the distribution of colonized 

hardbottom, with hotspots of abundance and biomass located on 

Margarita Reef and around El Palo reef (Figure 3.83). Very few blue tang 

were observed in nearshore mangroves and seagrasses (Figure 3.84).

A. coeruleus

a)

A. coeruleus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

A. coeruleus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.83. Maps of the interpolated (left map) and spatial (right map) distributions for blue tang (A. coeruleus): (a) abundance and (b) 

biomass.

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean A. coeruleus biomass (kg/100 m2)

0

1

2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean A. coeruleus density (#/100 m2)

Figure 3.84. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for blue tang (A. 

coeruleus).

Habitat type Habitat type

a) b)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

115 3 - Fish Communities

Goatfishes (Mullidae)

The smallest juveniles (<5 cm) represented a very 

small proportion (<6% of all Pseudupeneus maculatus

[spotted goatfish]) of all Mullidae (Figure 3.85). None 

of the smallest Mulloidichthys martinicus (yellow 

goatfish) were seen in the study area (Figure 3.85a). 

The largest proportion of all Mullidae were large 

juveniles and subadults, with a marked decline in the 

proportion of adults, with a very small proportion of 

the largest adults (>30 cm FL= 1% of the total). No P. 

maculatus larger than 30 cm FL were recorded and no 

M. martinicus larger than 35 cm FL were recorded in 

the study area (Figure 3.85). The maximum known for 

the species is 30 cm for P. maculatus and 39.4 for M. 

martinicus (Table 3.4).

P. maculatus

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

M. martinicus Frequency (%)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

P. maculatus Frequency (%)

Size class (cm)

a) b) Juveniles/subadults

Figure 3.85. Size frequency histogram for goatfish (Mullidae) in the southwest Puerto Rico study area. (a) yellow goatfish (M. martinicus) 

and (b) spotted goatfish (P. maculatus).

Size class (cm)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

116 3 - Fish CommunitiesYellow goatfish (Mulloidichthys martinicus)

Although M. martinicus was observed using all major 

habitat types it was less widespread and abundant than 

P. maculatus (Figures 3.86 and 3.87), with highest mean 

abundance over unvegetated sediments and colonized 

hardbottom (Figure 3.87a). Mean biomass was highest 

for colonized hardbottom. M. martinicus also utilized 

mangroves (Figure 3.87b). Yellow goatfish (Mulloidichthys martinicus) in St. John, USVI

a)

M. martinicus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

M. martinicus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.86. Maps of the interpolated (left map) and spatial (right map) distributions for yellow goatfish (M. martinicus): (a) abundance and 

(b) biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean M. martinicus biomass (kg/100 m2)

0

0.7

1.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean M. martinicus density (#/100 m2)

a) b)

Figure 3.87. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for yellow goatfish

(M. martinicus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

117 3 - Fish Communities

Spotted goatfish (Pseudupeneus maculatus)

P. maculatus was found in all major habitat types except 

mangroves and in all zones across the shelf (Figure 3.88). No 

distinct geographical hotspots were evident within the study area. 

Mean abundance was highest over seagrasses and then colonized 

hardbottom, while the reverse was true for mean biomass (Figure 

3.89).

P. maculatus

a)

P. maculatus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

P. maculatus biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.88. Maps of the interpolated (left map) and spatial (right map) distributions for spotted goatfish (P. maculatus): (a) abundance and 

(b) biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean P. maculatus biomass (kg/100 m2)

0

0.4

0.8

Colonized Macroalgae/ Unvegetated Mangrove

Mean P. maculatus density (#/100 m2)

a) b)

Figure 3.89. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for spotted 

goatfish (P. maculatus).

Habitat type

Hardbottom Seagrass Sediments

Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

118 3 - Fish CommunitiesJacks (Carangidae)

Highest mean Carangidae abundance was associated with 

colonized hardbottom habitat types (Figure 3.90a), with 

highest abundance found on or around the extensive area 

of colonized hardbottom on the mid and outer shelf zones 

(Figure 3.90a). The high biomass associated with larger 

schools of carangids and larger-bodied carangids were 

found at several locations over softbottom habitat types 

(seagrass and sand; Figure 3.90b) close to the edges of 

colonized hardbottom (Figure 3.90b). Very few carangids 

were observed in fringing mangroves and from nearshore 

environments in general (Figure 3.91). Yellow jack (Carangoides bartholomaei)

a)

Jack abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Jack biomass (g/100 m2)

0

0.1 - 1000

1000.1 - 5000

5000.1 - 10000

>10000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.90. Maps of the interpolated (left map) and spatial (right map) distributions for jack (Carangidae): (a) abundance and (b) biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean jack biomass (kg/100 m2)

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean jack density (#/100 m2)

a) b)

Figure 3.91. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for jack (Carangidae). Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

119 3 - Fish Communities

Two of the most common Carangidae 

species were Caranx crysos (blue runner) 

and Carangoides ruber (bar jack). C. 

crysos density and biomass were highest 

around unvegetated sediments. Very few 

were observed in colonized hardbottom 

and macroalgae/seagrass habitats, and 

absent in mangroves (Figure 3.92). C. 

ruber density and biomass were highest 

in colonized hardbottom habitats and very few recorded in softbottom habitats (sand and seagrasses; 

Figure 3.93). As with C. crysos, C. ruber were also absent from mangroves (Figure 3.93).

Blue runner (Caranx crysos) Bar jack (Carangoides ruber)

0

0.15

0.3

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. crysos biomass (kg/100 m2)

0

0.3

0.6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. crysos density (#/100 m2)

a) b)

Figure 3.92. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for blue runner 

(C. crysos).

Habitat type Habitat type

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. ruber biomass (kg/100 m2)

0

1

2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean C. ruber density (#/100 m2)

a) b)

Figure 3.93. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for bar jack (C. 

ruber).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

120 3 - Fish CommunitiesDamselfish (Pomacentridae)

Yellowtail damselfish (Microspathodon chrysurus)

Like most damselfish, M. chrysurus were very closely associated 

with colonized hardbottom habitat types, but exhibited a distribution 

primarily within the mid shelf zone (Figure 3.94). Mangroves were 

not used by this species and nearshore and lagoonal colonized 

hardbottom areas were rarely used. Mean abundance and biomass 

were very low over seagrasses and unvegetated sediments (Figure 

3.95).

Yellowtail damselfish (Microspathodon chrysurus)

a)

M. chrysurus abundance (100 m2)

0

1 - 10

10.1 - 25

>25

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

M. chrysurus biomass (g/100 m2)

0

0.1 - 100

100.1 - 500

500.1 - 1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.94. Maps of the interpolated (left map) and spatial (right map) distributions for yellowtail damselfish (M. chrysurus): (a) abundance 

and (b) biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean M. chrysurus biomass (kg/100 m2)

0

0.4

0.8

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean M. chrysurus density (#/100 m2)

a) b)

Figure 3.95. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for yellowtail 

damselfish (M. chrysurus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

121 3 - Fish Communities

Dusky damselfish (Stegastes adustus)

S. adustus were seen in all major habitat types, but were mostly 

observed over colonized hardbottom, with highest abundance 

found on the shallow water forereefs, reefcrests and patch reefs 

of the mid shelf and lagoonal zones (Figure 3.96). S. adustus were 

rarely seen on the deeper colonized hardbottom areas of the outer 

shelf zone. Lowest abundance and biomass were recorded for 

mangroves and unvegetated sediments (Figure 3.97).

 Dusky damselfish (Stegastes adustus) in St. Croix, USVI

a)

S. adustus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. adustus biomass (g/100 m2)

0

0.1 - 500

>500

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.96. Maps of the interpolated (left map) and spatial (right map) distributions for dusky damselfish (S. adustus): (a) abundance and 

(b) biomass.

0

0.03

0.06

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. adustus biomass (kg/100 m2)

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. adustus density (#/100 m2)

a) b)

Figure 3.97. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for dusky 

damselfish (S. adustus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

122 3 - Fish CommunitiesBeaugregory (Stegastes leucostictus)

S. leucostictus was observed in all major habitat types and 

was widely distributed in nearshore and lagoonal zones, 

with highest mean abundance and biomass in both fringing 

mangroves and mangrove cays (Figure 3.98). Comparatively 

few S. leucostictus were observed over deeper water 

colonized hardbottom on the mid and outer shelf zone. Lowest 

abundance and biomass were recorded for unvegetated 

sediments (Figure 3.99). Beaugregory (Stegastes leucostictus) juvenile in St. Croix, USVI

a)

S. leucostictus abundance (100 m2)

0

1 - 10

11 - 25

>25

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. leucostictus biomass (g/100 m2)

0

0.1 - 290

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.98. Maps of the interpolated (left map) and spatial (right map) distributions for beaugregory (S. leucostictus): (a) abundance and 

(b) biomass.

0

0.03

0.06

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. leucostictus biomass (kg/100 m2)

0

5

10

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. leucostictus density (#/100 m2)

a) b)

Figure 3.99. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for beaugregory

(S. leucostictus). 

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

123 3 - Fish Communities

Bicolor damselfish (Stegastes partitus)

S. partitus is found at approximately 39% of sites across the study area, 

with highest mean abundance and biomass associated with colonized 

hardbottom habitat types. Geographically, abundance is markedly 

higher on the colonized hardbottom sites located on the outer shelf 

zone and lowest for the very nearshore and lagoonal zones (Figure 

3.100). Lowest abundance and biomass were recorded for mangroves 

and unvegetated sediments (Figure 3.101).

Bicolor damselfish (Stegastes partitus)

a)

S. partitus abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. partitus biomass (g/100 m2)

0

0.1 - 500

>500

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.100. Maps of the interpolated (left map) and spatial (right map) distributions for bicolor damselfish (S. partitus): (a) abundance and 

(b) biomass.

0

0.03

0.06

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. partitus biomass (kg/100 m2)

0

4

8

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. partitus density (#/100 m2)

a) b)

Figure 3.101. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for bicolor 

damselfish (S. partitus).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

124 3 - Fish CommunitiesThreespot damselfish (Stegastes planifrons)

S. planifrons was well distributed across the colonized hardbottom 

habitat types of the mid shelf zone predominantly occurring in forereefs 

and reefcrest zones and in the most topographically complex areas of 

colonized pavement with sand channels on the mid and outer shelf 

zone (Figure 3.102). Lowest mean abundance and biomass of S. 

planifrons was recorded for mangroves (Figure 3.103).

Adult (left) and juvenile (right) threespot damselfish 

(Stegastes planifrons).

a)

S. planifrons abundance (100 m2)

0

1 - 10

11 - 25

26 - 50

>50

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. planifrons biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.102. Maps of the interpolated (left map) and spatial (right map) distributions for threespot damselfish (S. planifrons): (a) abundance 

and (b) biomass.

0

0.03

0.06

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. planifrons biomass (kg/100 m2)

Habitat type

0

2

4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. planifrons density (#/100 m2)

a) b)

Figure 3.103. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for threespot 

damselfish (S. planifrons).

Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

125 3 - Fish Communities

Cocoa damselfish (Stegastes variabilis)

S. variabilis exhibited a similar distribution as S. planifrons associated with 

topographically complex hardbottom, albeit observed at lower densities 

(Figures 3.104). Mean abundance and biomass were highest in colonized 

hardbottom and lowest in seagrasses. Of note was that S. variabilis 

abundance was higher in mangroves than seagrasses (Figure 3.105). Cocoa damselfish (Stegastes variabilis), St. 

John, USVI.

a)

S. variabilis abundance (100 m2)

0

1 - 10

>10

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. variabilis biomass (g/100 m2)

0

0.1 - 100

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.104. Maps of the interpolated (left map) and spatial (right map) distributions for cocoa damselfish (S. variabilis): (a) abundance 

and (b) biomass.

0

0.01

0.02

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. variabilis biomass (kg/100 m2)

0

1

2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. variabilis density (#/100 m2)

a) b)

Figure 3.105. Comparison of mean (+ SE) density and biomass by substrate type in southwest Puerto Rico for cocoa damselfish (S. 

variabilis).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

126 3 - Fish CommunitiesWrasses (Labridae)

Highest Labridae density and biomass were observed on 

the colonized hardbottom sites across the outer shelf zone, 

with several isolated abundance hotspots across colonized 

hardbottom sites in the midshelf zone (Figures 3.106 and 

3.107). 

Bluehead wrasse (Thalassoma bifasciatum)

a)

Wrasse abundance (100 m2)

0

1 - 10

11 - 50

51 - 100

>100

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Wrasse biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.106. Maps of the interpolated (left map) and spatial (right map) distributions for wrasse (Labridae): (a) abundance and (b) biomass.

>5000

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean wrasse biomass (kg/100 m2)

0

10

20

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean wrasse density (#/100 m2)

a) b)

Figure 3.107. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for wrasse 

(Labridae).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

127 3 - Fish Communities

Porgies (Sparidae)

Sparidae species were most abundant in fringing mangrove 

and mangrove islands, but were also found over colonized 

hardbottom habitat types across the mid- and outer shelf 

zones (Figures 3.108 and 3.109).

Saucereye porgy (Calamus calamus) in St. John, USVI

a)

Porgy abundance (100 m2)

0

1 - 10

>10

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

Porgy biomass (g/100 m2)

0

0.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.108. Maps of the interpolated (left map) and spatial (right map) distributions for porgy (Sparidae): (a) abundance and (b) biomass.

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean porgy biomass (kg/100 m2)

0

0.6

1.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean porgy density (#/100 m2)

a) b)

Figure 3.109. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for porgy 

(Labridae).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

128 3 - Fish CommunitiesOther species

Sharks and Rays

Sharks and rays were comprised of four species including three Dasyatis americana (southern stingray), 

four Ginglymostoma cirratum (nurse shark) and one Galeocerdo cuvier (tiger shark).

Sightings occurred in all major habitat types, either in or close proximity to seagrass beds across 

the shelf (Figure 3.110 and 3.111). Highest mean biomass was recorded for seagrasses/macroalgal 

beds and was lowest over unvegetated 

sediments. Relatively high abundance, 

but low biomass for mangroves indicates 

that most sharks and rays in mangroves 

were smaller individuals than those 

sighted over seagrasses and colonized 

hardbottom. Insufficient sightings 

precluded a description of spatial 

distributions (Figure 3.110).

Nurse shark (Ginglymostoma cirratum) 

Southern stingray (Dasyatis americana) 

Abundance (100 m2)

0

1 

Biomass (g/100 m2)

0

1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

a)

b)

Figure 3.110. Map of the spatial distributions for sharks and rays: (a) abundance 

and (b) biomass.

0

0.4

0.8

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean shark & ray biomass (kg/100 m2)

0

0.05

0.1

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean shark & ray density (#/100 m2)

a) b)

Figure 3.111. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for sharks and 

rays.

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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129 3 - Fish Communities

Queen triggerfish (Balistes vetula)

B. vetula was absent from nearshore and lagoonal areas, but 

was broadly distributed across colonized hardbottom habitat 

types of the mid and outer shelf zones (Figure 3.112). Highest 

abundance and biomass were recorded for topographically 

complex colonized pavement with sand channels. B. vetula

was absent from mangroves and very few were observed over 

unvegetated sediments across the study area (Figure 3.113).

 Queen triggerfish (Balistes vetula)

a)

B. vetula abundance (100 m2)

0

1 - 2

>2

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

B. vetula biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

>1000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.112. Maps of the interpolated (left map) and spatial (right map) distributions for queen triggerfish (B. vetula): (a) abundance and 

(b) biomass.

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean B. vetula biomass (kg/100 m2)

0

0.1

0.2

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean B. vetula density (#/100 m2)

a) b)

Figure 3.113. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for queen 

triggerfish (B. vetula).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

130 3 - Fish CommunitiesGreat barracuda (Sphyraena barracuda)

Highest mean abundance of S. barracuda was 

observed within both fringing mangroves of the 

mainland and mangrove cays (Figures 3.114a and 

3.115a). Highest biomass, however, were calculated 

for colonized hardbottom sites in the mid and outer 

shelf zones and softbottom sites in close proximity to 

colonized hardbottom (Figures 3.114b and 3.115b).

 Great barracuda (Sphyraena barracuda)

a)

S. barracuda abundance (100 m2)

0

1 - 7

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

b)

S. barracuda biomass (g/100 m2)

0

0.1 - 500

500.1 - 1000

1000.1 - 5000

>5000

Benthic HABITAT

Coral reef and colonized hardbottom

Submerged vegetation

Unconsolidated sediments

Figure 3.114. Maps of the interpolated (left map) and spatial (right map) distributions for great barracuda (S. barracuda): (a) abundance 

and (b) biomass.

0

0.2

0.4

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. barracuda biomass (kg/100 m2)

0

0.8

1.6

Colonized 

Hardbottom

Macroalgae/ 

Seagrass

Unvegetated 

Sediments

Mangrove

Mean S. barracuda density (#/100 m2)

a) b)

Figure 3.115. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for great 

barracuda (S. barracuda).

Habitat type Habitat type
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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131 3 - Fish Communities

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

B. vetula Frequency (%)

Juveniles/subadults

0

25

50

<5 5-10 10-15 15-20 20-25 25-30 30-35 >35

S. barracuda Frequency (%)

Size class (cm)

a) b)

Figure 3.116. Size frequency histogram for (a) queen triggerfish (B. vetula) and (b) great barracuda (S. barracuda) in the southwest Puerto 

Rico study area.

Size class (cm)

The largest proportion of the B. vetula population in the study area were subadults and small adults, 

with only a small proportion (5%) of the total comprising small juveniles and large adults were very 

infrequent (Figure 3.116a). The largest individual was estimated at 40 cm FL. 

S. barracuda reach sexual maturity at a minimum size of 66 cm FL, therefore juveniles represented 

the majority of the population surveyed in the study (Figure 3.116b). The maximum known size for the 

species is 200 cm TL and the largest in the study area was 150 cm FL.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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132 3 - Fish Communities3.4. Inter-annual trend in fish metrics (2001-2007)

Presented in this section is a synoptic overview of inter-annual changes in summary statistics (mean 

± SE) for 81 fish metrics at the level of species, family, trophic and community using data collected 

from the southwest Puerto Rico study area from 2001-2007. Between-year comparisons were tested 

for statistical significance using a non-parametric test (see Methods). The following results are based 

on values obtained from Table 3.5, where density values are reported as # individuals/100m2

, biomass 

reported in grams/100m2

, and richness (community metrics section only) reported as # species/100m2.

Table 3.5. Density and biomass (mean + SE) for selected fish species and families (2001-2007) for the southwest Puerto Rico study 

area. Colored text indicate significant (p<0.05) directional change from previous year. BLUE= increasing; RED= decreasing. Asterisk (*) 

indicate there was a significant difference when years were compared.

Fish variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Community metrics

Number of species * 10.41(0.77) 10.64 (0.55) 13.60 (0.58) 12.27 (0.53) 12.91 (0.58) 13.02 (0.61) 13.48 (0.62)

Total density (100 m2

) * 234.46 (46.68) 452.14 (122.44) 262.71 (77.61) 130.39 (29.20) 117.63 (19.02) 181.20 (61.01) 90.24 (9.07)

Biomass (g/100 m2

) * 1566.43 (246.60) 2850.21 (586.55) 2379.88 (275.44) 1661.84 (162.41) 2668.70 (347.93) 2690.22 (404.56) 2394.64 (232.87)

Herbivore density * 28.51 (3.16) 26.31 (2.28) 38.57 (2.64) 27.99 (2.05) 34.15 (2.71) 35.05 (2.73) 36.92 (2.87)

Herbivore biomass * 531.85 (110.30) 785.43 (122.25) 1091.05 (198.65) 830.05 (102.76) 1157.45 (158.35) 1125.46 (136.84) 1219.35 (172.45)

Piscivore density 3.37 (0.57) 4.63 (0.90) 4.30 (0.57) 3.15 (0.43) 4.37 (0.84) 6.72 (1.62) 3.97 (0.78)

Piscivore biomass 348.71 (85.19) 1065.54 (540.54) 546.22 (113.99) 327.79 (62.15) 760.95 (251.86) 527.03 (161.21) 399.50 (92.68)

Planktivore density 177.42 (44.63) 399.60 (121.44) 197.40 (76.74) 83.04 (27.93) 61.91 (17.73) 121.32 (59.80) 31.31 (8.24)

Planktivore biomass 153.94 (39.52) 215.01 (34.93) 210.35 (51.25) 139.65 (30.09) 258.23 (58.91) 188.76 (32.02) 171.17 (40.05)

Groupers - Serranidae

Grouper density * 0.14 (0.05) 0.19 (0.05) 0.32 (0.06) 0.31 (0.06) 0.34 (0.07) 0.44 (0.08) 0.46 (0.08)

Grouper biomass * 45.97 (25.44) 33.62 (11.62) 36.82 (10.30) 59.75 (14.27) 42.59 (10.31) 30.35 (6.43) 44.89 (11.15)

Graysby (C. cruentata) density * 0.07 (0.03) 0.10 (0.03) 0.23 (0.04) 0.18 (0.04) 0.22 (0.05) 0.31 (0.06) 0.31 (0.06)

Graysby (C. cruentata) biomass * 4.04 (2.69) 8.73 (3.15) 15.45 (3.72) 19.13 (5.25) 11.33 (2.88) 13.38 (3.44) 16.40 (4.18)

Coney (C. fulva) density 0.01 (0.01) 0.06 (0.04) 0.07 (0.03) 0.07 (0.03) 0.08 (0.03) 0.08 (0.03) 0.10 (0.05)

Coney (C. fulva) biomass 0.91 (0.91) 11.33 (7.31) 8.50 (4.10) 11.52 (4.84) 16.19 (6.67) 9.42 (3.62) 11.76 (5.29)

Red hind (E. guttatus) density 0.04 (0.02) 0.04 (0.01) 0.02 (0.01) 0.05 (0.02) 0.05 (0.02) 0.02 (0.01) 0.03 (0.01)

Red hind (E. guttatus) biomass 14.11 (7.46) 13.56 (5.60) 12.87 (7.68) 28.64 (11.03) 15.07 (6.27) 5.20 (2.96) 11.11 (4.92)

Snappers - Lutjanidae

Snapper density 2.41 (0.46) 3.38 (0.57) 2.85 (0.39) 2.24 (0.37) 2.00 (0.41) 5.32 (1.52) 3.39 (0.77)

Snapper biomass 162.97 (38.34) 286.98 (51.73) 212.25 (36.24) 145.93 (23.10) 195.71 (45.38) 245.74 (64.60) 209.61 (50.25)

Schoolmaster (L. apodus) density 1.31 (0.34) 2.25 (0.43) 1.80 (0.34) 1.22 (0.27) 1.12 (0.32) 2.22 (0.75) 1.73 (0.67)

Schoolmaster (L. apodus) biomass 90.80 (28.92) 155.65 (37.19) 77.99 (17.70) 59.13 (15.35) 71.26 (26.66) 85.08 (26.12) 82.01 (43.96)

