an · rpp than fished and unfished reefs in belize and saba. abundance of smaller scarids and...
TRANSCRIPT
COMMUNITY STRUCTURE, ABUNDANCE, AND BIOMASS OF FISHES ON A CARIBBEAN CORAL REEF,
SlAN KA'AN BlUSYHEKE KESEKVE, QUIN'L'ANA ROO, MEXICO: AN ANALYSIS BY DEPTH ZONE AND HABITAT
by
Scott B. Van Sant
With
Dr. Wes Tunnell, Jr., Dr. Quenton Dokken, Dr. Roy Lehman, and Dr. Kim Withers
Center for Coastal Studies Texas A&M University-Corpus Christi
6300 Ocean Drive, NRC 3200 Corpus Christi, Texas 78412
Sian Ka'an Series, No. 10 Center for Coastal Studies
Texas A&M University-Corpus Christi
July 2003
PREFACE
Texas A&M University-Corpus Christi has a long history of scientific research in the marine and coastal environments of Mkxico. Starting with research by Dr. Henry H. Hildebrand in the late 1950s on Alacrkn Reef and Laguna Madre de Tamaulipas to our more recent work during the 1970s, 1980s, and 1990s on the coral reefs and coast of Vcnicruz, we have been dedicated to studying tlic biodivcruity a ~ ~ c l r ~ ~ u i ~ i c : ecology of Mbxico and providing graduate research opportuilities in Mexico. Though distribution of thescs, disscrtations, tcchnical rcports, and scientific journal a
r
ticles, we have provided our research to Mexican scientists and natural resource managers.
Most recently, starting in 1996, we have established a long-term study site at Rancho Pedro Paila, near Boca Paila, in the northern part of the Sian Ka'an Biosphere Reserve in the state of Quintana Roo on the Caribbean side of the Yucatan Peninsula. In order to efficiently and effectively get our research to interested Mexican scientists, natural resource managers, and other interested persons, we have created the Sian Ka 'an Series. Since peer reviewed journal articles take one to three years to be published, this series will allow quick dissemination of the information. Additional copies may be obtained with instructions on the next page of the document.
John W. Tunnell, Jr., Ph.D. Director, Center for Coastal Studies and Professor of Biology at Texas A&M University-Corpus Christi
Sian Ka'an Series Center for Coastal Studies
Texas A&M U~liversily-Corpus Christi 6300 Ocean Drive, NRC 3200 Corpus C'hristi, 'l'exas 784 12
Phone: 361 -825-2736 Fax: 36 1-825-2770
Emnil: [email protected]
Title Price (USD)
No. 1 Keeney, Talitha S. 1999. Coral reef macroalgae in northern Sian Ka'an Biosphere Reserve, Quintana Roo, MCxico. M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus Christi, Texas. $7.00
No. 2 Milroy, Scott P. 1999. Effects of light availability on reef community structure of hermatypic corals within Sian Ka'an Biosphere Reserve, Quintana Roo, MCxico. M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus Christi, Texas. $7.00
No. 3 Hilbun, Nancy L. 2000. Distribution and abundance of echinoderms from Sian Ka'an Biosphere Reserve, Quintana Roo, MCxico. M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus Christi, Texas. $7.00
No. 4 Koltermann, Amy E. 2000. Ecological characterization of northwestern Caribbean ironshores, Sian Ka'an Biosphere Reserve, Quintana Roo, MCxico. M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus Christi, Texas. $7.00
t
No. 5 Tunnell, Kathryn D. 2001. Epibiont flora and fauna associated with two Rhizophora mangle forests, Veracruz and Quintana Roo, MCxico. . M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus ~hris t i , Texas. $7.00
No. 6 Campbell, Matthew D. 2001. A dry season analysis of larval and juvenile fish assemblages of the Sian Ka'an Biosphere Reserve, Quintana Roo, MCxico. M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus Christi, Texas. $7.00
No. 7 Childs, Catherine. 2002. Development of a natural resource conservation plan for Punta Allen peninsula, Sian Ka'an Biosphere Reserve, Quintana Roo, MCxico. M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus Christi, Texas. $7.00
No. 8 Bates, Thomas W. 2003. Locomotor behavior and habitat selection in intertidal gastropods from varying shore heights. M.S. Thesis. Biology P~agraoi, Texas A&M Urlivarsily-Qlrpus Chrisli, Corpus 67,00 Christi, Texas.
No. 9 Ledford, Chris. 2003. Comparison of coral species diversity and abundance between patch reefs and shallow reefs of the Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico. M.S. Thesis. Biology Program, Tcxas A&M TJnivcrsity-Corpils Christi, Cmpua Cluisti, Texas.
$7.00
No. 10 Van Sant, Scott. 2003. Community structure, abundance, and biomass of fishes on a Caribbean coral reef, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico: An analysis by depth zone and habitat. M.S. Thesis. Biology Program, Texas A&M University-Corpus Christi, Corpus Christi, Texas. $7.00
No. 11 Reed, Addie L. 2003. Implementation of a long-term coral reef monitoring plan, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico. M.S. Thesis. Biology Program, Texas A&M University- Corpus Christi, Corpus Christi, Texas. $7.00
Abstract
An urldcrwatcr visual ccnsus was conducted during May 1999 of' iishes alulig a fringing-banier reef site at Rancho Pedro Paila (RPP), Quintana Roo, Mexico, located within the Siai Ki'ari Biosphere Reserve. Analyses of abundance, mean length, dominance, and biomass of top ranking species, major families, and trophic guilds were produced for four depth zones and three habitat types. Community structure was analyzed by quantitatively comparing values of species richness, species diversity, and dominance for each depth zone and habitat type.
A total of 14,509 fishes were observed, representing 128 species and 35 Igmilies. The greatest abundance was found in the deep depth zone with the most complex habitat structure. Within each depth zone, values increased with increasing habitat complexity. Comparisons between each overall value for habitat were significantly different as well as when comparing the deep reef zone with the other depth zones. The greatest values for species richness and diversity were found at the shallow and deep reef within the most complex habitat type. Overall values for species richness were greatest for the deep reef zone, followed by the shallow reef, mid reef, and patch reef zone.
The reefs in the area of RPP, such as Boca Paila and Tampalam, are considered to be semi-protected because certain forms of fishing are restricted. However, results of visual census data taken during May 1999 suggests that abundance and biomass values for fishes targeted for fishing are low. Few or no large schools of snappers (Lutjanidae), grunts (Haemulidae), or goatfishes (Mullidae), were observed. During 1998, large groupers (Epinephelinae) were absent and no individuals of red grouper, Epinephelus morio, Nassau grouper, E. striatus, red hind, E. guttatus, or tiger grouper, Mycteroperca tigris were observed. Abundance of groupers was lower at RPP than fished and unfished reefs in Belize and Saba. Abundance of smaller scarids and acanthurids at RPP were higher than other fished and unfished reefs. The herbivorous acanthurid, Acanthurus coeruleus, ranked number one in abundance comprising 16.05% of the total number. A spawning aggregation and spawning event was documented for this species at the mid reef zone. Overall abundance at RPP would fall within the range of other Caribbean reefs from similar studies that were reportedly heavily and moderately fished. For the different depth zones, abundance values at RPP ranged from 55.64-1 10.78 (s80.60). Biomass values are comparable with other Caribbean reefs that were reportedly lightly fished. Biomass values for RPP ranged from 28.74-59.66 gm-2 (2=4.1.74).
Recommendations for protective measures include further protection and inclusion of coral reef areas as core zones; adherence to American Fisheries Society's recommendations for managing long-lived, slow growing reef fishes; and protection of spawning aggregations. This study and sampling design could be a model to quantitatively monitor long-term trends in community structure, diversity, and stocks as well as investigate factors that are altering these populations.
Un censo visual submarino de peces a lo largo del arrecife frangeante en el Rancho Pedro Paila (RPP), Quintana Roo, Mexico, localizado dentro de la Reserva de la Biosfera de Sian Ka'an sc llcv6 a cabo durantc cl mcs dc Mayo dc 1999. Analisis de abundancia, prv~lledio Je longitud, dominancia, y biomasa de las principales clasificaciones de espeices, familias importantes, y gremio trofico se realizaron en cuatro zonas de profundidad y tres tipos de habitat. La estructura de la comunidad se analiz6 comparando quantitativamente 10s valores de la riqueza y diversidad de especies, como tambikn dominancia en cada una de las zonas de profundidad y tip0 de habitat.
Un total de 14.509 peces se observaron, 10s cuales representan i28 especies y 35 Pdmilias. La mayor abundancia se encontr6 en la mas profunda de las zonas de profundidad, la cual present6 la estructura de habitat mas compleja. Dentro de cada zona de profundidad, 10s valores aumentaron a medida que la complejidad del habitat aumentaba. Las comparasiones entre el valor total de cada habitat fueron considerablemente diferentes, a1 igual que las comparasiones hechas de la zona mas profunda de el arrecife con el resto de zonas de profundidad. Los numeros mas altos de riqueza de especies fueron encontrados en la zona mas profunda del arrecife, seguidos por la zona del arrecife somero, arrecife medio, y zona del arrecife en parche.
Los arrecifes del area de RPP, como Boca Paila y Tampalam, se considerados semi- protegidos, porque ciertas fromas de pesca estan restringidas. Sin embargo, 10s datos del resultado de el censo visual tornados en Mayo de 1999 sugieren que 10s valores de abundancia y biomasa de 10s peces estudiados son bajos. Se observaron pocos o pequeiios bancos de peces snappers (Lutijanidae), roncos (Haemulidae), o salmonete (Mullidae). Durante 1998, grouper (Epinephelinae) grandes estuvieron ausentes, como tambikn lo estuvieron peces individuales de red groupers, Epinephelus morio, grouper de Nassau, E. striatus, red hind, E. guttatus, o tiger grouper, Mycteroperca tigris. La abundancia de groupers fue mas baja en RPP que en arrecifes de pezca y de no pezca en Belize y Saba. La abundancia de pequeiios scarids y acanthurids en RPP fue mas alta que en otros arrecifes de pezca y de no pezca. El hervivoro acanthurid, Acanthurus coerleus, clasific6 como numero uno en abundacia, comprendiendo un 16.05% de el ndmero total. Se document6 una agregacion reproductiva y un evento de desove de esta especie en la secci6n media del arrecife. En su totalidad, la abbdancia en RPP resultaria entre la gama de abundancia de otros arrecifes Caribeiios de acuerdo a otros estudios realizados que reportaron alta y moderada actividad pezquera. Para cada zona de profundidad, 10s valores de abundancia en RPP variaron desde 55.64-1 10.78 (%=80.60). Los valores de biomasa son comparables a otros arrecifes Caribeiios que reportaron activadad pezquera liviana. Los valores de biomasa de RPP variaron entre 28.74- 59.66 gm-2 (%=41.74).
Recomendaciones para tomar medidas protectoras incluyen protection adicional e inclusi6n de areas de arrecifes coralinos como zonas principales; adherencia a las recomendaciones de la American Fisheries Society para el manejo de peces de arrecifes de crecimiento lento y larga vida; y protecci6n de agregacion reproductiva. Este estudio y diseiio de muestreo puede ser utilizado como modelo para monitorear quantitativamente las tendencias a largo plazo de las estructuras de las comunidades y existencias, asi como tambikn para investigar 10s factores 10s que alteran estas poblaciones.
Table of Contents
Title Pagc. ......................................................................................... i
Abstract.. ............................................................................................ v
List of Tables. ......................................................................................... .ix
. . L i ~ t of Figures.. ..................................................................................... X I I
... Acknowledgments. .................................................................................. .xi11
Introduction. ........................................................................................ 1
....................................................................................... Study Area. .8
Methods and Materials. ........................................................................ 10
Results. ..................................................................................................... 16
Discussion.. ......................................................................................... 5 1
Literature Cited. ..................................... : ............................................... .62
Appendix. ............................................................................................... .68
Appendix I. Phylogenetic listing of all fish species (128) censused at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, hexico, May 1999, including frequency of occurrence, mean number (pN), and rank. ........................................................................... -68
Appendix 11. Species groubed by family (Randall, 1967), including length-weight conversion formulae used for estimating biomass (Bohnsack and Harper, 1982), used in this study at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.. ............................................................................................ 72
Appendix 111. Species grouped by trophic guild (Randall, 1967), including length-weight conversion formulae used for estimating biomass (Bohnsack and Harper, 1982), used in this study at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.. ............................................................................................. 74
Appendix IV. Frequency of occurrence, mean number (pN), rank iibundilr~ct! (N), and ovcrall rank for all species (128) cellsused at Rancho Pedro Paila, Sim Ka'm Biosphere Reservc, Quintana Roo, Mexico, May 1999. T,isted in order by rank abundance in descending order. . . . . . .. . . . , . . . , , , . . . . ... .... . ... .. ... .. .. .. ... .. . ... . . .. . ... . ... .. . . . ... . ... . . . ... . . . . ... . ...... .. .. . . ... . . .76
List of Tables
Table 1. Number of point count samples taken for cach depth zone and habitat type at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Qui~ilana Roo, Mexico, May 1999.. .......................................................... 14
Table 2. Total number of individuals (N), mean number of individuals (pN) (bold), standard deviation (a) of pN (italics), and range per observation (N Range) for depth zones and habitats sampled at Rancho Pedro Paila, Sian Ka' an Biosphere Reserve, Quintana. Roo, Mexico, May 1999 ................................................................................................................ 17
Table 3. Results of MANOVA and ANOVA showing level of . . .,- significance @-value) for all species, comparing number of individuals (N), log transformed mean number (pN), mean biomass (pB), mean for depth zones and habitats Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Significant values (W0.05) are in bold. Depth zones- D= deep reef, M= mid reef, S= shallow reef, P= patch reef. Habitats (1, 2, 3). ......................................................... 18
Table 4. Mean biomass (pB) (bold), standard deviation (a) (italics), and range per observation for depth zones and habitats sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.. ........................................................................................ 19
Table 5. Mean number of species (pS), and range for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve Quintana Roo, Mexico, May 1999.. ................................................................... 21
Table 6. Results of MANOVA and ANOVA showing level of significance for all species, comparing number of species @-value) (pS), and mean diversity (pH') for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. M= mid Significant values @<0.05) are in bold. Depth zones- D= deep reef,
.............................. reef, S= shallow reef, P= patch reef. Habitats (1, 2, 3).
