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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) 

A cooperative investigation between NOAAand the University of Puerto Rico 





** BRANCH y 



NOAA Technical MemorandunfNOS NCCOS 107 




Center for Coastal Monitoring 
and Assessment 



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 Pittman 1 - 2 - 3 , Sarah D Hile 1 3 , Christopher FG Jeffrey 1 - 3 , Randy Clark 1 , Kimberly Woody 1 , 
Brook D Herlach 1,3 , Chris Caldow 1 , Mark E Monaco 1 , Richard Appeldoorn 4 



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, Mayaguez, 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 




«*%®5?» 




Center for Coastal Monitoring 
and Assessment 



United States Department of 


National Oceanic and 


National Ocean Service 


Commerce 


Atmospheric Administration 




Gary Locke 


Jane Lubchenco 


Jack Dunnigan 


Secretary 


Administrator 


Assistant Administrator 













wW. 






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

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 multi- 
use 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 - RN 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. 




NOAA 

CORAL REEF 

CONSERVATION PROGRAM 



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

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) 

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 offish 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 m 2 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) 

The raw data were summarized by habitat type to examine differences in richness, abundance and 
CO 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 

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

CD 

> Major findings: 

"t± Diversity hotspots 

O • Highest coral cover was observed along the shelf edge southward of Terrumote I and around El Palo 
(D 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 m 2 ) 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 m 2 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 Pontes spp. were the most common stony corals, with M. annularis complex 
comprising 35% of total coral cover, and Pontes 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) 

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 



^ 



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



Total fish biomass, herbivore biomass, grouper abundance, parrotfish and wrasse abundance were 
CO significantly higher in 2007 than in 2001 



E 
E 



o 

CD 
X 

LU 



Grouper biomass in 2007 was significantly lower than in 2001 



tr\ * ^ n 'y ^' ^ va anc ' ^' aur °f rena t um increased in abundance consecutively for more than three years 

Uj between 2001 and 2007 

CD 

> • 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. /sen 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) 

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. 



page 
ix 



03 



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

Resumen 



Desde el 1999 el programa de Bio-geografia del Centro para Monitoreo y Evaluation Costera (CCMA- 

C BB por sus siglas en ingles) de la Administration National de Oceanos y Atmosfera (NOAA por sus 

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

tr\ y eva ' uar ' a condition del ambiente marino del suroeste de Puerto Rico. Este esfuerzo es parte del 

J ^ Programa de Conservation de Arrecifes de Coral (CRCP por sus siglas en ingles) bajo el Programa 

Q) National de Monitoreo de Ecosistemas de Arrecifes de Coral (NCREMP por sus siglas en ingles) de 

b > NOAA. Con el apoyo del NCREMP del CRCP el CCMA Neva a cabo el proyecto titulado "Monitoreo de 

~ti Ecosistemas de Arrecifes de Coral del Caribe" (CREM por sus siglas en ingles) cuyas metas incluyen: 

r2 (1 ) caracterizar espacialmente y monitorear la distribution, abundancia y tamano de la fauna asociada 

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

^ arenales y manglares); (2) relacionar esta information con datos de habitat a pequena escala y la 

|| | distribution y diversidad de tipos de habitat utilizando mapas de habitat benticos; (3) utilizar esta 

information 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 coleccion y distribution de datos de monitoreo. Los esfuerzos de monitoreo de la region de La 

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

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

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

En tiempos recientes, observaciones cientfficas y no-cientificas han indicado que la estructura y 
funcionamiento del ecosistema de arrecifes de coral de la region 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-explotacion pesquera, 
mortandades significativas del coral Acropora por enfermedades y eventos de blanqueamientos, el 
mas severo de estos en el 2005. El area es tambien un importante recurso recreativo donde se llevan 
a cabo actividades relacionadas con el mar incluyendo la navegacion, 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 dinamica temporal y la distribution espacial de la fauna marina. Este estudio provee 
datos ecologicos que sirven para evaluar la condition del ecosistema, apoyan la toma de decisiones en 
planificacion espacial (incluyendo la evaluation de la efectividad de estrategias de manejo) y permite 
identificar las necesidades de information futuras. La meta fundamental de este trabajo es lograr 
entender el ecosistema de arrecifes de coral y proveer information 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 tecnico contiene un analisis de los primeros siete anos (2001-2007) de censos 
de peces y caracterizacion bentica. El objetivo principal fue cuantificar cambios en la diversidad, 
abundancia, biomasa y estructura de tamahos de los peces y proveer information espacial explfcita de 
la distribution de especies claves y grupos espeefficos. Esta information sirve a su vez para comparar 
la estructura de la comunidad a lo largo del paisaje marino compuesto de multiples habitats, incluyendo 
los manglares de franja, arrecifes en distintas posiciones de la plataforma y areas de plataforma 
oceanica. 

Metodos: 

Para cada mision de muestreo semi-anual se seleccionaron sitios mediante un diseno aleatorio 
estratificado entre habitats de fondos duros y blandos (arenosos) segun el mapa de habitat bentico de 
NOAA (Menza et al., 2006). Los censos de peces se llevaron a cabo durante el dfa 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) 

banda de 25 por 4 metros durante 15 minutos. Todas las especies observadas se identificaron al nivel 
taxonomico mas bajo posible y las abundancias se cuantificaron en clases de tamanos. Para estimar 
la composicion del habitat bentico, cinco cuadratas de un metro cuadrado se colocaron al azar en 
el transecto y se cuantificaron los componentes bioticos y abioticos (por ejemplo cobertura de coral, 
algas, etc.). Para este analisis 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 habitat para examinar diferencias en la riqueza, abundancia y 
biomasa de peces en distintos habitats benticos y se mapearon a lo largo del area de estudio para 
examinar patrones en la distribucion de los atributos de comunidades y especies. Se resumieron cambios 
entre anos y por temporadas durante la duracion del estudio para identificar mudanzas significativos 
entre periodos de muestreo. La sfntesis de estos datos provee una base biologica mediante: 1) la 
caracterizacion espacial comprensiva de los recursos y la condicion del habitat y 2) los patrones de 
cambios en abundancias de peces y los componentes benticos de habitats en el area de estudio. 

Resultados sobresalientes: 

Centros de diversidad 

• La cobertura de coral mas alta se observo a lo largo del veril al sur de Cayo Turrumote I y en areas 
circundantes al Cayo El Palo. La cobertura de Acropora palmata (4. 1-1 0.4%) fue mayor en areas de 
Cayo El Palo mientras la cobertura del complejo de Montastraea annularis (25.1-30.3%) fue mayor 
al suroeste de Cayo Turrumote I 

• Puntosdealta riqueza de especies (10-14 especies por 1 00m 2 ) y diversidad (H = 1.7-2.5) de corales 
se observaron lejos de la costa en distintas areas de habitats de fondos duros 

• La rugosidad mayor (0.81-1) se observo en un area entre el Cayo Margarita y Cayo El Palo al igual 
que en Cayo Romero 

• Se detectaron areas donde la riqueza de especies de peces fue mas alta (41 especies por 100m 2 ) 
cerca del veril en habitat 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 herbfvoros coincidieron a lo largo del veril y alrededor del grupo de arrecifes 
de parche localizados entre el Cayo Margarita y Cayo El Palo 

Habitat bentico 

• Los habitats duros tuvieron una cobertura bentica dominada por algas con un promedio de 5% de 
cobertura de coral vivo a lo largo del area de estudio 

• Los habitats duros tuvieron una cobertura bentica dominada por algas con un promedio de 5% de 
cobertura de coral vivo a lo largo del area de estudio 

• Los corales mas comunes pertenecen a los generos Montastraea y Porites, donde M. annularis 
compuso el 35% de la cobertura total y Pontes astreoides se observo mas frecuentemente en 62% 
de los sitios de muestreo de habitats duros 

• Ejemplares del genero Acropora fueron observados muy pocas veces y entre estos Acropora 
cervicornis tuvo una distribucion espacial mas amplia que A. palmata 



^ 



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



El analisis temporal revelo que la cobertura de coral vivo vario significativamente entre algunos 
CO anos, pero en general declino a lo largo del muestreo (2001-2007) 



E 
E 



CD 
> 



Las rafces de mangles sumergidas proveen un sustrato bentico para la colonizacion de especies 
epffitas 



Peces 



Alo largo del area de estudio en La Parguera se identificaron peces de 210 especies perteneciente 
a 14 generos 



q • Las densidades mas altas se encontraron en los manglares donde los juveniles dominaron el 
Q) conjunto de peces 

X 

LU • Veinticinco de las 30 especies mas abundantes en los manglares tambien se observaron en los 
arrecifes de coral lo cual indica el uso de multiples tipos de habitats por estas especies. Ademas las 
tallas de peces en los manglares fueron menores a los arrecifes. Esto indica que estan ocurriendo 
cambios ontogenetics (a lo largo de su ciclo de vida) en los tipos de habitat para las familias roncos 
(Haemulidae), pargos (Lutjanidae), loros (Scaridae) y picuas (Sphyraenidae) 

• Basado en las tallas de los peces, los manglares aparentan tener una funcion de habitat intermedio 
para algunos roncos y pargos, donde las tallas menores estan asociadas a las hierbas marinas, las 
tallas intermedias en los manglares y el tamano promedio mayor se encuentra en los arrecifes de 
coral 

• Las especies mas comunes a lo largo de todo el paisaje marino de La Parguera fueron los loros 
pequenos Scarus iseriy 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 segun la lista de especies amenazadas de la Union Internacional para la Conservacion 
de la Naturaleza (UICN) 

• El avistamiento de varias especies de importancia comercial y de gran tamano ha disminuido 
sustancialmente durante los pasados 25 anos (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 
Batistes vetula 

• Los pargos mas grandes observados en La Parguera fueron de tallas menores al tamano maximo 
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 anos no se observo 
ningun ejemplar de Mycteroperca tigris ni Mycteroperca venenosa y solamente se observaron 
dos ejemplares de E. striatus, Mycteroperca bonaci y E. adscensionis mientras se cuantificaron 
unicamente 43 Epinephelus guttatus 

• Los meros de menor tamano, incluyendo C. cruentata (n=246) y C. fulva (n=81) fueron 
significativamente mas abundantes que los meros de gran tamano 



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

• Ninguna de las especies de meros observadas presento el tamano maximo reportado para su 
especie. Por ejemplo el tamano maximo observado para especies de Cephalopholis en La Parguera 
se estimo en 30 cm (largo horquilla) aunque la talla maxima 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 tamano maximo reportado en la literatura cientffica 

• La abundancia y biomasa de roncos (haemulidos) fue mas alta en los manglares cercanos a arrecifes 
de coral y hierbas marinas. Todas las especies comunes de roncos demostraron un cambio en la 
distribucion de tamanos que aumento en sitios cercanos al veril, por ejemplo la mayorfa de los 
juveniles se encontraron en lagunas cercanas a la costa y los adultos en areas 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 segregation ontogenetica 

• 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 herbivoros, al igual que la abundancia de meros, loros y labridos 
fueron significativamente mayores en 2007 que durante el 2001 

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

• Se observo un aumento consecutivo de mas de tres anos (entre 2001 y 2007) en la abundancia de 
C. fulva y S. aurofrenatum 

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

• La diferencia entre anos mas sobresaliente ocurrio entre 2003 y 2004 donde declinaron 65 metricas 
(aproximadamente 80% de todas las metricas), de las cuales cinco declinaron significativamente; 
seguido por un aumento de 70% de las metricas en el ano siguiente (2004-2005) 

• Se observaron densidades mas altas en verano que invierno para 65% de las 20 especies mas 
abundantes. Por ejemplo la densidad de Thalassoma bifasciatum y Chromis cyanea fue 50% mas 
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 
comparacion con verano 

• Los planctfvoros y picfvoros fueron mas abundantes durante temporada de verano en comparacion 
con invierno 

Recomendaciones: 

Para alcanzar un balance efectivo entre los usos del recurso marino por los humanos y los objetivos 
de conservation 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 multiples amenazas causando una 
baja cobertura de coral, mayor abundancia de algas y escasez de peces de gran tamano lo cual resulta 
en una mudanza de las especies dominantes. 



^ 



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

Los datos colectados por el programa de monitoreo de N-CREMP y la sfntesis 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 mas estudiadas del Caribe. Los resultados cientfficos al igual 
que los mapas proveen un borrador para comenzar un proceso de zonificacion marina que puede 
incluir areas de conservacion, reservas marinas de no pesca, y garantizar el acceso a lugares para 
actividades recreativas. Los lugares de mayor diversidad y distribution de especies o grupos claves 
han sido identificados y cartografiados. Ademas se ha evaluado la condition de las poblaciones de 
especies de importancia comercial lo cual provee un pre-requisito para el desarrollo de estrategias 
de manejo y planificacion 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 atencion especial por los manejadores. En una comparacion historica se destaca 
que la mayorfa de los peces de importancia comercial de gran tamano han declinado marcadamente 
en abundancia desde los anos 80. Las abundancias bajas de los meros y loros de gran tamano puede 
tener impactos a la funcion del ecosistema y deben recibir atencion prioritaria. 

El cambio en la composition bentica hacia mayor cobertura de algas en vez de corales en la mayorfa de 
la region esta asociado a una degradation del ecosistema de arrecife de coral. Los manglares son un 
habitat importante en esta region y tiene una funcion en el mosaico de habitats que estan 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 seran de mayor beneficio para la 
diversidad y la productividad. Se deben establecer metas para recuperar las poblaciones de peces 
de gran tamano y crear condiciones optimas de calidad de agua para ayudar a restaurar el habitat 
bentico y la arquitectura del arrecife de coral. La Reserva Natural de La Parguera es un ecosistema 
de arrecifes unico en Puerto Rico por su position 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 ecologica 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. 



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

Contents 

Executive Summary 
i Table of Contents 
ii 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 offish diversity, biomass and abundance 154 



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

3.7 A 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 offish 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 



page 
xvi 



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

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 



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

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. A3-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. NOAAdiver 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 Pontes 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) Pontes 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 



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

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),\ane 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 



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

Figure 3.32. Mean density for juvenile/subadult and adult by mapped habitat type for gray snapper 

(/_. 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 

(/_. mahogoni) 82 

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

(/_. mahogoni) 83 

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

and biomass 84 

Figure 3.37. Comparison of mean density and biomass by substrate type for lane snapper (/_. 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 



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

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 



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

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 (S. 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 (/_. 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 



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

Figure 3.126. Seasonal changes in mean goatfish biomass for: yellow (M. martinicus) and spotted goatfish 

(R maculatus) 151 

Figure 3.127. Seasonal changes in mean biomass for: queen triggerfish (S. 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 offish species richness across the seascapes of La Parguera 177 



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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) 



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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) 

1. Introduction 




La Parguera, Puerto Rico 



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. 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. 