Gray (L. griseus) density 0.27 (0.10) 0.46 (0.19) 0.22 (0.08) 0.49 (0.15) 0.18 (0.09) 1.88 (0.92) 0.76 (0.25)

Gray (L. griseus) biomass 19.51 (7.79) 46.62 (20.43) 40.27 (23.70) 33.29 (11.34) 46.68 (24.74) 92.97 (44.22) 46.26 (17.40)

Dog (L. jocu) density * 0.00 (0.00) 0.00 (0.00) 0.02 (0.01) 0.00 (0.00) 0.01 (0.01) 0.01 (0.01) 0.04 (0.02)

Dog (L. jocu) biomass * 0.00 (0.00) 0.00 (0.00) 6.91 (4.89) 0.00 (0.00) 2.02 (2.02) 1.92 (1.92) 12.87 (6.29)

Mahogany (L. mahogoni) density 0.02 (0.02) 0.04 (0.02) 0.02 (0.01) 0.01 (0.01) 0.14 (0.06) 0.04 (0.02) 0.04 (0.03)

Mahogany (L. mahogoni) biomass 0.20 (0.20) 1.11 (0.68) 5.01 (2.95) 0.80 (0.62) 3.21 (1.24) 3.21 (1.61) 4.64 (3.20)

Lane (L. synagris) density * 0.22 (0.08) 0.09 (0.03) 0.09 (0.03) 0.02 (0.01) 0.04 (0.02) 0.02 (0.01) 0.13 (0.07)

Lane (L. synagris) biomass * 6.57 (4.21) 3.66 (1.50) 7.02 (3.05) 1.49 (1.48) 3.69 (2.92) 0.64 (0.57) 2.65 (1.39)

Yellowtail (O. chrysurus) density * 0.58 (0.16) 0.54 (0.13) 0.66 (0.11) 0.48 (0.09) 0.51 (0.09) 0.97 (0.16) 0.66 (0.12)

Yellowtail (O. chrysurus) biomass * 45.89 (12.72) 79.94 (22.30) 56.42 (13.14) 49.03 (11.74) 68.66 (15.04) 61.56 (10.94) 61.12 (11.04)

Jacks - Carangidae

Jack density 0.88 (0.30) 1.23 (0.71) 1.71 (0.66) 0.75 (0.27) 3.32 (1.77) 2.86 (1.13) 0.28 (0.08)

Jack biomass 68.41 (43.43) 146.55 (75.20) 147.43 (79.48) 92.85 (45.91) 67.72 (28.87) 99.78 (36.61) 41.97 (28.16)

Blue runner (C. crysos) density 0.15 (0.15) 0.10 (0.07) 0.11 (0.08) 0.15 (0.11) 0.01 (0.01) 0.02 (0.01) 0.05 (0.04)

Blue runner (C. crysos) biomass 3.45 (3.45) 43.41 (33.41) 50.78 (39.12) 65.42 (43.22) 2.29 (2.29) 21.24 (17.93) 29.34 (27.25)

Horse-eye (C. latus) density * 0.25 (0.16) 0.01 (0.01) 0.02 (0.02) 0.03 (0.02) 0.02 (0.01) 0.01 (0.01) 0.00 (0.00)

Horse-eye (C. latus) biomass * 18.63 (15.34) 0.43 (0.43) 0.84 (0.66) 0.15 (0.14) 1.05 (0.86) 0.21 (0.21) 0.00 (0.00)

Bar (C. ruber) density 0.27 (0.09) 0.30 (0.12) 0.92 (0.38) 0.34 (0.12) 1.63 (0.66) 1.32 (0.59) 0.21 (0.06)

Bar (C. ruber)) biomass 4.69 (1.58) 11.65 (7.47) 10.78 (2.46) 9.40 (3.73) 24.16 (9.35) 15.40 (6.03) 5.37 (2.76)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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133 3 - Fish Communities

Table 3.5. Continued...

Fish variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Parrotfish - Scaridae

Parrotfish density * 11.54 (1.58) 12.22 (1.26) 19.56 (1.55) 12.40 (1.09) 12.59 (1.76) 15.51 (1.36) 17.47 (1.70)

Parrotfish biomass * 232.37 (50.12) 461.23 (78.82) 568.80 (79.61) 543.23 (71.00) 692.15 (99.02) 679.57 (82.55) 747.38 (129.55)

Striped (S. iseri) density 6.25 (1.20) 5.71 (0.83) 7.27 (0.96) 4.69 (0.63) 5.02 (1.35) 5.98 (0.77) 6.79 (1.34)

Striped (S. iseri) biomass 70.23 (15.88) 95.85 (23.39) 98.67 (17.14) 84.67 (12.00) 101.00 (14.69) 153.73 (25.73) 149.00 (26.30)

Princess (S. taeniopterus) density * 0.19 (0.11) 0.49 (0.16) 1.97 (0.65) 1.45 (0.32) 1.96 (0.41) 2.48 (0.41) 2.89 (0.46)

Princess (S. taeniopterus) biomass * 9.38 (6.85) 31.69 (7.85) 138.51 (53.87) 78.66 (20.46) 111.14 (20.29) 131.64 (24.48) 113.80 (19.11)

Greenblotch (S. atomarium) density * 0.27 (0.10) 0.72 (0.24) 0.73 (0.17) 0.54 (0.09) 0.13 (0.05) 0.30 (0.09) 0.23 (0.09)

Greenblotch (S. atomarium) biomass * 0.24 (0.11) 4.07 (2.48) 0.54 (0.15) 0.56 (0.14) 0.37 (0.31) 0.33 (0.11) 0.36 (0.23)

Redband (S. aurofrenatum) density * 0.90 (0.23) 2.10 (0.32) 2.46 (0.30) 2.65 (0.30) 2.97 (0.32) 4.27 (0.43) 4.08 (0.43)

Redband (S. aurofrenatum) biomass * 44.18 (9.63) 125.50 (22.47) 115.91 (17.71) 151.28 (20.03) 147.26 (19.24) 163.11 (20.92) 132.79 (15.49)

Bucktooth (S. radians) density * 1.90 (0.41) 1.24 (0.22) 3.45 (0.63) 1.29 (0.30) 0.64 (0.19) 0.38 (0.12) 0.99 (0.29)

Bucktooth (S. radians) biomass * 3.44 (0.98) 3.97 (1.00) 8.93 (2.46) 2.74 (1.17) 1.36 (0.71) 0.67 (0.26) 1.16 (0.42)

Stoplight (S. viride) density 1.02 (0.27) 0.99 (0.18) 1.50 (0.25) 1.11 (0.21) 1.03 (0.15) 0.99 (0.17) 1.18 (0.19)

Stoplight (S. viride) biomass 86.59 (28.51) 173.97 (42.13) 160.52 (34.59) 213.44 (43.77) 265.56 (63.36) 148.07 (28.99) 282.30 (83.74)

Grunts - Haemulidae

Grunt density 15.91 (4.03) 14.65 (2.98) 11.70 (2.07) 12.88 (2.98) 13.88 (3.62) 13.34 (5.42) 12.82 (4.75)

Grunt biomass 192.59 (86.01) 285.97 (72.06) 203.07 (62.32) 89.92 (16.28) 150.64 (37.44) 164.42 (40.39) 184.09 (43.56)

Tomtate (H. aurolineatum) density * 0.65 (0.60) 0.09 (0.05) 2.34 (0.90) 2.65 (0.91) 0.18 (0.10) 0.51 (0.29) 5.08 (4.19)

Tomtate (H. aurolineatum) biomass * 2.28 (1.46) 5.33 (2.40) 58.20 (42.48) 10.67 (4.54) 4.73 (2.28) 9.90 (3.52) 30.26 (22.40)

French (H. flavolineatum) density 9.26 (2.95) 7.64 (2.26) 4.18 (1.40) 3.46 (1.31) 4.17 (1.53) 2.11 (0.67) 1.98 (0.57)

French (H. flavolineatum) biomass 123.68 (61.36) 98.45 (29.42) 40.83 (10.95) 36.24 (10.39) 56.95 (22.64) 31.97 (8.51) 34.96 (11.71)

White (H. plumierii) density * 0.35 (0.20) 0.20 (0.06) 1.64 (0.61) 0.65 (0.30) 0.59 (0.17) 0.99 (0.39) 0.44 (0.14)

White (H. plumierii) biomass * 8.86 (3.99) 9.11 (3.83) 32.18 (10.02) 5.50 (2.07) 28.40 (9.87) 67.90 (28.49) 29.22 (7.30)

Bluestriped (H. sciurus) density * 3.33 (0.96) 4.63 (1.08) 2.34 (0.56) 0.83 (0.20) 0.94 (0.37) 0.97 (0.39) 0.99 (0.43)

Bluestriped (H. sciurus) biomass * 52.42 (25.91) 159.56 (52.29) 59.22 (16.26) 28.36 (6.36) 43.58 (19.29) 40.76 (12.72) 66.43 (27.09)

Surgeonfish - Acanthuridae

Surgeonfish density * 2.46 (0.48) 1.91 (0.32) 4.39 (0.71) 2.44 (0.28) 3.13 (0.36) 4.13 (0.66) 3.52 (0.51)

Surgeonfish biomass * 174.42 (61.08) 153.40 (35.74) 369.61 (141.82) 161.71 (26.79) 265.21 (40.17) 309.24 (56.32) 311.40 (64.73)

Ocean surgeon (A. bahianus) density * 1.33 (0.26) 1.09 (0.19) 2.29 (0.27) 1.25 (0.19) 1.89 (0.25) 2.79 (0.41) 1.83 (0.24)

Ocean surgeon (A. bahianus) biomass * 76.42 (23.07) 76.32 (18.87) 121.62 (20.14) 78.49 (13.03) 139.33 (22.76) 201.58 (27.99) 126.72 (17.10)

Doctorfish (A. chirurgus) density * 0.25 (0.09) 0.36 (0.13) 1.01 (0.31) 0.55 (0.12) 0.49 (0.11) 0.25 (0.08) 0.73 (0.29)

Doctorfish (A. chirurgus) biomass * 13.82 (5.88) 39.81 (20.94) 99.84 (39.18) 42.92 (18.93) 38.65 (12.71) 25.15 (10.72) 103.89 (53.90)

Blue tang (A. coeruleus) density * 0.88 (0.36) 0.45 (0.14) 1.05 (0.31) 0.64 (0.13) 0.75 (0.16) 1.07 (0.37) 0.93 (0.25)

Blue tang (A. coeruleus) biomass * 84.17 (53.65) 37.28 (12.44) 148.15 (97.84) 40.29 (8.67) 87.24 (20.37) 82.50 (34.03) 80.80 (23.90)

Other Families

Squirrelfish (Holocentridae) density * 1.85 (0.81) 0.76 (0.18) 0.72 (0.12) 0.54 (0.10) 0.65 (0.09) 1.18 (0.17) 0.97 (0.15)

Squirrelfish (Holocentridae) biomass * 107.47 (50.94) 79.52 (20.56) 68.96 (13.26) 59.90 (10.48) 65.30 (10.20) 99.11 (14.52) 85.95 (18.27)

Wrasse (Labridae) density * 8.87 (3.12) 8.84 (1.50) 11.66 (1.61) 8.18 (1.20) 10.56 (1.65) 13.69 (1.86) 12.67 (1.36)

Wrasse (Labridae) biomass * 50.02 (12.42) 59.96 (10.87) 63.92 (13.48) 28.30 (4.36) 98.59 (44.60) 47.17 (5.83) 64.04 (7.44)

Other species

Queen triggerfish (B. vetula) density * 0.03 (0.02) 0.05 (0.02) 0.02 (0.01) 0.02 (0.01) 0.03 (0.01) 0.12 (0.03) 0.18 (0.05)

Queen triggerfish (B. vetula) biomass * 18.46 (10.98) 26.60 (14.48) 7.62 (5.21) 4.18 (2.40) 22.05 (15.27) 48.81 (13.38) 71.73 (18.08)

Yellow goatfish (M. martinicus) density 0.54 (0.33) 0.14 (0.05) 0.13 (0.05) 0.08 (0.03) 0.19 (0.08) 0.30 (0.18) 0.71 (0.56)

Yellow goatfish (M. martinicus) biomass 40.64 (22.33) 19.28 (7.33) 28.48 (13.10) 7.48 (4.19) 31.44 (19.12) 17.20 (6.91) 31.72 (15.21)

Spotted goatfish (P. maculatus) density 0.28 (0.07) 0.74 (0.37) 0.36 (0.12) 0.13 (0.03) 0.31 (0.08) 0.33 (0.06) 0.29 (0.05)

Spotted goatfish (P. maculatus) biomass 13.55 (4.05) 29.82 (10.99) 18.29 (5.49) 5.57 (1.77) 17.23 (4.85) 13.72 (2.98) 16.89 (3.76)

Overall Comparisons

None of the fish metrics decreased for more than three consecutive years within the seven year monitoring 

period (2001-2007). Fifteen fish metrics (approximately 19%) exhibited at least three consecutive years 

of decline and 44 fish metrics (approximately 54%) exhibited at least two consecutive years of decline. 

All but one of the 15 metrics exhibiting a three year consecutive decline occurred prior to 2006. In 

contrast, 13 of the 15 metrics exhibiting a three year consecutive increase occurred from 2004-2007. 

Two fish metrics (C. fulva density and S. aurofrenatum density) exhibited more than three years of 

consecutive increase. Mean C. fulva density increased very slightly every year from 2001-2007, but 

was not statistically significant. Mean S. aurofrenatum density increased for five consecutive years from 

2001-2006 and although not greater in 2007 than 2006, mean density remained significantly higher 

than it was in years 2001 and 2002.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

134 3 - Fish CommunitiesA total of 65 (approximately 80%) metrics decreased 

from 2003 to 2004, five of which decreased 

significantly (p<0.05); followed by 70% of the metrics 

increasing the following year (2004-2005), with no 

significant increase. There were 108 (approximately 

62%) significant (p<0.05) increases as time 

progressed and 66 (approximately 38%) significant 

decreases. The highest number of significant 

differences (n=19) was found when comparing fish 

metrics between the first two years and the last year 

of the monitoring period between 2001 and 2007 and 

2002 and 2007. Fourteen metrics were significantly 

higher in 2007 than 2001 and five were significantly 

lower; and 16 metrics were significantly higher in 

2007 than 2002 and three were significantly lower. 

A summarized subset of metrics from Table 3.5 with 

significant differences in 2001/2007 and 2002/2007 

comparisons are shown in Tables 3.6 and 3.7. 

S. radians, A. chirurgus, H. flavolineatum and 

squirrelfish (Holocentridae) were the only metrics 

to have three consecutive years of decline for both 

metrics of mean density and biomass. S. radians

and A. chirurgus declined from 2003-2006 and H. 

flavolineatum and squirrelfish from 2001-2003. The 

other metrics that exhibited three years of consecutive 

decline include; total fish, planktivore, total snapper 

and L. apodus mean density from 2002-2005; and S. 

radians density and biomass, S. viride density and A. 

chirurgus mean biomass from 2003-2006.

3.4.1. Fish community metrics

A summarized subset of community metrics from Table 3.5 is presented in Table 3.8. 

Mean species richness increased from 2001-2003 followed by a decline in 2003-2004 and then 

increased for a period of three consecutive years (2004-2007). Mean species richness was significantly 

(p=0.0003) lower in 2001 than 2003 and 2007, and in 2002 than 2007. Mean fish density (all species 

combined) was highest in 2002 and lowest in 2007 and average total fish biomass was lowest in 2001 

Significantly higher in 2001 Significantly higher in 2007

Grouper biomass Species richness

Sparisoma radians abundance Total biomass

Sparisoma radians biomass Herbivore biomass

Caranx latus abundance Grouper abundance

Caranx latus biomass Cephalopholis cruentata abundance

Cephalopholis cruentata biomass

Wrasse abundance

Parrotfish abundance

Parrotfish biomass

Scarus taeniopterus biomass

Sparisoma aurofrenatum abundance

Sparisoma aurofrenatum biomass

Balistes vetula abundance

Balistes vetula biomass

Table 3.6. Inter-annual metrics with significant differences 

(p<0.05) in 2001 and 2007 comparisons.

Significantly higher in 2002 Significantly higher in 2007

Haemulon sciurus abundance Total species richness

Haemulon sciurus biomass Grouper abundance

Sparisoma radians biomass Cephalopholis cruentata abundance

Cephalopholis cruentata biomass

Lutjanus jocu abundance

Lutjanus jocu biomass

Scarus taeniopterus abundance

Scarus taeniopterus biomass

Sparisoma aurofrenatum abundance

Sparisoma aurofrenatum biomass

Surgeonfish abundance

Surgeonfish biomass

Acanthurus bahianus biomass

Wrasse abundance

Balistes vetula abundance

Balistes vetula biomass

Table 3.7. Inter-annual metrics with significant differences 

(p<0.05) in 2002 and 2007 comparisons.

Community variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Number of species * 10.4 (0.8) 10.6 (0.6) 13.6 (0.6) 12.3 (0.5) 12.9 (0.6) 13.0 (0.6) 13.5 (0.6)

Total density (100 m2

) * 234.5 (46.7) 452.1 (122.4) 262.7 (77.6) 130.4 (29.2) 117.6 (19.0) 181.2 (61.0) 90.2 (9.1)

Biomass (g/100 m2

) * 1,566.4 (246.6) 2,850.2 (586.6) 2,379.9 (275.4) 1,661.8 (162.4) 2,668.7 (347.9) 2,690.2 (404.6) 2,394.6 (232.9)

Herbivore density * 28.5 (3.2) 26.3 (2.3) 38.6 (2.6) 28.0 (2.1) 34.1 (2.7) 35.1 (2.7) 36.9 (2.9)

Herbivore biomass * 531.9 (110.3) 785.4 (122.3) 1,091.0 (198.6) 830.0 (102.8) 1,157.5 (158.3) 1,125.5 (136.8) 1,219.3 (172.5)

Piscivore density 3.4 (0.6) 4.6 (0.9) 4.3 (0.6) 3.1 (0.4) 4.4 (0.8) 6.7 (1.6) 4.0 (0.8)

Piscivore biomass 348.7 (85.2) 1,065.5 (540.5) 546.2 (114.0) 327.8 (62.2) 760.9 (251.9) 527.0 (161.2) 399.5 (92.7)

Planktivore density 177.4 (44.6) 399.6 (121.4) 197.4 (76.7) 83.0 (27.9) 61.9 (17.7) 121.3 (59.8) 31.3 (8.2)

Planktivore biomass 153.9 (39.5) 215.0 (34.9) 210.4 (51.3) 139.6 (30.1) 258.2 (58.9) 188.8 (32.0) 171.2 (40.0)

Table 3.8. Density and biomass (mean + SE) for community metrics (2001-2007) for the southwest Puerto Rico study area. Asterisks (*) 

indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up from Table 3.5 

where appropriate. Bold indicates either the lowest or highest mean for that metric.
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135 3 - Fish Communities

and highest the following year in 2002. Neither total fish density nor biomass had a significant difference 

between years of high and low means.

Mean density of herbivores exhibited three consecutive years of increase (2004-2007) and mean 

biomass of herbivores increased over two consecutive years (2001-2002) and then exhibited alternating 

directional change from 2002-2007. Mean herbivore density recorded from 2001-2007 was lowest in 

2002 (26.31 ± 2.28) and highest the following year in 2003 (38.57 ± 2.64). Mean biomass was lowest 

in 2001 (531.85 ± 110.30) and highest in 2007 (1219.35 ± 172.45). The high and low means for both 

herbivore metrics were significantly different (p<0.01).

Both mean piscivore density and biomass recorded from 2001-2007 was lowest in 2004, whereas 

mean density was highest in 2006 and mean biomass in 2002. Mean planktivore density was highest 

in 2002 and lowest in 2007 and mean biomass was lowest in 2004 and highest the following year in 

2005. Neither piscivore nor planktivore metrics were significantly different between years of highest and 

lowest.

 Checkered puffer (Sphoeroides testudineus) Squirrelfish (Holocentrus adscensionis) Blackbar soldierfish (Myripristis jacobus)

3.4.2. Taxonomic groups 

Large-body Groupers (Serranidae)

The species in this section include all large-bodied species from the Serranidae Family, Subfamily 

Epinephelinae (Cephalopholis, Epinephelus, Mycteroperca genus) that were recorded in the southwest 

Puerto Rico study area from 2001-2007. Grouper species included in the ‘total grouper’ metrics include 

C. cruentata, C. fulva, Epinephelus adscensionis (rock hind), E. guttatus and Epinephelus striatus

(Nassau grouper). The mean density and biomass for the most common grouper species recorded/

observed in the study area are summarized in Table 3.5 and subset Table 3.9.

Grouper variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Grouper (Serranidae) density * 0.1 (0.05) 0.2 (0.05) 0.3 (0.1) 0.3 (0.1) 0.3 (0.1) 0.4 (0.1) 0.5 (0.1)

Grouper (Serranidae) biomass * 46.0 (25.4) 33.6 (11.6) 36.8 (10.3) 59.8 (14.3) 42.6 (10.3) 30.4 (6.4) 44.9 (11.2)

Graysby (C. cruentata) density * 0.1 (0.03) 0.1 (0.03) 0.2 (0.04) 0.2 (0.04) 0.2 (0.05) 0.3 (0.1) 0.3 (0.1)

Graysby (C. cruentata) biomass * 4.0 (2.7) 8.7 (3.2) 15.5 (3.7) 19.1 (5.3) 11.3 (2.9) 13.4 (3.4) 16.4 (4.2)

Coney (C. fulva) density <0.01 (<0.01) 0.1 (0.04) 0.1 (0.03) 0.1 (0.03) 0.1 (0.03) 0.1 (0.03) 0.1 (0.05)

Coney (C. fulva) biomass 0.9 (0.9) 11.3 (7.3) 8.5 (4.1) 11.5 (4.8) 16.2 (6.7) 9.4 (3.6) 11.8 (5.3)

Red hind (E. guttatus) density 0.04 (0.02) 0.04 (0.01) 0.02 (0.01) 0.05 (0.02) 0.05 (0.02) 0.02 (0.01) 0.03 (0.01)

Red hind (E. guttatus) biomass 14.1 (7.5) 13.6 (5.6) 12.9 (7.7) 28.6 (11.0) 15.1 (6.3) 5.2 (3.0) 11.1 (3.9)

Table 3.9. Density and biomass (mean + SE) for selected grouper species (2001-2007) for the southwest Puerto Rico study area. 

Asterisks (*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up 

from Table 3.5 where appropriate. Bold indicates either the lowest or highest mean for that metric.

Only mean density and mean biomass for all groupers and C. cruentata exhibited significant differences 

between years. Mean density and biomass for C. cruentata were significantly (p<0.01) lower in 2001 

and 2002 than 2007. Mean density of total groupers was significantly lower (p=0.0046) in 2001 (0.14 ± 

0.05) than 2007 (0.46 ± 0.08), whereas total grouper biomass was significantly (p<0.05) higher in 2001 
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136 3 - Fish Communitiesthan 2007. All of the grouper metrics (except C. cruentata density) 

increased from 2006-2007, although not statistically significant. Four 

grouper metrics (C. cruentata and C. fulva density and biomass, and 

total grouper density) were lowest in 2001; and four grouper metrics 

(total grouper and C. cruentata biomass, and E. guttatus density and 

biomass) had their highest means in 2004.