Table 7. Mean species diversity (pH') for depth zones and habitats sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.. ................................................
Table 8. Overall number of individuals (N), percent of the total (%Tot), mean number (pN), mean length (pXL), mean biomass (pB), and standard deviations (sdpN, sdpXL, sdpB) for nine families at Rancho
Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. *Pomacanthidae/ Chaetodontidae (angelfishes/butterflyfishes) ............................................................... .25
Tablc 9. Mean number (pN) and mean biomass (pB) for iliile fanlilies for all habitats at Rancho Pedro Paila., Sian Ka'aii Riosphcre Resel-vc, Quintmn Roo, Mcxico, May 1999. Dcpth zones- P=patcl~ reef, S= shallow reef, M= mid reef, D= deep reef.. ............................................... 26
Table 10. Results of MANOVA and Analysis of ANOVA showing level of significance @-value) for nine families, comparing log transformed values for mean number (pN) and mean biomass (pB) for depth zones and habitats at Rancl~o Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Significant values (p<0.05) are in bold. Depth zones- D= deep reef, M= mid reef, S= shallow reef, P= patch reef. Habitats (1, 2, 3). ...................................................
Table 1 1. Mean number (pN) and mean biomass (pB) for nine families for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an
................................... Biosphere Reserve, Quintana Roo, Mexico, May 1999.. .30
Table 12. Overall number of individuals (N), percent of the total (%Tot), mean number (pN), mean length (pXL), mean biomass (pB), and standard deviations (sdpN, sdpXL, sdpB) for seven trophic guilds at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. SAF= sessile animal feeders.. ..................................... 38
Table 13. Mean number (pN) and mean biomass (pB) for all habitats for seven trophic guilds at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. P= patch reef, S= shallow reef, M= mid reef, D= deep reef, SAF= sessile animal feeders. .............................................................................. .3 9
Table 14. Results of MANOVA and ANOVA showing level of significance @-value) for seven trophic guilds, comparing log transformed values of mean number (pN) and mean biomass (pB) for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo,Mexico, May 1999. Significant values @<0.05) are in bold. Depth zones- D= deep reef, M= mid reef, S= shallow reef, P= patch reef. Habitats (1,2, 3). ...................
Table 15. Mean number (pN) and mean biomass (pB) for seven trophic guilds for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. SAF= sessile
....................................................................... animal feeders..
Table 16. Top 10 ranked species with values for rank abundance (N) for each depth zone observed at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. ........................................... .49
Table 17. Mean biomass, abundance, and number of species for all species obseived at Rancho Peclro Paila (RPP), Siarl Ka'an Biosphere Receive, Quinta~na Roo, Mexico, May 1999, with a comparison to similar studies with vnryiilg dcgrccs of fishing prcssurc. P= protected, SP= somewhat protected, T.P= ~lnprotected, F=fished, LF=lightly fished, MF=rnoderately fished, HF=heavily fished.. ........................................... .53
Table 18. Mean biomass and abundance for selected families observed at Rancho Pcdro Paila (:RPP), Sian Ka'rtn Biosphere Reserve, Quiiltnna Roo, Mexico, May 1999, with a comparison to similar studies with varying degrees of fishing pressure. P= protected, SP= somewhat protected, UP= unprotected, F=fished, LF=lightly fished, MF= moderately fished, HF=heavily fished.. ................................................. .56
Table 19. Mean biomass and abundance for trophic guilds observed at Rancho Pedro Paila (RPP), Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999, with a comparison to similar studies with varying degrees of fishing pressure. P= protected, SP= somewhat protected, UP= unprotected, F=fished, LF=lightly fished, MF= moderately fished, HF=heavily fished.. ................................................. .58
List of Figures
Figure 1. Map of the eastern portion of the Yucatan Peninsula and southern Quintana Roo, Mexico, showing the geographical position of the Sian Ka'an Biosphere Reserve and the locatioii of the study site at Rancho Pedro Paila (NOAA, NMFS, Office of Protected Resources, 2002, unpub. m.s.). lnscrt modified from Sanvicente-Aiiorve et al.
Figure 2. Imaginary cylinder used by stationary divers employing the Stationary Visual Census Technique (Bohnsack and Bannerot, 1986). Technique used in this study at Rancho Pedro Paila, Sian Ka'an
.................................... Biosphere Reserve, Quintana Roo, MX, May 1999.. 1 I
Figure 3. Mean number of individuals per observation (pN) for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth
................... zones- P= patch reef, S= shallow reef, M= mid reef, D= deep reef.. 18
Figure 4. Mean biomass (pB) for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth zones- P= patch Reef, S= shallow Reef, M= mid Reef, D= deep Reef.. ........................................ 20
Figure 5. Mean number of species (pS) for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth zones- P= patch reef, S= shallow reef, M= mid reef, D= deep reef.. ............................... 2 1
t
Figure 6. Mean species diversity (pH') for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth zones- P= patch reef, S= shallow reef, M= mid reef, D= deep reef.. .................................. ..23
Figure 7. Overall number of individuals (N), mean number (pN), mean length (pXL), and mean biomass (pB) for nine families at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.. . *Pomacanthidae/Chaetodontidae (angelfishes/butterflyfishes) ................................................................. ..26
Figure 8. Overall mean number (pN), mean length (pXL) and mean biomass (pB) for seven trophic guilds at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. SAF= sessile animal feeders. ..................................................................................... .3 8
xii
Acknowledgements
I would like to extend much thanks and appreciation to my major advisor, Dr.
Wes Tunnell for his continued moral siippnrt, field and travel support, reviews,
suggestions, good cheer, and endless kindness and patience throughout this process. I
greatly appreciate the support of Dr. Tunnell, Ms. Ronnie Emanuel, academic
advisor, and Dean Diana Marinez for going above and beyond for their support of my
completing academic requirements while living in and traveling to every comer of the
continent. I greatly appreciate the efforts of my dive partner, Mr. Devin Hayes, in the
field during all phases of the diving, boating, and sampling. I would like to thank my
thesis committee members, Dr. Jason Link for his helpful reviews, suggestions, and
his guidance with statistical analyses and SAS programs, Dr. Quenton Dokken, for
his suggestions and reviews, and Dr. Roy Lehman and Dr. Kim Withers for their
reviews. I am especially grateful for the tremendous assistance of Dr. Lance
Garrison, Mr. Truman Du, and Mr. Joe Liddle with statistical analyses and SAS
proghms. I would like to thank Dr. Jim Bohnsack and Dr. Callum Roberts for
helpful suggestions. I would like to thank Mr. Jeff Miller and Ms. Sue Hazlett for
assistance with data analysis and Ms. Kelly Neale for assistance with data entry.
Special thanks go out to Ms. Ten Frady and Mr. David Radosh for being supportive
of my thesis work and financial support for travel while working at the Northeast
Fisheries Science Center at Woods Hole, MA. I would like to thank Ms. Gloria
Krause for administrative assistance with long distance communications.
introduction
Until recently, the Caribbean coast of Mexico was among the most isolated in the
country. The coastal areas and coral reefs of Quintan Roo (Figure 1) have been impacted
by natural events, such as the high pressure cold fronts called nortes (Wells, 1988),
hurricanes (Rogers, 1993), coral bleaching and disease, and the Caribbean-wide mass
mortality of the sea urchin Diadema antillarium, an important keystone species (Lessios
et al., 1984), but human impact was minimal. This changed dramatically in the latter part
of the 2oth Century when the Mexican government developed Cancun as a destination
tourist resort. An international airport, extensive road system, and scores of beachfront
resort hotels were built to accommodate the growing number of tourists and cruise ships.
As a result, human impacts on the reefs along the eastern coast of Mexico are substantial.
The reefs off Veracruz are under stress from intense fishing, sedimentation, oil spills, and
many other perturbations (Tunnell, 1993). Oil has become the major industry in the
southern Gulf of Mexico since the discovery of the highly productive Campeche Bank oil
fieldr(Ferrk-Arnark, 1985). Recently the deforestation, development, and rapid
population growth has resulted in 60% of the land being cleared. Such deforestation can
lead to increased soil run-off, ultimately impacting coastal reefs (Ferrk-Amark, 1985). In
addition, the demand for beachfront resorts with sandy beaches has resulted in the
clearing of large areas of coastal mangroves, which are nursery areas for many reef fish.
The increase in vessel traffic also has potential for impacts on the coral reefs, ranging
from direct damage done by boat anchors and scuba divers, to indirect effects from the
dumping of sewage and garbage by cruise ships.
Fisheries here are also expanding rapidly, with the catch rate for the eastern coast
Figure 1. Map of the eastern portion of the Yucatan Peninsula and southern Quintana Roo, Mexico, showing the geographical position of the Sian Ka'an Biosphere Reserve and the location of the study site at Rancho Pedro Paila. (NOAA, NMFS, Office of Protected Resources, 2002, unpub. m.s.). Insert modified from Sanvicente-Aiiorve et al. (2002).
of Mexico at 0.25 million tons annually which accounts for about 20% of the national
catch (Ferrk-ArnarG 1985). Queen conch, Strornbus gigas, stocks have become depleted
md there is conccm that spiny lohstar, Pcrnlrlirus nrgrrs, is beirlg ovcrlished. Large reef
fishes such as groupers (Serranidae), known to form spawning aggregations at specific
sites and times of thc yeiir, are also vulr~crllhlc t.o intense fishing pressures. Of thc
aggregations of Nassau grouper, Epinephelus striatus, known and exploited for many
years by local fishermen, many have disappeared, possibly by overfishing (Colin, 1992;
Sadovy et al., 1994). In Belize a spawning aggregation of E. striatus has disappeared
from its traditional site in recent years (Carter et al., 1994). Similar trends have been
noted for grouper species in Jamaica, Florida and the Virgin Islands (Sadovy, 1999).
Recently, off southern Quintana Roo, disappearances and reduction in abundances have
been noted for aggregations of E. striatus (Aguilar-Perera and Aguilar-Davila, 1996).
Several grouper stocks of the southeastern U.S. and Caribbean are severely depleted and
have been recommended for protection under the Endangered Species Act (Huntsman,
1994). Intense fishing of spawning aggregations could have negative impacts on the
population size, sex ratio, genetic diversity, and behavior of the aggregating species
(Sedberry et al., 1993, unpub. m. s.).
Munro and Williams (1985) state the changes in species composition or relative
abundance of species in multispecies stocks depend largely on the level of intensity of
fishing, the relative catchabilities of the species, and the magnitude of the interactions
between species. Roberts (1995) reviewed recent data on the effect fishing has on coral
reefs and cited four main points. Reef fishing: (1) can lead to major direct and indirect
shifts in community structure; (2) reduces species diversity on reefs; (3) can result in the
loss of keystone species, which in turn can lead to major effects on reef processes; and
(4) rnay lead to loss of entire functional groups of species, resulting in impairnleilt of
potentially important ecosystem processes facilitated by those groups.
Several countries have taken action to protect their fisheries. In Belize, two
marine parks have been established (Carter, 1988), in the Cayman Islands only fishing by
hook and line is permitted, and in the U.S. Virgin Islands a permanent marine reserve was
established in 1998 to protect red hind (E. guttatus) aggregations (Beets and Friedlander,
1999). Such marine protected areas have been in existence for several decades and are
widely used to protect and manage marine habitats (Bjorklund, 1974; Polunin and
Roberts, 1993). Their increasing use stems fkom the growing need to both protect the
marine environment from human impact and to manage fisheries (e.g., Davis, 1989;
SEFSC, 1990; Roberts and Polunin, 1991; Roberts, 1995; Bohnsack, 1999). Bohnsack
(1992) states marine reserves are areas that are intended to prevent recruitment
overfishing and ensure the persistence of reef fish stocks and habitats from all
conshptive exploitation. It is assumed that if fishes that are normally targeted are
protected in reserves, then their numbers and sizes will increase. These effects are
expected to lead to increased egg production in reserves, and through planktonic
dispersal, replenishment of stocks in unprotected areas. Recently there have been
different assessments of the effects of marine protection on commercially important
fishes and communities (Polunin and Roberts, 1993; Jennings et al., 1995; Jennings and
Polunin, 1997; Roberts and Hawkins, 1997; Rogers and Beets, 2001). At times a
dramatic two-fold increase in biomass has been detected in even small protected areas
when compared to areas outside of them (Polunin and Roberts, 1994).