Diver collecting benthic habitat composition data 



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 km 2 ) 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 (Valdes-Pizzini, 1990; Aguiler-Perera et al., 2006). In 
addition, a small "no-take" marine reserve (7.6 km 2 ) has been proposed forTurrumote, south of Corral 
(Figure 1.1). 



o 
o 

O 



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

67°o'0"w eeww 




1 Margarita Reef 

2 El Palo 

3 Media Luna 

4 Isla Magueyes 

5 Enrique 

6 Corral 

7 Turrumote 

8 Romero 

I La Parguera Natural Reserve 
Max. Depth 47 m 



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. 



PRISM 1963 - 1995 Mean Annual Precipitation, Puerto Pico 




The Climate Source, Inc 
www. climate source, com 





Precipitation (nun) 




^■< 1,000 | | 1,200 


1,300 | | 1,500 - 1,600 | | 1,800 - 1,£>00 ^M 2,250 


■2,500 ^H 3,500 -4,000 


_U uooo- 1,100 | | 1,300 


1,400 | | 1,600 - 1,700 | | 1,900 - 2,000 ^M 2,500 


-3,000 | | > 4,000 


| | 1,100- 1,200 | | 1,400 


1,500 __^ 1,700 - 1,800 | | 2,000 - 2,250 __^ 3,000 


-3,500 



Map Created: November 2002 al^^ 6 2^^Z Kilometers Copyright (c) 2000 - 2002 OSU Spatial Climate Analysis Service 

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

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). 



o 
o 



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. 




O 



Mangrove islands of La Parguera, PR 




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



page 
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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) 

Tides 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). 



O 1.2 Ecosystem change and the impact of multiple stressors 

-i— ' 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/km 2 
and Rio Jacquas yielding 3,700 tonnes/km 2 (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-Rodrfguez, 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.brAA/EBens/ 
morelock/clnpho. htm#par. 



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



-1 1 

Terrestrial MAR (g cm" £ y"') 




0.01 0.02 0.03 0.04 

o -i ■ ' ' — 



0.05 0.06 



SURFACE MIXED LAYER 



I -A- 



I A- 

I A 1 

I * 1 

I A 1 

I A 1 

I A 1 

I A 1 



H 



Average MAR: 



| a | 0-39 +/- 0.02 g cm y" 

I A 1 



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■< 

m 

1960 > 



6 6 10 12 14 



Terrestrial % 



Figure 1.5. A) Aerial photograph showing sediment runoff in the 1980s along the developed shoreline at La Parguera. Source: http:// 
www.log.furg. brAVEBens/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 ai, 2008. 



In addition, disease and bleaching have led to substantial deterioration of 
decades. White plague II and Caribbean yellow band diseases (YBD) are 
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 etal., 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). 



coral reefs in the past two 
the most widespread and 




Bleached Siderastrea sidera colony 




Bleached Montastraea sp. colony 



page 
5 



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



Instantaneous Sea Surface 
Temperature (Oct 12, 2005) 



o 

O 



Aug-Oct 2005 

Whelan et al. 2007 

Clark et al. 2008 



Sep-Oct 2003 — 
Manzello et al. 2003 




SST(°C) DHW 



o> 




c 


c 


f 


(0 


o 


.Q 


CO 



.£2 






n 


CO 


(0 


O 


CO 




CO C/ 


E 


3 


M- 

o 


c 


2 

O 
Q. 


12 

c 

CD 


CD 
DC 


CD 



Aug-Sep1999 
Velasco et al. 2003 

Jul-Sep 1998 . 
Velasco et al. 2003 




Jun-Oct1995 _ 
Winter etal. 1998 



Sep-Oct 1990 h 
Goenaga & Canals 1990 



Aug-Dec 1987 - 
Goenaga etal. 1989 




20041 



-20031 



20021 



20011 



20001 



19991 



19981 



19971 



19961 



19951 



19941 



19931 



19921 



19911 



19901 



19891 



19881 



32 



30 



28.5 



26.7 



25 



15 



11 







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 space- 
time charts. Source: SST data processed 
by Varis Ransi of NOAA CCMA Remote 
Sensing Group. 



page 
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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) 

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 1 979 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., 1 991 ). 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). 




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. 



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



O 

o 



A 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 m 2 ) 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 aq uatic 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 




Zone 



| Backreef 

Bank/Shelf 

Bank/Shelf Escarpment 
| Fore reef 

Lagoon 

Reef Crest 

Shoreline Intertidal 
| Mangrove 
I Land 



Benthic HABITAT 



Coral Reef and Colonized Hardbottom 
Submerged Vegetation 
Uncolonized Hardbottom 
Unconsolidated Sediments 
| Mangrove 
Land 



Benthic TYPE 

Colonized Pavement 

Colonized Pavement with Sand Channels 

| Linear Reef 

| Spur and Groove Reef 

| Patch Reef (Aggregated) 

| Patch Reef (Individual) 

| Scattered Coral/Rock in 
Unconsolidated Sediment 

Reef Rubble Mud 

H Macroalgae | Mangrove 

Seagrass | Land 

Sand 

Benthic MODIFIER 

| Macroalgae/Patchy/10-50% 

Macroalgae/Patchy/50-90% 

Seagrass/Patchy/1 0-30% 

Seagrass/Patchy/30-50% 

Seag rass/Patchy/50-70% 

Seagrass/Patchy/70-90% 
| Seagrass/Continuous 
| Mangrove 
I Land 



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. 



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

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. 



o 

o 

O 



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 

J J 67°0'0"W 

surface. More details on the methods 1 

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 
etal.,2009). 











llttll 






<&*■ 






i 






/ 






^P 




/ 




jb^**** 


-;-=?- -—— ' 








*W Surface rugosity 








■ High 
B Low 








mLjSSSf^. 



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. 



page 
9 



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

1.4. References 

Acevedo, R. 5 J. Morelock, and R.A. Olivieri. 1989. Modification of coral reef zonation by terrigenous sediment 
q stress. Palaios 4(1 ): 92-1 00. 



o 



-Q Appeldoorn, R.S. 2008. Transforming reef fisheries management: application of an ecosystem-based approach 
q in the USA Caribbean. Environ. Conserv. 35: 232-241 . 

~£^ Aguilar-Pererra, A. 5 M. Scharer, M. Valdes-Pizzini. 2006. Marine protected areas in Puerto Rico: Historical and 
— current perspectives. 49(12): 961-975. 

I 

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-Rodriguez, I. 2006. Relationships between reef fish communities, water and habitat quality on coral 
reefs. M.S. Thesis. University Puerto Rico at Mayagiiez. Mayagiiez, 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. MIA- 
9C/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. 

Garcfa-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 



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

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, IP. 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. Mayaguez, 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 offish 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. 



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

C 

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■ ^^™ 

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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) 



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 offish 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. 




Figure 2.1. NOAA diver recording benthic habitat 
composition within the randomly placed 1 m 2 quadrat 
along the belt transect. 



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 m 2 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), long- 
spined 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 

r ° Figure 2.2. Marine debris. 



Table 2.1. Abiotic and biological variables measured to 
characterize benthic assemblages along fish transects in 
the southwest Puerto Rico study region. 




Benthic Biota 


Measurements 


Cover 

(%) 


Height Abund. 
(cm) (#) 


Abiotic 






Hardbottom 


X 


X 


Sand 


X 




Rubble 


X 




Fine sediment 


X 




Rugosity 
Water depth 
Biotic 






Corals (by species) 
Macroalgae 
Seagrass (by species) 
Gorgonians 

Sea rods, whips and plumes 

Sea fans 


X 
X 
X 

X 
X 


X 
X 

X X 
X X 


Encrusting form 
Sponges 
Barrel, tubes, rope, vase morph. 


X 
X 


X X 


Encrusting morphology 
Other benthic macrofauna 


X 




Anemonies and hydroids 


X 


X 


Tunicates and zoanthids 


X 




Macro-invertebrates 






Queen conch (by sexual maturity) 
Spiny lobster 
Long-spined urchin 




X 
X 
X 


Mangroves 
Prop roots 

Prop roots colonized by algae 
Prop roots colonized by sponges 
Prop roots colonized by other 




X 
X 
X 
X 



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




Figure 2.3. A selection of habitat types designated in the hierarchical classification scheme ofNOAA's benthic habitat map (Kendall et ai, 
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=1 58). 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. Scattered 
w/ sand Linear Patch coral/rock Bedrock/ 
channels reef reef in sand 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 



*Un known 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) 



Mapped habitat types 


Mapped 
Area (km 2 ) 


Vo Mapped 
Area 


# sites 
surveyed 


Area 

surveyed 

(km 2 ) 


% 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 



pavement and patch reefs (Table Table 2.3. The number of benthic habitat sites surveyed by mapped habitat type for the 
ox , , , , ,.„ southwestern Puerto Rico study region from 2001-2007. Mapped habitat categories are 

2.3). Hence observed differences from Kenda „ etaL (20 oi). 

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: 

T{Q r n)/n 

where Q. = quadrat /, and n is the total number of quadrats. Average site values were then used to 
calculate means and SE of measured variables per 100 m 2 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 non- 
parametric 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 



O 
Q. 

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o 



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

used to calculate means and SE of measured variables per 100 m 2 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): 

npi nv 

Ujuv = ^^^{Xiu.Xjv), \<ju<v<k 

i=i /=i 

Where 0, b)=1 if a < b, or if otherwise. The Jonckheere test Statistic (J) was then calculated as the sum 

of these U counts and was compared against a significance threshold (J a = 005 ) that was dependent on 

the number of sampling periods and number of sites surveyed within each sampling period (Wolfe and 

Hollander 1999). If J metrjc > J a = 005 , 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 overtime. 




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) 

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 



30 n 



20 



10 



n 



i 



L 



n . ^ . r-i 



s 



*? 



/ 



& 



& 



f 



& 



& 



& 



^ 



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). 



60 



40 



20 



e Turf algae 


a Macroalgae 


h Corals 


a Soft corals 


0CCA 


□ FAandCB 


□ Hydroids 


□ Sponges 





tfjm 



Jki. 



jSl 



Bedrock 



Pavement Linear Reef Patch Reef Reef Rubble 



Scattered 
Coral/Rock 



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) 



o 

Q. 

E 
o 

O 
o 



3 i 



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 Pontes 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 Pontes 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 
Pontes 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 




Diploria labyrintheformes (left) and Montastraea annularis complex (right) 



a^ 



n 



i 



A* 






r 



9 / 









r 



^r 



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. 



□ Montastraea 
a Pontes 
a Agaricia 
H Siderastrea 
Diploria 
Acropora 
B Colpophyllia 

□ Meandrina 
^" 2 - 

<5 f?3 mother 

> 
o 
o 

c 
ro 

0) 




Pavement 


Linear Reef 


Patch Reef 


Scattered 
Coral / Rock 


Bedrock 


(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. 

habitats (Figure 2.7). Interestingly, several genera were not 



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

observed on colonized bedrock habitats, 32 
but this pattern may have reflected the 
low number of colonized bedrock sites 
surveyed (n=2). 



24 



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 filiforme A 



^ 16 

> 
O 
O 

c 
re 

I 8 



ll 



n 



y 



y 



& 



>* 



o° T 



/>« ,o*' A*' f J>' ,,0° JT eP- ~ & 



J? 



+ f ^ ** * f f jf 

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. 

24 i 



CO 
O 

a. 

E 
o 

O 
o 



DO 
i 



8 12 



CD 




Thalassia 
testudinum 



Syringodium 
filiforme 



Halodule Halophila 

wrightii dec i pi ens 

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 

8 



*- 4 

9 

O 



<D 



ih. 



./ 



s 



\* 



V 



s 



? 

rv 



,<? 






# 









J- 



^ 



^ 



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) 



CO 

o 

E 
o 

O 

o 



(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 sub- 
aquatic 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 
help elucidate broad-scale patterns (e.g., 



_ 4 



Thalassia 
testudinum 



Syringodium 
filiforme 



Halodule 
wrightii 



Halophila 
decipiens 



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. 



18 



12 



-LL 



Macroalgae Detritus Seagrasses Prop Roots Sponges Soft Coral 

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. 



Maps of interpolated distributions are useful in that they 
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 




Diploria strigosa 



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




Live coral cover (%) 

<2.00 15.01-25 

2.01 -10 |^| 25.01 -51.S 
10.01 -15 



0d * °%v.# 



$ xxO • 



>•• O 



cro 



*% a 08 



a 



•at 
• 4 






o o 

5 ^o ° c 

cP x °o°Qb ^°°o $ 
o° n o» x 
o w 



Survey Sites N=572 




F/gure 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. 



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



Table 2.4. Summary statistics for hard coral 
surveys in the southwest Puerto Rico study area 


species found in 
from 2001-2007. 


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) 


Fa via frag urn 


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) 



(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 m 2 . 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). 




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) 



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. 





Col. Bedrock 


Col 


. Pavement 


Linear Reef 


Patch Reef 


Reef Rubble 


Scat. 


Coral/Rock in 




(n=2) 




(n=322) 






(n=100) 




(n=53) 


(n=1) 


Sand (n=94) 




Freq. % cover 


Freq. 


% cover 


Freq. 


% cover 


Freq. 


% cover 


Freq. % cover 


Freq. 


% cover 


Species 


occur Mean ± SE 


occur 


Mean 


±SE 


occur 


Mean ± SE 


occur 


Mean ± SE 


occur Mean ± SE 


occur 


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. ag ah cites 


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. frag His 




0.01 


<0.01 


<0.01) 
















A. lamarcki 




0.02 


0.02 


0.01) 


0.01 


<0.01 (<0.01) 












Agahcia 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. cylindhcus 




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. labyhnthiformis 




0.13 


0.07 


0.02) 


0.17 


0.06 (0.02) 


0.13 


0.09 (0.04) 




0.03 


0.02 (0.01) 


Diploha 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) 


1. rigida 




<0.01 


<0.01 


<0.01) 


0.01 


0.01 (0.01) 












1. 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. meandhtes 




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) 








0. 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. pontes 


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) 



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




P. astreoides cover (%) 

| <0.10 3.1 -5 

| 0.1-1 ^M 5.1 -9 

1.1 -3 



_l QQ ^- -v Survey Sites N=572 

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) Pontes astreoides. 



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. 