Mean grouper biomass was the only grouper metric that had 

consecutive years of increases and decreases and the only grouper 

metric to decrease significantly (p=0.0164) when comparing the start 

(2001) and finish of the monitoring data set (2007). Mean density and 

biomass of C. cruentata were significantly (p<0.01) higher in 2006 and 

2007 than in 2001; and significantly (p<0.01) higher in 2007 than 2002.

Mean density of C. fulva increased across all years from 2001-2007, 

although the consecutive change through years were not significant. 

The average C. fulva density recorded from 2001-2007 was lowest in 

2001 and the highest in 2007, and the average biomass was lowest in 

2001 and the highest in 2005. Neither metric had a significant difference 

between years of mean high or low. 

Mean density and biomass of E. guttatus did not exhibit any consecutive years of increase between 

2001 and 2007, but did show periods with two consecutive years of decrease. Mean density was lowest 

in 2003 and highest in 2003. Mean biomass was highest in 2004 and lowest in 2006. Neither metric had 

a significant difference between years of mean high or low.

Snappers (Lutjanidae)

Snapper metrics included all species from the Lutjanidae family that were recorded in the southwest 

Puerto Rico study area from 2001-2007. The snapper species included in the ‘total snapper’ metrics 

were mutton snapper (Lutjanus analis), L. apodus, blackfin snapper (Lutjanus buccanella), L. griseus, 

dog snapper (Lutjanus jocu), L. mahogoni, L. synagris, O. chrysurus and unidentified Lutjanus species. 

The mean density and biomass for the most common snapper species recorded/observed in the study 

area are summarized in Table 3.5 and subset Table 3.10.

E. guttatus

C. fulva

Snapper variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Snapper (Lutjanidae) density 2.4 (0.5) 3.4 (0.6) 2.8 (0.4) 2.2 (0.4) 2.0 (0.4) 5.3 (1.5) 3.4 (0.8)

Snapper (Lutjanidae) biomass 163.0 (38.3) 287.0 (51.7) 212.3 (36.2) 145.9 (23.1) 195.7 (45.4) 245.7 (64.6) 209.6 (50.2)

Lutjanus apodus density 1.3 (0.3) 2.3 (0.4) 1.8 (0.3) 1.2 (0.3) 1.1 (0.3) 2.2 (0.8) 1.7 (0.7)

Lutjanus apodus biomass 90.8 (28.9) 155.7 (37.2) 78.0 (17.7) 59.1 (15.4) 71.3 (26.7) 85.1 (26.1) 82.0 (44.0)

Lutjanus griseus density 0.3 (0.1) 0.5 (0.2) 0.2 (0.1) 0.5 (0.2) 0.2 (0.1) 1.9 (0.9) 0.8 (0.2)

Lutjanus griseus biomass 19.5 (7.8) 46.6 (20.4) 40.3 (23.7) 33.3 (11.3) 46.7 (24.7) 93.0 (44.2) 46.3 (17.4)

Lutjanus jocu density * 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) <0.01(<0.01) <0.01(<0.01) 0.04 (0.02)

Lutjanus jocu biomass * 0.0 (0.0) 0.0 (0.0) 6.9 (4.9) 0.0 (0.0) 2.0 (2.0) 1.9 (1.9) 12.9 (6.3)

Lutjanus mahogoni density 0.02 (0.02) 0.04 (0.02) 0.02 (0.01) 0.01 (<0.01) 0.1 (0.1) 0.04 (0.02) 0.04 (0.03)

Lutjanus mahogoni biomass 0.2 (0.2) 1.1 (0.7) 5.0 (3.0) 0.8 (0.6) 3.2 (1.2) 3.2 (1.6) 4.6 (3.2)

Lutjanus synagris density * 0.2 (0.1) 0.1 (0.03) 0.1 (0.03) 0.02 (0.01) 0.04 (0.02) 0.02 (0.01) 0.1 (0.1)

Lutjanus synagris biomass * 6.6 (4.2) 3.7 (1.5) 7.0 (3.0) 1.5 (1.5) 3.7 (2.9) 0.6 (0.6) 2.7 (1.4)

Ocyurus chrysurus density * 0.6 (0.2) 0.5 (0.1) 0.7 (0.1) 0.5 (0.1) 0.5 (0.1) 1.0 (0.2) 0.7 (0.1)

Ocyurus chrysurus biomass * 45.9 (12.7) 79.9 (22.3) 56.4 (13.1) 49.0 (11.7) 68.7 (15.0) 61.6 (10.9) 61.1 (11.0)

Table 3.10. Density and biomass (mean + SE) for selected snapper species (2001-2007) for the southwest Puerto Rico study area. 

Asterisks (*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up 

from Table 3.5 where appropriate. Bold indicates either the lowest or highest mean for that metric. 
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137 3 - Fish Communities

The only snapper metric to exhibit three consecutive years of increase 

was L. mahogoni biomass from 2004-2007. Mean total snapper density 

was highest in 2006 and lowest in 2005. Mean biomass for total snapper 

was highest in 2002 and lowest in 2003. Mean density of L. apodus was 

highest in 2002 and lowest in 2005 and mean biomass was highest in 

2002 and lowest in 2003. Mean density of L. griseus was highest in 2006 

and lowest in 2005 and mean biomass was highest in 2006 and lowest 

in 2001. Mean density of L. mahogoni was highest in 2005 and lowest in 

2004 and the average biomass was lowest in 2001 and the highest in 2003. None of these metrics had 

a significant difference between years of mean high or low.

Mean density and biomass of L. jocu was lowest in 2001, 2002 and 2004 

(0 ± 0 and 0 ± 0, respectively) and significantly highest in 2007 (0.04 

± 0.02 for density and 12.87 ± 6.289 for biomass). Mean density and 

biomass of L. synagris were significantly (p<0.01) higher in 2001 than 

2004 and 2006. Mean L. synagris density was highest in 2001 (0.22 ± 

0.08) and lowest in 2006 (0.02 ± 0.01) and mean biomass was highest 

in 2003 (7.02 ± 3.05) and lowest in 2006 (0.64 ± 0.57). Both L. synagris

and L. jocu metrics were significantly different (p<0.01) between years of 

high and low.

Mean biomass of O. chrysurus decreased from 2002-2004 and again 

from 2005-2007 after a brief increase from 2004-2005. Mean density of O. 

chrysurus was significantly (p<0.05) lower in 2002 than 2006; whereas, 

mean biomass was lower in 2002 than 2006 although not significant. 

Mean density of O. chrysurus was lowest in 2004 and highest in 2006; 

and mean biomass was lowest in 2001 and highest the following year in 

2002. None of these metrics had a significant difference between years 

of mean high or low.

Jacks (Carangidae)

Jacks include all species from the Carangidae family that were recorded in the southwest Puerto 

Rico study area from 2001-2007. The jack species included in the ‘total jack’ metrics are yellow jack 

(Carangoides bartholomaei), C. ruber, C. crysos, horse-eye jack (Caranx latus), black jack (Caranx 

lugubris), and unidentified Carangoides/Caranx species. The mean density and biomass for the most 

common jack species recorded/observed in the study area are summarized in Table 3.5 and subset 

Table 3.11.

L. mahogoni in St. Croix, USVI

Dog snapper (Lutjanus jocu) in St. John, USVI

O. chrysurus

Jack variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Jack (Carangidae) density 0.9 (0.3) 1.2 (0.7) 1.7 (0.7) 0.8 (0.3) 3.3 (1.8) 2.9 (1.1) 0.3 (0.1)

Jack (Carangidae) biomass 68.4 (43.4) 146.6 (75.2) 147.4 (79.5) 92.9 (45.9) 67.7 (28.9) 99.8 (36.6) 42.0 (28.2)

Caranx crysos density 0.1 (0.1) 0.1 (0.1) 0.1 (0.1) 0.2 (0.1) <0.01 (<0.01) 0.0 (0.0) 0.1 (0.0)

Caranx crysos biomass 3.5 (3.5) 43.4 (33.4) 50.8 (39.1) 65.4 (43.2) 2.3 (2.3) 21.2 (17.9) 29.3 (27.2)

Caranx latus density * 0.2 (0.2) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0)

Caranx latus biomass * 18.6 (15.3) 0.4 (0.4) 0.8 (0.7) 0.1 (0.1) 1.1 (0.9) 0.2 (0.2) 0.0 (0.0)

Carangoides ruber density 0.3 (0.1) 0.3 (0.1) 0.9 (0.4) 0.3 (0.1) 1.6 (0.7) 1.3 (0.6) 0.2 (0.1)

Carangoides ruber biomass 3.7 (1.6) 11.6 (7.5) 10.8 (2.5) 9.4 (3.7) 23.2 (9.4) 15.4 (6.0) 5.4 (2.8)

Table 3.11. Density and biomass (mean + SE) for selected jack species (2001-2007) for the southwest Puerto Rico study area. Asterisks 

(*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up from Table 

3.5 where appropriate. Bold indicates either the lowest or highest mean for that metric.
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138 3 - Fish CommunitiesMean density of total jacks was highest in 2005 and lowest in 2007; and the average biomass was 

highest in 2003 and the lowest in 2007. Mean density of C. ruber was highest in 2005 and lowest 

in 2007 and mean biomass was highest in 2003 and lowest in 2002. None of these metrics had a 

significant difference between years of mean high or low.

Mean density and biomass of C. crysos were the only jack metrics to 

increase from 2006-2007 and all others decreased. These two metrics 

were also the only jack metrics that did not have consecutive years 

of decrease, with mean biomass of C. crysos showing three years 

of consecutive increase (2001-2004). Mean density and biomass of 

C. crysos was highest in 2004 and lowest the following year in 2005. 

Neither metric had a significant difference between years of mean 

high or low.

Mean density of C. latus exhibited three years of consecutive decrease (2004-2007). C. latus density 

and biomass were significantly higher in 2001 than 2002, 2006 and 2007. Mean density and biomass 

were highest in 2001 (0.25 ± 0.16 and 18.63 ± 15.34, respectively) and the lowest in 2007 (0 ± 0 and 

0 ± 0, respectively). Both metrics had a significant difference (p=0.0157) between years of mean high 

or low.

Parrotfishes (Scaridae)

Parrotfish metrics included all species from the Scaridae family that were recorded in the southwest 

Puerto Rico study area from 2001 to 2007. The parrotfish species included in the ‘total parrotfish’ 

metrics were bluelip parrotfish (Cryptotomus roseus), rainbow parrotfish (Scarus guacamaia), S. iseri, 

S. taeniopterus, queen parrotfish (Scarus vetula), greenblotch parrotfish (Sparisoma atomarium), S. 

aurofrenatum, redtail parrotfish (Sparisoma chrysopterum), S. radians, S. rubripinne, S. viride, and 

unidentified Scarus and Sparisoma species. The mean density and biomass for the most common 

parrotfish species recorded/observed in the study area are summarized in Table 3.5 and subset Table 

3.12.

Fish diver and C. ruber

Parrotfish variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Parrotfish (Scaridae) density * 11.5 (1.6) 12.2 (1.3) 19.6 (1.5) 12.4 (1.1) 12.6 (1.8) 15.5 (1.4) 17.5 (1.7)

Parrotfish (Scaridae) biomass * 232.4 (50.1) 461.2 (78.8) 568.8 (79.6) 543.2 (71.0) 692.1 (99.0) 679.6 (82.6) 747.4 (129.5)

Scarus iseri density 6.3 (1.2) 5.7 (0.8) 7.3 (1.0) 4.7 (0.6) 5.0 (1.4) 6.0 (0.8) 6.8 (1.3)

Scarus iseri biomass 70.2 (15.9) 95.8 (23.4) 98.7 (17.1) 84.7 (12.0) 101.0 (14.7) 153.7 (25.7) 149.0 (26.3)

Scarus taeniopterus density * 0.2 (0.1) 0.5 (0.2) 2.0 (0.6) 1.4 (0.3) 2.0 (0.4) 2.5 (0.4) 2.9 (0.5)

Scarus taeniopterus biomass * 9.4 (6.8) 31.7 (7.9) 138.5 (53.9) 78.7 (20.5) 111.1 (20.3) 131.6 (24.5) 113.8 (19.1)

Sparisoma atomarium density * 0.3 (0.1) 0.7 (0.2) 0.7 (0.2) 0.5 (0.1) 0.1 (0.1) 0.3 (0.1) 0.2 (0.1)

Sparisoma atomarium biomass * 0.2 (0.1) 4.1 (2.5) 0.5 (0.2) 0.6 (0.1) 0.4 (0.3) 0.3 (0.1) 0.4 (0.2)

Sparisoma aurofrenatum density * 0.9 (0.2) 2.1 (0.3) 2.5 (0.3) 2.7 (0.3) 3.0 (0.3) 4.3 (0.4) 4.1 (0.4)

Sparisoma aurofrenatum biomass * 44.2 (9.6) 125.5 (22.5) 115.9 (17.7) 151.3 (20.0) 147.3 (19.2) 163.1 (20.9) 132.8 (15.5)

Sparisoma radians density * 1.9 (0.4) 1.2 (0.2) 3.5 (0.6) 1.3 (0.3) 0.6 (0.2) 0.4 (0.1) 1.0 (0.3)

Sparisoma radians biomass * 3.4 (1.0) 4.0 (1.0) 8.9 (2.5) 2.7 (1.2) 1.4 (0.7) 0.7 (0.3) 1.2 (0.4)

Sparisoma viride density 1.0 (0.3) 1.0 (0.2) 1.5 (0.2) 1.1 (0.2) 1.0 (0.2) 1.0 (0.2) 1.2 (0.2)

 Sparisoma viride biomass 86.6 (28.5) 174.0 (42.1) 160.5 (34.6) 213.4 (43.8) 265.6 (63.4) 148.1 (29.0) 282.3 (83.7)

Table 3.12. Density and biomass (mean + SE) for selected parrotfish species (2001-2007) for the southwest Puerto Rico study area. 

Asterisks (*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up 

from Table 3.5 where appropriate. Bold indicates either the lowest or highest mean for that metric.

Of all metrics analyzed, the parrotfish species exhibited the most significant difference for 

between year comparisons. The majority of the lowest means for parrotfish metrics occurred 

in 2001; whereas the majority of the highest means occurred in 2003. All parrotfish metrics, 
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139 3 - Fish Communities

except mean density of S. iseri and S. viride increased from 2001-2002. 

Mean density of total parrotfish increased from 2001-2003 followed by a 

significant (p<0.0001) decrease in 2004, and then increased consecutively 

until 2007, albeit not statistically significant. It is worthy to note that the 

mean density for total parrotfish in 2007 was lower than the mean density 

in 2003, before the significant decrease in 2003. Mean biomass was 

significantly (p<0.0001) lower in 2001 than all year from 2003-2007; and 

lower in 2002 than 2003. Mean density of total parrotfish was lowest in 

2001 (11.54 ± 1.58) and the highest in 2003 (19.56 ± 1.55); and mean biomass was lowest in 2001 

(232.37 ± 50.12) and highest in 2007 (747.38 ± 129.55). Both metrics had a significant difference 

(p<0.0001) between years of mean high or low.

Mean density of S. iseri exhibited three years of consecutive increase 

from 2004 to 2007, with highest density in 2003 and lowest the following 

year in 2003. Mean biomass was lowest in 2001 and the highest in 

2006. Neither metric had a significant difference between years of 

mean high or low. Mean density of S. taeniopterus exhibited three 

consecutive years of increase (2004 to 2007) and one of the highest 

number (n=10) of significantly different (p<0.0001) year comparisons 

indicative of high variability. Mean density of S. taeniopterus was 

highest in 2007 (2.89 ± 0.46) and significantly (p<0.0001) lowest in 

2001 (0.19 ± 0.11). Mean biomass of S. taeniopterus exhibited two consecutive years of increase, 

although neither were significant; however, mean biomass S. taeniopterus also had one of the highest 

number (n=11) of significantly different (p<0.0001) year comparisons. Mean biomass was lowest in 

2001 and highest in 2003 and not significantly different. Mean density and biomass of S. taeniopterus

was significantly (p<0.0001) higher between 2005 and 2007 than 2001 and 2002, but mean biomass 

was significantly (p<0.0001) higher in 2003 than all years from 2005-2007.

Mean density of S. atomarium was the only parrotfish metric that had two years of consecutive 

increase (2001-2003), followed by two years of decrease (2003-2005). Mean density and biomass of 

S. atomarium was significantly (p<0.0001) higher in 2004 than 2001, 2006, and 2007; and with 2003 

significantly higher than 2005 and 2007. Mean density of S. atomarium was highest in 2003 (0.73 ± 

0.17) and significantly (p<0.0001) lowest in 2005 (0.13 ± 0.05). Mean biomass was lowest in 2001 and 

highest the following year in 2002, and not significantly different.

Mean density of S. aurofrenatum increased from 2001-2006, but was not significantly different between 

years, whereas no consecutive years of change were found for mean biomass of S. aurofrenatum. Mean 

S. aurofrenatum density and biomass were highest in 2006 (3.27 ± 0.43 and 

163.11 ± 20.92, respectively) and significantly (p=<0.0001) lowest in 2001 

(0.90 ± 0.23 and 43.18 ± 9.63, respectively). Mean density and biomass of S. 

aurofrenatum were significantly higher from 2003 to 2007 than in 2001 and 

lower in 2002 than 2006 and 2007.

Mean density and biomass of S. radians decreased over three consecutive 

years from 2003-2006, with 2003-2004 exhibiting the only significant 

difference (p<0.0001). Mean biomass of S. radians exhibited the highest 

number (n=12) of significantly different (p<0.0001) year comparisons. S. 

radians density and biomass were significantly higher in 2001 and 2003 than all years from 2004-2007, 

with highest mean density and biomass recorded in 2003 (3.45 ± 0.63 and 8.93 ± 2.46, respectively) 

and significantly lowest in 2006 (0.38 ± 0.12 and 0.67 ± 0.26, respectively). 

Juvenile phase S. taeniopterus

Juvenile phase S. iseri

Terminal phase S. aurofrenatum
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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140 3 - Fish CommunitiesMean density of S. viride decreased over three consecutive years from 2003-2006 although the 

difference was not significant. From 2001-2005, mean density and biomass of S. viride showed an 

opposite directional change (i.e., when density increased, biomass decreased and vice versa). Mean 

density of S. viride was highest in 2003 and lowest in 2002 and mean biomass was highest in 2007 and 

lowest in 2001. Neither metric had a significant difference between years of mean high or low.

Haemulidae (Grunts)

Grunt metrics include all species from the Haemulidae family that were recorded in the southwest 

Puerto Rico study area from 2001-2007. The grunt species included in the ‘total grunt’ metrics were 

porkfish (Anisotremus virginicus), H. aurolineatum, caesar grunt (Haemulon carbonarium), smallmouth 

grunt (Haemulon chrysargyreum), H. flavolineatum, cottonwick (Haemulon melanurum), sailors choice 

(Haemulon parra), H. plumierii, H. sciurus and unidentified Haemulon species (most of which are 

juveniles). The mean density and biomass for the most common grunt species recorded/observed in 

the study area are summarized in Table 3.5 and subset Table 3.13.

Grunt variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Grunt (Haemulidae) density 15.9 (4.0) 14.6 (3.0) 11.7 (2.1) 12.9 (3.0) 13.9 (3.6) 13.3 (5.4) 12.8 (4.7)

Grunt (Haemulidae) biomass 192.6 (86.0) 286.0 (72.1) 203.1 (62.3) 89.9 (16.3) 150.6 (37.4) 164.4 (40.4) 184.1 (43.6)

Haemulon aurolineatum density * 0.6 (0.6) 0.1 (0.0) 2.3 (0.9) 2.7 (0.9) 0.2 (0.1) 0.5 (0.3) 5.1 (4.2)

Haemulon aurolineatum biomass * 2.3 (1.5) 5.3 (2.4) 58.2 (42.5) 10.7 (4.5) 4.7 (2.3) 9.9 (3.5) 30.3 (22.4)

Haemulon flavolineatum density 9.3 (2.9) 7.6 (2.3) 4.2 (1.4) 3.5 (1.3) 4.2 (1.5) 2.1 (0.7) 2.0 (0.6)

Haemulon flavolineatum biomass 123.7 (61.4) 98.4 (29.4) 40.8 (10.9) 36.2 (10.4) 56.9 (22.6) 32.0 (8.5) 35.0 (11.7)

Haemulon plumierii density * 0.4 (0.2) 0.2 (0.1) 1.6 (0.6) 0.7 (0.3) 0.6 (0.2) 1.0 (0.4) 0.4 (0.1)

Haemulon plumierii biomass * 8.9 (4.0) 9.1 (3.8) 32.2 (10.0) 5.5 (2.1) 28.4 (9.9) 67.9 (28.5) 29.2 (7.3)

Haemulon sciurus density * 3.3 (1.0) 4.6 (1.1) 2.3 (0.6) 0.8 (0.2) 0.9 (0.4) 1.0 (0.4) 1.0 (0.4)

Haemulon sciurus biomass * 52.4 (25.9) 159.6 (52.3) 59.2 (16.3) 28.4 (6.4) 43.6 (19.3) 40.8 (12.7) 66.4 (27.1)

Table 3.13. Density and biomass (mean + SE) for selected grunt species (2001-2007) for the southwest Puerto Rico study area. Asterisks 

(*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up from Table 

3.13 where appropriate. Bold indicates either the lowest or highest mean for that metric.

Of the selected grunt metrics, all of the individual species means, except total grunt metrics, exhibited 

a significant difference between years. Mean H. aurolineatum density and H. plumierii biomass were 

the only grunt metrics that did not have any consecutive years of decline. Overall, 2004 had the lowest 

means. Six of the metrics that decreased from 2003-2004, increased the following year (2004-2005).

Total grunt mean density was highly variable with several periods of two consecutive years of 

unidirectional change, which decreased from 2001-2003, increased from 2003-2005, then decreased 

from 2005-2007. None of these changes in the mean were statistically significant. Mean density of total 

grunt was highest in 2001 and lowest in 2003; and mean biomass was highest in 2002 and lowest in 

2003. Neither metric had a significant difference between years of mean high or low. The only times 

where mean total grunt density and biomass had the same directional change were from 2002-2003 

and 2004-2005. Total grunt biomass had three consecutive years of increasing means (2004-2007), 

although the changes were not significant. 