Reef fisheries, such as those of Quintana Roo, are veiy difficult to nianage due to
the large numbers of species caught, the kinds of gear used, and the number of places at
which calches are landed (Polunin and Roberts, 1993). Some of the coral reefs of this
area have, however, been protected as part of the Sian Ka'an Biosphere Reserve (Figure
I). Sian Ka'an was declared a national biosphere reserve by the Federal Government of
Mexico on January 20, 1986 (GutiCrrez-Carbonell and Bezaury-Creel, 1993). As a result,
an official management program for the Sian Ka'an was prepared. The program is
comprehensive. It identifies three management zones: 1) A core zone, which is a
natural, completely protected area; 2) a buffer zone, which surrounds the core zone and
allows for only minimal or low impact use; and 3) a transition zone, whch includes
human settlements and economic activities compatible with the conservation and
preservation of the reserve ecosystems. The management plan also contains 16
objectives, including protection, resource management, monitoring, environmental
restoration, archaeological and cultural protection and management, social development,
publl'c use (tourism), and infrastructure (SEMARNAP, 1993). Biological inventories
have been conducted, concentrating on identifying species composition in the various
habitats or identifying major habitat types (Navarro and Robinson, 1990; Olmsted and
Duran, 1990; Tangley, 1988), as well as recent quantitative ecological studies of coral
reef fishes (N6iiez Lara and Arias Gonzalez, 1998, Caballero and Schmitter-Soto, 2001,
Garduiio and Chavez, 2000).
Biosphere reserves are designed to carehlly incorporate limited and sustainable
-human activities into the planning and management of the area, but encompass areas
large enough to ensure maintenance of genetic, species, habitat, and ecosystem diversity
over time (Meffe and Carroll, 1994). Meffe and Carroll (1 994) summarize some of the
current problems at the Sian Ka'an Biosphere Reserve. Thcrc is little information on
spec:ies inler;iclinns or food webs, lalor is there sufficient data Sbr proposing infor~ned
quotas for fishing or other development activities, or for establishing a carrying capacity
for permanent inhabitants or t,ol~rists. Managers have yet to define the sizes and lacations
of the most critical habitats within the reserve, identify which species are most in danger,
or develop appropriate management programs for core areas. Sensitive areas such as
coral reefs are not included in the core areas, with the exception of a small portion near
the Cayo Culebra (Cayo Culebras) zone near the mouth of Ascension Bay (Figure 1).
Only the three core zones are well defined; buffer and transition zones do not yet
formally exist. The marine portions of the Sian Ka'an Biosphere Reserve were reviewed
and it was proposed that the boundaries be modified to include all of the reef system.
The three marine core zones were suggested to protect 12.3% of the marine environment
instead of the current 2.5% (Salazar-Vallejo et al., 1993). Chinchorro Bank, located 30
km o'ff of Quintana Roo's southern coast, was also suggested for further protection and
management (Aguilar-Perera and Aguilar-Dhvila, 1996). Protected coral reefs along the
shore and Chinchorro Banks (Banco Chinchorro) have little protection fiom fishing,
however Uaymil, Majahual, and Xcalak reefs (Arecifes de Uaymil, Majahual, y Xcalak)
further south along the reef tract are proposed for protection. Inaccessibility provides a
certain degree of protection, but at the same time hinders monitoring, research,
administration, and active protection. Enforcement remains inadequate for this expansive
area.
A descriptive biological inventory exists for many species in a variety of habitats
for the Sian Ka'an Biosphere Reserve (Navarro and Robinson, 1990). Biological
diversity studies include fish larvae of Ascension Bay (Vasquez Yeomans, 1990), yet
until recently few quantitative studies of rccf fish communities existed for this area.
Some of the studies of fishes for Quintana Roo include a study of fishes of nearshore and
inshore aquatic habitats of the Yucatan and Belize (~lvarez-~ui l len et al., 1986;
Schrnitter Soto and Gamboa Perez, 1996), a seasonal study of fish communities (Diaz-
Ruiz and Aguirre-Leon, 1993) and snappers (Diaz-Ruiz et al., 1996) inhabiting seagrass
and reef habitats off Cozumel, a study of marine fishes in rocky intertidal habitats
(Pamplona Salazar and Anguilar Rosas, 1992), and a study that investigated the effects
Hurricane Gilbert had on the coral reefs, fishes, and sponges at Cozumel (Fenner, 1991).
Other studies include a fish community study in seagrass beds (Garduiio and Chavez,
2000) and a reef fish diversity study on coral patches (Caballero and Schrnitter-Soto,
2001) along the coast of Quintana Roo. Recently studies have attempted to assess
anthropogenic effects; such as the effect fishing has on reef fishes or the effectiveness of
the riserve in restoring reef fish populations. Studies exist that have gathered data on
reef fish catch statistics and the socio-economic aspect of the demersal fishery (Basurto
Onegel, 1988), and recently there have been studies investigating effects fishing has on
reef fishes, such as spawning aggregations of Nassau grouper (Aguilar-Perera and
Aguilar-Davila, 1996), studies that compare trophic models for protected and unprotected
reefs (Arias-Gonzalez, 1998), and studies that investigate the relationship between
physical parameters and reef fish community structure (Nuiiez Lara and Arias Gonzalez,
The aim of the current study is to provide a quantitative assessment of the reef
fish community and economically important species using underwater visual censusing
techniques. From these data and comparisons with similar studies that have compared
fishes from reefs of varying fishing intensity (Bohnsack, 1982; Polunin and Roberts,
1993; Roberts, 1995; Sedberry et al., 1996), it may be possible to determine the effects
fishing has had on the community. This study provides a framework for long-term data
collection and monitoring of the reef fish community as well as any factors affecting the
community.
The objectives of this study at Rancho Pedro Paila (RPP), Sian Ka'an Biosphere
Reserve, Quintana Roo, were: (1) to produce a quantitative description of selected
members of the fish community including estimates of abundance, mean length, species
richness, species diversity, dominance, and biomass; (2) to produce a quantitative
description of the fish component grouped by seven trophic guilds and nine families; (3)
to compare the values in (1) for species and species groups by depth zone and by habitat
complexity; (4) to complete comparisons of data generated with comparable data from
~ t h e < ~ e o g r a ~ h i c regions; and (5) to make recommendations for management strategies to
maximize health, productivity, and sustainability of the fish community.
Study Area
The study site, Rancho Pedro Paila (RPP), is located on the eastern coast of the
Yucatan Peninsula, within the state of Quintana Roo, Mexico (Figure 1). It is located
- -t within the northern portion of the Sian Ka'an Biosphere Reserve lying between Tulurn to
the north and Boca Paila to the south (20" 02.552' N, 87' 28.052' W) (Tunnel1 et al.,
1993). The site is situated on a long, narrow sandbar, which extends 56 km south to
Punta Allen, at the north end of Ascension Bay. The reef crest is located approximately 1
km from the sandy shore of RPP with small spur and groove formations projecting
seaward.
The reefs in the area of RPP, such as Boca Paila and Tampalam, are considered to
be semi-protected because ccltaili forills or fisliilig are 1,estriclecl (Arias-Gonzalex, 1998).
This northeast section of the reef of the Sian Ka'an Biosphere Reserve is 100 km long
with varying development. A marine area of 37,000 ha, constituting mostly coral reefs
were added to the reserve in February 1998 (Arias-Gonzalez, 1998). Since its initiation,
some restrictions have been implemented such as the limited use of harpoons and nets.
However five fishing cooperatives and five independent license holders continue to
operate. They exploit mainly lobster and some have concessions to exploit coral crab,
shark, and use coral nets.
The Sian Ka'an Biosphere Reserve comprises an area of 528,000 ha, 408,000 ha
of which are terrestrial and 120,000 ha include lagoon or marine environments. The core
t
zone of the reserve comprises two terrestrial areas and one marine area. Where possible,
boundaries were defined to coincide with natural features. The reserve is bounded by the
Caribbean Sea and the barrier reef to a depth of 50 m in the east, the junction between the
marshes and semi-evergreen forests in the west, and the junction of Chetumal and .
Espiritu Santo Bays catchment basin in the south (19" 04' and 20'05' N, 87" 26' and 88"
03' E) (Tunnel1 et al., 1993).
The reserve lies on a partially emerged coastal limestone plane, which forms part
of the extensive barrier reef system along the eastern coast of Central America. From the
northern border of the reserve the coastal areas are mostly narrow sand bars enclosing a
system of brackish water lagoons and lakes within extensive mangrove swamps. The
barricr rccf and sand bars arc continuous with inten~~ittent rocky sl~ores ur puints, witil
they are interrupted by two large shallow bays. The geomorpl~ology from the Bay ol'
Espiritu Santo to Chetumal consists of long narrow beaches that are continuous with
narrow lagoons behind them that lack inlets to the sea. In the southern portion of the
reserve the reefs are somewhat hrther offshore and become increasiilgly better
developed. Southward of Punta Allen the sandy beaches are interrupted by two shallow
bays, Ascension Bay and the Bay of Espiritu Santo. The shoreline between the two bays
is formed of two sections of rocky shoreline. A 90 km fringing reef runs fairly
continuous along the coastline, interrupted at the mouth of Ascension Bay and becoming
more developed further south near the Belize border (Tunnel1 et al., 1993).
Materials and Methods
All samples used in this study were collected daily from May 9 through May 21,
1999: Preliminary sampling was conducted during May of 1998. Sampling consisted of
randomly chosen underwater visual census point counts using SCLrBA and the Stationary
Visual Census Technique of Bohnsack and Bannerot (1986). T h s method has proven
suitable for describing reef fish community structure, for making comparisons of fish
abundance and community structure, and for making comparisons of fish abundance and
community structure among sites and habitats (Bohnsack and Talbot, 1980; Bohnsack,
1982; Bohnsack and Bannerot, 1986; Clavijo et al., 1989). This method consists of a
diver listing the fish species seen in an imaginary cylinder of water (diameter=15 m), . . . . -
from the bottom to the surface, in a five-minute period at a randomly selected point
within a habitat at a given site (Figure 2). At each sampling point all species are recorded
within the cylinder for five minutes, while the diver rotates in one direction. After the
five minutes are up then the diver records the statistical information, not listing any new
species observed. Counts as well as lengths (to the nearest cm) were recorded, including
minimum, maximum, and mean leilgt11. Divers esti~llated fish 1e11gQs ur~clerwaler using
r=7.5 m ~ = n - r ~ h
h=depth (m) (*14.5 m)
Figure 2. Imaginary cylinder used by stationary divers employing the Stationary Visual Census Technique (Bohnsack and Bannerot, 1986). The technique used in this study at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, MX, May 1999.
their clipboard with a ruler attached to avoid errors due to magnification. Data was
recorded on underwater da1.a. sheets. Sampling was canductad during the day and
emphasizes only diurnally active, non-cryptic, reef associated species. As needed,
ichthyocide or anaesthetic, approved by permit (No. 3014, issued by SEMARNAT), was
used for species verification of young stages or difficult spccics. Names for species were
merged when two similar species were indistinguishable underwater. Species merged
include Kyphosus sectatrix/incisor (Bermuda/yellow chub) and Coryphopterus
personatus/hyalinus (masked/glass goby) (Appendix I).
Biomass estimates include the species belonging to the following families:
angelfisheshutterflyfishes (PornacanthidaelChaetodontidae), damselfishes
(Pomacentridae), groupers (Serranidae, subfamily Epinephelinae), goatfishes (Mullidae),
grunt (Haemulidae), parrotfishes (Scaridae), snappers (Lutjanidae), surgeonfishes
(Acanthuridae), wrasses (Labridae) and (Appendix 11). Angelfisheshutterflyfishes were
considered since they are known to be indicator species for coral reef health (Wantiez et #
al., 1995). The two families were combined since they are usually less abundant than
other families and all species :are members of the same trophic guild. Trophic level of
species was determined from published feeding studies (Randall, 1967) and reviews
(Kaufman and Ebersole, 1984) (Appendix 111). Trophic guilds selected include
benthivores (mobile and sessile invertebrates- crustaceans, molluscs, worms, and
echinoderms), herbivores, omnivores, pelagic piscivores (transient pelagic species),
piscivores (resident reef fishes), planktivores, and sessile animal feeders (attached soft-
bodied cnidarians, porifera, etc.). The weights of fishes were computed using regression
formulae to convert lengths to weights (Bohnsack and Harper, 1988) (Appendix 11,111).
When a species specific formula was absent, a morphologically similar species was
selected. Biomass values are defined as mass (g) x number per unit area (mZ).
Each species was ordered according to its overall rank abundance (Appendix N).
Rank abundance was calculated as frequency of occurrence x mean number of
iildividuals per obsenratio~l (pN). Tlle species wit11 the hipliest rarlk abundar~ct: wal;
ranked as 1. Species with equal rank abundance values were given the same rank. Rank
abundance is the same as the total number of individuals (N).
Unit area is determined from the area of the censused cylinder x the number of
sites censused (or collections). The area censused is the area (m2) for a cylinder, nr2,
where r=7.5 m (Figure 2). Abundance, biomass, and mean lengths values for species
groups were compared for each depth zone and habitat type.
Fish counts were made along four depth zones of the study site (Table 1). Depths
and habitats within each zone were determined from preliminary sampling during May
1998. The patch reef zone consists of various habitats from 1 m to 7 m, including reef
f
crest (Acropora palmata) and back reef habitats dominated by massive corals (Diploria
spp. and Montastrea spp.). The shallow reef zone consists of habitats fiom 3 to 10 m,
including spur and groove and low relief areas. The mid reef zone consists of habitats
from 10 to 20 m, including spur and groove, forereef slope, and low relief areas. The
deep reef site consists of similar habitats as the mid reef from 20 to 33 m.