Acropora cervicornis 



> '- ' 


rams*; ?-l w 

ifilBE 


%3PI * 1- .-■*. 






P9 



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




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. 



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. 




Acropora palmata 



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




Survey Sites N=572 



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. 



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. 

Macroalgae 




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




<5 25.1-40 

5.1 -15 |^| 40.1 -89.7 

15.1 -25 



Surx/ow Sitae N = mQ1 



b) 




Turf algae cover (%) 

<10 50.1-75 

10.1 -25 ^H 75.1 -93.8 
25.1 -50 



° x i° ^ 



O ~ v Q Q Q 



■S>A V 



Survey Sites N=572 




Sponge cover (%) 

<3 10.1-20 

3.1 -5 ^H 20.1 -30 

5.1 -10 



°8>°o °fc o/.« 



°*t 



D ^° °o °" 






1 km 
J I 






° 8 ° ° 






x^^^o" 



CCU 



Survey Sites N=572 



Figure 2. 1 7. Maps of the interpolated (left map) and spatial (right map) distributions for: (a) macroalgae, (b) turf algae, (c) soft coral cover 
and (d) sponges. 



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



CO 

o 

E 
o 

O 

o 



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 




Seagrass cover (%) 

<10 30.1-40 

10.1 -20 ^H 40.1 -96.5 
20.1 - 30 





T. testudinum cover (%) 

■ < 5 30.1 - 50 

| 5.1 -15 ^H 50.1 -96.5 

15.1 -30 



>*x * x \ xxx * 

M x \*Z ^ , 

x x X & 



Survey Sites N=521 





S. filiforme cover (%) 

| < 2 20.1 - 30 

| 2-10 ^H 30.1 -84.3 

10.1 -20 






°** XX ° * M,***** 

X X f 



Survey Sites N=521 



Figure 2.18. Maps of the interpolated (left map) and spatial (right map) distributions for: (a) seagrass, (b) Thalassia testudinum and (c) 
Syringodium filiforme. 



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

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. 



15 -, 



o 
o 



c 

CO 




10 



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, Pontes, Agaricia, and Siderastrea (Figure 

2.19, Table 2.6). 



'fl 


pU 


1 


JL 


-L 


I 


1 


1 


JL 


h 


X 


J 


_L 



4? <# 



«*W>« 



f 



«» <£ 



& 



# 



$ 



* 



# 



# 



Figure 2.1 9. 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. 



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. 



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 



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. 



Hardbottom habitat types 


N 


X 2 


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 



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



c/) 
O 



Q) 



32 



r? 24 



2 
o 
o 

15 

o 16 

o 

c 

CO 
0) 



□ Colonized Pavement 

a Colonized Pavement with Sand Channels 



1 



I 



I 



J 



I 



1 



IlJAI 



JL 



1 



Jj 



L 



4? *4 t* 4* jt 4* 



%* ^ ^ ^ 



J* 



5? 



.* 






£ 



S 



$ 



4* of 4* of 4* cy 

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). 



c 3 



I 



I 



i 



JL 



A I 



A, I 



1 



JL 



^ a N V * N fr & $S» ,& J> ,& & & & & & $ $ 

4* 4* <? 4* <* ^ %* «* J of 4? of 4? of 4* of 

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 



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 pair- 
wise 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 
(x 2 =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, PO.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). 



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



All five genera (Montastraea, Pontes, 
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). 



Table 2.9. Results of non-parametric AN OVA (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. 



Hard bottom Habitat 


Montastraea 


Pontes 


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. 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. 

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) Pontes 



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 



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



12 

































- 


□ Colonized Pavement 


- 




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a Colonized Pavement with Sand Channels 







J? 



24 - 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 Pontes 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 Diploha 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 Diploha 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, Pontes, Siderastrea, 
Diploha and Agaricia) in most habitats (Table 2.10 A-E). 



# v 4? <<? 4< 






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x 



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<& 



& 



.& 



<£ ^ c£ <£ 



& 






& 



4* • 



JP 



& 



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is* ^ ^sO ^ 



& 



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) Pontes spp. andc) Diploha 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) 
a) 4 



2 

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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 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). 

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 2.11. 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 



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. 



Algae types 


X 2 


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) 



™T Table 2. 13. Multiple non-parametric pair-wise comparisons of mean percent algae cover for: a) macroaigae, b) turf algae, c) cyanobacteria/ 
O filamentous algae and d) crustose coralline algae among sampling periods (2001-2007) in the southwest Puerto Rico study region. Green 
'p boxes with dots indicate significant pair-wise differences (p<0.05). 



o 


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c) Cyan 


obacteria / filamentous algae 


















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d) Cms 


tose coi 


alline al 


gae 
























Sum-07 



page 
34 



Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007) 
a) 50 

□ Macroalgae 



I 25 

c 



b) 



d) 



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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. 



O 
Q. 

E 
o 

O 
o 



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

2.3.4. Abundance and distribution of macroinvertebrates 

2.3 .4.1. Queen conch (Eustrombus gigasj 

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. 

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. 



■Hfi 


b _ 




InL ■'*•*' Aw\ 

■ 




• 


■■- r ;- 





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 





8 


89 




Mangrove 


1 


1 





1 






Hard 


1 


1 





1 




2005 


Soft 


2 


4 





4 


170 




Mangrove 


1 


1 





1 






Hard 


4 


4 


1 


5 




2006 


Soft 


11 


10 


5 


15 


179 




Mangrove 


















Hard 


4 


2 


2 


4 




2007 


Soft 


3 


1 


2 


3 


180 




Mangrove 


















Eustrombus gigas 

• immature • mature 



both 



Substrate type 



Hard bottom 


Unconsolidated 




sediment 


Submerged 




vegetation 


Mangrove 



Figure 2.25. Distribution map of queen conch (Eustrombus gigas,) in the southwest Puerto Rico study 
area. 




Immature Eustrombus gigas 



Substrate type 







1 



Hard bottom 

Submerged 
vegetation 



Unconsolidated 
sediment 

Mangrove 



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

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 m 2 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). 



b) 


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xx xx x x x x x 

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x x x x xx 

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x xx X ^ 




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xX x * 


1 km X x x X x > 




1 ' 





Mature Eustrombus gigas 

x • 1 • 2 



Substrate type 



Hard bottom 

Submerged 
vegetation 



Unconsolidated 
sediment 

Mangrove 



Figure 2.26. Distribution maps of a) immature and b) mature queen conch (Eustrombus gigas,) in 
the southwest Puerto Rico study area. 



a) 



10 



E 

o 



c 
as 
■o 

c 

as 



5) 



J 



fl 



2004 



2005 



U 



■ Hard 

□ Soft 

□ Mangrove 



I 



2006 



2007 



b) 



10 



0) 

o 

c 

OS 
T3 

c 

3 

■s 5 

to 

OJ 

5) 
ui 



■ Hard 

□ Soft 

□ Mangrove 



J 



2004 



2005 



2006 



2007 



Figure 2.27. Abundance of a) immature and b) mature queen conch (Eustrombus gigasj in the southwest Puerto Rico study area. 



O 
Q. 

E 
o 

O 
o 



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

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 2.1 5). 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). 



Table 2.15. Abundance of long-spined sea urchins (Diadema antillarumj observed 
within transects by year in the southwest Puerto Rico study area. 


Year 


Habitat 
type 


Sites Sites Total 
surveyed observed individuals 




2006 


Hard 


91 7 64 


2006 


Soft 


41 1 30 


2006 


Mangrove 


20 




2007 


Hard 


100 4 98 




2007 


Soft 


42 


2007 


Mangrove 


20 




Total 


314 12 192 





2.3.4.3. Caribbean spiny lobster (Pan ulirus argusj 

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). 



Table 2.16. Abundance of Caribbean spiny lobster (Panulirus argusj observed within 
transects by year in the southwest Puerto Rico study area. 



Year 


Habitat 


Sites 


Sites 


Total 


type 


surveyed 


observed 


individuals! 


2005 


Hard 


90 


2 


2 


2005 


Soft 


52 


1 


2 


2005 


Mangrove 


13 


2 


6 


2006 


Hard 


91 


4 


5 


2006 


Soft 


41 








2006 


Mangrove 


20 


1 


7 


2007 


Hard 


100 


4 


5 


2007 


Soft 


42 








2007 


Mangrove 


20 








Total 


469 


14 


27 




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 cm 2 with a total affected area of 1,054 cm 2 . 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. 




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) 

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 Pontes 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 m 2 ) 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 



O 
Q. 

E 
o 

O 
o 



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

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 m 2 ) 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) Dictyota sp. surrounding Dichocoenia stokesi 




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 over- 
fishing have reduced herbivory and have resulted in a phase-shift from coral-dominated to macroalgal- 
dominated reefs in the Caribbean (Lessiosetal., 1984; Gardner etal., 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. 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) 



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. CCAplay 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, 1 994). 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. 



■■-.'■■--■—^ 




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ffpvll 


Bife 


p. 

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<*^_ 


rlSt^ -ih"_ d 


% 



Crustose coralline algae 



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 majoroutbreak 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 Garcia- 
Sais et al., 2005, 2008; Weil et al., 2002). No major hurricane has hit 
Puerto Rico since 1 998, but per this study, reefs at depths less than 1 8 
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). 




Diseased S. sidera 



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. 

S. radians 




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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) 

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). 




Dense gorgonian habitat 



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 etal., 1981; Drew, 1983). 




T. testudinum 







S. filiforme 



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 



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

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 m 2 
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 near- 
shore 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. 




Sponges on mangrove prop root 




Tunicates on mangrove prop root 




Mussels on mangrove prop root 



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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) 

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. 





mm 




Colpophyllia natans 



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. 




Derelict trap 



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 



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

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. gigasj 

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 , 1 997), 
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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Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007) 

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 1 997; 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 m 2 , with highest density 
being two adult mature individuals per 100 m 2 . Additionally, the maximum number of conch observed 
during a sampling mission was 1 2 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 1 985 and April 1 986. Torres Rosado (1 987) visually surveyed 

40.81 ha of bottom substrate and observed 331 queen conch, with a density of 0.08 conch per 1 00 m 2 . 

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. 




E. gigas 



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 



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

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. antillarumj 

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 m 2 . 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. 




D. antillarum 



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. 



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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) 

2.4.6.3. Caribbean spiny lobster (P. argusj 

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, 1 985). 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 overtime, and a dedicated lobster- 
monitoring program will be needed to document long-term changes in 
lobster populations in La Parguera. 




P. argus 



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

2.5. References 

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Appeldoorn, R.S. 1992. Preliminary Calculations of Sustainable Yield for Queen Conch (Strombus Gigas) in 
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Appeldoorn, R.S. 1994. Queen conch management and research: status, needs, and priorities, pp. 301-320. 
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Gardner, T, I.M. Cote, J.A. Gill, A. Grant, and A. Watkinson. 2003. Long-term region-wide declines in Caribbean 
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Hoegh-Guldberg, O., P.J. Mumby, A.J. Hooten, R. S. Steneck, P. Greenfield, E. Gomez, CD. Harvell, PF. Sale, 
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Hollander, M. and D. Wolfe. 1999. Nonparametric Statistical Methods. New York: John Wiley & Sons. 

Hubbard, D.K., J.L. Sadd, A.I. Miller, L.P Gill, and R.F. Dill. 1981. The production, transportation, and deposition 
of carbonate sediments on the insular shelf of St. Croix, U.S. Virgin Islands, West Indies. Laboratory Contrib. No. 
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Hughes, T.P. 1994. Catastrophes, phase shifts and large scale degradation of a Caribbean coral reef. Science 
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Idjadi, J.A., S.C. Lee, J.F. Bruno, W.F. Precht, L. Allen-Requa, and P.J. Edmunds. 2006. Rapid phase-shift 
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Jeffrey, C.F.G. 2004. Benthic habitats, fish assemblages, and resource protection in Caribbean marine sanctuaries. 
Ph.D. Dissertation, University of Georgia. 145 pp. 

Kendall, M.S., M.E. Monaco, K.R. Buja, J.D. Christensen, C.R. Kruer, M. Finkbeiner, and R.A. Warner. 2001. 
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152 (on-line), http://ccma.nos.noaa.gov/products/biogeography/benthic/welcome.html. 

Kendall, M.S., J.D. Christensen, and Z. Hillis-Starr. 2003. Multi-scale data used to analyze the spatial distribution 
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Knowlton, N. 2001. The future of coral reefs. Proceedings of the National Academy of Sciences of the U.S.A. 
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Lessios, H.A. 1998. Mass mortality of Diadema antillarum in the Caribbean: what have we learned? Ann. Rev. 
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Lessios, H., D.R. Robertson, and J.D. Cubit. 1984. Spread of Diadema mass mortality through the Caribbean. 
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MacArthur, R.H., and R. Levins. 1964. Competition, habitat selection and character displacement in a patchy 
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McFarland, W.N., J.C. Ogden, and J.N. Lythgoe. 1979. The influence of light on the twilight migrations of grunts. 
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Matos-Caraballo, D. 2004. Comprehensive Census of the Marine Fishery of Puerto Rico, 2002. Commerical 
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Meyer, J.L., E.T. Schultz, and G.S. Helfman. 1983. Fish Schools: An Asset to Corals. Science 220: 1047-1049. 

Miller, J., E. Muller, C. Rogers, R. Waara, A. Atkinson, K. Whelan, M. Patterson, and B. Witcher. 2009. Coral 
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Morris, A., S. Chormanski and D. Die. 2004. Overview of the assessment history and the current management 
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Nagelkerken, I., G. van der Velde, M. W. Gorissen, G. J. Meijer, T van't Hof, and C. den Hartog. 2000. Importance 
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■ ^^™ 

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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) 



Chapter 3. Fish Communities, Groups and Species 



3.1. Introduction 

Previous studies offish 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 




Balloonfish (Diodon holocanthus) in turtle grass. 



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 
offish, 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 offish 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 NOAACCMABiogeography Branch (CCMA- 
BB) 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). 



CO 

CD 



E 

E 
o 

O 

CO 



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

While 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 m 2 ) 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. 







M/* 





Figure 3. 1. NOAA trained observer recording fish census 
data along the belt transect 



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




] Hard bottom 

| Mangroves 

J Seagrasses/macroalgae 

1 Sand or Mud 

# NOAA CCMA survey site 

Hi 



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 hard bottom 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 ;= 



1 - 



'Lm X ij - X ik 



^i-> ij ik. 



where x. is the abundance of the /th species in theyth 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. 