H. aurolineatum Smallmouth grunt (Haemulon chysargyreum) Juvenile grunts
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141 3 - Fish Communities

Mean biomass of H. aurolineatum exhibited three periods 

of two consecutive years of change, which increased from 

2001-2003, decreased from 2003-2005, then increased 

again from 2005-2007. Only the change from 2002-2003 was 

significant (p=0.0024). Additionally, mean H. aurolineatum

density increased significantly (p=0.0011) from 2002-2003 

and was significantly higher in 2003 than 2005. Mean density 

of H. aurolineatum was highest in 2007 and lowest in 2002 

and mean biomass was lowest in 2001 and the highest in 

2003. Mean density and biomass of H. flavolineatum was 

the only grunt metrics to have three consecutive years of 

decline (2001-2004). Mean density of H. flavolineatum was 

highest in 2001 and lowest in 2007; and the average biomass 

was highest in 2001 and the lowest in 2006. Neither H. aurolineatum and H. flavolineatum metrics have 

significant differences between years of mean high or low. 

Mean density of H. plumierii was lowest in 2002 (0.20 ± 0.06) and significantly (p=0.0423) highest the 

following year in 2003 (1.64 ± 0.61). Mean biomass of H. plumierii was lowest in 2004 and highest in 

2006. Mean density of H. sciurus exhibited three years of consecutive increases (2004-2007), although 

they were not significant. Mean density of H. sciurus and biomass were significantly (p<0.05) higher in 

2002 than all years from 2005-2007; and mean density was significantly (p=0.0002) higher in 2001 than 

2005. Mean density of H. sciurus was highest in 2002 and lowest in 2004; and the average biomass 

was highest in 2002 and lowest in 2003.

Surgeonfishes(Acanthuridae)

Surgeonfish metrics include all species from the Acanthuridae family that were recorded in the 

southwest Puerto Rico study area. The surgeonfish species included in the ‘total surgeonfish’ metrics 

were A. bahianus, A. chirurgus, A. coeruleus and unidentified Acanthurus species. The mean density 

and biomass for the most common surgeonfish species recorded/observed in the study area are 

summarized in Table 3.5 and subset Table 3.14. 

H. aurolineatum

Surgeonfish variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Surgeonfish (Acanthuridae) density * 2.5 (0.5) 1.9 (0.3) 4.4 (0.7) 2.4 (0.3) 3.1 (0.4) 4.1 (0.7) 3.5 (0.5)

Surgeonfish (Acanthuridae) biomass * 174.4 (61.1) 153.4 (35.7) 369.6 (141.8) 161.7 (26.8) 265.2 (40.2) 309.2 (56.3) 311.4 (64.7)

Acanthurus bahianus density * 1.3 (0.3) 1.1 (0.2) 2.3 (0.3) 1.3 (0.2) 1.9 (0.3) 2.8 (0.4) 1.8 (0.2)

Acanthurus bahianus biomass * 76.4 (23.1) 76.3 (18.9) 121.6 (20.1) 78.5 (13.0) 139.3 (22.8) 201.6 (28.0) 126.7 (17.1)

Acanthurus chirurgus density * 0.2 (0.1) 0.4 (0.1) 1.0 (0.3) 0.5 (0.1) 0.5 (0.1) 0.3 (0.1) 0.7 (0.3)

Acanthurus chirurgus biomass * 13.8 (5.9) 39.8 (20.9) 99.8 (39.2) 42.9 (18.9) 38.6 (12.7) 25.1 (10.7) 103.9 (53.9)

Acanthurus coeruleus density * 0.9 (0.4) 0.5 (0.1) 1.0 (0.3) 0.6 (0.1) 0.7 (0.2) 1.1 (0.4) 0.9 (0.3)

Acanthurus coeruleus biomass * 84.2 (53.6) 37.3 (12.4) 148.2 (97.8) 40.3 (8.7) 87.2 (20.4) 82.5 (34.0) 80.8 (23.9)

Table 3.14. Density and biomass (mean + SE) for selected surgeonfish species (2001-2007) for the southwest Puerto Rico study area. 

Asterisks (*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up 

from Table 3.14 where appropriate. Bold indicates either the lowest or highest mean for that metric.

The means for all surgeonfish metrics increased from 2002-2003 then 

decreased the following year from 2003-2003. The highest means for all 

of the surgeonfish metrics occurred in 2003, 2006 and 2007. The lowest 

means for all of the surgeonfish metrics occurred in 2001 and 2002. Density 

and biomass of all surgeonfish and A. coeruleus, as well as, mean density 

of A. chirurgus was significantly (p<0.05) lower in 2002 than 2006. Mean 

density of total surgeonfish was the only surgeonfish metric to increase over 

three consecutive years. Mean density and biomass of total surgeonfish Juvenile A. coeruleus
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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142 3 - Fish Communitieswas lowest in 2002 (1.91 ± 0.32 and 153.40 ± 35.74, respectively) and significantly (p<0.0001) higher 

the following year in 2003 (4.39 ± 0.71 and 369.61 ± 141.82, respectively). Surgeonfish density and 

biomass means were significantly (p<0.0001) lower in 2002 than in 2003, 2005, 2006 and 2007. 

Mean density and biomass of A. bahianus was highest in 2006 (2.79 

± 0.41 and 201.58 ± 27.99, respectively) and lowest in 2002 (1.09 

± 0.19 and 76.32 ± 18.87, respectively); however only biomass had 

a significant difference between years of mean high and low. Mean 

density of A. bahianus was significantly lower in 2004 than 2003 

and 2006 and significantly (p=0.0001) lower in 2002 than 2003 and 

2003. Mean biomass of A. bahianus was significantly lower in 2002 

than 2003, 2006 and 2007; and significantly higher in 2006 than 

2003. 

Mean density and biomass of A. chirurgus exhibited three 

consecutive years of decline. Mean density and biomass were 

significantly (p<0.05) higher in 2003 than 2006. Mean density 

was highest in 2003 and lowest in 2001 and mean biomass was 

lowest in 2001 (13.82 ± 5.88) and highest in 2007. Neither metric 

had a significant difference between years of mean high or low. 

Mean biomass of A. coeruleus was the only surgeonfish metric 

that didn’t have any consecutive years of increase. Mean density 

and biomass were significantly higher (p<0.05) in 2006 than 2002. 

Mean density was highest in 2006 (1.07 ± 0.37) and significantly 

(p=0.0261) lowest in 2002 (0.45 ± 0.14). Mean biomass was also 

lowest in 2002, but was highest the following year in 2003, although 

not statistically significant.

Other Families

Squirrelfish (Holocentridae) and wrasse (Labridae) were selected because they are locally abundant 

and geographically widespread families in the southwest Puerto Rico study area. The wrasses also 

make up a large part of the herbivore group. The species included in the ‘total squirrelfish’ metrics are 

squirrelfish (Holocentrus adscensionis), longspine squirrelfish (Holocentrus rufus), blackbar soldierfish 

(Myripristis jacobus), longjaw squirrelfish (Neoniphon marianus), reef squirrelfish (Sargocentron 

coruscum), and dusky squirrelfish (Sargocentron vexillarium). The species included in the ‘total wrasse’ 

metrics are Spanish hogfish (Bodianus rufus), C. parrae, dwarf wrasse (Doratonotus megalepis), slippery 

dick (Halichoeres bivittatus), yellowcheek wrasse (Halichoeres cyanocephalus), yellowhead wrasse 

(Halichoeres garnoti), clown wrasse (Halichoeres maculipinna), rainbow wrasse (Halichoeres pictus), 

blackear wrasse (Halichoeres poeyi), puddingwife (Halichoeres radiatus), hogfish (Lachnolaimus 

maximus), bluehead wrasse (Thalassoma bifasciatum), rosy razorfish (Xyrichtys martinicensis), pearly 

razorfish (Xyrichtys novacula) and green razorfish (Xyrichtys splendens). The mean density and 

biomass for these two common families recorded/observed in the study area are summarized in Table 

3.5 and subset Table 3.15.

A. bahianus

A. chirurgus

Other Families variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Squirrelfish (Holocentridae) density * 1.9 (0.8) 0.8 (0.2) 0.7 (0.1) 0.5 (0.1) 0.7 (0.1) 1.2 (0.2) 1.0 (0.2)

Squirrelfish (Holocentridae) biomass * 107.5 (50.9) 79.5 (20.6) 69.0 (13.3) 59.9 (10.5) 65.3 (10.2) 99.1 (14.5) 86.0 (18.3)

Wrasse (Labridae) density * 8.9 (3.1) 8.8 (1.5) 11.7 (1.6) 8.2 (1.2) 10.6 (1.7) 13.7 (1.9) 12.7 (1.4)

Wrasse (Labridae) biomass * 50.0 (12.4) 60.0 (10.9) 63.9 (13.5) 28.3 (4.4) 98.6 (44.6) 47.2 (5.8) 64.0 (7.4)

Table 3.15. Density and biomass (mean + SE) for additional selected families (2001-2007) for the southwest Puerto Rico study area. 

Asterisks (*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up 

from Table 3.5 where appropriate. Bold indicates either the lowest or highest mean for that metric.
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143 3 - Fish Communities

Mean density and biomass of total squirrelfish was highest in 2001 followed by three consecutive 

years of decrease (2001-2004) resulting in the lowest means in 2003. Neither metric had a significant 

difference between years of mean high or low. Squirrelfish density and biomass means were significantly 

(p<0.01) higher in 2006 than 2002 and 2003.

Mean density of total wrasse was highest in 2006 and lowest in 2004 and mean biomass was highest 

in 2005 and lowest in 2003. Neither metric had a significant difference between years of mean high and 

low. When comparing first and last years, mean density of wrasse was significantly (p=0.0014) higher 

in 2007 than 2001 and 2002.

Other Species

B. vetula, M. martinicus and P. maculatus were included in the summary metrics because they are 

commercially important fish species and exhibited significant inter-annual differences. The mean 

density and biomass for these species recorded/observed in the study area are summarized in Table 

3.5 and subset Table 3.16.

Mean density and biomass of B. vetula was highest in 2007 (0.18 ± 

0.05 and 71.73 ± 18.08, respectively) and lowest in 2004 (0.02 ± 0.01 

and 4.18 ± 2.40, respectively). Both metrics had a significant difference 

(p<0.0001) between years of mean high or low. Furthermore, mean 

density and biomass of B. vetula was significantly (p<0.0001) higher 

in 2007 than all years from 2001-2005 and were significantly higher in 

2006 than 2003 and 2003.

Like many other metrics, both goatfish species (M. martinicus and P. maculatus) had their lowest 

means in 2004, although no inter-year differences were statistically significant. Mean M. martinicus

density was highest in 2007 and lowest in 2004 and mean biomass was highest in 2001 and lowest 

in 2003. Both mean density and biomass of P. maculatus

was highest in 2002 and the lowest in 2004; and mean 

biomass was highest in 2002 and lowest in 2003. Neither 

M. martinicus nor P. maculatus metrics had a significant 

difference between years of mean high or low.

Other species, families and groups (such as sharks/rays, 

porgies, angelfish, butterflyfish, damselfish) were not 

included because mean density or biomass values were too 

low (i.e., small sizes or infrequent observations) to illustrate 

in the summary table.

Other species variable

2001 2002 2003 2004 2005 2006 2007

Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE) Mean (SE)

Balistes vetula density * 0.03 (0.02) 0.05 (0.02) 0.02 (0.01) 0.02 (0.01) 0.03 (0.01) 0.1 (0.03) 0.2 (0.1)

Balistes vetula biomass * 18.5 (11.0) 26.6 (14.5) 7.6 (5.2) 4.2 (2.4) 22.1 (15.3) 48.8 (13.4) 71.7 (18.1)

Mulloidichthys martinicus density 0.5 (0.3) 0.1 (0.05) 0.1 (0.1) 0.1 (0.03) 0.2 (0.1) 0.3 (0.2) 0.7 (0.6)

Mulloidichthys martinicus biomass 40.6 (22.3) 19.3 (7.3) 28.5 (13.1) 7.5 (4.2) 31.4 (19.1) 17.2 (6.9) 31.7 (15.2)

Pseudupeneus maculatus density 0.3 (0.1) 0.7 (0.4) 0.4 (0.1) 0.1 (0.03) 0.3 (0.1) 0.3 (0.1) 0.3 (0.1)

Pseudupeneus maculatus biomass 13.6 (4.1) 29.8 (11.0) 18.3 (5.5) 5.6 (1.8) 17.2 (4.9) 13.7 (3.0) 16.9 (3.8)

Table 3.16. Density and biomass (mean + SE) for additional selected species (2001-2007) for the southwest Puerto Rico study area. 

Asterisks (*) indicate there was a significant difference when years were compared. Values have been rounded to one decimal place up 

from Table 3.5 where appropriate. Bold indicates either the lowest or highest mean for that metric.

B. vetula

 Foureye butterflyfish (Chaetodon capistratus)
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144 3 - Fish Communities3.5. Seasonal patterns in 

fish metrics (2004-2007)

Inter-seasonal comparisons 

of fish metrics were based 

on data from 2004-2007 only 

when semi-annual surveys 

were standardized to just two 

specific periods representing 

winter (December-March) 

and summer (AugustSeptember). Mangroves 

were not surveyed during the 

August 2005 mission, thus 

the summer 2005 data are 

presented without mangrove 

data. Table 3.17 shows the 

winter and summer densities 

for the 20 most abundant fish 

species. 

Mean densities in summer were higher for 13 of the 20 most abundant fish. Not all means were very 

different, but greatest increase from winter to summer densities was recorded for T. bifasciatum, S. 

partitus, S. taeniopterus; C. personatus/hyalinus and C. cyanea. In contrast, seven species exhibited 

higher mean densities in winter, with highest difference recorded for S. iseri, H. aurolineatum and H. 

flavolineatum (Table 3.17).

3.5.1. Community

Overall, nine of the 12 community metrics (total and trophic groups) 

had lowest means in the winter sampling season. Five of those 

metrics occurred during the 2006 winter (total, herbivore and 

piscivore richness, and herbivore and planktivore biomass) and 

four occurred during the 2004 winter (total fish density, piscivore 

and planktivore density, and planktivore richness). Two metrics 

(mean fish biomass and piscivore biomass), however, had lowest 

means in summer 2007. Summer mean total fish density decreased 

consecutively from 2004-2007, while winter mean total density 

increased from 2004-2007 (Figure 2.117a). Total mean biomass 

was more highly variable alternating between winter and summer highs (Figure 2.117b). Mean species 

richness was higher during the summer months than winter for all years (Figure 2.117c).

Herbivore means in 2004 and 2006 exhibited a similar pattern; winter means were higher in 2004 and 

summer means were higher in 2006 for all herbivore metrics (Figure 3.118). Winter mean herbivore 

density increased from 2005-2007; however, were higher in the summer during that same time period 

(Figure 3.118a). Summer biomass exhibited consecutive years of decrease from 2004-2007 and from 

2004-2006 during the winter sampling period (Figure 3.118b). Winter mean herbivore biomass was 

higher in all years except for 2006. Mean herbivore richness also declined consecutively in the winter 

from 2004-2006 (Figure 3.118c). Winter herbivore richness means were higher from 2004-2005 and 

summer exhibited the highest means from 2006-2007 (Figure 3.118c).

Piscivore mean density and richness were highest during the summer sampling period and biomass was 

highest during the winter. Piscivore density was at least two times higher during the summer than winter 

Table 3.17. Winter and summer total and mean (+ SE) density for the 20 most abundant fish 

species (2004-2007) for southwest Puerto Rico study area. N=708 surveys (per 100 m2

).

Species Common name

Total

density

Winter

density

Winter mean 

density 

Summer

density

Summer 

mean density

Thalassoma bifasciatum bluehead wrasse 4068 1532 3.3 (0.9) 2536 7.3 (0.6)

Scarus iseri striped parrotfish 3989 2330 6.5 (0.4) 1659 3.8 (1.0)

Stegastes partitus bicolor damselfish 3376 1520 3.2 (0.6) 1856 5.3 (0.5)

Sparisoma aurofrenatum redband parrotfish 2489 1035 2.9 (0.3) 1454 3.2 (0.2)

Haemulon flavolineatum french grunt 2065 1200 3.3 (0.7) 865 2.5 (0.8)

Halichoeres garnoti yellowhead wrasse 1679 741 2.1 (0.3) 938 2.7 (0.3)

Scarus taeniopterus princess parrotfish 1573 595 1.7 (0.3) 978 2.8 (0.2)

Haemulon aurolineatum tomtate 1511 1047 2.9 (0.4) 464 1.3 (2.1)

Acanthurus bahianus ocean surgeon 1378 669 1.9 (0.2) 709 2.0 (0.2)

Coryphopterus personatus/

hyalinus glass/masked goby 1118 340 0.9 (0.6) 778 2.2 (0.3)

Lutjanus apodus schoolmaster 1118 606 1.7 (0.4) 512 1.5 (0.4)

Halichoeres bivittatus slipper dick 1086 603 1.7 (0.3) 483 1.4 (0.3)

Chaetodon capistratus foureye butterflyfish 1070 487 1.4 (0.1) 583 1.7 (0.1)

Nes longus orangespotted goby 1066 515 1.4 (0.3) 551 1.6 (0.3)

Coryphopterus

glaucofraenum bridled goby 1005 526 1.5 (0.2) 479 1.4 (0.3)

Stegastes leucostictus beaugregory 979 516 1.4 (0.2) 463 1.3 (0.2)

Stegastes planifrons threespot damselfish 835 410 1.1 (0.3) 425 1.2 (0.2)

Sparisoma viride stoplight parrotfish 763 365 1.0 (0.1) 398 1.1 (0.1)

Gnatholepis thompsoni goldspot goby 751 329 0.9 (0.2) 422 1.2 (0.2)

Chromis cyanea blue chromis 742 163 0.5 (0.5) 579 1.7 (0.1)

Planktivorous fish blue chromis (Chromis cyanea)
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145 3 - Fish Communities

0

80

160

2004 2005 2006 2007

Mean fish density (100 m-2)

Winter

0

3

6

2004 2005 2006 2007

Mean fish biomass (kg/100 m2)

Winter

Summer

0

8

16

2004 2005 2006 2007

Mean number of fish species (100 m-2)

Winter

a)

c)

b)

Figure 3.117. Seasonal change in mean (+ SE) fish: (a) density, (b) biomass and (c) species richness in the southwest Puerto Rico study

area.

0

24

48

2004 2005 2006 2007

Mean herbivore density (100 m-2)

Winter

0

1

2

2004 2005 2006 2007

Mean herbivore biomass (kg/100 m2)

Winter

Summer

0

4

8

2004 2005 2006 2007

Mean herbivore richness (100 m-2)

Winter

a)

c)

b)

Figure 3.118. Seasonal change in mean (+ SE) herbivore: (a) density, (b) biomass and (c) richness in the southwest Puerto Rico study area.

Summer

Summer

Summer

Summer
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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146 3 - Fish Communitiesfrom 2004-2006 (Figure 3.119a). Both winter and summer sampling periods increased from 2004-2005 

followed by decreasing means from 2005-2006, with summer continuing to decline from 2006-2007 

(Figure 3.119a). Piscivore mean biomass decreased consecutively from 2005-2007, whereas winter 

sampling periods were variable with peaks in 2005 and 2007 (Figure 3.119b). Mean piscivore species 

richness was higher in summer than winter for all years. Richness decreased consecutively during the 

winter from 2004-2006 and during the summer from 2005-2007 (Figure 3.119c).

0

3

6

2004 2005 2006 2007

Mean piscivore density (100 m-2)

Winter

0

1

2

2004 2005 2006 2007

Mean piscivore biomass (kg/100 m2)

Winter

Summer

0

1

2

2004 2005 2006 2007

Mean piscivore richness (100 m-2)

Winter

a)

c)

b)

Figure 3.119. Seasonal change in mean (+ SE) piscivore: (a) density, (b) biomass and (c) richness in the southwest Puerto Rico study area.

Summer

Summer

Comparison between trophic groups

Mean densities of herbivorous and planktivorous fish for winter and summer were a maximum of 

more than 30 times greater than the mean density of piscivores (Figures 3.118a, 3.120a and 3.119a). 

Herbivorous fish comprised approximately half of the mean total fish biomass for both winter and summer 

sampling periods (Figures 3.118b and 3.117b), although piscivore biomass appeared to follow the trend 

in total mean biomass more closely, with alternating years of increasing and decreasing of winter and 

summer values between years and between seasons (Figures 3.119b and 3.117b). The 2005 winter 

sampling period had the highest values for all biomass metrics except for herbivore biomass. The 2006 

winter sampling period had the lowest values for planktivore and herbivore biomass (Figures 3.120b 

and 3.118b) and summer of 2007 had the lowest for total biomass and piscivore biomass (Figures 

3.117b and 3.119b).

Mean richness of piscivores and planktivores (Figures 3.119c and 3.120c) followed the total trend 

values (Figure 3.117c) with the highest means occurring during the summer sampling period. Herbivore 

richness values were five times higher than piscivore richness during both winter and summer sampling 

periods (Figures 3.118c and 3.119c). Planktivore richness was approximately two times higher than 

piscivore richness (Figures 3.120c and 3.119c) and herbivore richness was approximately two and a 

half times higher than planktivore richness (Figures 3.118c and 3.120c).
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147 3 - Fish Communities

0

40

80

2004 2005 2006 2007

Mean planktivore density (100 m-2)

Winter

0

1

2

2004 2005 2006 2007

Mean piscivore biomass (kg/100 m2)

Winter

Summer

0

2

4

2004 2005 2006 2007

Mean planktivore richness (100 m-2)

Winter

a)

c)

b)

3.5.2. Taxonomic groups 

Overall family comparisons

The seasonal trends in biomass for four major fish families were examined including groupers, snappers, 

parrotfish and grunts.

Mean biomass for all four major families (grouper, snapper, parrotfish, grunts) was lower in the winter 

than summer, with lowest mean biomass for grouper, snapper and parrotfish during the 2006 winter 

sampling period (Figure 3.121). Mean biomass of grunts was highest during summer 2006 and lowest 

in winter 2005, although there was little difference between winter and summer in 2004, 2006 and 2007 

(Figure 3.121d). Grouper biomass was lowest in both winter and summer of 2006 (Figure 3.121a). 

Mean biomass of parrotfish was higher in the winter months except in 2006 (Figure 3.121c). Winter 

biomass values decreased from 2004-2006, then increased in 2007, whereas summer months were 

variable. In contrast, snapper mean biomass values were higher in the summer months (except 2005; 

Figure 3.121b). The highest and lowest mean biomass snappers occurred in 2005 (i.e., highest in the 

winter, lowest in summer).

Large-body Groupers (Serranidae)

The lowest mean biomass values for the three selected groupers (C. cruentata, C. fulva, E. guttatus) 

occurred during 2006 (Figure 3.122). C. cruentata and E. guttatus exhibited lowest mean biomass in 

winter 2006 and C. fulva in summer 2006 (Figure 3.122). Low mean biomass values for C. cruentata

and E. guttatus coincided during the same year and season (winter 2006) as mean biomass of total 

groupers (Figure 3.122a). The years of highest mean biomass, however, occurred at different times; 

C. cruentata in summer 2004, C. fulva in winter 2005 and E. guttatus in winter 2004 (Figure 3.122). 

Overall, there appeared to be little difference in mean biomass values for selected groupers throughout 

the years regardless of season.