Accuracy of length estimations were evaluated by comparing simultaneous point
counts taken by the two divers who collected all of the census data for both years. These
tests were done during May 1998, preliminary data collection, as well as the beginning of
the May 1999 data collection. Specific mechanics of the technique used were
Table 1. Number of point count samples taken for each depth zone and habitat type at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
Habitat types
Depth zones Depth range (m) 1 2 3 All
Patch Reef 1 - 7 15 15 15 45
Shallow Reef 3- 10 15 15 15 45
Mid-Reef 10-20 15 15 15 4 5
Deep Reef 20 - 33 15 15 15 45
All 1- 33 60 60 60 180
standardized and results of counts, species identification, and length estimation were
shown to be very similar. However, no correction factors based on measurements of
known standards were done. Diver bias error was reduced further since 70% of the data
was collected by the author.
The structural complexity of the substrate is known to have an important
influence on fish community structure (Luckhurst and Luckhurst, 1978; Kauhan and
Ebersole, 1984; Roberts and Ormond, 1987; Friedlander and Parrish, 1998). To control
variation in habitat characteristics for each area censused, visual estimates were made and
sketched of the dominant components of the benthos within each area sampled, including
percent cover of hard corals, gorgonians, sponges, sand, and substrata. The structural
complexity of the substrate were estimated on a 3 point scale and assigned as three
habitat types as follows:
(1) low relief (0-1 m) and sparse cover (< 25%);
(2) moderately complex with some overhangs or fissures, moderate relief (1 -3 m)
with widespread cover (25-50%); and
(3) very complex with caves, fissures, andor overhangs, high relief (> 3 m) with I
dense coral cover (> 50%).
Equal nuinbers of point counts were complctcd for eac11 of lltc Illree hi~bilat types
along the four depth zones. Each habitat contained 15 point counts per depth zone, a total
of 45 per depth zone, a total of 60 per habitat type, and a total of 180 overall (Table 1).
Additional measures of community structure, species divcrsity (H') and number
of species per census point (S) were calculated for collections by habitat and site. The
Shannon Weaver index (Shannon and Weaver, 1949) index combines both 'species
richness' or the number of species (S) and relative abundance, and was calculated with
the following formula:
Wherepi= ni/N, or the proportion of the total sample belonging to fish species "i". N=
. the total number of individuals of all fish species in the sample, and ni = the number of
individuals of each fish species "i" in the sample. Where s= the total number of species.
Diversity (H') was calculated according to the method of Pielou (1969).
Although the data is balanced, it is non-normal displaying positive skewedness.
To stabilize unequal variances and heteroscedasticity a log transformation (Y=LOG
(X+l)) was performed for all values of mean biomass (yB) and mean number of
individuals per observation (yN). The transformed values displayed low to moderate
levels of heteroscedasticity and non-normality was reduced. Under these conditions the
V statistic (Pillai's Trace) MANOVA (multiple analysis of variance) followed by
multiple comparison procedures using a Bonferroni adjustment are suggested (Johnson
and Field, 1993). The Bonferroni adjusts for the number of tests by dividing a! by N tests
for the significance level. A MANOVA procedure was performed contrasting pB and
pN with dep1.h zones and habitats for all species combined, families, and trophic groups.
This was followed by a two-way ANOVA (anlrtysis of' variance) comparing cach
response variable. The same MANOVA and ANOVA procedures without
transformations were performed comparing and contrasting mean number of species ($3)
and mean species diversity (pH') for all species. A large nurse shark (Ginglyostoma
cirratum) was removed from the data during analysis of biomass by trophic guild since it
was an extreme outlier and skewed biomass values (>3 x lo6 gm-2).
Results
Rank Abundance.-A total of 14,509 fishes were observed, representing 128 species and
35 families (Appendix I, N ) . Species were ordered according to rank abundance
(frequency of occurrence x pN) (Appendix I, N) . Acanthurus coeruleus (blue tang) was
the top ranked species comprising 16.05% of the total number of individuals. The top 10
rankdd species comprise 66.33%, the top 20 ranked species comprise 79.55%, the top 35
comprise 90.23%, the top 50 comprise 96.45%, and the top 60 ranked species comprise
98.59% of the total number of individuals. Sixty-eight additional species were rare and
comprised 1.41 % of the total number of individuals, yet comprise 53.13% of the total
number of species. The top ranked 35 species encompassed 13,089 individuals, 90.23%
by number and 27.34% of the species. .
Abundance and Biomass by Depth Zones and Habitat Types.-Mean number of
individuals per observation (pN) increased with increasing depth and complexity of
habitat type (Table 2). When comparing combination of N and pN for every depth zone
Table 2. Total number of individuals (N), mean number of individuals (pN) (bold), standard deviation (a) of pN (italics), and range per observation (N Range) for depth zones and habitats sampled at Rancho Pedro Paila, Sian Ka' an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
Habitat Depth Zone Type Patch reef Shallow reef Mid reef Deep reef All
1 N 47 1 757 559 671 2458 CtN 31.40 50.47 37.27 44.73 40.97 u 14.64 24.49 15.84 34.49 24.26
N Range 13-66 25-1 30 1 1-73 13-1 54 11-154
2 IV 90 1 1062 942 1960 4865
Ct N 60.07 70.73 62.80 130.67 81.07 u 30.43 40.30 62.15 65.32 58.17
N Range 29-122 34- 1 86 30-283 42-230 29-283
3 N 1132 1441 2259 2354 71 86
)-IN 75.47 96.07 150.60 156.93 11 9.77 u 42.45 49.10 127.81 83.42 88.10
N Range 38-165 39-1 92 30-470 25-297 25-470
All N 2504 3260 3760 4985 14509
Ct N 55.64 72.42 83.56 11 0.78 80.60 u 35.74 42.79 94.43 79.36 70.06
N Range 13-1 65 25-31 2 1 1-470 14-297 1 1-470
and habitat, it reveals that all values increase within each depth zone with increasing
habitat complexity (1 -3) (Table 2, Figure 3). Mid and deep reef zones at the forereef
showed the greatest increases in pN, especially for habitat type 2 and 3. For patch and
shallow reef zones pN increased steadily with increasing complexity, and for mid and
deep reef zones it was more dramatic. Mean number (pN) was greater with increasing
habitat complexity within each depth zone as well as combined for all depth zones. All
values were higher for shallow reefs when compared to patch reefs. The deep reef had
high and similar values for habitat 2 and 3. There was a significant effect of habitat,
zone, and interactions between zone and habitat (Table 3). Comparisons made for pN
Mean Number of Individuals per Observation (pN) for Depth Zones and Habitats
I8O.O0 1
PI P2 P3 P S1 S2 S 3 S MI M2 M3 M Dl D2 D3 D 1 2 3 ALL
Figure 3. Mean number of individuals per observation (pN) for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth zones- P= patch reef, S= shallow reef, M= mid reef, D= deep reef.
Table 3. Results of MANOVA and ANOVA showing level of significance (p-value) for all species, comparing log transformed values for mean number (pN) and mean biomass (pB) for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Significant values (p<0.05) are in bold. Depth zones- D= deep reef, M= mid reef, S= shallow reef, P= patch reef. Habitats (1,2,3).
ALL
ALL HAB ZONE HAB*ZONE 1 -2 2-3 1-3 D-M D-P D-S M-P M-S P-S
MANOVA ANOVA yN*uB cr N cr B
<0.0001 <0.0001
between each habitat type were found to be statistically significant (p< 0.05) (3>2>1)
(Tablc 3). Comparisons of I IN between depth zoiies sliowed a sigilificarlt (pc 0.05)
difference between all zones and the deep reef (D>P, D>M, D>S), as well as between
shallow and patch (S>P), but no significant difference between the mid reef when
compared to the shallow and patch reef zone (M-S, M-P). Mean number of individuals
(pN) for each habitat, combining all depth zones, shows a significant (p< 0.05) increase
with increasing complexity (3>2>1) (Table 2, 3).
There was no significant difference between depth zones for overall mean
biomass (pB) (Table 3). Values for pB were highest at the mid reef zone, followed by
the patch, shallow, and deep reef zone (M>P>S>D) (Table 4, Figure 4). There is a
Table 4. Mean biomass (pB) (bold), standard deviation (a) (italics), and range per observation for depth zones and habitats sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
Habitat Depth Zone type Patch reef Shallow reef Mid reef Deep reef All
1 PB 28.33 19.65 15.31 11.76 18.76 u 4 7.39 20.16 1 7.94 10.82 27.80
Range 0.147-1 92.52 4.72-83.93 1.96-61.23 1.06-37.97 0.147-1 92.52
PB 60.57 24.86 25.04 27.93 34.60 u 86.77 23.95 30.34 16.02 49.32
Range 6.02-274.21 9.62-103.63 8.09-127.67 4.72-61.47 4.72-274.21
3 CIB 55.25 47.09 138.62 46.52 71.87 u 58.63 35.12 156.51 29.92 93.03
Range 5.75-1 75.57 10.36-1 56.1 5 9.85-494.38 8.51 -1 04.64 5.75-494.38
All CI B 48.05 30.53 59.66 28.74 41.74 u 66.39 29.14 106.74 24.70 66.39
Range 0.147-274.21 4.72-1 56.1 5 1.96-494.38 1.06-104.64 0.147-494.64
Mean Biomass (pB) for Depth Zones and Habitats
P I P2 P3 P S1 S2 S3 S M I M 2 M 3 M D l D2 03 D 1 2 3 ALL
Figure 4. Mean biomass (pB) for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth zones- P= Patch Reef, S= Shallow Reef, M= Mid Reef, D= Deep Reef.
general increase in pB values for all individuals for each depth zone with increasing
habitat complexity. An exception to this is the patch reef zone where habitat 2 exceeds
habitat 3. There is a significant (p<0.05) difference between habitats for pB (3>2>1)
(Table 3,4).
Species Richness and Diversity.-Values for mean species richness or mean number of
species (pS), within depth zone, show an increase with increasing habitat complexity
(Table 5, Figure 5). The highest values for pS overall are for the deep reef, followed by
the shallow reef, mid reef, and the patch reef zone (D>S>M>P). There was no significant
Table 5. Mean number of species (pS), and range for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
- - --
Habitat Depth Zone Type Patch reef Shallow reef Mid reef Deep reef All
1 CIS 9.00 17.53 12.67 14.87 13.52 u 2.27 3.87 4.24 4.75 4.93
Range 4-1 3 1 1-24 5-1 8 7-24 4-24
ct s 14.27 20.00 18.20 24.27 19.1 8 u 3.15 4.33 2.01 6.04 5.44
Range 9-1 9 12-28 15-21 15-36 9-36
3 ct s 17.1 3 22.80 23.87 24.87 22.1 7 u 3.83 4.35 4.58 4.55 5.20
Range 1 1-24 17-32 17-33 1 6-31 11-33
All P s 13.47 20.1 1 18.24 21.33 18.29 u 4.59 4.63 5.92 6.84 6.29
Range 4-24 1 1-32 5-33 7-36 4-36
Mean Number of Species (pS) for Depth Zones and Habitats 30.00 I
P2 P3 P S1 S2 S3 S MI M2 M3 M D l D2 D3 D 1 2 3 ALL
Figure 5. Mean number of species (pS) for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth zones- P= patch reef, S= shallow reef, M= mid reef, D= deep reef.
difference between deep and shallow (D-S) and between mid and shallow reef zones (M-
S) (Table 6). All comparisons between habitats (3>2>1) and depth zones ( U W , U>M,
M>P) and interactions were significant (p<0.050) (Table 6). A comparison of pS for
each habitat for all depth zones reveals that values are highest for the shallow reef for
habilal 1 (Sl), deep reef for habitat 3. (W), and dccp rccf for habitat 3 (D3). For habitat
3, pS increases with depth. Values and ranges for pS become more similar with
increasing habitat complexity.
Table 6 . Results of MANOVA and ANOVA showing level of significance (p-value) for all species, comparing values for mean number of species (pS) and mean species diversity (pH) for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Significant values (p<0.05) are in bold. Depth zones- D= deep reef, M= mid reef, S= shallow reef, P= patch reef. Habitats (1, 2,3).
ALL MANOVA ANOVA ANOVA pS*uH' u S uH'
ALL <0.0001 0.0089 HAB ZONE HAB*ZON E 1-2 '
2-3 1-3 D-M D-P D-S M-P M-S P-S
Values for species diversity, measured here as mean species diversity (pHr),
increase for the patch and deep reef zone with increasing habitat complexity. The
shallow reef zone shows a reverse trend for habitat complexity and for the mid reef the
value for habitat 2 is slightly greater than for 3 (Table 7, Figure 6). Values for the
Table 7. Mean species diversity (pH') for depth zones and habitats and sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
Habitat Depth Zones types Patch rcct Shallow reef . . Mid reef Deep reet - All
1 IJ H' 1.87 2.48 2.06 2.25 2.17 D 0.32 0.35 0.37 0.32 0.40
Range 1.16-2.36 1.74-2.90 1.44-2.69 1.73-2.89 1.16-2.90
2 IJH' 2.12 2.47 2.44 2.30 2.33 D 0.46' 0.46 0.46 0.52 0.48
Range 1.30-2.68 1.54-3.08 1.08-2.77 1.25-2.90 1.08-3.08
3 P H' 2.23 2.40 2.25 2.41 2.32 u 0.42 0.40 0.88 0.36 0.55
Range 1.31-2.85 1.67-2.88 0.77-3.18 1.48-2.90 0.77-3.1 8
ALL IJ H' 2.07 2.45 2.25 2.32 2.27 u 0.43 0.39 0.62 0.40 0.49
Range 1.16-2.85 1.54-3.08 0.77-3.1 8 1.25-2.90 0.77-3.18
Mean Species Diversity (pH') for Depth Zones and Habitats
P I P2 P3 P S1 S2 S3 S M I M2 M3 M D l D2 D3 D 1 2 3 ALL
Figure 6. Mean species diversity (pH') for depth zones and habitats (1-3, all) sampled at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Depth zones- P= patch reef, S= shallow reef, M= mid reef, D= deep reef.
shallow and deep reef zones are higher and similar for habitats 3, while values for patch
and mid are similar for habitat 3. Clomparing pH' values for each habitat type across
depth zones, for habitat 1 and 2, shallow reef values are highest followed by the deep,
mid, and patch reef zone (S 1>D 1>M1 >PI). For habitat 2, shallow reef values are
highest followed by iiiid, deep, and the palcl~ rccIxonc: (SZ>MZ>D2',P2). For habitat 3,
deep reef pH' values are highest followed by shallow, mid, and the patch reef zone
(D3 S3>M 2). Overall comparisons between habitats are insignificant and values for
habitat 2 and 3 are similar (2 S>1) (Table 6). There are significant differences between
deep and patch (D>P) (p<0.0121) and between shallow and patch reef zones (S>P)
(p<0.0002) (Table 6).