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

An ANOSIM test, a multivariate version of Analysis of Variance (ANOVA; Primer v6; Clarke and Warwick, 
1 994), 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 

E 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 offish assemblage composition being: R<0.25 = barely separable; R>0.5 = overlapping, 
O but clearly different and R>0.75 = well separated. 

O 

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 
M and thematic levels are delineated in an ecologically meaningful way for fish. For instance: Do fish 
LL 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 
tf\ 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 inter- 
habitat 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 (December- 
March) and summer field seasons (August) and then plotting means (± SE) using bar charts. 



2D Stress: 0.16 









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

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). 




Habitat Structure 

±SOFT 
▼ HARD 

MANGROVE 



CO 
CD 



E 

E 
o 

O 

CO 



CO 



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. 



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 

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. 




Jenkinsia sp. and schoolmaster (Lutjanus apodus) 



Mapped habitat type 



Patch Reef Patch Reef 



Colonized Col. Pav. w/ 
Pavement Sand Chan. 



Seagrass Macroalgae 



Patch Reef (Individual) 

Patch Reef (Aggregate) 

Linear Reef 

Colonized Pavement 

Col. Pavement w/ Sand Channels 

Seagrass 

Macroalgae 

Unconsolidated Sediments 

Mangrove 



0.18 

0.05 
-0.03 
0.13 



-0.06 



0.07 



0.16 


0.10 


0.29 


0.25 


0.43 




0.26 


0.34 
0.21 


0.74 
0.56 


0.49 
0.40 


0.78 


0.09 


0.24 


0.71 


0.28 


0.89 


0.86 


0.89 


0.85 


0.90 


0.47 



0.02 



CO 

CD 



E 

E 
o 

O 

CO 



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

reef 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. 



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 



Continuous Patchy 
70-90% 



Continuous >90% 
Patchy 70-90% 
Patchy 50-70% 
Patchy 30-50% 
Patchy 10-30% 
Sand (Uncon. sed.) 



Patchy 
50-70% 



Patchy 
30-50% 



Patchy 
10-30% 



0.28 








0.52 


0.17 






0.18 


0.27 


0.58 
0.47 




0.36 


0.34 


0.01 


0.46 


0.05 


0.04 


0.11 



0.13 



Fish assemblages of the sparser 30-50% seagrass coverwere notsignificantlydifferentthan 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. 




60 



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

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 offish biomass and unvegetated sediments supported 
lowest mean fish biomass (Figure 3.4b). Similarly, highest fish species density (number of species ^ 




Total fish abundance (100 m 2 ) 

■ <10 Benthic HABITAT 

11-50 Coral reef and colonized hardbottom 

51 - 200 Submerged vegetation 

| >200 Unconsolidated sediments 




Total fish biomass (g/100 m 2 ) 

^H <1000 Benthic HABITAT 

1001 - 5000 Coral reef and colonized hardbottom 

5001 - 10000 Submerged vegetation 

I >10000 Unconsolidated sediments 




Fish species 


richness (100m 2 ) 









>30 




■■ 1 








Benthic HABITAT 


■■ 6-10 








Coral reef and colonized hardbottom 


^H 11 -20 








Submerged vegetation 


^M 21 - 30 








Unconsolidated sediments 




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. 



CO 

CD 



E 

E 
o 

O 

CO 



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



a) 1500 

E 



= 750 

> 

c 

*«- 
o 



E 



E 



b) 6n 



— 3 



E 
o 
!5 



■,■,■,■,■,■ 




-M 



I . . 



JL 



^ ^ 



fy o> 



• • 



./ V 



^ 



Habitat type 



Habitat type 

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 m 2 ) 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 m 2 , 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 m 2 . 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 



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. 



Species/Fa 




i 


Habitats 
used 




Mean (±SD) Fork Length, cm 




Mang 


rove 


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 apod us 


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) 



62 



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

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). 




Herbivorous surgeonfish (Acanthurus species). 



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 




Figure 3.6. Maps of the interpolated (left map) and spatial (right map) distributions for herbivore: (a) abundance, (b) biomass and 
(c) species richness. 



CO 

CD 



E 

E 
o 

O 

CO 



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



a) 



50 



25 



a) 



24 



12 - 



JL 



n 



Colonized 
Hard bottom 



b) 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hard bottom 



£ 

■i 1 



b) 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



r~—\ 




Colonized 
Hard bottom 



c) 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



U 10 



c) 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



II n n r 



4 n 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Habitat type 



Habitat type 



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. 



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. 



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). 



64 



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




Piscivore fish abundance (100m 2 ) 
Benthic HABITAT 



I -10 

II -50 
51 -100 
>100 



Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 




r^a^p^ joe,. . • W ° »„ o » 

1 km <P o o° ~ Q § 

1 ' o ft° ^bo o o n o ° 



o §> 




Figure 3.9. Maps of the interpolated (left map) and spatial (right map) distributions for piscivore: (a) abundance, (b) biomass and (c) species 
richness. 



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). 




Piscivorous fish, cero mackrel (Scomberomorus regalis). 



CO 

CD 



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

In the mangroves, planktivorous fish abundance was dominated by frequently observed schools of 
small-bodied silvery fish from families including Clupeidae, Atherinidae and Engraulidae (Figure 3.1 Oa). 
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). 



a) 



CO £ 

</> 

LL I 

CD 
I 9 



1200 n 



600 




b) 



0.4 



E 
o 
5 0.2 



Q. 

C 




Planktivorous species, Creole wrasse {Clepticus parrae). 



c) 



n 




Jenkinsia species in mangroves. 



Colonized 
Hardbottom 



Macroalgae/ Unvegetated 

Seagrass Sediments 



Mangrove 



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. 



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




Planktivore fish abundance (100m 2 ) 



o 

I -10 

II -50 
51 -100 
>100 



Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



1 km 



• • * $ • * •• •• 



si c 




Planktivore 

o 

^H 1 -5 
^H 6-10 



species richness (100m 2 ) 

Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



Figure 3.11. Maps of the interpolated (left map) and spatial (right map) distributions for planktivore: (a) abundance, (b) biomass and (c) 
species richness. 



CO 

CD 



E 

E 
o 

O 

CO 



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

3.3.2. 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 m 2 and 1.4 kg/100 m 2 , 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 m 2 ). Biomass was highest over colonized pavement on the mid and outershelf (Figure 3.11b). 



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 Fish Base (http://www. 
fishbase.org). TL=total length; FL= fork length. Refer to Appendix 
C for detailed information. 



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 (Ojeda 

Serrano, 2007). 







Approx. size 
class at first 


Max. known 

c ; 7 a* Tl 


Max. size 
observed in 


Species 




maturity* 




PR, FL 


Batistes 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 apod us 


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 martinicu 


s 15-20 


39.4 


35 


Pseudupeneus maculati 


js 15-20 


30 


30 


Scarus iseri 


10-15 


35 


35 


Scarus taeniopterus 


10-15 


35 


35 


Sparisoma aurofrenatun 


7 10-15 


28 


35 


Sparisoma rubripinne 


-- 


47.8 


40 


Sparisoma viride 


15-20 


64 


50 


Sphyraena barracuda 


-- 


200 


150 



a) 



0.8 -i 



o 0.4 - 




b) 



0.1 



o 0.05 



i±i 



Macroalgae/ Unvegetated Mangrove 

Seagrass Sediments 



Colonized Macroalgae/ Unvegetated 

Hardbottom Seagrass Sediments 



Mangrove 



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) 




Grouper biomass (g/100 m 2 ) 

Benthic HABITAT 

I 0.1 - 1000 Coral reef and colonized hardbottom 

>1000 Submerged vegetation 

Unconsolidated sediments 



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. 



a) 



so -r- 



il 25 



c) 



£ 25 



3 



Juveniles/subadults 




50 -f 



<5 



5-10 10-15 15-20 20-25 25-30 30-35 >35 
Size class (cm) 



Juveniles/subadults 



b) 



iles/subadults 




n 



5-10 



10-15 15-20 20-25 25-30 
Size class (cm) 



30-35 



>35 



JZL 



5-10 



>35 



10-15 15-20 20-25 25-30 30-35 
Size class (cm) 

Figure 3. 14. Size frequency histogram for select grouper (Serranidae) in the southwest Puerto Rico study area, (a) graysby (C. cruentataj, 
(b) coney (C. fulvaj and (c) red hind (E. guttatusj. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Graysby (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 m 2 (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 biomass (g/100 m 2 ) 

Benthic HABITAT 

7} 0.1 - 500 Coral reef and colonized hardbottom 

>500 Submerged vegetation 

Unconsolidated sediments 



* °° ° o 

rP°° 

O O 



1 km 



oco # ^ 

° - 8 ° o 

O 9 •0?>C«0 90 ° 

gf ^o^&o 9 • q@ oo. o 



_ 



'<$$&«> o c 

© o° ^o 
o © ° 



F/gi/re 3.75. Maps of f/?e interpolated (left map) and spatial (right map) distributions for graysby (C. cruentataj; (a) abundance and (b) 
biomass. 



a) 



0.6 n 



0.3 



b) 



0.1 



£ 
o 

£ 0. 



05 



_Q_ 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3. 16. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for graysby (C. 
cruentataj. 



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

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). 



0.4 -i 



o 

o 

T— 

"55 

c 

<D 

4S 



■ Juveniles/Subadults 
□ Adults 



20.7 



20.7 



15.0 








16.4 



16.0 



10.8 



9.1 



2.4 2.4 

Mi 







10.6 



7.4 







4.3 
1.1 



3.8 2.1 



Colonized Colonized Col. pav. Linear Macro- Mangrove Patch Reef Scat coral/ Seagrass Uncon. 

bedrock pavement sand chan. reef algae reef rubble rock sand sediment 



Fish size classes 

Q) Juvenile <5 - 10 cm 
Subadult 10 -15 cm 
£ Adult >15 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 




J . • • • * 



6> 






•vv 






1 km 

I I 






CO 



E 

S 

o 

CO 



CO 



Figure 3.17. Top: Mean density (+SE) for juvenile/subadult and adult by mapped habitat type for graysby (C. cruentataj. 
Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for graysby 
(C. cruentataj in the southwest Puerto Rico study area. 



page 



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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) 

Coney (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 biomass (g/100m 

Benthic HABITAT 

1 . 500 Coral reef and colonized hardbottom 
500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 



Figure 3. 18. Maps of the interpolated (left map) and spatial (right map) distributions for coney (C. fulvaj; (a) abundance and (b) biomass. 



a) 



v.z - 


















0.1 ■ 








■ 








1 1 1 1 



b) 



0.1 i 



0.05 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove Colonized Macroalgae/ Unvegetated Mangrove 

Hardbottom Seagrass Sediments 

Figure 3. 19. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for coney (C. fulvaj. 



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

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). 



0.5 i 








■ Juveniles/Subadults 


CM 








□ Adults 


E 




18.0 






o 

o 

J* 












fulva density 

o 
en 












4.3 


6 


6.3 












£ 

o 

IE 






6.1 


3.0 

i 




















1.4 

T 


I 


1 












n . 







■ 




.1 












oo 


U 1 


Colonized Colonized Col. pav 


Linear 


Macro- Mangrove Patch 


Reef Scat coral/ Seagrass Uncon. 


bedrock pave me 


mt sand cha 


n. reef 


algae reef 


rubble rock sand sediment 


Fish size classes 










Juvenile <5 - 10 cm 








4 *v>> *t\ 


Subadult 10 -15 cm 








y *- 




£ Adult >1 5 










Benthic HABITAT 










Coral reef and 
colonized hardbottom 










Submerged vegetation 










Unconsolidated sediments 






• • • * • • 


.* * . ••..••- 






1 k 

1 l 


n 




# ♦• # •• 


/•%v * 
















w m 









CO 
CD 



E 

E 
o 

O 

CO 



CO 



Figure 3.20. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for coney (C. fulvaj. Number above 
error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for coney (C. fulvaj in the 
southwest Puerto Rico study area. 



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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) 

Red 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 3.21 b). 










i- 






M 



Red hind {Epinephelus guttatus) 




E. guttatus biomass (g/100m 2 ) 

Benthic HABITAT 

1 . 500 Coral reef and colonized hardbottom 
^■500.1-1000 Submerged vegetation 

^H >iooo 



Unconsolidated sediments 



Figure 3.21. Maps of the interpolated (left map) and spatial (right map) distributions for red hind (E. guttatusj; (a) abundance and (b) 
biomass. 



a) 



0.2 n 



0.1 - 



3 
UJ 

c 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



1= 

o 



0.1 n 



0.05 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.22. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for red hind (E. 
guttatusj. 



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

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). 



0.2 



o 
o 

3* 

"</3 

c 

<D 

s 



0.1 



3 

uj 

c 

CO 



12.7 



■ Juveniles/Subadults 
D Adults 



2.7 2.7 



8.5 



ri 



0.5 



4.0 



3.0 







2.1 



ii 







Colonized Colonized Col. pav. 
bedrock pavement sand chan. 



Fish size classes 

Subadult 10 - 20 cm 

£ Adult >20 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 



Linear 
reef 



Macro- 
algae 



Mangrove 



Patch 
reef 



Reef 
rubble 



Scat coral/ Seagrass 
rock sand 



Uncon. 
sediment 



CO 
CD 



E 

E 
o 

O 

CO 



CO 




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. guttatusj in 
the southwest Puerto Rico study area. 



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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) 

Snappers (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 m 2 ) 



o 

I -10 

II -50 
51 - 100 
>100 



Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 









3 o ## 






o o 

"- ; § c oc 





• 8** 

q, o 



o « 



a^o? 




.°. # .°*f - / • S °. °„"# <W $/<& • **&~°« 



• •• ° 



*° »x>o 8 o a 



9 6 J o c 
o 



o O, 



o o^ 



•o^ 



o* *° 



o o. 



Snapper biomass (g/100 m 2 ) 



^H 0.1 - 500 Benthic HABITAT 

500.1 - 1000 Coral reef and colonized hardbottom 

1000.1-5000 Submerged vegetation 

I >5000 Unconsolidated sediments 

Figure 3.24. Maps of the interpolated (left map) and spatial (right map) distributions for snapper (Lutjanidae): (a) abundance and (b) 
biomass. 



1 km 










Q- ; O 



a) 20 



10 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



b) 



1.2 



0.6 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.25. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for snapper species 
(Lutjanidae). 