Figure 3.120. Seasonal change in mean (+ SE) planktivore: (a) density, (b) biomass and (c) richness in the southwest Puerto Rico study 

area.

Summer

Summer
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148 3 - Fish Communities

Figure 3.121. Seasonal change in mean (+ SE) fish biomass in the southwest Puerto Rico study area for: (a) groupers, (b) snappers, (c) 

parrotfish and (d) grunts.

0

0.5

1

2004 2005 2006 2007

Mean grouper biomass (kg/100 m2)

Winter

0

0.5

1

2004 2005 2006 2007

Mean snapper biomass (kg/100 m2)

Winter

Summer

0

0.5

1

2004 2005 2006 2007

Mean parrotfish biomass (kg/100 m2)

Winter

a)

c)

b)

0

0.5

1

2004 2005 2006 2007

Mean grunt biomass (kg/100 m2)

Winter d)

0

0.05

0.1

2004 2005 2006 2007

Mean C. cruentata biomass (kg/100 m2)

Winter

0

0.5

1

2004 2005 2006 2007

Mean C. fulva biomass (kg/100 m2)

Winter

Summer

0

0.05

0.1

2004 2005 2006 2007

Mean E. guttatus biomass (kg/100 m2)

Winter

a)

c)

b)

Summer

Summer Summer

Figure 3.122. Seasonal change in mean (+ SE) grouper biomass in the southwest Puerto Rico study area for: (a) graysby (C. cruentata), 

(b) coney (C. fulva) and (c) red hind (E. guttatus).

Summer

Summer
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149 3 - Fish Communities

Snappers (Lutjanidae)

Highest mean biomass for L. griseus and O. chrysurus was recorded in winter 2005 and L. apodus

had the highest mean biomass in summer 2004 (Figure 3.123). The lowest mean biomass for all three 

species occurred during the winter sampling period; L. apodus and L. griseus in winter 2004 and O. 

chrysurus in winter 2006. It is likely that had mangroves been sampled in summer 2005 that L. griseus

biomass would be higher for that period.

0

0.1

0.2

2004 2005 2006 2007

Mean L. apodus biomass (kg/100 m2)

Winter

0

0.1

0.2

2004 2005 2006 2007

Mean L. griseus biomass (kg/100 m2)

Winter

Summer

0

0.1

0.2

2004 2005 2006 2007

Mean O. chrysurus biomass (kg/100 m2)

Winter

a)

c)

b)

Figure 3.123. Seasonal change in mean (+ SE) snapper biomass in the southwest Puerto Rico study area for: (a) schoolmaster (L. apodus), 

(b) gray snapper (L. griseus) and (c) yellowtail snapper (O. chrysurus).

Summer

Summer

Parrotfish (Scaridae)

All four species exhibited a comparatively low mean biomass during the summer 2007 sampling period 

(Figure 3.124). For S. aurofrenatum and S. viride, winter 2006 was the period with lowest mean biomass 

during the time series (Figure 3.124c,d). Mean biomass of S. iseri was the highest in winter 2007 and 

the lowest in summer 2004 (Figure 3.124a), whereas S. taeniopterus had the exact opposite trend, 

having the highest mean biomass in summer 2004 and the lowest mean in winter 2007 (Figure 3.124b). 

Overall, S. iseri biomass values were higher during the winter sampling periods than summer, except 

during 2006 (Figure 3.124a). S. taeniopterus had higher means in the summer sampling period, except 

in 2005. Mean biomass of S. viride was higher in the winter season from 2004-2007, except for 2006 

(Figure 3.124d). In 2005, mean biomass for S. viride in 2005 differed very little between seasons. Mean 

biomass of S. viride during the winter sampling period decreased from 2004-2006, then increased in 

2007; while values during the summer sampling period increased from 2004-2005 then decreased from 

2005-2007 (Figure 3.124d).
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150 3 - Fish Communities

0

0.25

0.5

2004 2005 2006 2007

Mean S. iseri biomass (kg/100 m2)

Winter

0

0.25

0.5

2004 2005 2006 2007

Mean S. taeniopterus biomass (kg/100 m2)

Winter

Summer

0

0.25

0.5

2004 2005 2006 2007

Mean S. aurofrenatum biomass (kg/100 m2)

Winter c)

0

0.25

0.5

2004 2005 2006 2007

Mean S. viride biomass (kg/100 m2)

Winter d)

Summer

Summer

a) b)

Summer

Figure 3.124. Seasonal change in mean (+ SE) parrotfish biomass in the southwest Puerto Rico study area for: (a) striped parrotfish (Sc. 

iseri), (b) princess parrotfish (Sc. taeniopterus), (c) redband parrotfish (Sp. aurofrenatum) and (d) stoplight parrotfish (Sp. viride).

Grunts (Haemulidae)

Two of the three selected grunt species (H. flavolineatum and H. plumierii) had the highest biomass 

mean occur during the summer 2006 sampling period; whereas, the highest mean biomass for H. sciurus

during the 2004 winter sampling period (Figure 3.125). None of the selected grunt species shared the 

same sampling period for lowest mean biomass. Summer biomass values for H. flavolineatum were 

higher than all winter values except for 2007 (Figure 3.125a). Additionally, the highest and lowest mean 

biomass values for H. flavolineatum occurred during the same year. From 2004-2006 H. flavolineatum

winter biomass values were less than 0.02 kg/100m2

 and summer biomass values were greater than 

0.02 kg/100m2

; the highest winter biomass and lowest summer biomass values occurred in 2007 (Figure 

3.125a).

The highest H. plumierii biomass values for winter and summer sampling periods were in 2006, and 

the lowest biomass values were recorded in 2004; with summer 2004 having the lowest biomass 

values across all years (Figure 3.125b). H. sciurus mean biomass values were higher during the winter 

sampling periods for all years except for 2006 (Figure 3.125c). The lowest mean biomass for H. sciurus

was during the summer 2005 sampling period. 
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151 3 - Fish Communities

0

0.1

0.2

2004 2005 2006 2007

Mean H. flavolineatum biomass (kg/100 m2)

Winter

0

0.1

0.2

2004 2005 2006 2007

Mean H. plumierii biomass (kg/100 m2)

Winter

Summer

0

0.1

0.2

2004 2005 2006 2007

Mean H. sciurus biomass (kg/100 m2)

Winter

a)

c)

b)

Figure 3.125. Seasonal change in mean (+ SE) grunt biomass in the southwest Puerto Rico study area for: (a) French grunt (H. flavolineatum), 

(b) white grunt (H. plumierii) and (c) bluestriped grunt (H. sciurus).

Summer

Summer

Other species

The two goatfish (Mullidae) species recorded in the southwest Puerto Rico study area were M. martinicus

and P. maculatus. There appeared to be little change in mean biomass from 2004-2007 for either 

goatfish species, with the exception of winter 2007 for M. martinicus (Figure 3.126a). The highest and 

lowest means for M. martinicus occurred during the winter sampling seasons, with the highest mean 

biomass recorded in winter 2007 and the lowest in winter 2004 (Figure 3.126). P. maculatus exhibited 

an opposite pattern, with the highest and lowest means occurring during the summer months, highest 

in summer 2006 and the lowest in summer 2004 (Figure 3.126b).

0

0.1

0.2

2004 2005 2006 2007

Mean M. martinicus biomass (kg/100 m2)

Winter a)

0

0.1

0.2

2004 2005 2006 2007

Mean P. maculatus biomass (kg/100 m2)

Winter

Summer

b)

Figure 3.126. Seasonal change in mean (+ SE) goatfish biomass in the southwest Puerto Rico study area for: (a) yellow goatfish (M. 

martinicus) and (b) spotted goatfish (P. maculatus).

Summer
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152 3 - Fish CommunitiesMean biomass for B. vetula was highest in summer 2007 and lowest in summer 2004 (Figure 3.127a) 

and appears to have increased gradually in both winter and summer seasons over the time series. 

Mean biomass for S. barracuda was highly variable between seasons and years, with very low values 

(<0.0001 kg/100m2) in the winters of 2004 and 2006 and summer of 2007. The biomass means for 

the winter sampling period were variable with alternating increase and decrease, while biomass in the 

summer sampling period were more consistent from 2004-2006 (Figure 3.127b).

0

0.1

0.2

2004 2005 2006 2007

Mean B. vetula biomass (kg/100 m2)

Winter a)

0

1

2

2004 2005 2006 2007

Mean S. barracuda biomass (kg/100 m2)

Winter

Summer

b)

Figure 3.127. Seasonal and inter-annual (2003-2006) change in mean (+ SE) biomass in the southwest Puerto Rico study area for: (a) 

queen triggerfish (B. vetula) and (b) great barracuda (S. barracuda).

3.6. Historical comparison of species occurrence between 1980-1981 and 2001-2007

Kimmel (1985) used timed underwater visual census techniques to survey the fish communities of the 

La Parguera region associated with 21 biotopes across the insular shelf (e.g., mangroves, seagrasses, 

high relief shoreward reefs, high relief shelf edge reefs, etc.). Kimmel identified over 250 species in 168 

samples between 1980 and 1981. The visual fast count method was used of 50 minute timed surveys, 

with data recorded within 10 minute survey periods. Eight locations were surveyed within each biotope. 

These data do not provide comparable quantitative samples from which densities can be calculated 

and compared with CCMA-BB data, but if large differences occurred in the frequency of occurrence for 

particular species in 1980 compared with 2001-2007, then this is likely to be indicative of real change 

due to the spatially comprehensive across shelf sampling inherent in both studies.

In 1980-1981, rainbow parrotfish (S. guacamaia) was 

observed by Kimmel at 50% of samples from the eight 

mangrove sites surveyed (mean abundance 2.79); 

25% of high relief shoreward coral reef samples (mean 

abundance 3.36); 38% of low relief coral reef (mean 

abundance 2.75); 31% of low relief shoreward reefs 

(mean abundance 6.10) and 6% of high relief shelf edge 

reefs (mean abundance 0.15). The data reveal that in 

the early 1980s, the species occurred widely across 

the shelf from mangroves to shelf edge habitat types. 

In contrast, only two individuals of S. guacamaia were 

observed between 2001 and 2007 from 1,167 samples 

in the La Parguera region, thus confirming the rarity of 

the species in the region and supporting its status as 

vulnerable according to the IUCN red list of threatened 

species (Figure 3.128).

Summer

Figure 3.128. A spearfisherman with a 9.78 kg rainbow 

parrotfish (Scarus guacamaia) caught in Puerto Rico 

(exact location unknown) on January 29th 2005. Source: 

International Underwater Spearfishing Association.
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153 3 - Fish Communities

Midnight parrotfish (S. coelestinus) was less frequently observed 

in 1980 and 1981 than rainbow parrotfish with sightings at 19% 

of high relief shoreward reef samples (mean abundance 0.95); 

19% low relief apron reef samples (mean abundance 0.52); 

25% low relief shoreward reef samples (mean abundance 1.67); 

19% high relief shelf edge samples (mean abundance 0.94). In 

contrast, between 2001 and 2007, no midnight parrotfish were 

observed in the study area even though the range of habitat 

types sampled included those where the species was observed 

in 1980 and 1981. The status of this species urgently requires 

evaluation in the region.

In 1980 and 1981, Nassau grouper (E. striatus) was observed at 44% of high relief outer reef samples 

(mean abundance 1.23); 25% high relief shoreward reef samples (mean abundance 0.57); 6% low 

relief apron samples (0.07); 25% low relief shoreward reef samples (mean abundance 0.89); 50% of 

high relief shelf edge samples (mean abundance 1.80). In contrast, between 2001 and 2007 only two 

individual Nassau grouper were observed in the La Parguera study area indicating that populations 

have been depleted since 1980-81 and are not showing signs of recovery even though the species has 

been legally protected as a “Species of Concern” since 1991 under the Endangered Species Act.

In 1980 and 1981, M. tigris was observed only at 17% of high relief shelf edge reef samples (mean 

abundance 0.73). Between 2001 and 2007, M. tigris were not recorded on any transects in the study 

area suggesting that the population of this species may have declined further since 1980-1981. Historical 

comparison of other grouper species suggest that E. adscensionis was more frequently sighted over 

coral reefs in 1980-1981 (17.9%) than in 2001-07 (0.4%); C. fulva at 36.4% in 1980-1981 versus 7.8% 

in 2001-2007; E. guttatus at 64.7% in 1980-1981 versus 7.5% in 2001-07; and C. cruentata at 58.8% 

in 1980-1981 versus 29% in 2001-2007.

In 1980 and 1981, O. chrysurus an important species to regional fisheries, was observed at 78.4% of 

samples across hard, softbottom and mangroves, with highest abundance over seagrasses and low 

relief shoreward reef sites. In contrast, although O. chrysurus were the most abundant snapper in the 

study area in 2001-2007, they were observed at only 27.4% of all sites sampled. A recent published 

analyses using a portion of the 2001-2007 data for southwest Puerto Rico has shown that yellowtail 

snapper is one of several overfished species based on the body-length distribution and spawning 

potential (Ault et al., 2008). Kimmel’s data do not provide size estimates for the 1980-1981 period, 

thus it is not possible to tell the proportion of adults and juveniles. Based on the most recent and bestavailable data, it appears that the population size may have declined in the past 25 years and thus this 

species should be carefully monitored and protected from fishing if it is to recover. In addition, declines 

were found for other snapper species. L. jocu was observed at 8.3% of all samples across the shelf 

in 1980-81 and at 0.8% in 2001-2007. L. synagris, a fished species, occurred at 12.2% of samples in 

1980-1981 and 3.9% in 2001-2007. L. mahogoni was observed at 47.4% of the coral reef samples in 

1980 versus 2.9% in 2001-2007.

In 1980 and 1981, B. vetula, a popular local food fish, was observed at 20% of coral reef samples, with 

highest abundance at high relief offshore reefs. Although similar habitat preferences were observed in 

2001-2007, the percent occurrence had declined by half to approximately 10% occurrence across the 

shelf.

Furthermore, in 1980-1981, S. planifrons (an indicator species of complex and live coral cover) occurred 

at 46% of all coral reef samples across the shelf, with highest abundance and occurrence at high and 

low relief shoreward reefs (91% and 81% respectively) and high relief outer reefs (50%). During 2001-

Midnight parrotfish (Scarus coelestinus) Source: Kevin Eddy
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154 3 - Fish Communities2007, the percentage occurrence of S. planifrons, however, was lower, with S. planifrons present in 

31% (n=167) of hardbottom samples (n=541). To determine whether these differences are an artifact of 

the different sampling designs or a real shift due to fishing and changes in the structural complexity of 

coral reefs and the decline in live coral cover will require further studies.

3.7. Summary of results

3.7.1. Fish assemblage composition

• When comparing fish assemblage composition between major mapped habitat types (softbottom, 

hardbottom and mangroves), greatest difference was found between mangroves and hardbottom 

habitat types. The higher abundance of Haemulidae (grunts), Lutjanidae (snappers) and smallbodied schooling fish (Jenkinsia spp. and Atherinomorus spp.) in mangroves contributed most to 

dissimilarities. Thus, the results indicated that benthic habitat maps classified at the coarse thematic 

resolution of mangroves, hard and softbottom are ecologically meaningful and can be used to 

determine spatial differences in fish community composition.

• At finer levels of the hierarchical classification scheme (i.e., finer thematic resolution such as linear 

reefs, patch reefs, etc.), fish communities were not easily separated by mapped habitat types. This 

suggests that other characteristics including topographic complexity may need to be incorporated 

into mapped classes to better predict the distribution of fish communities.

• At the finest thematic resolution where softbottom habitat types were classified by the amount of 

seagrass cover, fish assemblages associated with continuous seagrasses were most distinct when 

compared with assemblages associated with lower proportions of seagrass cover, with a possible 

threshold effect at the level of >90% seagrass cover.

3.7.2. Multi-habitat use

• Our multi-habitat surveys of fish revealed that 12 of the 30 most abundant fish species observed 

in mangroves have also been observed over seagrasses, coral reefs and unvegetated sediments. 

Twenty-five of 30 species were observed in both mangroves and coral reefs indicating a high level 

of multi-habitat use by common fish species in the La Parguera region.

• For approximately 50% of the 30 most abundant fish in mangroves, fish body length was markedly 

smaller in mangroves than individuals of the same species on coral reefs, indicative of size dependent 

ontogenetic habitat shifts, particularly for Haemulidae, Lutjanidae, Scaridae and Sphyraenidae 

(barracuda).

• Based on body size, mangroves appeared to function as an intermediate habitat type for some 

grunts and snappers, with smallest fish associated with seagrasses, larger fish in mangroves and 

the largest mean length recorded for fish on coral reefs.

3.7.3. Spatial patterns of fish diversity, biomass and abundance

• Across the La Parguera study area a total of 210 fish species were identified to species level, with at 

least another 14 fishes identified only to genera.

• Highest species richness of 41 fish per 100 m2 area was recorded near the eastern shelf edge over 

colonized pavement with sand channels.

• Hotspots of high fish species richness, high fish biomass and high herbivore abundance and richness 

co-occurred along the shelf edge, particularly along the central to eastern edge of the surveyed area 

and around the complex of patch reefs between Margarita Reef and the El Palo area.

• Highest fish abundance was associated with mangroves, both inshore and around offshore mangrove 

cays, with the assemblages composed mostly of juveniles indicating that mangroves of the La 

Parguera region are an important resource for juvenile fish.
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155 3 - Fish Communities

• Although S. guacamaia is thought to have a high dependence on mangroves and coral reefs, only 

two individuals were observed between 2001 and 2007 in the La Parguera region confirming the 

rarity of the species in the U.S. Caribbean and its status as vulnerable according to the IUCN red 

list of threatened species. No midnight or blue parrotfish were sighted within the surveyed areas 

between 2001 and 2007. 

• The small to medium-bodied parrotfish species, (S. iseri and S. aurofrenatum) were the most 

commonly occurring fish across the seascapes at La Parguera.

• O. chrysurus was the most abundant and widespread snapper species and the 11th most commonly 

occurring fish species (738 individuals) in the La Parguera study area found at 27.4% (n=320) of 

survey sites.

• The largest snappers seen in the study area were markedly smaller than the maximum known size 

for the species, particularly L. apodus, O. chrysurus and L. griseus.

• Small-bodied groupers were more abundant than large-bodied grouper and included C. cruentata

(n=246) and C. fulva (n=81). Abundance and biomass of small-bodied grouper species was highest 

along the shelf edge, with a maximum density of seven grouper recorded at one site (100 m2).

• None of the grouper species observed had attained the maximum known size for their species, 

with maximum length for Cephalopholis species in the La Parguera region estimated at 30 cm FL 

compared with a maximum known for C. fulva of 41 cm TL and 43 cm TL for C. cruentata. The largest 

E. guttatus was approximately 50% of the maximum recorded size.

• From a total of 1,167 surveys (572 from hardbottom) over seven years, no M. tigris or M. venenosa

were observed and only two E. striatus, two M. bonaci, two E. adscensionis and 43 E. guttatus were 

observed, even though a known spawning aggregation for E. guttatus, E. adscensionis, M. tigris and 

M. venenosa exists along the shelf edge in southwest Puerto Rico.

• Five sharks (two species) and three stingrays were the only sharks and rays observed within transects 

at La Parguera between 2001 and 2007.

3.7.4. Habitat and ontogenetic space use patterns

• O. chrysurus juveniles occurred in 10 of the 11 habitat types, with highest abundance over colonized 

hardbottom. Juveniles showed no spatial segregation from adults, but adults were more frequently 

observed in the mid and outer shelf zones than in lagoonal areas.

• Parrotfish (Scaridae) exhibited high spatial heterogeneity in distributions across the shelf, with 

highest abundance over the most topographically complex hardbottom. Juveniles and adults cooccurred across the study area.

• S. taeniopterus exhibited a preference for mid and outer shelf zones with highly contiguous coral 

reef, while striped parrotfish occurred in all zones across the shelf.

• Grunt (Haemulidae) abundance and biomass was highest in the nearshore and offshore mangroves 

in close proximity to coral reefs and seagrass beds. All common species of grunt showed a strong 

across shelf size-dependent distribution, with the majority of juveniles in lagoonal nearshore areas 

and adults in deeper mid- and outer-shelf zones. Juveniles and adults did also co-occur at several 

sites across the shelf indicating some flexibility in the strategy for ontogenetic segregation.

• Damselfish (Pomacentridae) species S. planifrons, M. chrysurus and S. variabilis abundance is 

highest on topographically complex coral reefs and can be used as a good indicator of complex coral 

reefs.

• B. vetula, a fished species in the La Parguera region, exhibited highest abundance and biomass in the 

topographically complex colonized pavement with sand channels habitat type of the outer shelf zone.

• S. barracuda exhibited a strong size segregated spatial distribution, with juveniles confined to 

nearshore lagoonal areas and most adults found in the mid and outer shelf zones.
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156 3 - Fish Communities3.7.5. Temporal patterns in fish abundance and biomass

• None of the fish metrics decreased for more than three consecutive years within the seven year 

monitoring period (2001-2007), but 14 metrics decreased for at least three consecutive years prior 

to 2006.

• Grouper biomass in 2007 was significantly lower than in 2001.

• Only C. fulva and S. aurofrenatum increased in abundance consecutively for more than three years 

between 2001 and 2007.

• Total fish biomass, herbivore biomass, grouper abundance, parrotfish and wrasse abundance were 

significantly higher in 2007 than in 2001.

• Total snapper density and L. apodus density decreased from 2002 to 2005.

• The most striking inter-annual difference occurred between 2003 and 2004, whereby 65 metrics 

(approximately 80% of all metrics) decreased, with five decreasingly significantly; followed by 70% 

of metrics increasing the following year (2004-2005).

• Sixty-five percent of the 20 most abundant fish exhibited higher density in summer than in winter. 

T. bifasciatum and C. cyanea mean density was more than 50% higher in summer. However, H. 

aurolineatum, H. flavolineatum and S. iseri were more abundant in winter.

• Mean biomass for grouper, snapper, parrotfish and grunts was lower in the winter than summer.

• Planktivores and piscivores were more abundant in summer than winter.

• Comparison of the number of times a species was sighted across the shelf at La Parguera in 1980-

81 versus 2001-07 indicated that the population sizes of several fished species have declined 

substantially over the past 25 years. These include S. guacamaia, S. coelestinus, E. striatus, M. 

tigris, E. adscensionis, E. guttatus, C. fulva, E. cruentata, O. chrysurus, L. synagris, L. mahogoni, L. 

jocu and B. vetula. These are all relatively large-bodied and important food fish for the region.

3.8. Discussion

3.8.1. Fish community composition and mapped habitat types

Benthic habitat maps are often produced for multiple purposes including characterizing the distribution, 

abundance and diversity of biological resources to support decision making in marine management. For 

instance, benthic maps often play a central role in site identification and prioritization for marine protected 

areas, delineating essential fish habitat in ecosystem-based management and zoning in marine spatial 

planning, but rarely are the classifications evaluated for their meaningfulness to key faunal groups. 