Families.-Nine families of reef fishes, include diurnally active reef residents and
ecologically and economically important species, were selected to describe abundance,
distribution, and habitat preference (Appendix I, 11). These families represent 83 species, t
12,633 individuals and 87.07% of the total number of individuals sampled.
The labrids (wrasses) are the dominant family comprising 12 species, 29.45% of
the total number (N=4273, pN=5.80) (Appendix I, 11, Table 8, Figure 7). Clepticus
parrai (creole wrasse) individually ranks number 2 (N=1282, pN=29.81), followed
closely by Thalassoma bifasciatum (bluehead wrasse) which ranks number 4 (N=1274,
pN=10.53) and Halichoeres garnoti (yellowhead wrasse) which ranks number 5 (N=975,
pN=6.63) (Appendix IV). The top two ranking labrids are planktivorous and comprise
17.62% of the total number of individuals. The labrids dominate in all three habitats with
pN values that increase with habitat complexity, and are significantly (p<0.0046) greater
'l'able 8. Overall number of individuals (N), percent of the total (%Tot), mean nilmher (pN) , mean length (pXL), lneall bionlass (pB), and standard deviations (sdpN, sdpXL, sdpB) for nine families at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. *Pomacanthidae/Chaetodontidae (angelfishes/butterflyfishes).
Family N %Tot p N pXL pB sdpN sdpXL sdpB
'Po~nacar ~ltiidae/ 439 3.03 1.81 12.03 3.46 I . 6.27 6.61 Chaetodontidae
Pomacentridae 2524 17.40 15.98 7.53 0.89 18.37 1.87 0.97 (damselfishes)
Mullidae 182 1.25 2.25 11.69 0.74 1.89 5.22 1.40 (goatfishes)
Epinephelinae 214 1.47 1.89 20.54 7.48 1.06 15.05 30.73 (groupers)
Haemulidae 420 2.89 3.13 18.53 2.87 2.12 2.77 5.23 (grunts)
Scaridae 1286 8.86 7.48 15.97 5.57 5.35 5.49 7.64 (parrotfishes)
Lutjanidae (snappers)
Acanthuridae 2915 20.09 17.25 13.84 12.87 46.65 3.12 52.43 (surgeonfishes)
t
Labridae 4273 29.45 24.28 11.16 2.27 17.59 6.27 4.54 (wrasses)
between 3 and 1 for pN (3>1) (Table 9, Table 10). For depth zones, labrids dominate
overall for pN at the deep, shallow, and patch reef zone and are the second most
dominant group at the mid reef zone (Table 11). Values for pN are greatest at the deep,
shallow, and patch, followed by the mid reef zone (D>S>P>M). Values for pN are
significantly (Pc0.05) greater between deep and mid and between deep and patch (D>M,
D>P). Values for pN are also significantly (p<0.05) greater between shallow and mid
Overall Mean Number (pN), Mean Length (vXL), and Mean Biomass (pXL) for Families
Figure 7. Overall number of individuals (N), mean number (pN), mean length (pXL), and mean biomass (pB) for nine families at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Poma~Chaeto=Pomacanthidae/Chaetodontidae.
Table 9. Mean number (pN) and mean biomass (pB) for nine families for all habitats at Rancho Pedro Paila, Sian Ka'an Biosphere ReservC, Quintana Roo, Mexico, May 1999. Depth zones- P=patch reef, S= shallow reef, M= mid reef, D= deep reef.
All Habitat 1 Habitat 2 Habitat 3 All Family p N p B p N p B pN p B p N pB Pomacanthidael Chaetodontidae 2.09 1.62 3.38 4.47 3.64 4.55 1.81 3.46 Pomacentridae 5.50 0.351 18.59 0.932 18.88 1.13 15.98 0.89 Mullidae 2.49 0.78 1.81 0.74 2.25 0.77 2.25 0.74 Epinephelinae 1.67 10.35 1.97 14.35 1.95 6.84 1.89 7.48 Haemulidae 1.82 1.48 2.74 2.24 3.90 3.88 3.13 2.87 Scaridae 4.94 2.95 7.36 4.97 9.79 8.51 7.48 5.57 Lutjanidae 2.03 2.14 2.25 1.98 4.28 4.77 3.22 3.42 Acanthuridae 6.27 2.25 12.54 5.74 30.66 28.63 17.25 12.87 Labridae 16.89 1.98 23.95 2.75 31.99 2.11 24.28 2.27
Table 10. Results of MANOVA and ANOVA showing level of significance (p-value) for nine families, comparing log transformed values for mean number (yN) and mean biomass (yB) for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphcrc Reserve, Quintana Roo, Mexico, May 1999. Sigllificarlt values (p<0.05) are in bold. Depth zones- D= deep reef, M= mid reef, S- shallow rccf, 1'- patch reef. Habitats (1, 2, 3).
FAMILIES POMACANTHIDAEI MAN OVA ANOVA ANOVA - CHAETODONTIDAE IJN*PB IJN p 6 ALL <0.0001 0.4602 HA6 ZONE HAB*ZONE 1 -2 2-3 1-3 D-M D-P D-S M-P M-S P-S
POMACENTRIDAE ALL HA6 ZONE HAB*ZONE 1-2 2-3 1-3 D-M D-P '
D-S M-P M-S P-S
MANOVA IJN*IJB
ANOVA IJ N
<0.0001 <0.0001 <0.0001 <0.0001 <0.0001 0.2091 <0.0001 <0.0001 <0.0001
ANOVA IJ 6
<0.0001 0.0004 <0.0001 0.1 107 0.0018 0.2085 <0.0001 0.0005 0.0008
MANOVA ANOVA ANOVA MULLIDAE IJN*IJB IJN IJ 6 ALL 0.0465 0.0022 HA6 0.3861 0.3059 0.9974 ZONE <0.0001 0.0254 <0.0001 HAB*ZONE 0.3641 0.8305 0.4593 1-2 0.1 420 0.1362 0.9490 2-3 0.7654 0.5895 0.9920 1-3 0.41 65 0.31 53 0.9547 D-M 0.0028 0.751 6 0.0062 D-P 0.1 944 0.0693 0.21 26 D-S 0.2090 0.2624 0.0767 M-P 0.0008 0.1 150 0.1 232 M-S <0.0001 0.1324 <0.0001 P-S 0.0047 0.0026 0.0022
Table 10. Continued.
P
MANOVA ANOVA ANOVA EPINEPHELINAE IJN*l.rB LJN IIB ALL 0.1082 0.0326 HAB ZONE HAB*ZON E 1 -2 2-3 1 -3 D-M D-P D-S M-P M-S P-S
MANOVA ANOVA ANOVA HAEMULIDAE uN*uB UN II B ALL <0.0001 0.0043 HAB <0.0001 CO.0001 0.0009 ZONE 0.0066 0.1344 0.0507 HAB*ZONE 0.0220 0.0021 0.0722 1-2 0.1835 0.0983 0.0678 2-3 0.0002 0.0001 0.01 52 1-3 <0.0001 <0.0001 0.0004 D-M 0.51 84 0.2566 0.4070 D-P 0.0564 0.021 5 0.0220 D-S 0.0087 0.1 706 0.6255 M-P 0.2430 0.1453 0.0929 M-S 0.0299 0.7986 0.1 874 P-S ' 0.0082 0.21 06 0.0080
MANOVA ANOVA ANOVA SCARIDAE l.rN*lJB IJ N l.r 6 ALL <0.0001 ~0.0001 HAB ZONE HAB*ZONE 1-2 2-3 1-3 D-M D-P D-S M-P M-S P-S
Table 10. Continued.
MANOVA ANOVA ANOVA LUTJAN IDAE IJN*IJB IJN P B ALL 0.0186 0.0431 HAB 0.0061 0.0007 0.0035 ZONE 0.1 640 0.2571 0.5026 HAB*ZONE 0.36'1 3 0.6953 0.6780 1 -2 0.9828 0.8744 0.8532 2-3 0.0021 0.0005 0.0021 1-3 0.0282 0.0082 0.021 5 D-M 0.3853 0.2067 0.5558 D-P 0.3003 0.1396 0.1 502 D-S 0.0366 0.0778 0.9236 M-P 0.5772 0.5942 0.3202 M-S 0.3552 0.5928 0.6380 P-S 0.1 636 0.9062 0.1 822
MANOVA ANOVA ANOVA ACANTHURIDAE ALL HAB ZONE HAB*ZONE 1 -2 2-3 1-3 D-M D-P D-S M-P M-S ' P-S
MANOVA ANOVA ANOVA LABRIDAE IJN*IJB IJN IJ B ALL 0.0002 0.271 4 HAB ZONE HAB'ZONE 1-2 2-3 1-3 D-M D-P D-S M-P M-S P-S
Table 1 1. Mean number (1tN) and mean biomass (pB) for nine families for depth zones and habitats at Ranchu PeJlu Paila, Siau Ra'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
Patch Reef Zone Habitat 1 Habitat 2 Habitat 3 Al I Family p N pB p N pB pN pB pN p B Pomacanthidael Chaetodontidae 1.33 0.623 2.33 4.95 2.33 4.87 2.00 3.48 Pomacentridae 1 .OO 0.017 4.88 0.513 4.79 0.461 3.55 0.33 Mullidae 3.24 0.520 2.60 1.48 3.50 0.800 3.24 0.930 Epinephelinae 1.33 37.69 2.67 54.43 1.20 1 .OO 1.73 34.28 Haemulidae 1 .OO 0.349 1.33 1.16 4.29 6.32 2.21 2.61 Scaridae 5.08 2.38 8.40 3.91 12.07 6.34 8.51 4.21 Lutjanidae 2.00 1.50 1.50 0.902 3.08 3.1 1 2.19 1.84 Acanthuridae 6.86 2.73 15.80 8.21 22.87 13.05 15.18 7.99 Labridae 11.14 1.76 16.53 1.58 18.64 1.28 15.44 1.54
Shallow Reef Zone Habitat 1 Habitat 2 Habitat 3 All Family pN p B pN )-I B p N pB pN pB Pomacanthidael Chaetodontidae 2.20 2.92 2.00 8.45 2.38 4.90 2.19 5.42 Pomacentridae 7.53 0.766 12.27 1.13 14.67 1.24 11.49 1.04 Mullidae 1.33 0.050 1.25 0.068 1.50 0.113 1.36 0.077 Epinephelinae 1.62 1.18 1.50 1.20 2.1 1 11.67 1.74 4.68 Haemulidae 2.50 2.59 2.91 2.60 3.20 3.15 2.87 2.78 Scaridae 6.53 4.86 7.27 5.80 8.33 10.35 7.38 7.00 Lutjanidae 2.00 1.39 1.78 2.33 3.71 6.97 2.50 3.56 Acanthuridae 9.07 1.98 10.20 2.41 5.69 1.90 8.32 2.10 Labridae 20.67 1.76 19.27 1.27 49.33 2.28 29.62 1.77
Mid deef Zone Habitat 1 Habitat 2 Habitat 3 All Family p N pB p N p B p lV pB pN p B Pomacanthidael Chaetodontidae 2.40 0.224 3.93 2.72 3.86 3.83 3.40 2.26 Pomacentridae 5.93 0.235 10.27 0.536 15.80 1.09 10.67 0.619 Mullidae 2.83 2.02 1.40 0.739 2.56 1.92 2.26 1.56 Epinephelinae 1.92 1.20 1.50 1.13 2.00 3.45 2.17 1.93 Haemulidae 1.63 1.20 2.38 2.00 4.53 3.41 2.85 2.20 Scaridae 4.46 2.58 7.40 4.69 6.64 8.68 6.17 5.31 Lutjanidae - 2.00 4.02 1.67 1.59 4.93 4.23 2.87 3.28 Acanthuridae 6.42 3.32 19.93 10.70 88.06 97.03 38.14 37.02 Labridae 15.14 1.29 13.27 1.43 15.47 2.16 14.63 1.63
Table 1 1. Continued. - Deep Reef Zone Habitat 1 - Habitat 2 Habitat 3 All Family p N pB pN p B p N PB pN IJB - Pornacanlhidoc/ Chaetodontidae 2.42 2.72 5.27 1.78 6.00 4.59 4.56 3.03 Pomacentridae 7.53 0.387 46.93 1.55 40.27 1.75 31.58 1.23 Mullidae 2.17 0.52 2.00 0.670 1.43 0.276 1.87 0.48/ Epinephelinae 1.92 1.33 1.50 0.65 2.00 I .51 1.81 1.16 Haemulidae 2.14 1.79 4.33 3.21 3.57 2.65 3.35 2.55 Scaridae 3.69 1.97 6.36 5.48 12.13 8.66 7.39 5.37 Lutjanidae 2.13 1.64 4.07 3.09 5.42 4.78 3.87 3.1 7 Acanthuridae 2.73 0.960 4.21 1.64 6.00 2.53 4.31 1.71 Labridae 21.00 3.13 46.71 6.71 44.20 2.71 37.30 4.18
and between shallow and patch reef zones (S>P, S>M).