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

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. 



a) 



50 



Juveniles/subadults 



25 



% 

o 



b) 



25 



c 
o 

o 



25 







so -r 



niles/subadults 



25 - 



<5 5-10 1 



0-15 15-20 20-25 
Size class (cm) 



25-30 30-35 



>35 




50 -r 



Juveniles/subadults 




d) 



50 n 



10-15 15-20 20-25 25-30 
Size class (cm) 

niles/subadults 



30-35 



£ 25 - 



I 



5-10 10-15 15-20 20-25 25-30 30-35 >35 
Size class (cm) 

©I __ Juveniles/subadults 




□ - 



10-15 15-20 20-25 25-30 
Size class (cm) 



30-35 >35 



<5 



>35 



5-10 10-15 15-20 20-25 25-30 30-35 
Size class (cm) 

Figure 3.26. Size frequency histogram for select snapper (Lutjanidae) in the southwest Puerto Rico study area, (a) schoolmaster (1. 
apodusj, (b) gray snapper (L. griseusj, (c) mahogany snapper (1. mahogonij, (d) lane snapper (1. synagrisj and (e) yellowtail snapper (O. 
chrysurusj. 



CO 

CD 



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

Schoolmaster (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). 



CO 




L. apodus 




L. apodus biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

1000.1 - 5000 Unconsolidated sediments 

^H >5000 

Figure 3.27. Maps of the interpolated (left map) and spatial (right map) distributions for schoolmaster (1. apodusj; (a) abundance and (b) 
biomass. 



a) 



8. 



o 

& 



14 



b) 



E 
o 

1 

o 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



0.8 n 



0.4 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.28. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for schoolmaster 
(L. apodusj. 



78 



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

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. apoduswere 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). 



14 -i 
















■ Juveniles/Subadults 








96.2 


D Adults 


CM 

E 






1 




o 






n 




o 










^™ 










% 




















> 










*j 










"35 










c 










o 7 ■ 










T3 










CO 










5 










■c 










o 










a 










rc 










J 










c 










<0 










0) 










§ 






■29.5 




n . 


91 3.6 7.5 2.3 


9.0 40 


in a™ 


5.3 ia 3j? 2.1 1.1 3 - 2 


Colonized Colonized Col. pav. 


Linear 


Macro- Mangrove Patch 


Reef Scat coral/ Seagrass Uncon. 


bedrock pavement sand chan. 


reef 


algae reef 


rubble rock sand sediment 




L. apodus 




#^ L 


%CW^x 


Fish sizG classes 








Juvenile <5 - 15 cm 








Subadult 15-20 cm 






£ Adult >20 


^^ 


^:- 















Benthic HABITAT 


S m 


• # 


f 




^^m Coral reef and 

colonized hardbottom 


• 


• 


• 

• 




Submerged vegetation 
Unconsolidated sediments 


• 
• 




• 


* . • 

• • 




f 


• 
• 


• 
• 


i 




1 km 

1 I 




• 

1 

• 


• 
• 









CO 
CD 



E 

E 
o 

O 

CO 



CO 



Figure 3.29. Mean density (+SE) for juvenile/subadult and adult by mapped habitat type for schoolmaster (L. apodusj. Number 
above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for schoolmaster (L. 
apodusj in the southwest Puerto Rico study area. 



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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) 

Gray 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) 





L. griseus biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

1000.1 - 5000 Unconsolidated sediments 

^H >5000 

Figure 3.30. Maps of the interpolated (left map) and spatial (right map) distributions for gray snapper (1. griseus,); (a) abundance and (b) 
biomass. 



a) 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.4 



0.2 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.31. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for gray snapper 
(1_. griseusj. 



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

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 1 1 habitat types (Figure 3.32). 



o 
o 



* 



c 


S 

Q> 



rc 
o 













48.7 

n 








■ Juveniles/Subadults 

■ Adults 


50.0 

T 




















1 


1.8 0.1 


0.5 


20 1.0 





1^ 


1.8 o 








2-1 1.1 1-7 

•--=== 1— =*= 1 i 



Colonized Colonized Col. pav. Linear 
bedrock pavement sand chan. reef 



Macro- Mangrove Patch 
algae reef 



Reef Scat coral/ Seagrass Uncon. 
rubble rock sand sediment 



CO 
CD 



E 
E 

o 
O 

CO 



CO 




Fish size classes 

Q Juvenile <5 - 15 cm 
Subadult 15 -20 cm 
Adult >20 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 



Figure 3.32. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for gray snapper (L. griseusj. 
Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for gray 
snapper (1_. griseusj in the southwest Puerto Rico study area. 



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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) 

Mahogany 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 biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

Submerged vegetation 
Unconsolidated sediments 



Figure 3.33. Maps of the interpolated (left map) and spatial (right map) distributions for mahogany snapper (L. mahogonij; (a) abundance 
and (b) biomass. 



a) 



0.4 n 



b) 



0.2 - 



0.02 i 



o 
!5 

"5 
o 

o 



0.01 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.34. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for mahogany 
snapper (1. mahogonij. 



82 



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

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). 



0.4 i 










■ Juveniles/Subadults 












D Adults 


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o 

3* 












mahogoni density 

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5.0 




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rubble 


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1 km 

1 l 





























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CD 



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E 
o 

O 

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CO 



Figure 3.35. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for mahogany snapper (L. 
mahogonij. Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult 
for mahogany snapper (L. mahogonij in the southwest Puerto Rico study area. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Lane 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 biomass (g/100 m 2 ) 
Benthic HABITAT 



0.1 -500 



Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



Figure 3.36. Maps of the interpolated (left map) and spatial (right map) distributions for lane snapper (L. synagrisj; (a) abundance and (b) 
biomass. 



a) 






0.3 i 



0.15 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.02 n 



w 
w 

CO 

E 

E 0.01 



_ri_ 



_ti 



Colonized Macroalgae/ Unvegetated Mangrove 

Hardbottom Seagrass Sediments 



Figure 3.37. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for lane snapper 
(1_. synagrisj. 



84 



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

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). 



CO 
CD 



E 

E 
o 

O 



1.2 -I 








■ Juveniles/Subadults 


^ 








D Adults 


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Fish size classes 




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colonized hardbottom 


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• 
• 




L^—~ ^ 


Submerged vegetation 


• 


• 




• 


Unconsolidated sediments 


1 km 

1 I 




• i 


• 



CO 



CO 



Figure 3.38. Mean density (+SE) for juvenile/subadult and adult by mapped habitat type for lane snapper (L. synagrisj. 
Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for lane 
snapper (1_. synagrisj in the southwest Puerto Rico study area. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Yellowtail snapper (Ocyurus chrysurus) 
0. 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 
■ o 

^M 0.1 -io 

^M 10.1 -20 



abundance (100 m 2 ) 
Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



■" PC 



o # £ o 9 o 



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O • o G ' 



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c ic~ 




O. chrysurus biomass (g/100m 2 ) 

Benthic HABITAT 

1 - 500 Coral reef and colonized hardbottom 
500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 



Figure 3.39. Maps of the interpolated (left map) and spatial (right map) distributions for yellowtail snapper (O. chysurusj; (a) abundance 
and (b) biomass. 



a) 



3 



1.2 n 



0.6 - 



b) 




o 

c 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



0.2 n 



0.1 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.40. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for yellowtail 
snapper (O. chysurusj. 



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

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 1 1 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). 



CO 
CD 



E 

E 
o 

O 

CO 



18 i 


100.0 




■ Juveniles/Subadults 










n Adults 




E 












o 
o 












chrysurus density 
















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fi . 







29.7 39 -° 
^21.6 ^-,21.1 


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Colonized Colonized Col. pav. 


Linear Macro- Mangrove Patch 


Reef Scat coral/ Seagrass Uncon. 


bedrock pavement sand chan 


reef algae reef 


rubble rock sand 


sediment 


Fish size classes 

£ Juvenile <5 - 15 cm 
£ Subadult 15-20 cm 


O. chrysut 


ftp )Qc, (L ^ J9 •* 








£ Adult >20 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 


• 

*. 4 








Submerged vegetation 
Unconsolidated sediments 




• • • • 
• % • • •• 
• • •• •• s 

• • • 
• • • - •• 


• 






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1 km 

1 I 


• 


• 















CO 



F/gi/re 3.41. Mean density (+ SE) forjuvenile/subadult and adult by mapped habitat type for yellowtail snapper (O. chysurusj. 
Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for yellowtail 
snapper (O. chysurusj in the southwest Puerto Rico study area. 



page 



CO 

CD 



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

Parrotfish (Scaridae) 

Scaridae abundance follows closely the distribution of colonized 
hardbottom, although theabundance 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 



E 

E 
o 

O 

■ _ 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 





Parrotfish abundance (100 m 2 ) 





Benthic HABITAT 


^H 1 


Coral reef and colonized hardbottom 


^M 11 -25 


Submerged vegetation 


^M 26-50 


Unconsolidated sediments 


^H >50 





o 




n 







Parrotfish biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

1000.1 -5000 Unconsolidated sediments 

■1 >5000 

Figure 3.42. Maps of the interpolated (left map) and spatial (right map) distributions for parrotfish (Scaridae): (a) abundance and (b) 
biomass. 



a) 



C 
0) 

■a 



c 



24 



12 



b) 



E 
o 
'E 



o 



1.2 i 



0.6 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Un vegetated 
Sediments 



Mangrove 



Macroalgae/ 
Seagrass 

Figure 3.43. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for parrotfish species 
(Scaridae). 



. _ JuYenUes/siikadiUtS- 




b) 



50 y* 



liles/subadults 



25 - 



10-15 15-20 20-25 25-30 
Size class (cm) 



30-35 



>35 




10-15 15-20 20-25 25-30 
Size class (cm) 



30-35 >35 



d) 



50 -r 



Juveniles/subadults 



£ 25 

c 
c 
.5. 

1 



5-10 10-15 15-20 20-25 25-30 30-35 >35 
Size class (cm) 




Q 



5-10 10-15 15-20 20-25 25-30 30-35 
Size class (cm) 



>35 



10-15 15-20 20-25 25-30 30-35 



>35 



Size class (cm) 

Figure 3.44. Size frequency histogram for select parrotfish (Scaridae) in the southwest Puerto Rico study area, (a) striped parrotfish (S. 
iserij, (b) princess parrotfish (S. taeniopterusj, (c) redband parrotfish (S. aurofrenatumj, (d) yellowtail parrotfish (S. rubripinnej and (e) 
stoplight parrotfish (S. viridej. 



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

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). 



CO 

CD 



E 

E 
o 

O 

CO 



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

Striped 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. 




S. iseri abundance (100 m 2 ) 

Benthic HABITAT 

1-10 Coral reef and colonized hardbottom 

11-25 Submerged vegetation 

26 - 50 Unconsolidated sediments 

■ >50 




i» y«» •• • . • *? * ' 



i 






} n°0 



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•A • 





S. /sen biomass (g/100 m 2 ) 

Benthic HABITAT 
I 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 






* •.. -J& .V. •*! */•*.*••* • . *'V. % *.; ' 

■ ' o*/;.. •oo.* 6 */'*^ o a **: • ' 

• ••• • * * ♦• V% • 



• •-. 



/-.% 



Figure 3.45. Maps of the interpolated (left map) and spatial (right map) distributions for striped parrotfish (S. iserij; (a) abundance and (b) 
biomass. 



a) 



10 i 



b) 



E 
o 
'E 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



0.4 



0.2 



Colonized 
Hardbottom 



X±l 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.46. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for striped 
parrotfish (S. iserij. 



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

S. /ser/ 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). 



CO 
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0) 

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100.0 



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□ Adults 



74.5 




25.6 



35.1 








Colonized Colonized Col. pav. Linear 
bedrock pavement sand chan. reef 



Macro- Mangrove Patch 
algae reef 



Reef Scat coral/ Seagrass Uncon. 
rubble rock sand sediment 



CO 



CO 




Fish size classes 

Juvenile <10 cm 
• Adult >10 



Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 



page 



CO 

CD 



E 

E 
o 

O 

CO 



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

Princess 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) 





S. taeniopterus abundance (100 m 2 ) 
Benthic HABITAT 



I -10 

II -25 
26-50 
>50 



Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 





S. taeniopterus biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

^M 1000.1 -5000 
^H >5000 



Unconsolidated sediments 



D O 



••• & «° m •• • 






F/gi/re 3.48. Maps of the interpolated (left map) and spatial (right map) distributions for princess parrotfish (S. taeniopterusj; (a) abundance 
and (b) biomass. 



a) 



CO 

C 



E 

o 



0.4 



0.2 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.49. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for princess 
parrotfish (S. taeniopterusj. 



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

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). 



o 
o 

T— 

3* 

"c/3 

c 
o 

s 



CD 
CO 

c 

CO 

o 



36.9 



29.7 



27.0 



■ Juveniles/Subadults 
□ Adults 



48.8 



36.6 



2O.0 






12.8 



23.6 



2.4 4.9 







10.9 
T 10.6 

I .. I 



4.3 3.2 



0.4 0.4 



Colonized Colonized Col. pav. 
bedrock pavement sand chan. 



Linear 
reef 



Macro- 
algae 



Mangrove 



Patch 
reef 



Reef 
rubble 



Scat coral/ Seagrass Uncon. 
rock sand sediment 



CO 
CD 



E 
E 

o 
O 

CO 



CO 



Fish size classes 

Juvenile <10 cm 
# Adult >10 



Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 




1km )(■ 9 gp4 .•• • 



Figure 3.50. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for princess parrotfish (S. 
taeniopterusj. Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile and adult for 
princess parrotfish (S. taeniopterusj in the southwest Puerto Rico study area. 



page 



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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) 

Red band 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.) 




CO 




. aurofrenatum biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 



Figure 3.51. Maps of the interpolated (left map) and spatial (right map) distributions for redband parrotfish (S. aurofrenatumj; (a) 
abundance and (b) biomass. 



a) 



■D 

i 



£ 



E 
o 
!o 
S 



0.2 



J_ 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.52. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for redband 
parrotfish (S. aurofrenatumj. 



94 



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

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). 



o 
o 

3* 

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c 

<D 



2! 
S 

c 

CO 





■ Juveniles/Subadults 
□ Adults 



83.6 



13.1 



78.2 



63.1 



68.5 

J. 



67.0 



81.0 

I 



67.3 



7.3 
2.4 ^ ™ 0.6 



30.9 

S34.0 
T 
■— •— ■— ^™ 



1.3 



4.3 



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Colonized Colonized Col. pav. 
bedrock pavement sand chan. 



Linear 
reef 



Mangrove 



Patch 
reef 



Reef 
rubble 



Scat coral/ Seagrass Uncon. 
rock sand sediment 



Fish size classes 

Juvenile <10 cm 
# Adult >10 



Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 




Figure 3.53. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for red band parrotfish (S. 
aurofrenatumj. Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile and adult for 
redband parrotfish (S. aurofrenatumj in the southwest Puerto Rico study area. 