In Australia, Ward et al. (1999) examined the utility of habitat maps as surrogates for biodiversity and 

concluded that the mapped classes were good surrogates for biodiversity. This project evaluated the 

utility of the NOAA benthic habitat map classes for predicting differences in fish community composition 

across multiple habitat classes at different levels of a hierarchical map classification. It was expected 

that fish communities would differ significantly among habitat types due to the perceived structural 

differences between habitat types observed during aerial photo interpretation and via underwater 

observations, which originally resulted in the delineation and classification of distinct map classes.

In fact, our characterization and analysis revealed statistically significant compositional differences in 

the fish assemblages only when samples were grouped at the coarsest thematic resolution (softbottom, 

hardbottom and mangroves). Of all pairwise habitat comparisons, fish communities associated with 

mangroves and fish associated with hardbottom habitats showed highest dissimilarity (least overlap). 

This was primarily due to the higher abundances of grunts and of small-bodied schooling species 

such as Atherinids, Clupeids and Engraulids within mangroves compared with hardbottom areas. 

Similarly, Chittaro et al. (2005) in Belize found low similarity between the fish community composition 

of mangroves and coral reefs and the authors interpreted this as indicative of low connectivity between 
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the two habitat types. Community similarity/dissimilarity, however, should not be used as a measure 

of relative connectivity since it focuses on assemblage level patterns and not individual species 

movements. For example, high exchange between habitat types can occur for specific species, yet the 

community compositions for the two habitat types can still be very dissimilar.

At finer thematic resolutions considerable overlap was detected in fish community composition, 

particularly within hardbottom habitat types, where high similarity occurred between geomorphologically 

different classes such as patch reefs, linear reefs and colonized pavement. Although differences 

sometimes occurred they were not statistically significant and thus we conclude that the assemblage 

composition did not respond strongly to the different biophysical structures observed in the aerial 

photography and subsequently delineated by the cartographer. With regard to the use of habitat maps 

for predicting diversity patterns, the coarsest levels of the classifications scheme also performed best. 

In an earlier study focusing on predicting fish species richness in southwest Puerto Rico, Pittman 

et al. (2007b) coupled Biogeography Branch fish survey data with seascape structure from the 

benthic habitat map to examine between-habitat differences 

in fish species richness. The results from a simple regression 

tree model revealed that the greatest difference in fish species 

richness was found between softbottom and hardbottom sites, 

with pairwise comparisons between habitat types within the 

softbottom grouping and between habitat types within the 

hardbottom grouping being mostly non-significant. Some of the 

overlap in community similarity may be explained by high withinhabitat type variability making habitat types less distinct as has 

been described in similar studies elsewhere in the Caribbean 

(Harborne et al., 2008; Mumby et al., 2008). Recent work in St 

Croix (U.S. Virgin Islands) has demonstrated that both the spatial 

resolution of a benthic map and the thematic resolution selected 

for a study can influence the model of fish-habitat relationships 

(Kendall and Miller, 2008; 2009).

Interestingly though, at the finest thematic resolution, where softbottom habitat types were classified 

by the amount of seagrass cover our study did reveal some differences. Fish assemblages associated 

with continuous seagrasses were most distinct when compared with assemblages associated with 

samples from several sites with incrementally lower proportions of seagrass cover, suggesting that a 

threshold effect may be occurring at the level of >90% seagrass cover. Very few studies have examined 

fish communities across a gradient in seagrass cover to identify threshold effects in the faunal-seagrass 

relationship. In seagrass beds of Moreton Bay (Australia) a gradual decline in resident fish abundance 

was detected, along spatial gradients in seagrass cover, until approximately 15–20% seagrass cover, 

beyond which many abundant species were absent (Pittman et al., 2004). In the Caribbean (Virgin 

Islands, Florida Keys, and Turks and Caicos), an examination of fish communities on coral reefs along a 

spatial gradient in seagrass cover revealed that fish diversity and 

abundance increased from 0 to 20–30% seagrass coverage then 

plateaued at 40% indicating a threshold like response (GroberDunsmore, 2005). Critical thresholds imply that absence, loss, 

or degradation of habitat can have deleterious effects on the 

distribution and population dynamics of some species, and are 

therefore crucially important to studies predicting the impact 

of environmental change in tropical marine systems. Further 

studies are required to determine if a threshold effect occurs in 

the relationship between fish and seagrass cover since other 

factors could influence the patterns along a gradient including 

Baitfish in mangrove habitat

Colonized hardbottom habitat
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto RIco): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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158 3 - Fish Communitiesthe surrounding habitat types (i.e., proximity to mangroves and coral reefs), water quality and the type 

of seagrasses. It is likely that the very sparse seagrasses function more like unvegetated sand in that 

they have a more similar structure and refuge function, sediment chemistry, food resources and other 

conditions than do highly vegetated areas with dense and long-leaved seagrasses and muddy organic 

rich sediments.

3.8.2. Multi-habitat utilization

Although direct evidence of habitat connectivity cannot be directly inferred from our underwater surveys, 

the current report does demonstrate that many fish species use multiple habitat types in the La Parguera 

region. By the term “use” we are referring to space use i.e., the fish were observed over specific habitat 

types and were thus associated with them. Our multi-habitat surveys of fish revealed that 12 of the 30 

most abundant fish species observed in mangroves also used 

seagrasses, coral reefs and unvegetated sediments. As many 

as 25 of 30 species were observed in both mangroves and 

coral reefs, indicating a high level of multi-habitat use by many 

of the most common fish species. Additional analyses revealed 

a strong linear relationship (r2

=0.77) between the number 

of habitat types used by a species and its total abundance 

across the region. As such, fish species that have evolved 

to use all habitat types (seascape generalists) are also the 

most abundant species across the region. These seascape 

relationships require further study and need to be evaluated 

relative to the implications for resource management.

For some species, distinctive size distribution patterns across habitat types were revealed suggesting that 

combinations of habitat types may provide a sequence of ontogenetic stepping stones from settlement 

habitat to adult habitat. Based on body size, mangroves appeared to function as an intermediate 

habitat type for some grunts and snappers, with smallest fish associated with seagrasses, larger fish 

in mangroves and the largest mean length recorded for fish on coral reefs. This size-specific habitat 

use pattern has also been observed for grunts and snappers in Florida (Serafy et al., 2007), Belize 

(Mumby et al., 2004) and Puerto Rico (Appeldoorn et al., 1997; Aguilar-Perera and Appeldoorn, 2007; 

2008). The strong evidence for multiple habitat use in tropical marine fish requires that we move away 

from a single habitat approach when studying ecological relationships toward a seascape approach 

that considers mosaics of habitat types (Pittman and McAlpine, 2003). The problem is pervasive and 

in the scientific literature multi-habitat fish species are routinely assigned a single habitat descriptor 

based on a perceived habitat association regardless of their movements and resource requirements. 

For example, a multi-habitat species is often separately classified as a “seagrass fish species”, a “coral 

reef fish species”, and a “mangrove fish species” depending on the habitat surveyed and the focus of 

the research project. While single habitat descriptors may be appropriate for habitat specialists that are 

resident within a single habitat type, they are inappropriate 

for species that are capable of using more than one habitat 

type (habitat generalists) and particularly misleading for 

individuals that are critically dependent on multiple habitat 

types through daily home range movements and ontogenetic 

habitat shifts. Single habitat discriptors are also routinely 

misused when applied to assemblages and communities (i.e., 

coral reef fish community, mangrove fish community) since 

habitat association is typically based on only a snapshot of 

their entire space-use patterns captured during relatively finescale surveys. The fact that some species may have a critical 

Mangrove habitat

Seagrass habitat
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159 3 - Fish Communities

dependence on one or a specific combination of habitat types, while others have adapted generalist 

strategies able to use a wide range of habitat types at all life stages adds significant complexity to the 

conventional view on fish-habitat relationships and strategies for delineation of essential fish habitat 

(EFH) and design of marine protected areas (MPAs).

3.8.3. Spatial patterns of fish diversity, biomass and abundance

Hotspots of high fish species richness, high fish biomass and high herbivore abundance and richness 

occurred along the shelf edge, particularly along the central to eastern edge of the surveyed area. 

Edges are often associated with elevated productivity and diversity since species from either side of 

the edge or from adjacent patch types intermingle. The shelf edge is the boundary between shallower 

shelf environments and the deeper water slope areas creating heterogeneous habitats. A higher 

diversity of planktivorous species is typical of shelf edges due to stronger currents and upwelling 

carrying plankton rich waters along and over the edge. In addition, the shelf edge is characterized by 

high topographic complexity of the seafloor, which provides higher surface area for habitat and more 

abundant refuge space for a larger number and diversity of species including large-bodied fish such as 

grouper. Additional diversity and abundance hotspots were identified around the complex of patch reefs 

between Margarita Reef and the El Palo area. The recent availability of LiDAR bathymetry (airborne 

laser altimetry) has allowed us to develop a 3-dimensional surface for the La Parguera seafloor. 

Examination of the 3-dimensional model of bathymetry derived from hydrographic LiDAR data reveals 

a complex aggregation of patch reefs over the seafloor around the El Palo reef. It is likely that this high 

structural complexity together with the geographical location on the shelf near extensive seagrasses 

and mangroves supports the diverse and productive fish assemblages observed. The shelf edge areas 

and the El Palo region also exhibited the highest species richness of herbivorous fish, an assemblage 

characteristic that can have beneficial effects on coral reef structure and function. Experimental studies 

by Burkepile and Hay (2008) demonstrated that multiple species of herbivorous fish lowered macroalgal 

abundance by 54–76%, enhanced cover of crustose coralline algae by 52–64%, increased coral cover 

by 22%, and prevented coral mortality.

The most abundant fish species were the small-bodied parrotfish 

species (S. iseri and S. aurofrenatum), confirming that herbivores 

numerically dominate the fish communities across the shelf at 

La Parguera. This could be indicative of a phase shift towards 

algae dominated seascapes. Further historical comparisons need 

to be carried out to determine if the abundance and distribution 

of these species has changed over the years and whether shifts 

in the numerically dominant species have occurred relative to 

environmental changes.

The highest abundance and biomass of parrotfish was recorded over the most topographically complex 

hardbottom across the study area, but species-specific distributions were evident too. For example, S. 

taeniopterus exhibited a preference for mid- and outer- shelf zones with highly contiguous coral reef, 

while striped parrotfish occurred in all zones across the shelf. In northeast St. Croix too, S. taeniopterus

exhibited higher abundance across the deeper extensive colonized pavement areas seaward of Buck 

Island than did S. iseri. This appears to be related to a preference for structurally complex areas in 

deeper water. 

3.8.4. Habitat and ontogenetic space use patterns

As discussed above, many species associated with coral reef ecosystems utilize multiple habitat 

types, often with very different biophysical structure (seagrasses, mangroves, coral reefs, etc.) and 

species composition. For some species the habitat types used can also be in geographically distinct 

zones across the shelf that may or may not correspond with changes in water depth. The patterns in 

S. iseri
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160 3 - Fish Communitiesacross shelf distributions for specific species and the influence of size on these patterns was shown 

by Christensen et al. (2003) using the first year of data collected by the Biogeography Branch and was 

highlighted by Aguilar-Perera and Appeldoorn (2008). Christensen et al. (2003) indicated that habitat 

type was more important than cross-shelf location in determining the distributional patterns among 

fishes in the study area and Aguilar-Perera and Appeldoorn (2008) found a consistent spatial pattern 

with fish assemblages (based on the most abundant species) grouping into clusters associated with 

mangroves, seagrasses, shallow coral reefs and deeper water coral reefs, with densities and diversity 

highest over coral reefs than any other habitat type.

Building on this work through analysis of a greater number of samples with more extensive spatial 

coverage collected over seven years we show some distinctive spatial patterns both across the shelf 

and by habitat for juveniles and adults. Some key species exhibited spatial segregation between 

distribution patterns of juveniles and adults, while for other species juveniles and adults co-occurred 

at the same sites, habitat types and zones. Grunts (Haemulidae) showed a strong across shelf sizedependent distribution, with the majority of juveniles in lagoonal nearshore areas and adults in deeper 

mid- and outer-shelf zones. Juveniles and adults, however, also co-occured at several sites across the 

shelf indicating some flexibility in the dominant strategy of ontogenetic across-shelf segregation.

O. chrysurus juveniles occurred in 10 of the 11 habitat types, indicating that the juveniles are very 

generalist users of the seascape. Further research would be required to determine the relative importance 

of different habitat types on growth and survival. Although densities differ, adults and juveniles co-exist 

across the shelf at the same sites, habitats and zones and thus no geographically distinct nearshore 

nursery area was evident for O. chrysurus in southwest Puerto Rico as has been defined elsewhere 

in the Caribbean and referred to as “nursery species” by Nagelkerken (2000). Nagelkerken et al. 

(2000, 2001) defined “nursery species” as those species with high densities of juveniles in nearshore 

habitat types such as mangroves and seagrasses and with an almost complete absence of juveniles 

on the coral reefs where the adults were more abundant. Aguilar-Perera and Appeldoorn (2007) also 

showed that several species considered as nursery species by Nagelkerken in Curacao could not 

be considered as such in southwest Puerto Rico including H. flavolineatum, H. sciurus, H. plumerii, 

L. apodus and O. chrysurus. In the current study, juveniles of these species were recorded at higher 

densities in mangroves than elsewhere, but juveniles were also observed on mid and outer shelf edge 

zones. The geographical differences across the Caribbean may reflect real differences in habitat use 

strategies or may reflect differences in predation pressure or the amount of survey effort between 

studies. Furthermore, densities of fish in space are highly influenced by habitat structure and location 

and can be a misleading indicator of habitat quality and should not be used alone when assigning 

relative value to a specific habitat types (van Horne, 1983).

In addition, juvenile and adult parrotfish co-occurred across the study area. For these ubiquitous 

species where all life stages can co-occur, it is conceivable that although nearshore areas may be 

beneficial in supporting high abundance across the region, nearshore areas may not be critical to the 

maintenance of populations in the region. In contrast, some species showed geographically distinctive 

zones for juveniles and adults in line with the concepts defined by the nursery hypothesis. For example, 

S. barracuda exhibited a strong size segregated spatial distribution, with 

juveniles confined to nearshore lagoonal areas and most adults found in 

the mid and outer shelf zones. Other species, showed no segregation of 

juveniles and adults, but instead exhibited a strong apparent preference 

for the most topographically complex colonized hardbottom areas. For 

instance, damselfish (Pomacentridae) species S. planifrons, M. chrysurus 

and S. variabilis were distributed over the most topographically complex 

coral reefs and their presence and possibly abundance may be useful as 

a good indicator of complex coral reefs. B. vetula also exhibited highest S. planifrons
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161 3 - Fish Communities

abundance and biomass in the topographically complex colonized pavement with sand channels habitat 

type of the outer shelf zone.

3.8.5. Size structure and shifts in predators for vulnerable species

Declines in maximum body size and changes in the abundance of size classes over time can be 

indicative of fishing pressure. O. chrysurus, a highly prized food fish in the Caribbean was the 11th 

most commonly occurring fish species (738 individuals) in the La Parguera study area found at 27.4% 

of survey sites. The largest fish was estimated at 40 cm FL, therefore, none of these individuals had 

achieved anywhere near the maximum known size for the species (86.3 cm TL) and only 4% of all 

fish observed were more than 30 cm FL. For this species too, historical comparison is required to 

determine if changes in the size structure of the population have occurred. This is particularly important 

to determine whether yellowtail snapper should be listed as overfished by NOAA’s National Marine 

Fisheries Service. Based on life history characteristics, O. chrysurus is classified as a species with 

relatively slow population doubling time (4.5-14 years) and low resilience to fishing (Froese and Pauly, 

2010).

Also of note was that small-bodied groupers were markedly 

more abundant than large-bodied groupers. With the exception 

of red hind, the large-bodied groupers were exceedingly rare 

across the shelf. From a total of 1,167 surveys over seven 

years, no M. tigris or M. venenosa were observed and only 

two E. striatus and two M. bonaci were observed. The largest 

E. guttatus was approximately 50% of the maximum recorded 

size and this species should receive close attention with 

regard to its population size structure and viability in the La 

Parguera region. In contrast, abundance of C. cruentata was an order of magnitude higher across the 

La Parguera study area than red hind and C. fulva were also recorded at higher densities than all of the 

larger-bodied grouper species combined. Furthermore, C. cruentata was the only grouper species that 

appeared to be increasing in density from 2002 to 2007. 

Elsewhere in the Caribbean and Florida, shifts in the dominant predators have been attributed to fishing 

pressure, since large-bodied predators such as groupers are particularly vulnerable to fishing and when 

removed from the ecosystem are replaced with smaller, often less-targeted predators (Pauly et al., 

1998). Studies by Chiappone in fished and unfished areas in Florida showed that the abundance and 

biomass of small-bodied groupers, including C. fulva, were higher in areas where fishing had reduced 

the large-bodied groupers (e.g., Epinephelus spp., Mycteroperca spp.). Our own studies in northeast 

St Croix, USVI. (Pittman et al., 2008) reported only four individuals of the very large-bodied and highly 

vulnerable grouper species, compared with 697 of the low to moderate vulnerability grouper (413 C. 

fulva, 231 E. guttatus and 53 C. cruentata). Comparison between fish monitoring surveys around 

Buck Island (St. Croix, U.S. Virgin Islands) conducted in 1979 (Gladfelter and Gladfelter, 1980) and 

a geographically similar subset of CCMA-BB monitoring surveys conducted between 2001 and 2006 

reveal that while E. striatus, M. venenosa and M. tigris existed at low densities in 1979 they were absent 

between 2001 and 2006. In contrast, E. guttatus and C. fulva have increased in density significantly 

from 0.08 individuals/100 m2

 of red hind in 1979 to 0.18/100 m2 in 2001-2006; and from 0.01/100 m2 of 

C. fulva in 1979 to 0.30/100 m2 in 2001-2006 (Table 12 in Pittman et al., 2008).

The consequences of such shifts may have cascading effects through the biological community. 

Studies by Stallings (2008, 2009) have shown that coney are more voracious predators of newly settled 

fish than are some of the larger-bodied species such as E. striatus and have been found to have a 

more significant impact on the recruitment to patch reefs for a wide range of common fish species. 

The ecological implication of an increase in small-bodied grouper on the fish community of the U.S. 

Nassau grouper (Epinephelus striatus)
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162 3 - Fish CommunitiesCaribbean is currently unknown. Research is needed to determine if a shift in predators has occurred in 

the La Parguera region and then to determine what the ecological consequences are for the population 

and community dynamics of the region.

3.8.6. Temporal patterns in fish abundance and biomass

Our semi-annual long-term monitoring data revealed a distinctive seasonal pattern in the abundance 

of fish in southwest Puerto Rico with 65% of the 20 most abundant fish exhibiting higher densities in 

summer than winter. For two small bodied planktivorous fish species, T. bifasciatum and C.cyanea, the 

mean density was more than 50% higher in summer and this may reflect seasonal differences in food 

availability for planktivorous fish. Studies by Youngbluth (1980) in Jobos Bay, Puerto Rico found peaks 

in zooplankton biomass and copepod densities from August-October.

It is likely that the largest changes in abundance of large-bodied fish happened before the current 

CCMA-BB monitoring program (i.e., pre-2001). Synoptic analyses of trends over the entire sampling 

period showed that none of the fish metrics decreased for more than three consecutive years within the 

seven year monitoring period (2001-2007), but 14 species decreased for three consecutive years prior 

to 2006. The upturn from 2006-2007 could be an indication of a new trend and requires continuation 

of long-term monitoring to elucidate on future trends. This apparent increase at the end of the study 

period is also apparent among many other fish metrics with 13 of 15 metrics increasing consecutively 

over three years showing an increase in the latter half of the study period between 2004 and 2007. 

Total fish biomass, herbivore biomass, grouper abundance, parrotfish and wrasse abundance were 

significantly higher in 2007 than in 2001. S. aurofrenatum exhibited more than three consecutive years 

of increase. This pattern could benefit the health of coral reefs in the area since redband parrotfish 

are known to consume large quantities of several major macroalgal species from the genera Dictyota, 

Halimeda, Lobophora, Sargassum, Haloplegma, Kallymenia and Codium species (Burkepile and Hay, 

2008). Dictyota and Lobophera spp. have been implicated in phase shifts from coral dominated reefs 

to macroalgal dominated reefs, thus an increase in a key consumer of Dictyota spp. and Lobophora 

variegata may play an important role in controlling macroalgal biomass. Furthermore, these fast growing 

macroalgal species also inhibit coral recruitment (Kuffner et al., 2006).

For the small-bodied grouper species, mean biomass was significantly higher in 2007 than it was in 

2001. This increase could be due to increased food availability or due to the relatively low abundance 

of large piscivorous fish (i.e., low predation pressure). For example, the large-bodied groupers showed 

no increase over seven years and total snapper density and L. apodus density decreased from 2002-

2005. Similarly, release of coney and graysby populations due to shifts in the dominant piscivorous fish 

(i.e. large groupers) on the reef have been documented elsewhere in the Caribbean (Stallings, 2008).
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Pittman, S.J., J. Christensen, C. Caldow, C. Menza, and M. Monaco. 2007b. Predictive mapping of fish species 

richness across shallow-water seascapes of the U.S. Caribbean. Ecol. Model. 204: 9-21.

Pittman, S.J., S.D. Hile, C.F.G. Jeffrey, C. Caldow, M.S. Kendall, M.E. Monaco, and Z. Hillis-Starr. 2008. Fish 

assemblages and benthic habitats of Buck Island Reef National Monument (St. Croix,U.S. Virgin Islands) and the 

surrounding seascape: A characterization of spatial and temporal patterns. NOAA Technical Memorandum NOS 

NCCOS 71. Silver Spring, MD. 96 pp. 

Pittman, S.J., B. Costa, and T. Battista. 2009. Using Lidar bathymetry and boosted regression trees to predict the 

diversity and abundance of fish and corals. J. Coast. Res. S53: 27-38.

SAS Institute. 2006. SIGMAPLOT© Software. SAS Institute Inc., Cary, NC, USA.

Serafy, J.E., M. Valle, C.H. Faunce, and J.G. Luo. 2007. Species-specific patterns of fish abundance and size 

along a subtropical mangrove shoreline: an application of the delta approach. Bull. Mar. Sci. 80: 609-624.
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Stallings, C.D. 2008. Indirect effects of an exploited predator on recruitment of coral-reef fishes. Ecology 89: 

2090-2095. 

Stallings, C.D. 2009. Fishery-independent data reveal negative effect of human population density on Caribbean 

predatory fish communities. PLoS One 4(5): e5333.

van Horne, B. 1983. Density as a misleading indicator of habitat quality. J. Wildl. Manage. 47(4): 893-901.

Ward, T.J., M.A. Vanderklift, A.O. Nicholls, and R.A. Kenchington.1999. Selecting marine reserves using habitats 

and species assemblages as surrogates for biological diversity. Ecol. Appl. 9: 691-698. 