The acanthurids (surgeonfishes) are represented by three species and are the
second most dominant family, comprising 20.09% (N=2915, pN=17.25) of the total
number of individuals (Appendix 11, IV, Table 8, Figure 7). Acanthurus cueruleus (blue
tang) individually ranks number 1 (N=2328, pN= 19.73), comprising 16.05% of the total. t
Acanthurus bahianus (ocean surgeonfish) ranks number 7 (N=538, pN=3.74), comprising
3.71% of the total. Acanthurids not only have the highest value for pN but also for pB
(12.87), and a medium value for pXL (13.84). Acanthurids show a significant difference
for pN and pB values between depth zones, habitats, and interactions between habitats
and zones (Table 10). Both values increase with habitat complexity and are significantly
(p<0.0009) greater between habitat 3 and 1 (3>1) (Table 9, 10). Values for pB are all
significantly (p<0.05) different between each habitat (3>2>1). Values for pN are all
significantly @<0.05) less between the deep reef with all other zones (D<S, D<M, D<P).
The pomacentrids (damselfishes) are the third most dominant family, comprising
12 species and 17.40% of the total number (N=2524, pN=15.98) (Appendix 11, IV, Table
8, Figure 7). The planktivorous species, Chromis cyaneus (blue chromis) and the
omnivore Stegastes partitus (bicolor damselfish) dominate the pomacentiidae at 77.62%
(Appendix IV). Chromis cyaneus ranks number 3, at N=1275, pN=l3.7 1) and "$lt!g(~stcl,~
partitus ranks number 6 (N=685, pN=6.46) (Appendix IV). Due to their small sizes
(pXL=7.53) pomacentrids have low values for pB (0.890) (Table 8). The pomacentrids
show significant effects of habitat, zone and interactions. Pomacentrids have very high
and similar values for pN at habitat 2 (pN=18.59) and habitat 3 (pN=18.88) (Table 9).
There were high values for pN for the shallow reef (pN=l1.49), mid reef (pN=10.67),
and deep reef zone (pN=3 1.58) (Table 1 1). There were no significant differences
between shallow and mid reef zones and between habitats 2 and 3 but there was
significant differences (pc0.05) for the other zones and habitats (Table 10).
The scarids (parrotfishes) comprise 14 species at 8.86% of the total number
(N=1286, pN=7.48) with Sparisoma aurofrenatum (redband parrotfish), dominating the
scarids at 40.51% (N=521, pN=7.48) and ranking 8 overall. (Appendix II, IV, Table 8,
Figure 7). Scarus iseri (striped parrotfish) is the next most abundant scarid comprising
2 1 35% (N=28 1, pN=3.3 9) of the scarids and ranking 1 1 overall. Parrotfish have the
third highest value for pB (7.48) and pXL (15.97) (Table 8). Values for pN and pB
increase with increasing habitat complexity (3>2>1) (Table 9). The values for pN and
pB for the patch, shallow, and mid reef zone increase with increasing habitat complexity
(Table 11). The differences of pN and pB between habitat 1 and 2 (2>1) and between 1
and 3 (3>1) are significantly (p<0.05) different (Table 10). There are no significant
differences between depth zones for pN and pB values.
The pomacanthids/chaetodontids (angelfishes/butterflyfishes) comprise nine
species and 3.03% (N=439, pN=1.8 1) of the total number (Appendix 11, IV, Table 8).
'l'hey arc doiniilated by the cliaetoclontid Chuetodon cupis~rutus (foureye butterflyfish),
comprising 34.40% (N=15 1, pN=2.29) of the family and ranking 17 overall. The
pornacanthid Holocunthus tricolor (rock beauty) compnscs 3 1.89% (N=140, pN=1.84)
and ranks 18 overall. Both species combined comprise 66.29% of the family by number.
The pomacanthids/chaetodontids have similar values with acanthurids for pXL (12.03)
and a medium value for pB (3.46). The differences of pN between habitat 1 and 2 (2>1)
and between 1 and 3 (3>1) are significantly (p<0.05) different. All comparisons by depth
zone for pN are significant (p<0.05) with the exception of patch and shallow which have
very similar values.
The haemulids (grunts) were not abundant and only comprise 2.89% of the total
number (N-420, pN=3.13) (Appendix 11, IV, Table 8). Of the nine species of grunt
observed, Haemulonflavolineatum (French grunt) was dominant at 25.24% (N=106, t
pN=1.43) and a rank of 24 overall. Haemulon plumieri (white grunt) (N=84, pN=1.18)
and Anisotremus virginicus (porkfish) (N=84, pN=1.3 1) each rank 30 overall and
comprise 20.00% each or 40% total. Haemulon sciurus (bluestripe grunt) comprise
16.67% of the haemulids (N=70, pN=1.46) and a rank of 35 overall. Grunts have the
third highest pXL (18.53) but a low pB (2.87). The three species of Haemulon above . -
were observed frequently but only solitary or in a small group, no large schools were
observed. No sizeable schools were observed for any species of grunt. There was an
increase in values for pN and pB with increasing habitat complexity and significant
differences (p<0.05) between 2 and 3 (3>2) and between 1 and 3 (3>1) (Table 9, 10).
Values for pN and pB increased by depth zone but values for the shallow and mid reef
zone were very similar and insignificant. Thcrc was a significant (pC0.02 15, F0.0220)
difference between values for pN and pB between the deep and patch reef zones (D>P)
and a significant (p<0.0080) difference for pB between the patch and the shallow reef
zone (S>P).
The lutjanids (snappers) comprise six species, 2.62% of the total (N=380,
pN=2.62) with Ocyurus ch ysurus (yellowtail snapper) dominating the lutjanids at
66.58% (N=253,2.78) and ranked number 12 overall (Appendix 11, IV, Table 8).
Lutjanus mahogoni (mahogany snapper) was not abundant but ranks second to 0.
ch ysurus at 19.2 1 % (N=73, pN=2.15) of the lutjanids and ranked 34 overall (Table 8).
Snappers have the second highest value for pXL (20.35) and a medium value for pB
(2.51), similar to the pomacanthids/chaetodontids. With the possible exception of 0.
chrysurus, no sizeable schools of snapper were observed. The highest values for pN and
pB were for habitat 3 and there were significant (Pc0.05) differences between habitats 3
and 2 (3>2) and between 3 and 1 (3>1) (Table 9, 10). Although snappers had higher
values at the deep zone, the values at the other three zones were very similar and
insignificant.
The epinephelins (groupers) comprised six species and had a low value for
abundance (N=214, pN=1.47) but the highest value for pXL (20.54) and the second to
highest value for pB (7.48) (Appendix 11, IV, Table 8). Epinephelus fulvus (coney) is a
small species and ranks 22 overall, and comprised 54.21% of the groupers (N=l16,
pN=1.57). Epinephelus cruentatus (graysby) is a small species that comprised 29.91% of
the groupers and was low in abundance (N=64, pN=1.23). The medium sized E. guttatus
(red hind) was very low in abundance (N=18, yN=1.13) as well as the large sized
Adj~ctnrop~rcn: hontzci (black g o ~ ~ p e r ) (N-12, yN-1.20). Groupcrs showcd a significailt
(p<0.05) difference for yB values for all zones compared with the patch reef zone (P>S,
P>M, P>D) (Table 10, 11). The yB value for the patch reef zone was 34.28 and all other
values for yN and yB were low.
The illullids (goatfishes) comprise two species, 1.25% of the total (N=l82,
yN=2.25) and Pseudupeneus maculatus (spotted goatfish) is dominant and comprises
71.97% (N=13 1, yN=1.82) and ranks 19 overall, while Mulloidichthys martinicus
(yellow goatfish) only comprises 35.69% (N=51, yN=3.40) and ranks 42 (Appendix 11,
IV, Table 8). Goatfishes exhibited a medium range for yXL (1 1.69) and the lowest value
for yB (0.74). Goatfishes were not abundant and were observed only in small groups; no
large schools were observed. There were no significant differences for yN and yB
between habitats (Table 10). The highest value for yN (3.24) was at the patch reef zone
and the highest value for yB (1.56) was at the mid reef. There was a significant
(p<Or0062, p<0.0001) difference for yB between the mid and deep reef (M>D) and
between the mid and the shallow reef (M>D). There is a significant difference for yN
(p<0.0026) and yB (p<0.0022) between patch and shallow reef zones (P>S).
Mean number of individuals (yN) and biomass (yB) are compared for families
(Appendix I, 11) for each category of depth zone and habitat type (Table 8). Beginning
with the habitats overall (Table 9), labrids dominate each habitat, followed by acanthurids
and pomacentrids. At habitat 2 labrids dominate, followed by pomacentrids and
acanthurids. Acanthurids have the highest values for yN, and the values for yN and yB
increase with increasing habitat complexity. Labrids are second highest in yN, which
increases with habitat complexity. Labrids have low values for pB, which decreases with
increased habitat complexity. Like the labrids, the mullids have higher values for pN for
habitat type 1 with low values for pB. Most of the species of labrids and mullids found
in this zone are epibenthic feeders (benthivores) that prefer sand bottom and low reef
areas found in habitat 1. Lower values for pB, especially in habitat 1, are due to a large
number of smaller juvenile fishes in the patch reef zone. Groupers have high values for
pB and low values for pN, as evidenced by several large black grouper (Mycteroperca
bonaci) seen in the low relief areas of habitat 1 and 2.
For the shallow reef zone habitats pN and pB values for families are compared
(Table 10, Figure 8). The acanthurids and labrids have the higher pN values, with the
acahthurids lower value at habitat 3. Values for pB are low for these families probably
due to a larger number of juveniles in these shallow low relief habitats and the lack of
large adult schools. The hghest value for pN are for the labrids at habitat 3, however
they have a very low pB mostly due to the large number of planktivorous bluehead
wrasie (Thalassoma bifasciatum) and juvenile parasite cleaning stages. Pomacentrids,
scarids, haemulids show an increase in pN and pB with increasing habitat complexity.
Epinephelins and lutjanids have higher pN values at habitat 3 and a high pB value.
For the mid reef zone habitats pN and pB values for families are compared (Table
11, Figure 10). Acanthurids have the highest values for pN and pB, which increase with
increasing habitat complexity. Very high values at habitat 3 were due to a large
spawning aggregation observed. Pomacentrids and labrids have higher pN and pB
values at habitat 3, due to the dominance of the planktivorous blue chromis (Chromis
cyaneus) and creole wrasse (Clepticus parrai) off of the forereef slope. Mullids have
higher values at habitat 1, low relief sandy areas. Scarids pN values decrease and pB
values increase with increasing habitat complexity. This was due to larger individuals
seen on the high relief forereef areas while smaller more numerous foraging groups
occurred on low relief areas. Lutjanids and haemulids values are relatively low but have
a higher pN value at habitat 3. Lutjanids have a higher pB value at habitat 1 and
haemulids have a higher pB value at habitat 2. Groupers have relatively low values as
well, but have higher values at habitat 3.
For the deep reef zone habitats pN and pB values for families are compared
(Table 11). Pomacentrids and labrids dominate the deep reef zone and have high values
for pN but low values for pB. This is due to the dominance of the two planktivorous
species mentioned above. Acanthurids have relatively low values for pN and pB, yet
they dominated all of the other depth zones for pN. Lutjanids and scarids increase in pN
and pB with increasing habitat complexity, with relatively high values for habitat type 3.
Pomacanthids/chaetodontids increase slightly for values in pN with increasing habitat
$
complexity.
Trophic Structure.-To analyze the reef fish community based on trophic structure, all
species were assigned to seven major trophic guilds (Appendix 111). The top three trophic
guilds were herbivores, planktivores, and benthivores, comprising 80.44% of the total
number of individuals (Table 12, Figure 8).
The planktivores are the dominant trophic guild, comprising 32.75% of the total
numbers (N=4752) with the highest value for pN (29.89) and the next to lowest value for
Table 12. Overall number of individuals (N), percent of the total number of individuals (%Tot), mean number (yN), mean length (yXL), mean biomass (yB), and standard deviations (sdyN, sdyXL, sdyB) for seven trophic guilds at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. SAF= sessile animal feeders.
Trophic Guild Benthivores Herbivores Omnivores Pelagic Piscivores Piscivores Planktivores SAF
35 1 Overall Mean Number (pN), Mean Length (WL), and Mean Biomass (pN) for Tro~hic Guilds
Benthivores Herbivores Omnivores Pelagic Piscivores Planktivores SAF Piscivores
Figure 8. Overall mean number (yN), mean length (yXL), mean biomass (yB), and standard deviations for seven trophic guilds at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. SAF= sessile animal feeders
pB (2.95) and pXL (8.05) (Table 12, Figure 8). Of the 10 top ranking species four are
planktivores, comprising 29.22% (Clepticus parrai, Chromis cyaneus, ~ a l a s s o m a
bifasciatum, and Inermia vittata) (Appendix 111, IV). All of these species concentrate at
the forereef slope areas of the mid and deep reef zone where the zooplankton supply is
abundant. An exception is Thalassoma bifasciatum which was abundant in every zone
and habitat and where the juvenile stages also feed as parasite cleaners. All comparisons
for yN and yB by habitat, depth zone, and interactions between habitat and zone are
significant for planktivores. There is a sharp and significant (p<0.05) increase in yN and
yB withincreasing habitat complexity (3>2>1) (Table 13, 14). There is a significant
(p<0.05) difference for yN and yB between zones as well, with the highest values at the
deep reef, followed by the shallow, mid, and patch reef zone (D>S>M>P) (Tablel4, 15).