CO 



E 

E 
o 

O 



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

Bucktooth 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). 




CO 




PT V .¥« 



S. radians abundance (100 m 2 ) 
Benthic HABITAT 

1-10 Coral reef and colonized hardbottom 

11-25 Submerged vegetation 

>25 Unconsolidated sediments 



S. radians biomass (g/100 m 2 ) 

Benthic HABITAT 

I 0.1 - 370 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. radiansj; (a) abundance 
and (b) biomass. 



a) 



CO 

C 



b) 



0.02 i 



0.01 - 



1 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.55. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for bucktooth 
parrotfish (S. radiansj. 



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

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). 



a) 



0.4 n 



C 

c 



0.2 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 




Terminal phase yellowtail parrotfish (Sparisoma rubripinne) in St. 

Croix, USVI. 



b) 



Z 0.04 i 



o 
'E 

c 
,c 
.a 



0.02 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.56. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for yellowtail 
parrotfish (S. rubripinnej. 




S. rubripinne abundance (100 m 2 ) 
Benthic HABITAT 

1-10 Coral reef and colonized hardbottom 

>10 Submerged vegetation 

Unconsolidated sediments 



b) 




3T ; 


™ * 


^^P^* 






9 








JL 




• 




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1 km 




























IDW interpolation using 11 S7 samples 



S. rubripinne biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 



>1000 



Unconsolidated sediments 




Figure 3.57. Maps of the interpolated (left) and spatial (right) distributions for yellowtail parrotfish (S. rubripinnej; (a) abundance and (b) 
biomass. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Stoplight 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 





viride abundance (100 m 2 ) 
Benthic HABITAT 



1 -10 
>10 



Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



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>£ o . .oS. . °. » .' 



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Wr/de biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

■ 1000.1 -5000 

■ >5000 



Unconsolidated sediments 



1 km 




o °o ° °«* # # • • fe % *& 5^°V i ^ |«'oo. o cP" <§ifc •" " o\ 






F/gure 3.58. Maps of tf?e interpolated (left map) and spatial (right map) distributions for stoplight parrotfish (S. viridej; (a) abundance and 
(b) biomass. 



a) 



CO 

C 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.6 



0.3 



CO 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



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) 

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). 



o 
o 

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c 

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s 

c 



65.5 



■ Juveniles/Subadults 

■ Adults 



100.0 



I 3 '" 1 48.0 

I T 46.0 

50.0 32.4 I j T 

ill . . £■ 



149.1 



2.6 



25.5 

Ik 



3.2 3.2 



Colonized Colonized Col. pav. Linear 
bedrock pavement sand chan. reef 



Macro- 
algae 



Mangrove 



Patch 
reef 



Reef Scat coral/ Seagrass Uncon. 
rubble rock sand sediment 



Fish size classes 

Q Juvenile <5 - 10 cm 
Subadult 10 - 15 cm 
Adult >15 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 




1 km •• W * * V * •/. # * •• 



CO 
CD 



E 

E 
o 

O 

CO 



CO 



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. viridej in the southwest Puerto Rico study area. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Grunts (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 3.61 a 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). 

r^ i ■- i-ii ii -- ■- / i ■ i Assemblage of tomtates (Haemulon aurolineatum). 

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. 





Grunt abundance (100 m 2 ) 

Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 





I -10 

II -25 
26-50 
>50 



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Grunt biomass (g/100 m 2 ) 

Benthic HABITAT 

0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

1000.1 - 5000 Unconsolidated sediments 

>5000 



Figure 3.61. Maps of the interpolated (left map) and spatial (right map) distributions for grunt (Haemulidae): (a) abundance and (b) biomass. 



100 



Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007) 
a ) 50 n b ) 0.8 n 



25 



3 



0.4 



Colonized 
Hard bottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



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). 



Juveniles/subadults 




iles/subadults 



c) 



25 




50 -r 



5-10 10-15 15-20 20-25 25-30 30-35 

Size class (cm) 

Juveniles/subadults _^ 



>35 



hi 



d) 



25 



I I 



5-10 10-15 15-20 20-25 25-30 

Size class (cm) 



30-35 



>35 



I I 



5-10 10-15 15-20 20-25 25-30 30-35 

Size class (cm) 



>35 



50 -i 



Juveniles/subadults 



5-10 10-15 15-20 20-25 25-30 

Size class (cm) 



30-35 



>35 



Figure 3.63. Size frequency histogram for select grunts (Haemulidae) in the southwest Puerto Rico study area, (a) tomtate (H aurolineatumj, 
(b) French grunt (H flavolineatumj, (c) white grunt (H. plumierii,) and (d) bluestriped grunt (H. sciurus,). 



CO 

CD 



E 

E 
o 

O 

CO 



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

Tomtate (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 





H. aurolineatum abundance (100 m 2 ) 
Benthic HABITAT 



I -10 

II -25 
26-50 
>50 



Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



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IDW interpolatk 



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H. aurolineatum biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 



1000.1 -5000 
>5000 



Unconsolidated sediments 




Figure 3.64. Maps of the interpolated (left map) and spatial (right map) distributions for tomtate (H aurolineatumj; (a) abundance and (b) 
biomass. 



a) 

^ 12 



c 
o 



b) 



0.1 



E 
o 

a £ 005 



■ 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.65. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for tomtate (R 
aurolineatumj. 



102 



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

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). 



CO 
CD 



E 

E 
o 

O 



o 
o 



CO 

c 
a> 

s 

CO 
CD 

! 

CtJ 



■ Juveniles/Subadults 

■ Adults 

18.1 



7.3 




5.5 



10.8 



5.8 







I 6.5 

8.5 4.7 

1 ■ l°- 4 ■ I 1 - 1 . 



Colonized Colonized Col. pav. 
bedrock pavement sand chan. 



Linear 
reef 



Macro- Mangrove Patch 
algae reef 



Reef Scat coral/ Seagrass Uncon. 
rubble rock sand sediment 



Fish size classes 

Q Juvenile <5 - 10 cm 
Subadult 10 -15 cm 
% Adult >1 5 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 




CO 



CO 



Figure 3.66. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for tomtate (H aurolineatumj. 
Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for tomtate 
(H aurolineatumj in the southwest Puerto Rico study area. 



CO 

CD 



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

French 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). 



CO 




French grunt {Haemulon flavolineatum) 




Figure 3.67. Maps of the interpolated (left map) and spatial (right map) distributions for French grunt (R flavolineatumj; (a) abundance and 
(b) biomass. 



a) 

L 30 



3* 



15 



b l 0, 



0.2 



I 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.68. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for French grunt 
(H flavolineatumj. 



104 



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

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). 



30 



o 
o 



10 

c 

CD 

I 

*«* 
CO 
CD 
,C 

1 



c 

CO 
CD 



15 



71.8 



■ Juveniles/Subadults 

■ Adults 



24.37.2 



27.3 



16.0 



23.0 



Ik. 



2.4 o 



10.6 



16.4 4.3 




16.0 



Colonized Colonized Col. pav. 
bedrock pavement sand chan. 



Linear 
reef 



Macro- 
algae 



Mangrove 



Patch 
reef 



Reef 
rubble 



Scat coral/ Seagrass Uncon. 
rock sand sediment 



CO 
CD 



E 
E 

o 
O 

CO 



CO 



Fish size classes 

Q Juvenile <5 - 10 cm 
Subadult 10 -15 cm 
£ Adult >1 5 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 




• V 

••• 






>»• 






1 km 

J 






• *•• .. ..• . 

# • 







•• • 






Figure 3.69. Mean density (+ SE) forjuvenile/subadult and adult by mapped habitat type for French grunt (H flavolineatumj. 
Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for French 
grunt (H. flavolineatumj in the southwest Puerto Rico study area. 



CO 

CD 



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

White 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. 



CO 




White grunt {Haemulon plumierii) 




H. plumierii biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 

Figure 3.70. Maps of the interpolated (left map) and spatial (right map) distributions for white grunt (R plumieriij; (a) abundance and (b) 
biomass. 



a) 



3 i 



1.5 - 



| 



I 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



b) 



0.2 



o 



a: 



0.1 



^ ^ 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.71. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for white grunt 
(H plumieriij. 



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

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. 



o 
o 

"35 

£ 
0) 
T3 2 

: £ 

Q) 

E 

a 

c 

Q) 



14.6 



■ Juveniles/Subadults 

■ Adults 



18.4 






1 2.4 



8.5 



12.6 7.5 



10.8 







£n rfi 



8.0 



5.0 



14.1 



i 



73 12.7 







Ik 



1.1 3.2 



Colonized Colonized Col. pav. Linear Macro- Mangrove Patch 

bedrock pavement sand chan. reef algae reef 



Reef Scat coral/ Seagrass Uncon. 
rubble rock sand sediment 



CO 
CD 



E 

s 

o 

CO 



CO 



Fish size classes 

Q Juvenile <5 - 10 cm 
£ Subadult 10 -15 cm 
£ Adult >1 5 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 



H. plumerii 




Figure 3. 72. Mean density (+ SE) forjuvenile/subadult and adult by mapped habitat type for white grunt (H plumieriij. Number 
above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for white grunt (H 
plumieriij in the southwest Puerto Rico study area. 



CO 

CD 



CO 



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

Bluestriped 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) 





H. sciurus biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

^M 1000.1 -5000 
^H >5000 



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. 



a) 



14 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.6 



o 
!5 



0.3 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3. 74. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for bluestriped 
grunt (H sciurusj. 



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

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. 



14 



o 
o 

>> 

"35 

c 

0) 
"D 
</> 

2 

3 

o 
to 



c 

(0 

CD 



■ Juveniles/Subadults 
□ Adults 



75.0 



50.0 



2.7 4.210.3 3.014.0 



4.9 



25.6 7 - 2 32 

1 . mil _ . ■ 



8.5 



8.1 



10.9 



1.1 3.2 



Colonized Colonized Col. pav. 
bedrock pavement sand chan. 



Linear 
reef 



Macro- 
algae 



Mangrove 



Patch 
reef 



Reef 
rubble 



Scat coral/ Seagrass Uncon. 
rock sand sediment 



Fish size classes 

Q Juvenile <5 - 10 cm 
Subadult 10 -15 cm 
£ Adult >15 

Benthic HABITAT 

Coral reef and 
colonized hardbottom 

Submerged vegetation 

Unconsolidated sediments 



CO 
CD 



E 
E 

o 
O 

CO 



CO 




Figure 3.75. Mean density (+ SE) for juvenile/subadult and adult by mapped habitat type for bluestriped grunt (R sciurusj. 
Number above error bar indicates percent occurrence. Bottom: Spatial distribution of juvenile, subadult and adult for 
bluestriped grunt (H sciurusj in the southwest Puerto Rico study area. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Surgeonfish (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). 





.j& m**% 


^Jtt^ 




*L <• ■ 


^■^fjk.% 


~> 


■ 


«~ 




_K&.~ 


> 


SP* 




• ; 


l 




I? 


"•' .-; 


r&k 



Assemblage of blue tangs (Acanthurus coeruleus). 




Surgeonfish biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

1000.1 - 5000 Unconsolidated sediments 

^H >5000 

Figure 3.76. Maps of the interpolated (left map) and spatial (right map) distributions for surgeonfish (Acanthuridae): (a) abundance and (b) 
biomass. 



a) e 



E 

o 



b) 



£ 
o 
!5 



0.6 -i 



0.3 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Unvegetated 
Sediments 



Mangrove 



Macroalgae/ 
Seagrass 

Figure 3. 77. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for surgeonfish 
(Acanthuridae). 



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

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). 



a) 



so -r 



Juveniles/subadults 



Juveniles/subadults 




10-15 15-20 20-25 
Size class (cm) 



25-30 30-35 



>35 




CO 
CD 



E 
E 

o 
O 

CO 



CO 



5-10 



10-15 15-20 20-25 25-30 
Size class (cm) 



30-35 



C) 



50 -r 



Juveniles/subadults 



25 



M 



<5 



5-10 



10-15 



15-20 20-25 
Size class (cm) 



25-30 30-35 



>35 



Figure 3. 78. Size frequency histogram for surgeonfish (Acanthuridae) in the southwest Puerto Rico study area, (a) ocean surgeon (A. 
bahianusj, (b) doctorfish (A. chirurgusj and (c) blue tang (A. coeruleus). 



page 



CO 

CD 



E 

E 
o 

O 

CO 



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

Ocean 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). 



xArf 












A. bahianus abundance (100 m 2 ) 
Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 





I -10 

II -25 
26-50 
>50 



(HO 






oo «^co- ^ 

* o 
# . c as, cr o ~-q o A> 

o • •!<*• •• o «° ^.r>°P 



<^*o 






.».. . .°o o0, .»° °° % Jq °tii?*»\« • .° @ © oo. °<P <#*» o 

1km 




A bahianus biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 100 Coral reef and colonized hardbottom 

100.1 - 500 Submerged vegetation 

500.1 - 1000 Unconsolidated sediments 

^M >iooo 

Figure 3.79. Maps of the interpolated (left map) and spatial (right map) distributions for ocean surgeonfish (A. bahianusj; (a) abundance 
and (b) biomass. 



a) 



E 

o 

o 



3 
C 

■5 



n 



Colonized 
Hardbottom 



Macroalgae/ Unvegetated 

Seagrass Sediments 



Mangrove 



b) 



0.4 n 



£ 
o 
!5 



0.2 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



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,). 



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

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. chirurgus biomass (g/100 m 2 ) 
Benthic HABITAT 



0.1 - 500 
500.1 -1000 
1000.1 - 5000 
>5000 



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. 



a) 



1 n 



3 



0.5 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.2 -i 



o 
!5 



c 



0.1 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.82. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for doctorfish (A. 
chirurgusj. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Blue 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 abundance (100 m 2 ) 





Benthic HABITAT 


^H 1 


Coral reef and colonized hardbottom 


^H 11 -25 


Submerged vegetation 


^M 26-50 


Unconsolidated sediments 


j^H >50 







A. coeruleus biomass (g/100 m 2 ) 
■ o Benthic HABITAT 



0.1 - 500 
500.1 -1000 
1000.1 - 5000 
>5000 



Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



Boo 






::: :;- : -.: O 



° .°f 



O GO qj 



8 .5c 

o o » 



•<£P # 



■ •. ... ■-• •, 






o« ..;- 



#tf 






Figure 3.83. Maps of the interpolated (left map) and spatial (right map) distributions for blue tang (A. coeruleusj; (a) abundance and (b) 

biomass. 

a) b) 



T3 

3 



0.2 n 



£ 
o 
!5 



0.1 



_L 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.84. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for blue tang (A. 
coeruleusj. 