Youngbluth, M.J. 1980. Daily, seasonal and annual fluctuations among zooplankton populations in an unpolluted 

tropical embayment. Est. Coast. Mar. Sci. 10: 265-287.
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166 3 - Fish Communities
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

167Appendix A

Appendix A

Species/Species group Fish Family

USVI % of 

total landings 

US Caribbean

Grunts Haemulidae 0.47

Groupers Serranidae 13.91

Goatfish Mullidae 0.99

Parrotfish Scaridae 5.83

Lane snapper (L. synagris) Lutjanidae 0.03

Yellowtail snapper (O.chrusurus) Lutjanidae 2.89

Triggerfishes Balistidae 29.68

Squirrelfishes Holocentridae 4.84

Mutton snapper (L. analis) Lutjanidae 0.13

Other snappers Lutjanidae 1.04

Hogfish Labridae 1.06

Trunkfish Ostraciidae 0.08

Table A1. Finfish landings as a proportion of the total finfish landings 

reported for the U.S.Caribbean in 1980. Listed are the most commonly 

landed species and species groups. Data from the Caribbean Fisheries 

Management Council (CFMC, 1985).

CFMC (Caribbean Fisheries Management Council). 1985. Fishery Management Plan, Final Environmental Impact Statement and Draft Regulatory Impact Review for the Shallow 

Water Reef Fish Fishery of Puerto Rico and the U.S. Virgin Islands. San Juan, PR. 179.pp. 

http://www.caribbeanfmc.com/
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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168Appendix BAppendix B

Table B1. Fish species list and summary data on occurrence, abundance and biomass (2001-2007) for the southwest La Parguera, 

Puerto Rico study region.

Family

Species name Common name

%

occurrence

Total

occurrence

Total

abundance

Mean abundance Total

biomass, g

Mean biomass, g

 (+ SE) (+ SE)

Acanthuridae

Acanthurus bahianus ocean surgeonfish 41.9 489 2125 1.8 (0.10) 139867.3 119.9 (8.0)

Acanthurus chirurgus doctorfish 15.6 182 630 0.54 (0.07) 63940.7 54.8 (11.5)

Acanthurus coeruleus blue tang 24.9 291 963 0.83 (0.10) 93316.7 80.0 (17.5)

Acanthurus UNK SURGEONFISH sp 0.2 2 3 <0.01 (<0.01) 2.3 (<0.01) (<0.01)

Apogonidae

Apogon binotatus barred cardinalfish 0.3 3 3 <0.01 (<0.01) 1.4 (<0.01) (<0.01)

Apogon maculatus flamefish 0.3 4 9 (<0.01) (<0.01) 4.1 (<0.01) (<0.01)

Apogon quadrisquamatus sawcheek cardinalfish 0.4 5 10 (<0.01) (<0.01) 4.6 (<0.01) (<0.01)

Apogon townsendi belted cardinalfish 0.3 3 5 <0.01 (<0.01) 2.3 (<0.01) (<0.01)

Apogon UNK CARDINALFISH sp 0.2 2 2 <0.01 (<0.01) 0.92 (<0.01) (<0.01)

Astrapogon stellatus conchfish 0.1 1 2 <0.01 (<0.01) 1.0 (<0.01) (<0.01)

Atherinidae

Atherinomorus UNK SILVERSIDE sp 1.4 16 19533 16.7 (6.0) 5155.0 4.4 (1.6)

Aulostomidae

Aulostomus maculatus trumpetfish 4.0 47 48 0.04 (<0.01) 4554.5 3.9 (0.71)

Balistidae

Balistes vetula queen triggerfish 5.1 60 77 0.07 (0.01) 34089.5 29.2 (4.9)

Melichthys niger black durgon 2.4 28 193 0.17 (0.05) 69209.4 59.3 (17.5)

Belonidae

Belonidae UNK NEEDLEFISH Family sp 0.2 2 5 <0.01 (<0.01) 84.7 0.07 (0.07)

Tylosurus crocodilus crocodilus houndfish 0.1 1 4 <0.01 (<0.01) 164.7 0.14 (0.14)

Blenniidae

Ophioblennius macclurei redlip blenny 2.4 28 92 0.08 (0.02) 252.3 0.22 (0.05)

Bothidae

Bothus lunatus peacock flounder 0.3 3 3 <0.01 (<0.01) 393.4 0.34 (0.33)

Bothus ocellatus eyed flounder 0.3 3 3 <0.01 (<0.01) 12.4 0.01 (<0.01)

Bothus UNK LEFTEYE FLOUNDER sp 0.3 4 4 <0.01 (<0.01) 18.5 0.02 (<0.01)

Callionymidae

Paradiplogrammus bairdi lancer dragonet 1.5 17 21 0.02 (<0.01) 14.9 0.01 (<0.01)

Carangidae

Carangoides bartholomaei yellow jack 0.3 3 29 0.02 (0.02) 12943.0 11.1 (10.6)

Caranx crysos blue runner 1.8 21 94 0.08 (0.03) 38365.9 32.9 (11.5)

Caranx latus horse-eye jack 1.2 14 40 0.03 (0.01) 2371.5 2.0 (1.4)

Caranx lugubris black jack 0.2 2 2 <0.01 (<0.01) 4059.9 3.5 (2.6)

Carangoides ruber bar jack 12.3 144 861 0.74 (0.15) 13993.1 12.0 (2.2)

Caranx UNK JACK sp 0.1 1 2 <0.01 (<0.01) 205.5 0.18 (0.18)

Decapterus macarellus mackerel scad 0.9 11 428 0.37 (0.19) 13947.1 12.0 (4.9)

Decapterus UNK SCAD sp 0.3 4 260 0.22 (0.12) 8884.3 7.6 (3.9)

Selar crumenophthalmus bigeye scad 0.3 3 162 0.14 (0.11) 17212.0 14.7 (10.4)

Trachinotus goodei palometa 0.1 1 2 <0.01 (<0.01) 1021.8 0.88 (0.88)

Carcharhinidae

Galeocerdo cuvier tiger shark 0.1 1 1 <0.01 (<0.01) 92976.9 79.7 (79.7)

Chaenopsidae

Acanthemblemaria aspera roughhead blenny 0.1 1 1 <0.01 (<0.01) 0.20 (<0.01) (<0.01)

Acanthemblemaria maria secretary blenny 2.3 27 43 0.04 (<0.01) 8.6 (<0.01) (<0.01)

Acanthemblemaria spinosa spinyhead blenny 0.5 6 6 (<0.01) (<0.01) 1.2 (<0.01) (<0.01)

Acanthemblemaria UNK TUBE BLENNY sp 2.1 24 35 0.03 (<0.01) 7.0 (<0.01) (<0.01)

Chaenopsis limbaughi yellowface pikeblenny 1.0 12 12 0.01 (<0.01) 13.6 0.01 (<0.01)

Chaenopsis ocellata bluethroat pikeblenny 1.1 13 29 0.02 (0.01) 75.5 0.06 (0.03)

Chaenopsis UNK PIKEBLENNY sp 0.4 5 6 (<0.01) (<0.01) 4.0 (<0.01) (<0.01)

Emblemaria pandionis sailfin blenny 0.9 10 11 (<0.01) (<0.01) 2.2 (<0.01) (<0.01)

Emblemariopsis UNK BLENNY sp 0.6 7 8 (<0.01) (<0.01) 1.6 (<0.01) (<0.01)

Chaetodontidae

Chaetodon capistratus foureye butterflyfish 42.2 493 1526 1.3 (0.06) 23585.6 20.2 (1.5)

Chaetodon ocellatus spotfin butterflyfish 0.5 6 6 (<0.01) (<0.01) 159.1 0.14 (0.07)

Chaetodon sedentarius reef butterflyfish 0.3 4 4 <0.01 (<0.01) 26.2 0.02 (0.01)

Chaetodon striatus banded butterflyfish 5.4 63 93 0.08 (0.01) 2358.6 2.0 (0.37)

Prognathodes aculeatus longsnout butterflyfish 0.2 2 2 <0.01 (<0.01) 13.8 0.01 (0.01)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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169Appendix B

Table B1 cont...

Family

Species name Common name

%

occurrence

Total

occurrence

Total

abundance

Mean abundance Total

biomass, g

Mean biomass, g

 (+ SE) (+ SE)

Cirrhitidae

Amblycirrhitus pinos redspotted hawkfish 0.6 7 7 (<0.01) (<0.01) 5.7 (<0.01) (<0.01)

Clupeidae

Clupeidae UNK HERRING Family sp 2.2 26 5897 5.1 (2.04) 2084.2 1.8 (1.4)

Jenkinsia UNK HERRING sp 6.9 80 130548 111.9 (23.5) 6268.8 5.4 (1.1)

Congridae

Heteroconger longissimus brown garden eel 1.2 14 342 0.29 (0.11) 12233.7 10.5 (4.6)

Dactylopteridae

Dactylopterus volitans flying gurnard 0.1 1 1 <0.01 (<0.01) 5.1 (<0.01) (<0.01)

Dasyatidae

Dasyatis americana southern stingray 0.3 3 3 <0.01 (<0.01) 1163.2 1.0 (0.70)

Diodontidae

Diodon holocanthus balloonfish 4.4 51 76 0.07 (0.01) 3691.2 3.2 (1.3)

Diodon hystrix porcupinefish 2.0 23 28 0.02 (<0.01) 26105.0 22.4 (7.6)

Echeneidae

Echeneis naucrates sharksucker 1.1 13 18 0.02 (<0.01) 4559.2 3.9 (2.4)

Echeneis neucratoides whitefin sharksucker 1.1 13 15 0.01 (<0.01) 1558.3 1.3 (0.42)

Engraulidae

Engraulidae UNK ANCHOVIES Family sp 0.2 2 1160 0.99 (0.87) 181.7 0.16 (0.14)

Ephippidae

Chaetodipterus faber atlantic spadefish 0.2 2 59 0.05 (0.05) 4301.6 3.7 (3.5)

Gerreidae

Eucinostomus gula silver jenny 3.8 44 109 0.09 (0.02) 1129.8 0.97 (0.20)

Eucinostomus melanopterus flagfin mojarra 8.2 96 468 0.40 (0.07) 674.1 0.58 (0.15)

Eucinostomus UNK MOJARRA sp 0.5 6 20 0.02 (<0.01) 37.8 0.03 (0.02)

Gerres cinereus yellowfin mojarra 9.3 108 397 0.34 (0.05) 3095.8 2.7 (0.43)

Ginglymostomatidae

Ginglymostoma cirratum nurse shark 0.3 4 4 <0.01 (<0.01) 30007.4 25.7 (20.0)

Gobiidae

Bathygobious soporator frillfin goby 0.3 4 8 (<0.01) (<0.01) 36.5 0.03 (0.02)

Bollmannia boqueronensis white-eye goby 0.8 9 14 0.01 (<0.01) 9.8 (<0.01) (<0.01)

Coryphopterus dicrus colon goby 4.2 49 91 0.08 (0.01) 59.8 0.05 (<0.01)

Coryphopterus eidolon pallid goby 1.3 15 24 0.02 (<0.01) 15.8 0.01 (<0.01)

Coryphopterus glaucofraenum bridled goby 28.1 328 1758 1.5 (0.15) 1292.5 1.1 (0.13)

Coryphopterus lipernes peppermint goby 1.8 21 91 0.08 (0.02) 59.7 0.05 (0.01)

Coryphopterus personatus/

hyalinus

masked/glass goby 6.0 70 1299 1.1 (0.24) 851.4 0.73 (0.16)

Microgobius UNK GOBY sp 0.1 1 4 <0.01 (<0.01) 0.85 (<0.01) (<0.01)

Ctenogobius saepepallens dash goby 9.4 110 1307 1.1 (0.24) 856.7 0.73 (0.16)

Ctenogobius stigmaticus marked goby 0.3 3 26 0.02 (0.02) 17.0 0.01 (0.01)

Elacatinus chancei shortstripe goby 1.8 21 32 0.03 (<0.01) 8.0 (<0.01) (<0.01)

Elacatinus dilepis orangesided goby 2.0 23 59 0.05 (0.02) 14.8 0.01 (<0.01)

Elacatinus evelynae sharknose goby 20.1 235 605 0.52 (0.05) 151.6 0.13 (0.01)

Elacatinus saucrus leopard goby 0.2 2 8 (<0.01) (<0.01) 2.0 (<0.01) (<0.01)

Coryphopterus UNK GOBY sp 0.1 1 2 <0.01 (<0.01) 1.3 (<0.01) (<0.01)

Gnatholepis thompsoni goldspot goby 19.5 228 1068 0.92 (0.09) 388.3 0.33 (0.09)

Gobiidae UNK GOBY Family sp 1.7 20 227 0.19 (0.08) 148.8 0.13 (0.05)

Gobiosoma grosvenori rockcut goby 0.2 2 2 <0.01 (<0.01) 1.3 (<0.01) (<0.01)

Lophogobius cyprinoides crested goby 0.7 8 21 0.02 (<0.01) 139.0 0.12 (0.10)

Microgobius carri seminole goby 3.4 40 240 0.21 (0.06) 66.8 0.06 (0.02)

Microgobius signatus microgobius signatus 0.1 1 1 <0.01 (<0.01) 0.21 (<0.01) (<0.01)

Elacatinus UNK GOBY sp 0.1 1 1 <0.01 (<0.01) 0.25 (<0.01) (<0.01)

Nes longus orangespotted goby 15.3 179 1502 1.3 (0.13) 4391.5 3.8 (0.48)

Oxyurichthys stigmalophius spotfin goby 1.5 18 48 0.04 (0.01) 135.8 0.12 (0.04)

Priolepis hipoliti rusty goby 0.4 5 5 <0.01 (<0.01) 1.9 (<0.01) (<0.01)

Grammatidae

Gramma loreto fairy basslet 7.5 88 503 0.43 (0.07) 251.8 0.22 (0.04)

Haemulidae

Anisotremus virginicus porkfish 6.5 76 121 0.10 (0.01) 7769.2 6.7 (1.3)

Haemulon aurolineatum tomtate 9.1 106 2016 1.7 (0.68) 21637.9 18.5 (7.5)

Haemulon carbonarium caesar grunt 1.8 21 46 0.04 (0.01) 2236.6 1.9 (0.50)

Haemulon chrysargyreum smallmouth grunt 1.4 16 115 0.10 (0.05) 1865.6 1.6 (0.53)
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170Appendix BTable B1 cont...

Family

Species name Common name

%

occurrence

Total

occurrence

Total

abundance

Mean abundance

 (+ SE)

Total

biomass, g

Mean biomass, g

 (+ SE)

Haemulon flavolineatum French grunt 27.2 317 5114 4.4 (0.58) 65514.5 56.1 (8.4)

Haemulon macrostomum Spanish grunt 0.5 6 6 (<0.01) (<0.01) 305.0 0.26 (0.14)

Haemulon parra sailors choice 2.3 27 72 0.06 (0.02) 2186.9 1.9 (0.62)

Haemulon plumierii white grunt 15.1 176 846 0.72 (0.13) 31558.1 27.0 (5.1)

Haemulon sciurus bluestriped grunt 18.2 212 2238 1.9 (0.23) 75884.2 65.0 (10.2)

Haemulon UNK GRUNT sp 10.0 117 5104 4.4 (1.07) 2045.1 1.8 (0.41)

Holocentridae

Holocentrus adscensionis squirrelfish 5.7 67 127 0.11 (0.02) 16165.8 13.9 (2.8)

Holocentrus rufus longspine squirrelfish 28.1 328 784 0.67 (0.07) 65778.2 56.4 (5.1)

Myripristis jacobus blackbar soldierfish 3.8 44 113 0.10 (0.02) 10040.1 8.6 (2.4)

Neoniphon marianus longjaw squirrelfish 0.7 8 9 (<0.01) (<0.01) 223.3 0.19 (0.08)

Sargocentron coruscum reef squirrelfish 0.2 2 3 <0.01 (<0.01) 76.0 0.07 (0.06)

Sargocentron vexillarium dusky squirrelfish 0.7 8 10 (<0.01) (<0.01) 190.9 0.16 (0.08)

Kyphosidae

Kyphosus sectatrix chub (bermuda/yellow) 0.3 3 3 <0.01 (<0.01) 126.4 0.11 (0.10)

Labridae

Bodianus rufus Spanish hogfish 3.0 35 47 0.04 (<0.01) 4071.8 3.5 (0.88)

Clepticus parrae creole wrasse 2.0 23 506 0.43 (0.17) 14832.7 12.7 (6.6)

Doratonotus megalepis dwarf wrasse 0.3 3 4 <0.01 (<0.01) 0.92 (<0.01) (<0.01)

Halichoeres bivittatus slippery dick 27.1 316 1989 1.7 (0.14) 12810.1 11.0 (1.2)

Halichoeres cyanocephalus yellowcheek wrasse 0.1 1 1 <0.01 (<0.01) 0.30 (<0.01) (<0.01)

Halichoeres garnoti yellowhead wrasse 27.7 323 2127 1.8 (0.14) 13213.9 11.3 (0.91)

Halichoeres maculipinna clown wrasse 8.7 101 242 0.21 (0.03) 1504.9 1.3 (0.25)

Halichoeres pictus rainbow wrasse 0.6 7 25 0.02 (0.01) 198.4 0.17 (0.09)

Halichoeres poeyi blackear wrasse 14.7 171 547 0.47 (0.05) 2376.2 2.0 (0.24)

Halichoeres radiatus puddingwife 3.4 40 57 0.05 (<0.01) 1096.4 0.94 (0.26)

Lachnolaimus maximus hogfish 3.0 35 43 0.04 (<0.01) 3396.5 2.9 (0.69)

Thalassoma bifasciatum bluehead 34.1 398 5995 5.1 (0.40) 12013.7 10.3 (1.1)

Xyrichtys martinicensis rosy razorfish 3.7 43 586 0.50 (0.27) 1671.7 1.4 (0.89)

Xyrichtys novacula pearly razorfish 0.3 3 6 (<0.01) (<0.01) 20.1 0.02 (0.01)

Xyrichtys splendens green razorfish 8.3 97 388 0.33 (0.05) 1815.1 1.6 (0.32)

Labrisomidae

Labrisomus nuchipinnis hairy blenny 0.2 2 2 <0.01 (<0.01) 8.9 (<0.01) (<0.01)

Malacoctenus aurolineatus goldline blenny 0.5 6 10 (<0.01) (<0.01) 6.8 (<0.01) (<0.01)

Malacoctenus boehlkei diamond blenny 0.7 8 8 (<0.01) (<0.01) 1.9 (<0.01) (<0.01)

Malacoctenus macropus rosy blenny 4.3 50 107 0.09 (0.02) 72.5 0.06 (0.01)

Malacoctenus triangulatus saddled blenny 2.1 25 34 0.03 (<0.01) 11.7 0.01 (<0.01)

Malacoctenus UNK SCALY BLENNY sp 0.3 4 4 <0.01 (<0.01) 1.4 (<0.01) (<0.01)

Malacoctenus versicolor barfin blenny 0.1 1 2 <0.01 (<0.01) 0.48 (<0.01) (<0.01)

Lutjanidae

Lutjanus analis mutton snapper 0.3 4 4 <0.01 (<0.01) 825.3 0.71 (0.47)

Lutjanus apodus schoolmaster 18.8 219 1974 1.7 (0.19) 103463.1 88.7 (11.3)

Lutjanus buccanella blackfin snapper 0.1 1 2 <0.01 (<0.01) 3.0 (<0.01) (<0.01)

Lutjanus cyanopterus cubera snapper 0.1 1 1 <0.01 (<0.01) 2814.4 2.4 (2.4)

Lutjanus griseus gray snapper 8.1 94 741 0.63 (0.15) 56346.6 48.3 (9.6)

Lutjanus jocu dog snapper 0.8 9 13 0.01 (<0.01) 4253.9 3.6 (1.3)

Lutjanus mahogoni mahogany snapper 2.2 26 54 0.05 (0.01) 3221.6 2.8 (0.75)

Lutjanus synagris lane snapper 3.9 46 89 0.08 (0.01) 4069.8 3.5 (0.83)

Lutjanus UNK SNAPPER sp 1.0 12 43 0.04 (0.02) 228.7 0.20 (0.15)

Ocyurus chrysurus yellowtail snapper 27.4 320 738 0.63 (0.05) 71432.6 61.2 (5.5)

Malacanthidae

Malacanthus plumieri sand tilefish 3.7 43 58 0.05 (<0.01) 11013.5 9.4 (2.4)

Microdesmidae

Ptereleotris helenae hovering goby 3.6 42 100 0.09 (0.02) 109.6 0.09 (0.04)

Monacanthidae

Cantherhines macrocerus american whitespotted filefish 0.2 2 7 (<0.01) (<0.01) 321.3 0.28 (0.21)

Cantherhines pullus orangespotted filefish 0.9 10 10 (<0.01) (<0.01) 963.0 0.83 (0.31)

Monacanthus ciliatus fringed filefish 1.2 14 14 0.01 (<0.01) 56.9 0.05 (0.02)

Monacanthus tuckeri slender filefish 1.5 18 20 0.02 (<0.01) 58.6 0.05 (0.02)

Mugilidae

Mugil cephalus striped mullet 0.1 1 3 <0.01 (<0.01) 485.2 0.42 (0.42)
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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171Appendix B

Table B1 cont...