However for yB at the mid and deep reef zone the values are the same and insignificant
(M=S). Planktivores dominate overall by habitat and depth zone including the deep and
shallow reef zone as well as habitat 2 and 3.
The herbivores are a close second in dominance to the planktivores, comprising
31.52% of the total number (N=4473, yN=25.84) (Table 12, Figure 8). Herbivores have
the h;ghest value for yB (19.49) and the third highest value for yXL (17.22). The
Table 13. Mean number (pN) and mean biomass (pB) for all habitats for seven trophic guilds at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. P= patch reef, S= shallow reef, M= mid reef, D= deep reef, SAF= sessile animal feeders.
Trophic Guild Benthivores Herbivores Omnivores Pelagic Piscivores Piscivores Plan ktivores SAF
Habitat 1 Habitat 2 Habitat 3 All
pN pB pN pB pN p B pN pB 14.43 3.47 13.12 4.65 16.48 6.26 14.68 4.79 11.55 5.29 21.85 12.33 43.27 40.10 25.84 19.49 3.83 0.127 6.95 0.251 6.88 0.260 6.56 0.23 3.53 10.50 14.03 11.67 3.28 20.14 8.13 16.22 2.54 10.12 3.31 11.25 5.41 8.49 3.94 8.18 9.14 0.680 28.30 2.23 44.61 5.06 29.89 2.95 2.19 1.66 3.76 4.53 3.91 4.53 3.60 3.50
Table 14. Results of MANOVA and ANOVA showing level of significance (p-value) for seven trophic guilds, comparing log transformed values of mean number (pN) and mean biomass (pB) for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999. Significant values (p<0.05) are in bold. Depth zones- D= deep reef, M= mid reef, S= shallow reef, P= patch reef. Habitats (1, 2, 3).
TROPHIC GUILDS MANOVA AIVOVA ANOVA
BENTHIVORES uN*uB u N uB ALL 0.01 28 0.0009 HAB ZONE HAB*ZOP 1-2 2-3 1 -3 D-M D-P D-S M-P M-S P-S
HERBIVORES ALL HAB ZONE HAB*ZONE 1 -2 2-3 1 -3 D-M D-P ' D-S M-P M-S P-S
MANOVA ANOVA ANOVA uN*uB uN u B
<0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 0.0002 0.0086 <0.0001 0.001 0 0.0812 0.0003 0.0002 <0.0001 0.001 7 0.0042 0.0004 <0.0001 <0.0001 <0.0001 0.0020 0.0006 0.0009 ~0.0001 0.0001 0.0350 0.0010 0.0004 0.0244 0.01 33 0.6736 0.1 943 0.0965 0.9415 0.2480 0.6929 0.7246 0.8843
MANOVA ANOVA ANOVA OMNIVORES uN*vB IJ N uB ALL <0.0001 0.2004 HAB ZONE HAB*ZONE 1 -2 2-3 1-3 D-M D-P D-S M-P M-S P-S
Table 14. Continued.
PELAGIC MANOVA AIVOVA ANOVA PlSClVORES uN*clB uN u B ALL 0.0460 0.7282 HAB 0.0981 0.2955 0.3704 ZONE 0.0707 0.0438 0.3593 HAB*ZONE 0.8901 0.4025 0.9452 1 -2 0.3461 0.2598 0.81 58 2-3 0.0230 0.1 467 0.1764 1-3 0.6336 0.9881 0.4053 D-M 0.21 54 0.9974 0.1288 D-P 0.0955 0.0394 0.0902 D-S 0.2804 0.2244 0.1268 M-P 0.0181 0.01 16 0.9036 M-S 0.2077 0.1 356 0.9340 P-S 0.4605 0.241 3 0.8238
MANOVA ANOVA ANOVA PISCIVORES uN*uB u N u B ALL <0.0001 0.0129 HAB 0.0002 <0.0001 0.0255 ZONE 0.0008 0.0229 0.4342 HAB*ZONE 0.0221 0.2213 0.0429 1-2 0.331 4 0.1 691 0.7782 2-3 0.0033 0.0008 0.0284 1-3 <0.0001 <0.0001 0.01 58 D-M 0.8635 0.6249 0.9416 D-P 0.0001 0.0076 0.1 542 D-S 0.0138 0.0312 0.5273 M-P 0.0006 0.0260 0.1403 M-S 0.0540 0.1 171 0.4863 P-S 0.21 24 0.4077 0.3957
MANOVA ANOVA ANOVA PLANKTIVORES uN*uB uN uB ALL <0.0001 <0.0001 HAB ZONE HAB*ZON E 1-2 2-3 1-3 D-M D-P D-S M-P M-S P-S
Table 14. Continued.
SESSILE ANIMAL MANOVA ANOVA ANOVA. FEEDERS IJN*IJB IJN ~1 B ALL <0.0001 0.41 95 HAB ZONE HAB*ZON E 1 -2 2-3 1-3 D-M D-P D-S M-P M-S P-S
herbivores are composed of 23 species, including all acanthurids, scarids, and seven
unrelated species from other families (Appendix I, 111). Of the 10 top ranking species,
three are herbivores (Acanthurus coeruleus, A. bahianus, and Sparisoma aurofrenatum),
comprising 23.34% of the total number. For habitats, herbivores are the second most
domkate group for pN for each habitat and have the highest value for yB at habitats 2
and 3. All comaprisons of pN and yB between each habitat are significantly (p<0.05)
different (3>2>1) (Table 13, 14). Herbivores are one of the dominant groups for each
depth zone for pN and yB and completely dominate at the mid reef zone (M>P>S>D).
All comparisons of yN and pB of the deep depth zone between the other three zones are
significantly (p<0.05) less (D<P, D<S, D<M) (Table 14). Herbivores had significantly
lower values at the deep reef zone.
The next most abundant trophic guild is the benthivores, which is the most
Table 15. Mean number (pN) and mean biomass (pB) for seven trophic groups for depth zones and habitats at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
Patch Reef Zone Habitat 1 Habitat 2 Habitat 3 All
Trophic Guild pN p B p N pB pN pB p N pB Benthivores 13.60 4.26 13.20 4.15 14.60 7.21 13.80 5.21 Herbivores 11.55 5.52 21.85 17.97 43.26 27.21 26.02 16.90 Pelagic Piscivores 8.80 8.91 20.71 28.05 6.67 27.39 12.06 21.45 Omnivores 1.00 0.017 2.71 0.193 3.42 0.272 2.38 0.161 Piscivores 1.71 32.73 2.78 36.89 3.62 7.28 2.70 25.63 Plan ktivores 3.50 0.103 8.00 0.108 11.50 0.379 7.67 0.197 SAF 1.33 0.623 3.67 5.08 3.18 4.13 2.73 3.28
Shallow Reef Zone Habitat 1 Habitat 2 Habitat 3 All
Trophic Guild p N pB p N pB p N pB pN pB Benthivores 10.40 2.72 11.87 3.47 15.73 2.88 12.67 3.36 Herbivores 17.93 7.32 22.40 8.90 16.87 15.94 19.06 10.72 Omnivores 4.80 0.220 5.07 0.302 5.15 0.294 5.01 0.272 Pelagic Piscivores 2.00 14.58 32.67 15.40 1.60 17.78 12.09 15.92 Piscivores 2.69 1.93 2.38 2.54 5.07 14.47 3.38 6.31 Plan ktivores 13.33 0.856 15.73 0.422 51.47 2.78 26.84 1.35 SAF 2.60 3.07 2.17 8.56 2.62 5.55 ' 2.46 5.73
. .
Mid Reef Zone Habitat 1 Habitat 2 Habitat 3 All Tro hic Guild N B I N B N B Benthivores 15.27 4.26 9.47 4.15 13.53 7.21 12.76 5.21 Herbivores 10.54 5.67 28.27 15.55 95.20 105.38 44.67 42.20 Omnivores 4.64 0.017 6.29 0.193 7.87 0.272 6.27 0.161 ~ e l a d c Piscivores 1.33 8.91 1.25 28.05 3.57 27.39 2.05 21.45 Piscivores 2.84 32.73 2.85 36.89 6.93 7.28 4.21 25.63 Planktivores 5.73 0.103 12.47 0.108 23.36 0.379 13.85 0.197 SAF 2.40 0.623 3.93 5.08 3.86 4.13 3.40 3.28
Deep Reef Zone Habitat 1 Habitat 2 Habitat 3 All
Trophic Guild pN pB p N p B p N p B pN pB Benthivores 18.47 3.93 17.93 7.62 22.07 7.56 19.49 6.37 Herbivores 6.14 2.67 11.07 6.89 20.20 11.86 12.47 7.14 Omnivores 4.86 0.1 15 13.73 0.316 11.07 0.255 9.89 0.229 Pelagic Piscivores 2.00 0.526 1.50 0.33 1.29 5.65 1.60 2.17 Piscivores 2.93 2.08 5.27 3.60 6.00 5.05 4.73- 3.58 Plan ktivores 14.00 1.34 77.00 7.64 92.13 14.92 61.04 7.97 SAF 2.42 2.72 5.27 1.78 6.00 4.59 4.56 3.03
speciose group and comprises 45 species, including all haemulids and mullids, most
labrids, and several other families, such as sparids, balistids, holocentrids, serranids,
ostraciids, tetraodontids, and others (Appendix I, 111). Benthivores comprise 18.2 1% of
the total number (N=2642, pN= 14.68) with medium values for pXL (15.04) and pB
(4.79) (Table 12, Figure 8). Halichoeres garnoti was the only benthivorous species that
ranked in the top 10, comprising 6.72% of the total. Benthivores were dominant by pN
for habitat 1 and were third overall for habitats 2 and 3 (Table 13). Values for pN were
significantly (p<0.0415) different when comparing habitats 2 and 3 (3>2) and for pB
when comparing habitats 1 and 2 ( 2 ~ 1 ) (p<0.0100) and habitats 2 and 3 (3>2) (p<0.0001)
(Table 14). Values for pN were significantly (p<0.005) greater when comparing the deep
reef with with the other three depth zones (D>P, D>S, D>M). Values for pB were
significantly (p<0.0044) greater when comparing the deep reef with the shallow reef zone
(DBS).
The next four trophic guilds (pelagic piscivores, piscivores, omnivores, and
sessile animal feeders) are lower in abundance (N=2541) and represent 17.51% of the
total number (Table 12, Figurk 8). The piscivores comprise 4.16% of the total number
(N=603, pN=3.94) and the second highest value for pXL (20.27) and the third highest
value for pB (8.18). The piscivores comprise 16 species, including all of the
epinephelins, lutjanids, aulostomids, and synodontids (Appendix 111, IV). Ocyurus
chrysurus is the dominant piscivore and ranks 12, comprising 41.96% of all piscivores
(N=253, pN=2.78) (Appendix IV). For habitats, values for pN and pB increase with
increasing habitat complexity (3>2>1) and these values at habitat 3 are significantly
(p<0.05) greater for habitat 3 when compared to 2 (3>2) and to 1 (3>1) (Table 13, 14).
The pelagic piscivores are transient, non-resident reef fishes, and were placed in a
separate trophic guild. This trophic guild includes four species, including two carangids,
a scombrid, and a sphyraenid (Appendix I, 111). They comprised 3.42% of the total
number (N=496, pN=8.13) and the highest value for pXL (24.69) and the second highest
value for pB (16.22) (Table 12, Figure 8). Caranx ruber was the only species of
piscivore that ranked in the top 10, comprising 2.32% of the total (Appendix IV). For pN
and pB values, pelagic piscivores had high values in the shallow and patch reef zones and
in habitat 2, but low values in the deep reef zone (Table 15). They also had low pN
values but high pB values in the deep reef zone and for habitats 1 and 2 (Table 13).
None of the comparisons by habitat or depth zone were significant (Table 14).
The omnivores are comprised of only seven species, including five pomacentrids,
a clinid, and a balistid (Appendix I, 111) and comprise 6.69% of the total number (N=971,
pN=6.56) with the lowest values for pXL (6.12) and pB (0.230) (Table 12, Figure 8).
Stegastes partitus was the only species of omnivore that ranked in the top 10, comprising
4.72% of the total number and 70.55% of the omnivores (N=685, pN=6.46) (Appendix
IV). For zones, the values for pN increase with depth (D>M>S>P) and are significantly
(pC0.05) different between the deep reef and the patch and shallow reef zones (D>P,
D>S) as well as between the mid reef and the patch reef zone (M>P) (Table 14, 15). The
mid and deep reef have higher and similar pN values (4.73,4.21) and the mid and patch
reef have equal pB values (25.63).
The sessile animal feeders (SAF) comprise 3.25% of the total, and have the
lowest values for abundance (N=47 1, pN=3.60), but medium values for pXL (1 1.63) and
pB (3.50) (Table 12, Figure 8). SAF comprise 11 species, including all
pomacanthids/chaetodontids, as well as Abudefduf saxatzlis (Pomacentridae) and Aluterus
scriptus (Balistidae) (Appendix 111, I). SAF values for pN are significantly (p<0.0060,
p<0.0004) greater between habitat 3 and 1 (3> 1) and between habitat 2 and 1 (2>1)
(Table 13, 14). Values for pN are significantly (pc0.05) greater between the deep reef
and patch reef zone (D>P) and between the deep reef and shallow reef zone @>S) as
well as between the mid and shallow reef zone (M>S) (Table 14, 15). The highest value
for pN is at the deep reef zone (4.56) and for pB it is at the shallow reef zone (5.73).