38* 



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

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). 




C/) 
CD 



P. maculatus 



CO 



a) 



50 -i* 



Juveniles/subadults 



25 







b) 



50 t 



Juveniles/subadults 



u- 25 - 



<5 5-10 10-15 15-20 20-25 25-30 30-35 >35 

Size class (cm) 




<5 5-10 10-15 15-20 20-25 25-30 30-35 >35 

Size class (cm) 



Figure 3.85. Size frequency histogram for goatfish (Mullidae) in the southwest Puerto Rico study area, (a) yellow goatfish (W\. martinicusj 
and (b) spotted goatfish (P. maculatusj. 



page 



CO 



E 

E 
o 

O 



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



Yellow 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 



CO 




M. martinicus abundance (100 m 2 ) 
Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 





I -10 

II -25 
26-50 
>50 




M. martinicus biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

^■500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 




Figure 3.86. Maps of the interpolated (left map) and spatial (right map) distributions for yellow goatfish (M. martinicusj; (a) abundance and 
(b) biomass. 



a) 



1.4 i 



§ 0.7 

o 

c 



b) 

J. 0.1 



n ^ 



o 

to 

3 
O 

c 



0.05 



Colonized Macroalgae/ Unvegetated 

Hardbottom Seagrass Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.87. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for yellow goatfish 
(M. martinicusj. 



116 



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

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 





P. maculatus 

o 
^M 1 -io 

^M 11 -25 
^H 26-50 
^H >50 



abundance (100 m 2 ) 
Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 



1 km 






Q. 



3° c -or x 






1 °°^ 



„<*> 



o 6> n 




P. maculatus biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

^■500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 




Figure 3. 88. Maps of the interpolated (left map) and spatial (right map) distributions for spotted goatfish (P. maculatusj; (a) abundance and 
(b) biomass. 



a) 



q: 

c 



0.8 n 



co 0.4 

3 



b) 



q: 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



0.1 



0.05 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.89. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for spotted 
goatfish (P. maculatusj. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Jacks (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). 



<4a ^ 




\ 


«» 




r 'i^k. 






* 


'"'- ^v. 






y 


- 





Yellow jack (Carangoides bartholomaei) 




Jack abundance (100 m 2 ) 

Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 





I -10 

II -25 
26-50 
>50 



> c °5> 

' o o "*aBoo» oo ° ^E <£°°fP o °° *• °^ 



^ 



qOo ° o 



1 km 



o o c 



°CDO 



^oo 



Ho® 



c£ oc 




Jack biomass (g/100 m 

Benthic HABITAT 

0.1 - 1000 Coral reef and colonized hardbottom 

1000.1 - 5000 Submerged vegetation 

5000.1 - 10000 Unconsolidated sediments 

^M >ioooo 



Figure 3.90. Maps of the interpolated (left map) and spatial (right map) distributions for jack (Carangidae): (a) abundance and (b) biomass. 



a) 



b) 



0.4 i 



£ 
o 
5 



0.2 - 



d 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



Figure 3.91. Comparison of mean (+ SE) density and biomass by habitat type in the southwest Puerto Rico study area for jack (Carangidae). 



118 



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

Two of the most common Carangidae 

species were Caranxcrysos (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) 



CO 
CD 



E 

E 
o 

O 

CO 



a) 



0.6 n 



0.3 ■ 



s 



o 

c 



b) 



0.3 



o 
!5 



fr 



0.15 



CO 



Colonized 
Hardbottom 



Macroalgae/ Unvegetated Mangrove 

Seagrass Sediments 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.92. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for blue runner 
(C. crysosj. 



a) 



2 -i 



1 - 



CD 

■§ 



b) 



0.1 



E 
o 
!5 

5*. 

CD 
j3 



0.05 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



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,). 



page 



CO 

CD 



E 

E 
o 

O 

CO 



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

Damselfish (Pomacentridae) 

Yellowtail damselfish (Microspathodon chrvsurus) 
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) 





M. chrysurus biomass (g/100 m 2 ) 

Benthic HABITAT 
| 0.1 - 100 Coral reef and colonized hardbottom 

100.1-500 Submerged vegetation 

500.1 -1000 Unconsolidated sediments 



Figure 3.94. Maps of the interpolated (left map) and spatial (right map) distributions for yellowtail damselfish (W\. chrysurusj; (a) abundance 
and (b) biomass. 



a) 



i 



0.8 i 



0.4 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 

_ 0.1 
E 



E 
o 
'E 



3 



0.05 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



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,). 



120 



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

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 





S. adustus biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

>500 Submerged vegetation 

Unconsolidated sediments 



Figure 3.96. Maps of the interpolated (left map) and spatial (right map) distributions for dusky damselfish (S. adustusj; (a) abundance and 
(b) biomass. 



a) 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.06 



o 



c 



0.03 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.97. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for dusky 
damselfish (S. adustusj. 



CO 



E 

E 
o 

O 



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

Beaugregory (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 






, ■«.■-> 



CO 




S. leucostictus abundance (100 m 2 ) 
Benthic HABITAT 

1-10 Coral reef and colonized hardbottom 

11-25 Submerged vegetation 

>25 Unconsolidated sediments 




S. leucostictus biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 290 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. leucostictusj; (a) abundance and 
(b) biomass. 



a) 



5t 



10 



b) 
a- 0.06 

E 

o 



n 



0.03 



Colonized 
Hardbottom 



Macroalgae/ Unvegetated 
Seagrass Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.99. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for beaugregory 
(S. leucostictusj. 



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

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) 





S. partitus biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

>500 Submerged vegetation 

Unconsolidated sediments 



o # 
•o 



°® ° S> o 



Figure 3. 100. Maps of the interpolated (left map) and spatial (right map) distributions for bicolor damselfish (S. partitusj; (a) abundance and 
(b) biomass. 



a) 



CO 

c 



b) 



0.06 i 



n 






0.03 - 






Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.101. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for bicolor 
damselfish (S. partitusj. 



CO 

CD 



E 

E 
o 

O 

CO 



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

Threespot 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). 



*>&?*& 




S. planifrons abundance (100 m 2 ) 





Benthic HABITAT 


^M 1 -io 


Coral reef and colonized hardbottom 


^H 11 -25 


Submerged vegetation 


^H 26-50 


Unconsolidated sediments 


^H >50 






OQO Q Q% o . 



3 O 




* © @ o © ^ o o 



? ooo o6> 



o <9^o 



1 km 



o o c 



o o o ©« 



_^ 



_, 9y 






C bo O c 



, @ & 






61 6 







S. planifrons biomass (g/100 m 2 ) 
Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

>1000 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. 



a) 



* 

> 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



b) 

E 

o 









0.06 n 



0.03 



^ r 1 ! 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



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,). 



124 



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



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. 





S. variabilis abundance (100 m 2 ) 
Benthic HABITAT 

| 1 . 10 Coral reef and colonized hardbottom 

>10 Submerged vegetation 

Unconsolidated sediments 



9 



3 °lM<S>$o* ° o o° ° p o \|c- o. ° o- > . -„ c : o f o. 



•c^^S 



,, „o c 



° °o o cftf to a 






* • •* 



- 

..;' 



oo^%° ooo c 

°n$ * 8> O ( 






o" <9 



> 



o o w 



o 



• o o 
Oqo 



lO 



3° ° r to o c 



Qo o 




S. variabilis biomass (g/100 m 2 ' 
Benthic HABITAT 




0.1 -100 



Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 






o o o* ^M^m • ULA * oQ> u o c 



..>■■• 



1 km 



@ o o 
Oqo 



°l oS o o o c 



::o 






oo 



F/gi/re 3.704. Maps of f/?e interpolated (left map) and spatial (right map) distributions for cocoa damselfish (S. variabilis,); (a) abundance 
and (b) biomass. 



a) 



S 
5 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.02 



0.01 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.105. Comparison of mean (+ SE) density and biomass by substrate type in southwest Puerto Rico for cocoa damselfish (S. 
variabilis,). 



CO 

CD 



E 

E 
o 

O 

CO 



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

Wrasses (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) 





Wrasse abundance (100 m 2 ) 

Benthic HABITAT 

| 1 . 10 Coral reef and colonized hardbottom 

11-50 Submerged vegetation 

51 - 100 Unconsolidated sediments 

■ >100 




Wrasse biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

1000.1 -5000 Unconsolidated sediments 

"1 >5000 




^H >5000 ' ■ ' 

Figure 3. 106. Maps of the interpolated (left map) and spatial (right map) distributions for wrasse (Labridae): (a) abundance and (b) biomass. 



a) 20 



10 



b) 0.2 



0.1 



n 



h 



n 



Colonized Macroalgae/ Unvegetated Mangrove Colonized Macroalgae/ Unvegetated Mangrove 

Hardbottom Seagrass Sediments Hardbottom Seagrass Sediments 

Figure 3.107. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for wrasse 
(Labridae). 



126 



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

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 




Porgy biomass (g/100 m 2 ) 

Benthic HABITAT 

| 0.1 - 1000 Coral reef and colonized hardbottom 

>1000 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. 



a) 



1.2 



0.6 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



E 
o 



0.1 -, 



0.05 - 



1 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.109. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for porgy 
(Labridae). 



CO 

CD 



E 

E 
o 

O 

CO 



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

Other 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 un vegetated 

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) 





o °o o 
° o 



0<9 £ ° 8 „ o o%®c 



o <ri 



°<§> ° o 
oo 
o o r 



%H>Te 



0"t)Q 

o ° 
o @ 






o o ° 

°oo 






if 



°°n ® 



Abundance (100 m 2 ) 


Biomass (g/100 m 2 ) 


Benthic HABITAT 





o 


Coral reef and colonized hardbottom 


^H 1 


^H 1 - 1000 


Submerged vegetation 




^H >iooo 


Unconsolidated sediments 



Southern stingray {Dasyatis americana) 



Figure 3.110. Map of the spatial distributions for sharks and rays: (a) abundance 
and (b) biomass. 



a) 



0.1 



0.05 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Un vegetated 
Sediments 



Mangrove 



b) 



0.8 i 



0.4 - 



n 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.111. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for sharks and 
rays. 



128 



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



Queen triggerfish (Batistes 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). 



. 



yM 




Queen triggerfish (Balistes vetula) 




B. vetula biomass (g/100 m 2 ) 

Benthic HABITAT 

0.1 - 500 Coral reef and colonized hardbottom 

500.1-1000 Submerged vegetation 

>1000 Unconsolidated sediments 



Figure 3. 112. Maps of the interpolated (left map) and spatial (right map) distributions for queen triggerfish (B. vetulaj; (a) abundance and 
(b) biomass. 



a) 



0.2 



0.1 - 



. 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



b) 



0.2 n 



0.1 - 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.113. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for queen 
triggerfish (B. vetulaj. 



CO 



E 

E 
o 

O 



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

Great 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) 




CO 




S. barracuda biomass (g/100 m 2 ) 
Benthic HABITAT 

Coral reef and colonized hardbottom 
Submerged vegetation 
Unconsolidated sediments 





0.1 - 500 

500.1 -1000 

1000.1 - 5000 

>5000 



Figure 3.114. Maps of the interpolated (left map) and spatial (right map) distributions for great barracuda (S. barracudaj; (a) abundance 
and (b) biomass. 



a) 



1.6 -i 



b) 



0.8 



0.4 n 



c/) 

</) 

E 
o 

5 0.2 

3 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Colonized 
Hardbottom 



Macroalgae/ 
Seagrass 



Unvegetated 
Sediments 



Mangrove 



Figure 3.115. Comparison of mean (+ SE) density and biomass by substrate type in the southwest Puerto Rico study area for great 
barracuda (S. barracudaj. 



130 



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



a) 



iles/subadults 




b) 



50 n 



u- 25 



3 



CO 



10-15 




25-30 30-35 



>35 



15-20 20-25 25-30 30-35 >35 <5 5 " 1U 10 " 15 15 " 20 20 " 25 

Size class (cm) Size class (cm) 

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. 

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. 



C/) 
CD 



E 

E 
o 

O 

C/) 



CO 



page 



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

3.4. 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 

Efor 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/1 00m 2 , biomass 
reported in grams/1 00m 2 , and richness (community metrics section only) reported as # species/1 00m 2 . 



CO 



CO 



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 
Mean (SE) 


2002 
Mean (SE) 


2003 
Mean (SE) 


2004 
Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
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 m 2 ) * 


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/1 00 m 2 )* 


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 (/_. 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 (/.. 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) 



132 



Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007) 
Table 3.5. Continued... 





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) 


Red band (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) 


Red band (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) 






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. bah i an us) 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. 



CO 

CD 



E 

E 
o 

O 

CO 



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



A 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. 



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





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 




Batistes vetula abundance 




Batistes vetula biomass 



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



Significantly higher in 2002 Significantly higher in 2007 



Haemulon sciurus abundance 
Haemulon sciurus biomass 
Sparisoma radians biomass 



Total species richness 
Grouper abundance 
Cephalopholis cruentata abundance 
Cephalopholis cruentata biomass 
Lutjanus jocu abundance 
Lutjanusjocu 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 
Batistes vetula biomass 



3.4.1. Fish community metrics 

A summarized subset of community metrics from Table 3.5 is presented in Table 3.8. 

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. 



Community variable 


2001 
Mean (SE) 


2002 
Mean (SE) 


2003 
Mean (SE) 


2004 
Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
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 m 2 )* 


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/1 00 m 2 )* 


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) 



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 



134 



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

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. 

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. 





2001 


2002 


2003 


2004 


2005 


2006 


2007 


Grouper variable 


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) 



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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E. guttatus 



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

than 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. 




C. fulva 



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. 

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. 



2001 2002 2003 2004 2005 2006 2007 
Snapper variable 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) 





136 



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

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 l mahogoni m st. croix, usvi 

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 ± and ± 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. 




Dog snapper (Lutjanusjocu) in St. John, USVI 



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. 




O. chrysurus 



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. 



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. 



Jack variable 




2001 
Mean (SE) 


2002 
Mean (SE) 


2003 
Mean (SE) 


2004 
Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
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 
Carangoides ruber biomass 


0.3(0.1) 
3.7(1.6) 


0.3(0.1) 
11.6(7.5) 


0.9 (0.4) 
10.8(2.5) 


0.3(0.1) 
9.4 (3.7) 


1.6(0.7) 
23.2 (9.4) 


1.3(0.6) 
15.4(6.0) 


0.2 (0.1) 

5.4 (2.8) 



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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) 

Mean 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. 