Family

Species name Common name

%

occurrence

Total

occurrence

Total

abundance

Mean abundance

 (+ SE)

Total

biomass, g

Mean biomass, g

 (+ SE)

Mullidae

Mulloidichthys martinicus yellow goatfish 6.6 77 330 0.28 (0.10) 28166.3 24.1 (4.9)

Pseudupeneus maculatus spotted goatfish 16.6 194 412 0.35 (0.06) 19362.4 16.6 (2.2)

Muraenidae

Gymnothorax funebris green moray 0.3 4 4 <0.01 (<0.01) 5129.5 4.4 (3.3)

Gymnothorax miliaris goldentail moray 0.2 2 2 <0.01 (<0.01) 28.4 0.02 (0.02)

Gymnothorax moringa spotted moray 0.7 8 8 (<0.01) (<0.01) 1593.1 1.4 (0.55)

Ophichthidae

Myrichthys breviceps sharptail eel 0.1 1 1 <0.01 (<0.01) 421.9 0.36 (0.36)

Myrichthys UNK SNAKE EEL sp 0.2 2 2 <0.01 (<0.01) 75.4 0.06 (0.06)

Opistognathidae

Lonchopisthus micrognathus swordtail jawfish 1.6 19 47 0.04 (0.01) 187.1 0.16 (0.07)

Opistognathus aurifrons yellowhead jawfish 4.9 57 138 0.12 (0.02) 508.6 0.44 (0.09)

Opistognathus macrognathus banded jawfish 0.2 2 3 <0.01 (<0.01) 8.0 (<0.01) (<0.01)

Opistognathus UNK JAWFISH sp 0.1 1 3 <0.01 (<0.01) 7.9 (<0.01) (<0.01)

Opistognathus whitehursti dusky jawfish 0.1 1 1 <0.01 (<0.01) 3.9 (<0.01) (<0.01)

Ostraciidae

Acanthostracion polygonius honeycomb cowfish 0.1 1 1 <0.01 (<0.01) 0.48 (<0.01) (<0.01)

Acanthostracion quadricornis scrawled cowfish 0.3 4 4 <0.01 (<0.01) 25.0 0.02 (0.02)

Lactophrys bicaudalis spotted trunkfish 0.5 6 6 (<0.01) (<0.01) 3298.8 2.8 (2.3)

Lactophrys trigonus trunkfish 0.1 1 1 <0.01 (<0.01) 75.4 0.06 (0.06)

Lactophrys triqueter smooth trunkfish 1.3 15 15 0.01 (<0.01) 1112.6 0.95 (0.31)

Lactophrys UNK TRUNKFISH sp 0.1 1 1 <0.01 (<0.01) 0.83 (<0.01) (<0.01)

Paralichthyidae

Syacium UNK SAND FLOUNDER sp 0.3 4 5 <0.01 (<0.01) 208.2 0.18 (0.18)

Pomacanthidae

Holacanthus ciliaris queen angelfish 2.8 33 41 0.04 (<0.01) 9250.8 7.9 (2.0)

Holacanthus tricolor rock beauty 2.8 33 37 0.03 (<0.01) 3202.3 2.7 (0.64)

Pomacanthus arcuatus gray angelfish 7.8 91 133 0.11 (0.01) 110334.9 94.5 (13.8)

Pomacanthus paru French angelfish 1.5 17 24 0.02 (<0.01) 10979.9 9.4 (3.9)

Pomacentridae

Abudefduf saxatilis sergeant major 10.4 121 488 0.42 (0.05) 6720.3 5.8 (1.5)

Abudefduf taurus night sergeant 0.3 3 3 <0.01 (<0.01) 218.4 0.19 (0.12)

Chromis cyanea blue chromis 6.1 71 992 0.85 (0.16) 2994.2 2.6 (0.45)

Chromis multilineata brown chromis 3.5 41 327 0.28 (0.08) 1773.2 1.5 (0.45)

Microspathodon chrysurus yellowtail damselfish 10.8 126 398 0.34 (0.05) 29812.5 25.5 (3.8)

Stegastes adustus dusky damselfish 6.9 81 1187 1.0 (0.17) 12003.6 10.3 (1.8)

Stegastes diencaeus longfin damselfish 7.1 83 489 0.42 (0.08) 7110.6 6.1 (1.0)

Stegastes leucostictus beaugregory 26.1 305 2135 1.8 (0.15) 12900.3 11.1 (0.95)

Stegastes partitus bicolor damselfish 38.8 453 4378 3.8 (0.26) 6202.1 5.3 (0.90)

Stegastes planifrons threespot damselfish 15.2 177 1718 1.5 (0.15) 23525.8 20.2 (2.5)

Stegastes variabilis cocoa damselfish 19.7 230 699 0.60 (0.05) 5594.0 4.8 (0.38)

Priacanthidae

Heteropriacanthus cruentatus glasseye snapper 0.4 5 16 0.01 (0.01) 1326.0 1.1 (0.70)

Priacanthus arenatus bigeye 0.1 1 2 <0.01 (<0.01) 245.1 0.21 (0.21)

Scaridae

Cryptotomus roseus bluelip parrotfish 19.6 229 948 0.81 (0.08) 4341.4 3.7 (0.49)

Scarus guacamaia rainbow parrotfish 0.2 2 2 <0.01 (<0.01) 23.3 0.02 (0.01)

Scarus iseri striped parrotfish 45.0 525 6977 6.0 (0.39) 128555.1 110.2 (7.9)

Scarus taeniopterus princess parrotfish 21.1 246 2034 1.7 (0.16) 108626.2 93.1 (10.7)

Scarus UNK PARROTFISH sp 0.3 4 5 <0.01 (<0.01) 297.9 0.26 (0.25)

Scarus vetula queen parrotfish 1.1 13 27 0.02 (<0.01) 5005.8 4.3 (1.9)

Sparisoma atomarium greenblotch parrotfish 13.6 159 504 0.43 (0.05) 1125.6 0.96 (0.38)

Sparisoma aurofrenatum redband parrotfish 42.8 499 3408 2.9 (0.14) 152788.8 130.9 (7.3)

Sparisoma chrysopterum redtail parrotfish 6.8 79 154 0.13 (0.02) 28634.8 24.5 (6.1)

Sparisoma radians bucktooth parrotfish 23.8 278 1625 1.4 (0.13) 3716.3 3.2 (0.48)

Sparisoma rubripinne yellowtail parrotfish 4.4 51 124 0.11 (0.02) 16550.8 14.2 (4.4)

Sparisoma UNK PARROTFISH sp 1.5 18 30 0.03 (<0.01) 67.4 0.06 (0.02)

Sparisoma viride stoplight parrotfish 30.1 351 1314 1.1 (0.08) 229172.5 196.4 (19.8)

Sciaenidae

Equetus lanceolatus jackknife fish 0.1 1 1 <0.01 (<0.01) 0.08 (<0.01) (<0.01)
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172Appendix BTable B1 cont...

Family

Species name Common name

%

occurrence

Total

occurrence

Total

abundance

Mean abundance

 (+ SE)

Total

biomass, g

Mean biomass, g

 (+ SE)

Equetus punctatus spotted drum 0.4 5 5 <0.01 (<0.01) 652.4 0.56 (0.27)

Odontoscion dentex reef croaker 0.3 4 46 0.04 (0.03) 251.6 0.22 (0.14)

Pareques acuminatus highhat 1.6 19 26 0.02 (<0.01) 276.0 0.24 (0.17)

Scombridae

Scomberomorus regalis cero 0.9 11 14 0.01 (<0.01) 23748.6 20.4 (7.5)

Scorpaenidae

Scorpaena plumieri spotted scorpionfish 0.6 7 7 (<0.01) (<0.01) 984.8 0.84 (0.44)

Scorpaena UNK SCORPIONFISH sp 0.1 1 1 <0.01 (<0.01) 9.3 (<0.01) (<0.01)

Serranidae

Alphestes afer mutton hamlet 0.2 2 2 <0.01 (<0.01) 14.7 0.01 (<0.01)

Cephalopholis cruentata graysby 13.8 161 246 0.21 (0.02) 15443.2 13.2 (1.5)

Cephalopholis fulva coney 3.8 44 81 0.07 (0.01) 12244.5 10.5 (2.0)

Diplectrum bivittatum dwarf sand perch 0.5 6 13 0.01 (<0.01) 17.8 0.02 (<0.01)

Diplectrum formosum sand perch 0.1 1 1 <0.01 (<0.01) 0.34 (<0.01) (<0.01)

Epinephelus adscensionis rock hind 0.2 2 2 <0.01 (<0.01) 12.9 0.01 (<0.01)

Epinephelus guttatus red hind 3.6 42 43 0.04 (<0.01) 16774.9 14.4 (2.7)

Epinephelus striatus Nassau grouper 0.2 2 2 <0.01 (<0.01) 1331.3 1.1 (0.91)

Hypoplectrus aberrans yellowbelly hamlet 0.9 10 11 (<0.01) (<0.01) 56.8 0.05 (0.02)

Hypoplectrus chlorurus yellowtail hamlet 9.9 115 157 0.13 (0.01) 1294.1 1.1 (0.14)

Hypoplectrus guttavarius shy hamlet 0.9 11 12 0.01 (<0.01) 64.8 0.06 (0.02)

Hypoplectrus indigo indigo hamlet 0.9 11 12 0.01 (<0.01) 226.0 0.19 (0.08)

Hypoplectrus nigricans black hamlet 1.8 21 22 0.02 (<0.01) 163.4 0.14 (0.04)

Hypoplectrus puella barred hamlet 8.9 104 141 0.12 (0.01) 553.2 0.47 (0.06)

Hypoplectrus unicolor butter hamlet 5.5 64 75 0.06 (<0.01) 550.3 0.47 (0.08)

Hypoplectrus UNK HAMLET sp 4.4 51 67 0.06 (<0.01) 101.9 0.09 (0.03)

Liopropoma rubre peppermint basslet 0.1 1 1 <0.01 (<0.01) 5.8 (<0.01) (<0.01)

Mycteroperca bonaci black grouper 0.2 2 2 <0.01 (<0.01) 2744.4 2.4 (2.1)

Rypticus bistrispinus freckled soapfish 0.1 1 1 <0.01 (<0.01) 5.8 (<0.01) (<0.01)

Rypticus saponaceus greater soapfish 0.2 2 2 <0.01 (<0.01) 164.0 0.14 (0.10)

Serraniculus pumilio pygmy sea bass 0.3 3 3 <0.01 (<0.01) 1.1 (<0.01) (<0.01)

Serranus baldwini lantern bass 7.1 83 253 0.22 (0.03) 216.4 0.19 (0.03)

Serranus tabacarius tobaccofish 5.0 58 107 0.09 (0.02) 482.8 0.41 (0.18)

Serranus tigrinus harlequin bass 8.7 101 150 0.13 (0.01) 726.2 0.62 (0.08)

Serranus tortugarum chalk bass 3.9 45 325 0.28 (0.07) 138.7 0.12 (0.03)

Serranus UNK SEABASS sp 1.2 14 26 0.02 (<0.01) 25.7 0.02 (0.01)

Sparidae

Archosargus rhomboidalis sea bream 3.3 39 135 0.12 (0.03) 7890.1 6.8 (1.8)

Calamus calamus saucereye porgy 1.4 16 18 0.02 (<0.01) 2597.3 2.2 (0.73)

Calamus penna sheepshead porgy 0.1 1 1 <0.01 (<0.01) 8.3 (<0.01) (<0.01)

Calamus pennatula pluma 4.0 47 61 0.05 (<0.01) 14349.1 12.3 (3.5)

Calamus UNK PORGY sp 0.2 2 3 <0.01 (<0.01) 77.7 0.07 (0.05)

Diplodus holbrooki spottail pinfish 0.1 1 5 <0.01 (<0.01) 518.3 0.44 (0.44)

Sphyraenidae

Sphyraena barracuda great barracuda 10.4 121 209 0.18 (0.02) 160161.4 137.2 (32.3)

Sphyraena picudilla southern sennet 0.2 2 54 0.05 (0.04) 39372.0 33.7 (28.0)

Stromateidae

Stromateidae UNK BUTTERFISH Family sp 0.5 6 16 0.01 (<0.01) 6.3 (<0.01) (<0.01)

Syngnathidae

Cosmocampus elucens shortfin pipefish 0.1 1 1 <0.01 (<0.01) 1.2 (<0.01) (<0.01)

Hippocampus reidi longsnout seahorse 0.1 1 1 <0.01 (<0.01) 0.04 (<0.01) (<0.01)

Synodontidae

Synodus intermedius sand diver 5.2 61 63 0.05 (<0.01) 3476.1 3.0 (0.85)

Tetraodontidae

Canthigaster rostrata sharpnose puffer 25.0 292 484 0.41 (0.03) 1072.9 0.92 (0.10)

Sphoeroides spengleri bandtail puffer 3.1 36 40 0.03 (<0.01) 993.5 0.85 (0.32)

Sphoeroides testudineus checkered puffer 4.4 51 78 0.07 (0.01) 6249.7 5.4 (1.1)

Triglidae

Triglidae UNK searobin family 0.1 1 1 <0.01 (<0.01) 21.5 0.02 (0.02)

Tripterygiidae

Enneanectes UNK TRIPLEFIN sp 0.3 3 3 <0.01 (<0.01) 1.2 (<0.01) (<0.01)
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173Appendix C

Appendix C

Species Common name

Approx. size 

class first 

maturity*

Juv./subadult size 

class (cm)

Max.

known 

size*, TL

Max. size

 observed 

PR, FL

Serranids

Cephalopholis cruentata graysby 15-20 <15 42.6 30

Cephalopholis fulva coney 15-20 <15 41 30

Epinephelus guttatus red hind 20-25 <20 76 40

Lutjanids

Lutjanus apodus schoolmaster 20-25 <20 67.2 45

Lutjanus griseus gray snapper 25-30 <25 89 65

Lutjanus mahogoni mahogany snapper 15-20 <15 48

Lutjanus synagris lane snapper 20-25 <20 89 65

Ocyurus chrysurus yellowtail snapper 20-25 <20 86.3 40

Haemulids

Haemulon aurolineatum tomtate 15-20 <15 25 25

Haemulon flavolineatum French grunt 15-20 <15 30 30

Haemulon plumierii white grunt 15-20 <15 53 30

Haemulon sciurus bluestriped grunt 15-20 <15 46 35

Scarids

Scarus iseri striped parrotfish 10-15 <10 35 35

Scarus taeniopterus princess parrotfish 10-15 <10 35 35

Sparisoma aurofrenatum redband parrotfish 10-15 <10 28 35

Sparisoma viride stoplight parrotfish 15-20 <15 64 50

Other species

Mulloidichthys martinicus yellow goatfish 15-20 <15 35.4 25

Pseudupeneus maculatus spotted goatfish 15-20 <15 27.8 25

Balistes vetula queen triggerfish 20-25 <20 47.1 30

Sphyraena barracuda great barracuda n/a n/a 200 130

Table C1. Summary information on selected species from five key fish families 

showing maximum size observed in the SW Puerto Rico study region compared with 

maximum known size for the species and the proportion of juveniles found based on 

1167 samples from 2001-2007. * Maximum known fish size and size at first maturity 

are from FishBase (http://www.fishbase.org). TL=total length; FL= fork length.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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174Appendix DAppendix D

Methods

To assist in monitoring coral reef ecosystem resources and to achieve a better understanding of fish-habitat 

relationships in the U.S. Caribbean, CCMA-BB developed a fish and macro-invertebrate monitoring protocol 

to provide precise, fishery-independent and size-structured survey data, needed to comprehensively assess 

faunal populations and communities (Menza et al., 2006). In addition, a complementary benthic composition 

survey was also developed to support studies of fish-habitat relationships. These data collection activities and 

analytical products are core components of NOAA’s Coral Reef Conservation Program (CRCP) implemented 

through CCMA-BB’s CREM project. CREM protocols were created primarily to quantify long-term changes in 

fish species and assemblage diversity, abundance, biomass and size structure and to compare these metrics 

between areas inside and outside of Marine Protected Areas (MPAs). A stratified random sampling design was 

used to optimize the allocation of samples and allow rigorous inferences to the entire study area. Three strata 

were selected based upon: 1) the study objectives; 2) parsimony in the approach; and 3) results from statistical 

analyses of variance (Menza et al., 2006). The “hard” stratum comprised bedrock, pavement, rubble and coral 

reefs. The “soft” stratum comprised sand, seagrasses and macroalgal beds. The “mangrove” stratum comprised 

the seaward edge of mangrove habitat able to be surveyed with these underwater methods (Figure 1.8, pg. 8).

Field survey methods

This report uses underwater census data collected from 2001 to 2007. Survey missions occurred each year 

during all seasons and were standardized to Winter and Summer starting in 2005 (Table 2.2, pg. 14). This data 

set is part of a broader ongoing monitoring study that began in 2001, with over 1,000 transects surveyed thus 

far throughout the La Parguera study region. There are two complementary components to the biological field 

methods: (1) benthic habitat composition surveys and (2) fish surveys.

Benthic habitat composition surveys

To conduct benthic habitat surveys, an observer places a 1 m2 quadrat divided into 100 (10 x 10 cm) smaller 

squares (1 square = 1% cover) at five randomly pre-selected locations along the transect, such that a quadrat 

is placed once somewhere within every 5 m interval along the transect (see schematic below). Percent cover is 

estimated within the quadrat in a two-dimensional plane perpendicular to the observer’s line of vision.

Schematic of fish transect with random habitat quadrat placement.

4m

0m 5m 10m 15m 20m 25m

1 m2 quadrat

Information recorded includes:

Habitat structure (e.g., colonized hardbottom, spur and groove, patch reef, pavement) - based on the habitat 

types used in the benthic habitat maps (Kendall et al., 2002; Figure 1.8 pg. 8), until 2004, after which habitat 

structure was classified only to hard, soft and mangrove.

Abiotic footprint - defined as the percent cover (to the nearest 1%) of sand, rubble, hardbottom, fine sediments 

and other non-living bottom types within a 1 m2

 quadrat. 
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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175Appendix D

Biotic footprint - defined as the percent cover to the nearest 1% of algae, seagrass, upright sponges, gorgonians 

and other biota and to the nearest 0.1% for live, bleached and recently dead/diseased coral within a 1 m2 quadrat.

Transect depth profile - the depth at each quadrat position. Depth is measured with a digital depth gauge and 

rounded up or down to the nearest foot.

Maximum canopy height - for each biota type, height of soft structure (e.g., gorgonians, upright sponges, seagrass, algae) is recorded to the nearest 1 cm.

Hardbottom rugosity - measured by placing a 6-m chain at two randomly selected start positions ensuring no 

overlap along 25-m belt transect. The chain is placed such that it follows the relief along centerline of the belt 

transect. Two divers measure the straight-line horizontal distance covered by the chain.

Proximity of structure - on seagrass and sand sites, the habitat diver records the absence or presence of reef or 

hard structure within 3 m of the belt transect. 

Table 2.1 (pg. 13) provides a list of measured variables. The habitat observer also counts queen conch (Eustrombus gigas), long-spined sea urchins (Diadema antillarum) and Caribbean spiny lobster (Panulirus argus). 

Mangrove habitat data

At mangrove sites, an observer swims close to the prop roots and surveys as far into the mangroves as possible, 

up to 2 m and then out to the edge of the mangrove overhang such that the total area surveyed is still 100 m2

. In 

this case, some of the survey may necessarily fall on seagrass habitat. This is allowed as the mangrove habitat 

is defined as a transition zone habitat. In addition to the habitat data collected above, further mangrove data are 

collected including number of prop roots, number of prop roots colonized by algae,number of prop roots colonized by sponges and nujmber of prop roots colonized by other (tunicates, anemones, zooanthids, etc).

Fish surveys

Fish surveys were conducted along a 25 m long by 4 m wide belt (100 m2) using a fixed survey duration of 15 

minutes (Figure 3.1, pg. 56). The fixed duration of 15 minutes standardizes the samples collected to facilitate 

between-site comparisons. The number of individuals per species is recorded in 5 cm size class increments up 

to 35 cm using the visual estimation of fork length. Individuals greater than 35 cm are recorded as an estimate 

of the actual fork length to the nearest centimeter.

Macroinvertebrates counts

Queen conch

The abundance of immature and mature queen conch (Eustrombus gigas) was assessed and quantified within 

the 25 x 4 m belt transects used for fish surveys. The maturity of each conch was determined by the presence 

(mature) or absence (immature) of a flared lip (pg. 46). Conch were included in the survey protocol from August 

2004 onward.

Caribbean spiny lobster

Abundance of Caribbean spiny lobsters (Panulirus argus) was reported for the period 2005 to 2007. Lobster 

sightings were recorded during fish and benthic composition surveys (i.e., within the 100 m2

 survey unit area). 

Lobsters were recorded if seen, but without active searches of holes or crevices. 

Long-spined sea urchins

Long-spined sea urchins (Diadema antillarum) were counted within the 25 x 4 m belt transect during 2006 and 

2007. No measurements of size or estimates of maturity were collected.

Marine debris data

Type of marine debris within 25 x 4 m belt transect was noted. The size of the marine debris and the area of affected habitat is also recorded along with a note identifying any flora or fauna that colonized the debris. Marine 

debris data collection began in 2007.
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

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176Appendix DObserver training

Observers were trained and tested in the identification of species/groups for both fish and habitat surveys by 

pairing inexperienced and experienced observers in the water and comparing data. Fish size estimation training 

was carried out in situ by estimating lengths of model fish of various shapes and sizes.

Data management

All fish and benthic habitat survey data were quality assessed before storage on an online relational database. 

All survey data were stored with a unique identification number and a geographical coordinate to facilitate spatial 

analyses. The database (including metadata) that provides detailed field methods are available online: http://

ccmaserver.nos.noaa.gov/ecosystems/coralreef/reef_fish/protocols.html.

Although the 1m-square-quadrat remained the basic method of choice for habitat data collection, overtime, 

changes in data collection methods were made for some habitat variables and several additional variables were 

added. These changes were deemed necessary to capture more precise information and as many variables as 

possible to explain better the observed variability in reef fish assemblage metrics. Detailed information on all 

changes to the protocols for collecting habitat data in Puerto Rico can be found at: http://ccmaserver.nos.noaa.

gov/ecosystems/coralreef/reef_fish/protocols.html. A brief list is included below:

Over time, some changes were made to the stratified random site selection process as follows: 1) Habitat strata 

initially consisted of hard bottom, sand, seagrass and mangrove. Sand and seagrass strata were subsequently 

combined into one soft bottom strata. This action was taken after the February 2002 mission. 2) In addition to 

the habitat strata, Puerto Rico originally contained three strata representing levels of protection from waves and 

currents. These strata were the Bank Shelf, Outer Lagoon and Inner Lagoon. This was changed beginning with 

the December 2002 mission to simply Protected and Unprotected. After the January 2005 mission, strata of 

Protected and Unprotected was removed leaving only habitat strata. 3) A small subset of sites were resampled 

during each mission through June 2002. These station names contain the letter ‘P’ indicating they are permanent 

stations. 

In 2007, algae data collection changed from identification of each alga to the genus level to grouping algae into 

six morphological groups: macro, turf, crustose, filamentous, rhodolith, and cyanobacteria for more efficient data 

collection. 

References

Menza, C., J. Ault, J. Beets, C. Bohnsack, C. Caldow, J. Christensen, A. Friedlander, C. Jeffrey, M. Kendall, J. 

Luo, M.E. Monaco, S. Smith, and K. Woody. 2006. A guide to monitoring reef fish in the National Park Service’s 

South Florida/Caribbean Network. NOAA Technical Memorandum NOS NCCOS 39. Silver Spring, MD. 166 pp. 

http://ccma.nos.noaa.gov/news/feature/FishMonitoring.html. 

Kendall, M.S., C.R. Kruer, K.R. Buja, J.D. Christensen, M. Finkbeiner, R. Warner, and M.E. Monaco. 2002. Methods used to map the benthic habitats of Puerto Rico and the U.S. Virgin Islands. NOAA Technical Memorandum 

152. Silver Spring, MD. http://ccma.nos.noaa.gov/products/biogeography/usvi_pr_mapping/manual.pdf 
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007)

page

177Appendix E

Appendix E

Figure E1. Spatial predictive map of fish species richness across the seascapes of La Parguera developed 

from the relationship between the number of species observed via underwater fish surveys and the underlying 

topographic complexity derived from chain rugosity measurements and historical depth soundings. The predictive 

map was then overlaid on the more recent LiDAR bathymetry. Modeling was performed using a regression tree 

as documented in Pittman et al. (2007a) with a 75% overall map accuracy.


United States Department of Commerce

Gary Locke

Secretary

National Oceanic and Atmospheric Administration

Jane Lubchenco

Acting Undersecretary of Commerce for Oceans and Atmosphere

National Ocean Service

Jack H Dunnigan

Assistant Administrator
Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico) - NOAA Technical Memorandum NOS NCCOS 107