Fish diver and C. ruber 

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 ± and 
± 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. 

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. 



Parrotfish variable 




2001 
Mean (SE) 


2002 
Mean (SE) 


2003 
Mean (SE) 


2004 
Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
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) 



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, 



138 




Juvenile phase S. taeniopterus 




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

except mean density of S. /sen 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 Juvenile P hase s - iseri 

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 43.1 8 ± 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). 




Terminal phase S. aurofrenatum 



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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) 

Mean 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. 

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. 



Grunt variable 




2001 
Mean (SE) 


2002 
Mean (SE) 


2003 
Mean (SE) 


2004 
Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
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) 



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

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. 




H. aurolineatum 



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. 

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. 



Surgeonfish variable 




Mean (SE) 


Mean (SE) 


2003 
Mean (SE) 


2004 
Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
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) 



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 



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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) 

was 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. bahianuswas 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. 




A. bahianus 




A. chirurgus 



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. 

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. 



Other Families variable 




2001 
Mean (SE) 


2002 
Mean (SE) 


2003 
Mean (SE) 


2004 
Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
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 " 


c 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) 





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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) 

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. 

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. 





Mean (SE) 


Mean (SE) 


Mean (SE) 


Mean (SE) 


2005 
Mean (SE) 


2006 
Mean (SE) 


2007 
Mean (SE) 


Other species variable 


Batistes 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) 


Batistes 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) 



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. 




B. vetula 



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. Foureye butterflyfish (Chaetodon capistratus) 




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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) 



Table 3. 1 7. 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 m 2 ). 









Total 


Winter 


Winter mean 


Summer 


Summer 




mmon name 


density 


density 


density 


density 


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) 



3.5. 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 (August- 
September). 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 




Planktivorous fish blue chromis (Chromis cyanea) 



144 



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



a) 160 i 




T 




b) 6, 






■ Winter 
□ Summer 


E 

o 
o 




E 

o 
o 




T 


Mean fish densit) 

00 

o o 


^T 


I 

i — 1 L — 


ii 


J- 

Mean fish biomass 

o w 


1 
i 




h 



2004 



2005 



2006 



2007 



C) 



16 



2004 



2005 



2006 



2007 



Figure 3.11 
area. 



2004 2005 2006 2007 

7. Seasonal change in mean (+ SE) fish: (a) density, (b) biomass and (c) species richness in the southwest Puerto Rico study 



CD 



a) 48 



•o 24 
!5 



b) 2-, 



rh 



(0 

E 
.2 1 

-Q 

§ 

'E 



m 



■ Winter 
□ Summer 



_L 



c) 



2004 



2005 



2006 



2007 



2004 



2005 



2006 



2007 




2004 2005 2006 2007 

Figure 3.118. Seasonal change in mean (+ SE) herbivore: (a) density, (b) biomass and (c) richness in the southwest Puerto Rico study area. 



CO 

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CO 



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

from 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). 

b) 



c) 




2 n 



■ Winter 
□ Summer 



2004 



2005 



JL 



2006 



2007 



2004 2005 2006 2007 

Figure 3.119. Seasonal change in mean (+ SE) piscivore: (a) density, (b) biomass and (c) richness in the southwest Puerto Rico study area. 

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 3.11 7b), 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). 



146 



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



a) 




b) 2n 




.2 1 

CD 



■ Winter 
□ Summer 



ihhl 



c) 



2004 



2005 



2006 



2007 



2004 



2005 



2006 



2007 



n 



2004 2005 2006 2007 

Figure 3.120. Seasonal change in mean (+ SE) planktivore: (a) density, (b) biomass and (c) richness in the southwest Puerto Rico study 
area. 

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. 



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



■ Winter 
□ Summer 




2004 2005 2006 2007 2004 2005 2006 2007 

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. 



a) o.i 



E 
o 
5 0.05 




C) 01 



b) 



0.5 



■tl 



■ Winter 
□ Summer 



2004 



2005 



2006 



2007 



2004 



2005 



2006 



2007 



3 



0.05 - 



n 



I 



2004 2005 2006 2007 

Figure 3. 122. Seasonal change in mean (+ SE) grouper biomass in the southwest Puerto Rico study area for: (a) graysby (C. cruentataj, 
(b) coney (C. fulvaj and (c) red hind (E. guttatusj. 



148 



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

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. 



a) 0.2 



o 



0.1 - 




C) 0.2 



0.1 



b) 0.2 



o 



0.1 



■ Winter 
□ Summer 



fl 



2004 



2005 



2006 



2007 



2004 



2005 



2006 



2007 




i in 



2004 2005 2006 2007 

Figure 3. 123. Seasonal change in mean (+ SE) snapper biomass in the southwest Puerto Rico study area for: (a) schoolmaster (L. apodusj, 
(b) gray snapper (1. griseusj and (c) yellowtail snapper (O. chrysurusj. 



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. /sen 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 3.1 24b). 
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). 



CO 

CD 



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



CO 




b) 0.5 



co 0.25 



0.5 i 


20 


04 


2005 20 


06 


2007 


0.25 ■ 
■ 


II 


f 


1 n 


f 


1 

1 



d) 



o 0.25 



■ Winter 
□ Summer 




1 1 11*1. 



■ 



2004 



2005 



2006 



2007 




2004 2005 2006 2007 2004 2005 2006 2007 

Figure 3.124. Seasonal change in mean (+ SE) parrotfish biomass in the southwest Puerto Rico study area for: (a) striped parrotfish fSc. 
iserij, (b) princess parrotfish fSc. taeniopterusj, (c) redband parrotfish fSp. aurofrenatumj and (d) stoplight parrotfish fSp. viridej. 



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/1 00m 2 and summer biomass values were greater than 
0.02 kg/1 00m 2 ; 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. 



a) 



Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007) 
0.2 



o 



0.1 - 



C) 




0.2 



b) 0.2 



0.1 



ii, 



3 

a: 



i 



■ Winter 
□ Summer 



ILL 



2004 



2005 



2006 



2007 



2004 



2005 



2006 



2007 



0.1 - 



1^ 



±> 



2004 2005 2006 2007 

Figure 3. 125. Seasonal change in mean (+ SE) grunt biomass in the southwest Puerto Rico study area for: (a) French grunt (H . flavolineatumj, 
(b) white grunt (H. plumieriij and (c) bluestriped grunt (H. sciurusj. 



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). 



CO 
CD 



E 

E 
o 

O 

CO 



CO 



a) 0.2 -I 



E 

o 



E 
o 
!5 

V) 

3 
O 

c 
£ 



0.1 



b) 0.2 



~ 




o 

V) 

to 

3 



c 



0.1 



■ Winter 
□ Summer 



ih 



2004 



2005 



2006 



2007 



2004 



2005 



2006 



2007 



Figure 3.126. Seasonal change in mean (+ SE) goatfish biomass in the southwest Puerto Rico study area for: (a) yellow goatfish (M. 
martinicusj and (b) spotted goatfish (P. maculatusj. 



CO 

CD 



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

Mean 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 
(O.0001 kg/1 00m 2 ) 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). 



a) 



0.2 n 



CO 



0.1 



b) 2 -| 



UQ 



3 



■ Winter 
□ Summer 



i 



■ 



2004 2005 2006 2007 2004 2005 2006 2007 

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. vetulaj and (b) great barracuda (S. barracudaj. 

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). 




Figure 3.128. A spearfisherman with a 9.78 kg rainbow 
parrotfish fScarus guacamaiaj caught in Puerto Rico 
(exact location unknown) on January 29th 2005. Source: 
International Underwater Spearfishing Association. 




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

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 reqion. 

° Midnight parrotfish (Scarus coelestinus) Source: Kevin Eddy 

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 1 980-1 981 . 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 best- 
available 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- 



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

2007, the percentage occurrence of S. planifrons, however, was lower, with S. planifrons present in 
31 % (n=1 67) 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. 

E 
E 



3.7. Summary of results 
O 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 small- 
bodied 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 

?0 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 offish 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 offish diversity, biomass and abundance 

• Across the La Parguera study area a total of 21 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 m 2 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. 



154 



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

• 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 1 1 th 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 1 of the 1 1 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 co- 
occurred 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. 



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

q\\ 3.7.5. 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. /sen 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 1 980- 
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. sthatus, 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 



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

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. 




Baitfish in mangrove habitat 



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 within- 

habitat 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 offish communities on coral reefs along a 
spatial gradient in seagrass cover revealed that fish diversity and 
abundance increased from to 20-30% seagrass coverage then 
plateaued at 40% indicating a threshold like response (Grober- 
Dunsmore, 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 




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the 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 offish 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 (r 2 =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. 




Mangrove habitat 



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 fine- 
scale surveys. The fact that some species may have a critical 

Seagrass habitat 




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

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 offish 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 



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across 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 size- 
dependent 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 offish 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 Home, 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 




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

abundance and biomass in the topographically complex colonized pavement with sand channels habitat 
type of the outer shelf zone. 




Nassau grouper {Epinephelus striatus) 



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 1 979 they were absent 
between 2001 and 2006. In contrast, E. guttatus and C. fulva have increased in density significantly 
from 0.08 individuals/100 m 2 of red hind in 1979 to 0.18/100 m 2 in 2001-2006; and from 0.01/100 m 2 of 
C. fulva in 1979 to 0.30/100 m 2 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. 



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

Caribbean 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. 



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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 

O 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 (1 980) in Jobos Bay, Puerto Rico found peaks 

in zooplankton biomass and copepod densities from August-October. 



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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., C.A. McAlpine, and K.M. Pittman. 2004. Linking fish and prawns to their environment: A hierarchical 
landscape approach. Mar. Ecol. Prog. Ser. 283: 233-254. 

Pittman, S.J., C. Caldow, S.D. Hile, and M.E. Monaco. 2007a. Using seascape types to explain the spatial 
patterns offish in the mangroves of southwest Puerto Rico. Mar. Ecol. Prog. 348: 273-274. 

Pittman, S.J., J. Christensen, C. Caldow, C. Menza, and M. Monaco. 2007b. Predictive mapping offish 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 offish 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 offish abundance and size 
along a subtropical mangrove shoreline: an application of the delta approach. Bull. Mar. Sci. 80: 609-624. 



164 



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

Stallings, CD. 2008. Indirect effects of an exploited predator on recruitment of coral-reef fishes. Ecology 89: 
2090-2095. 



CO 
CD 



Stallings, CD. 2009. Fishery-independent data reveal negative effect of human population density on Caribbean 
predatory fish communities. PLoS One 4(5): e5333. 

van Home, B. 1983. Density as a misleading indicator of habitat quality. J. Wildl. Manage. 47(4): 893-901. 



E 
E 



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. 



CO 



CO 



CO 

CD 



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



E 

E 
o 

O 

CO 



CO 



page 
166 



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

Appendix A 



Table A1. Fin fish landings as a proportion of the total fin fish 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). 



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 (/.. synagris) 


Lutjanidae 


0.03 


Yellowtail snapper (O.chrusurus) 


Lutjanidae 


2.89 


Triggerfishes 


Balistidae 


29.68 


Squirrelfishes 


Holocentridae 


4.84 


Mutton snapper (/.. analis) 


Lutjanidae 


0.13 


Other snappers 


Lutjanidae 


1.04 


Hogfish 


Labridae 


1.06 


Trunkfish 


Ostraciidae 


0.08 


CFMC (Caribbean Fisheries Management Council). 1985. Fishery Management Plan, Fi- 
nal 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/ 



< 

X 

C 

Q_ 

< 



page 
57 



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

Appendix B 



Table B1. Fish species list and summary data on 
Puerto Rico study region. 


occurrence, 


abundance and biomass 


(2001-2007) for the southwest La Parguera, 


Family 




% 


Total 


Total 


Mean abundance 


Total 


Mean biomass, g 


Species name 


Common name 


occurrence occurrence 


abundance 


(±SE) 


biomass, g 


(±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 


blackjack 


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) 
Table B1 cont... 



Family 




% 


Total 


Total 


Mean abundance 


Total 


Mean biomass, g 


Species name 


Common name 


occurrence 


occurrence 


abundance 


(± 


SE) 


biomass, g 


(±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 


MO JARRA 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 


oranges potted 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) 



Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007) 
Table B1 cont... 



Family 




% 


Total 


Total 


Mean abundance 


Total 


Mean biomass, g 


Species name 


Common name 


occurrence 


occurrence 


abundance 


(± 


SE) 


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 




















Bod i an us 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 


oranges potted 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) 
Table B1 cont... 



Family 




% 


Total 


Total 


Mean abundance 


Total 


Mean biomass, g 


Species name 


Common name 


occurrence 


occurrence 


abundance 


(± 


5E) 


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) 



Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): Spatial and temporal patterns in fish and benthic communities (2001-2007) 
Table B1 cont... 



Family 




% 


Total 


Total 


Mean abundance 


Total 


Mean biomass, g 


Species name 


Common name 


occurrence 


occurrence 


abundance 


(±SE) 


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 


grays by 


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) 


Serrani cuius 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) 



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



Appendix C 



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 Fish Base (http://www.fishbase.org). TL=total length; FL= fork length. 







Approx. size 


Juv./sub- 


Max. 


Max. size 






class first 


adult size 


known 


observed 


Species 


Common name 


maturity* 


class (cm) 


size*, TL 


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 


red band 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 


Batistes vetula 


queen triggerfish 


20-25 


<20 


47.1 


30 


Sphyraena barracuda 


great barracuda 


n/a 


n/a 


200 


130 



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

Appendix 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 

Q) to provide precise, fishery-independent and size-structured survey data, needed to comprehensively assess 

O faunal populations and communities (Menza et al., 2006). In addition, a complementary benthic composition 

O 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 m 2 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 




1 m 2 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 m 2 quadrat. 



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

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 m 2 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, sea- 
grass, 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 (Eustrom- 
bus 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 m 2 . 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 colo- 
nized 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 m 2 ) 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. 

Macroin vertebrates 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 m 2 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 af- 
fected habitat is also recorded along with a note identifying any flora or fauna that colonized the debris. Marine 
debris data collection began in 2007. 



CD 

Q_ 

Q_ 



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

Observer 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. Meth- 
ods 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) 

Appendix E 



18 i:: Q'0"N- 




■18°0'0"N 



LU 
X 

C 
CD 
Q_ 
Q. 

< 



Predicted fish species richness 

| |<12 

| 12-15 

|>15 



67 C 0'0"W 




°Q- ¥ 500 Meters 



Adapted from Pittman et al. (2007a) 

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. 



page 




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 




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