Values for pN and pB of trophic guilds are compared for each reef zone habitat
(Table 15). Planktivores have the highest values for pN in all areas, especially for the
shallow and deep reef zones in habitat type 2 and 3. Both omnivores and planktivores
have relatively low pB values since most species in these groups are small. Benthivores
have a higher pN value in habitat type 1 for all depth zones. Omnivores show a distinct
preference for mid and deep reef zones and their values for pN and pB increase with
increasing habitat complexity. Herbivores, for the most part, have high values for pN
and ;B and increase with habitat complexity. Values are higher in the patch and mid reef
zone. Sessile animal feeders (SAF) have higher values for pN at all habitat 2 and 3 sites
with a higher pB at patch and shallow reef zones versus mid and deep reef zones.
Piscivores show an increase in values at the deep reef zone with increasing habitat
complexity. Piscivores have relatively high values at the type 3 habitat types for all depth
zones and the type 2 habitat for the deep depth zone.
A comparison of pN and pB for trophic guilds was made of habitats for each
depth zone. Starting with the patch reef zone (Table 1 9 , benthivores pN values decrease
with increasing habitat complexity while pB increases. Herbivores values for pN and pB
increase with increasing habitat complexity. Planktivores have the highest values for pN,
which increase with habitat complexity, but have low pB values. SAF have relatively
high values for pN and pB at habitat 2 and 3, but low values at habitat 1. Piscivores have
higher values for habitat 3 and for pN at habitat 1, and low values for pN and pB at
habitat 2 and pB at habitat 1. Omnivores have low values, especially for habitat 1,
however values increase with habitat complexity.
For the shallow reef zone habitats pN and pB values for families are compared
(Table 15). Planktivores have the highest values for pN and with their numbers doubling
at habitat 3 compared with 1 and 2. Biomass values are low, especially at habitat 2.
Piscivores had low values for pN and pB for habitat 1 and 2, with all values increasing
with habitat complexity. Benthivores had very similar values for pN at habitat 1 and 2,
and low values for pB, but higher value for pN at habitat 3. Herbivores had similar
values for pN and a higher value for pB at habitat 3. SAF had low values for all
variables except high pB values at habitat 3 and highest at habitat 2. fi
Values for pN and pB, for trophic guilds, will be compared at mid reef zone
habitats (Table 15). Herbivores have high values for pN and pB and these values
increase with habitat complexity. The values for pN and pB at habitat 3 are the highest
values of any depth zone and habitat for any trophic guild due to the large concentration
of blue tang (Acanthurus coeruleus) seen there during a spawning aggregation.
Planktivores have the next highest pN values. Values for pN and the relatively low
values for pB increase with habitat complexity. Values for pN for benthivores decrease
with increasing habitat complexity with higher values of pB at habitat 3. Values for pN
for omnivores are relatively high and for pB are low but both values increase with habitat
complexity. Piscivores have low values at habitat 2 and low pN value at habitat 1 with a
higher pB. SAF have low values with a higher pN at habitat 2 and pB at habitat 3.
Comparisons of trophic guilds at deep depth zone habitats reveal dominance of
this zone by planktivores (Table 15). Planktivores have high values for pN and low
values for pB, both of which increase with habitat complexity. Greater biomass values at
habitat 3 reveal greater numbers of larger planktivorous species such as creole wrasse
(Clepticusparrai). Omnivores have the next higher values for pN. Values for pN
increase with habitat complexity, as do pB values, which are very low. Values at habitat
3 are virtually the same. Piscivores and herbivores values increase with increasing
habitat complexity. Sessile animal feeder pN values increase slightly with increasing
habitat complexity, while pB values are low and lowest at habitat 2. Benthivores
decrease in pN with increasing habitat complexity. Biomass values are low, similar and
slightly higher at habitat 2 and 3.
species Dominance by Depth Zones. -The patch reef zone had the lowest overall
abundance (N=2504), significantly (p<0.05) different from the mid and deep reef zones
(Table 3, 16). The herbivorous acanthurid, Acanthurus coeruleus, comprising 22.00%
(N=55 I), and ranked number 1. Of the top ten in the patch reef zone, five were
herbivores, comprising 39.90%. These include A. bahianus, Scarus iseri, Sparisoma
aurofrenatum, and Kyphosus sectatrix/incisor. Two benthivorous labrids, Halichoeres
garnoti and H. bivittatus, comprise 10.14% of the number of individuals for the patch
reef zone. Only one planktivorous labrid, Thalassoma bifasciatum, ranks number 2,
comprising 9.50% (N=238). Two pelagic piscivorous carangids, Caranx bartholomaei
Table 16. Top 10 ranked species with values for rank abundance (N) for each depth zone observed at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
Patch Reef Shallow Reef Mid Reef Deep Reef 2504 3260 3760 4985
Acanthurus coeruleus Thalassoma bifasciatum Acanthurus coeruleus Chromis cyaneus 551 664 1567 908
Thalassoma bifasciatum Clepticus parrai Halichoeres garnoti Clepticus parrai 238 258 242 868
Halichoeres garnoti Halichoeres garnoti Stegastes partitus Inermia vittata 130 218 222 408
Acanthurus bahianus Caranx ruber Chromis cyaneus Halichoeres garnoti 125 209 180 385
Caranx bartholomaei Chromis cyaneus Clepticus parrai Stegastes partitus 125 187 153 3 77
Scarus iseri Acanthurus coeruleus Thalassoma bifasciatum Thalassoma bifasciatum 124 179 150 223
Halichoeres bivittatus Acanthurus bahianus Sparisoma Sparisoma 124 157 aurofienatum aurofienatum
130 158
Sparisoma Sparisoma aurofienatum Acanthurus bahianus Acanthurus bahianus aurofienatum 128 117 139
102
taranx ruber Stegastes partitus Ocyurus chiysurus Ocyurus chiysurus 98 85 62 122
Kyphosus sectatrid Sparisoma viride Scarus iseri Coiyphopterus incisor 81 56 personatus/hyalinus
9 7 109
and C. ruber comprise 8.91 %.
Planktivorous fishes were important in the shallow reef zone where large foraging
groups of T. fasciatum were observed picking zooplankton and numerous schools in
cleaning stations were observed (Table 16). All top ranking planktivorous species in the
shallow zone comprised 34.02% and included, Thalassoma bifasciatum, ranks number 1,
comprising 20.37% (N=664), Clepticus parrai (N=258), and Chromis cyaneus (N=l87).
Top ranked herbivorous species comprised 16.72% (N=545) and included A. coeruleus,
A. bahianus, S. aurofrenatum, and S. viride. A benthivorous labrid, H. garnoti (N=2 18),
and an omnivorous pomacentrid, P. partitus (N=85), ranked here as well. One schooling
pelagic piscivore, C. ruber, occurred here (N=209) and ranked number 4.
The mid reef was dominated by A. coeruleus, a herbivorous acanthurid,
comprising 41.68% of all individuals in this zone. Up to 400 individuals were observed
during one five minute census, and approximately 2000 were seen during one dive. This
group spawning event was observed during the late afternoon on May 1 1, 1999 at 16:45
CT, which followed full moon 11 days and was between the last quarter and new moon.
The main group swam en masse of approximately 500 individuals 3-9 m off of the
bottom at a depth of 15-2 1 m, back and forth transecting the outer fingers of the forereef
slope, over a distance of approximately 100 m. Sometimes the group was spread out into
two or more groups but would often merge. On several occasions, one group at a time of
I
six to 10 individuals would break off from the main group to undergo spawning by
making quick rushes from approximately 10 m above the bottom to within 5 m of the
surface to release milt and eggs into the water column.
Other herbivores observed at the mid reef include A. bahianus (N=157), and the
scarids, S. aurofrenatum and S. iseri. All of these top ranking herbivores comprised
49.73%. Planktivores were important to the mid reef as well, with the top ranking
species comprising 14.49% of the zone (N=545). The planktivores included C. cyaneus,
C. parrai, i? bifasciatum, and 0. chrysusrus. The benthivore, H. garnoti, ranked number
2 (N=242) and the omnivore, P. partitus, ranked number 3 (N=222).
The deep reef was dominated by planktivorous species; C. cyaneus, a labrid, C.
parrai and an inermid, Inermia vittata as well as T. bifasciatum, 0 . chlysurus; and the
gobiid Coryphopterus personatus/hyalinus. These top ranking species comprised 52.92% t
(N=2638) of the zone. Inermia vittata and C. parrai formed dense schools and moved
swiftly along the forereef. Chromis cyaneus and T. bifasciatum formed dense slow
moving schools above the reef. Ocyurus chrysurus roved across the reefs in numerous
small groups. Covphopterus personatus/hyalinus are very small gobiids that hovered in
small schools just above the substrate. The benthivore, H. garnoti, ranked number 4
(N=385) and the omnivore, P. partitus, ranked number 5 (N=377). The herbivores were
less important in this zone, with S. aurofrenatum (N=158) and A. bahianus (139),
comprising 5.96% of the deep reef zone.
Discussion
As with many studies in the region (Sedberry et al., 1996; Arias Gonzalez, 1998;
~t i f igz Lara and Arias Gonzhlez, 1998), the greatest abundance was found in the deep
depth zone with the most complex habitat structure, particularly the forereef slope areas
at the deep and mid depth zones. Within each depth zone, values increased with
increasing habitat complexity. Within each habitat only the most complex habitat (3)
showed an increase in values with an increase in depth. Comparisons between each
overall value for habitat were significantly different as well as when comparing the deep
reef zone with the other depth zones (Table 3). Biomass values were significantly greater
with increasing habitat complexity but not for depth zone (Table 3,4, Figure 4). These
results coincide with findings on a study of nearby reefs investigating physical
parameters, such as topographical complexity, depth, percentage encrusting coral, and
vertical relief (Nufiez Lara and Arias Gonzhlez, 1998). Their findings revealed that
topographical complexity was the most important contributor explaining fish species
abundance and composition pattern variations for reefs in the central Mexican Caribbean.
The greatest values for species richness and diversity were found at the deep and
shallow reef and especially at habitat 3 (Table 5, 7, Figure 5, 6). Overall values for .
species richness were greatest for the deep reef zone, followed by the shallow reef, mid
reef, and patch reef zone (D>S>M>P). These findings coincide with similar studies on
nearby reefs that had greater species richness and diversity at reef crest and slope habitats
versus back reef habitats. Within each depth zone, species richness increased with
increasing habitat complexity while diversity showed no significant difference between
habitats and very similar values for habitat 2 and 3. Species richness was significantly
different between habitats as well as between all zones except between the deep and
shallow reef zone (Table 6). Values for diversity were higher at the shallow, followed by
the dkep, then mid and patch reef zone (S>D>M>P). Values for species diversity were
significantly different between patch and shallow (P<S) and patch and deep (P<D). The
mid reef zone had lower values for diversity due to the greatest dominance by one
species, A. coeruleus at 41.68% (Table 16). The deep and shallow reefs had high
dominance but high abundance and number of species, so diversity will be high. The
patch had lower abundance and number of species, so diversity was lower. For other
Mexican Caribbean reefs, species richness and diversity were found to be greater where
there was the greatest development and complexity regardless of whether the reefs were
protected or unprotected (Arias Gonzhlez, 1998). The coral reefs found further south
along Quintana Roo, such as Majahual reef, occur over a wider continental shelf, which
favors the growth of large reef structures. The species richness and diversity were greater
at Majahual compared with semi-protected reefs at Boca Paila and Tampalam reefs
located near FWP. Values at Boca Paila were greater than Tampalam as reef development
and complexity was greater. Values at all sites, protected and unprotected in Belize were
much lower than for this study (Table 17).
Table 17. Mean biomass, abundance, and number of species for all species observed at Rancho Pedro Paila (RPP), Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999, with a comparison to similar studies with varying degrees of fishing pressure. P= protected, SP= somewhat protected, UP= unprotected, F=fished, LF=lightly fished, MF=moderately fished, HF=heavily fished.
Protection1 Fishing Mean number
Study SiteIRegion Pressure Biomass (m2) Abundance of species
This study, 4 1.74 80.6 18.29 1999 RPP SPLF (28.74-59.66) (55.64-1 10.78) (13.47-21.33) Van Sant, 1998, pers. 88.63 observ./data RPP SPLF (63.70-128.46) 20.88-22.00
Sedbeny et al., Tres Cocos, Barrier 1996 , Reef, Reef Cut UP/HF
Glovers, Atoll, Forereef Slope UPIMF Lighthouse Reef, Atoll, Forereef ' Slope P/UF
Hol Chan, Barrier Reef, Reef Cut P/UF
Polunin and Hol Chan, Barrier Roberts, 1993 Reef, Reef Cut PAJF 77-340
Saba SP/LF 27
Labrids, acanthurids, and pomacentrids dominate the reef in abundance and
comprise 66.94% in total number (Appendix 111, IV, Table 8). All three families are very
abundant at all four depth zones. No haemulids, epinephelins, mullids, or
chaetodontids/pomacanthids ranked 10 or less in any of the depth zones. No large
schools of haemulids or lutjanids were observed. Only one species of snapper ranked in
the top 10; Ocyurzu chrysurus (yellowtail snapper). 'l'his species is more abundant in the
forereef slope where smaller individuals feed primarily on zooplankton and larger
individuals prey on small planktivorous fishes and benthic crustaceans and worms
(Randall, 1967). This reduction in the abundance of carnivores, such as epinephelins,
lutjanids, and haemulids with an increase in abundance in herbivores, such as acanthurids
and scarids has been shown for fished reefs in this region (Sedbeny et al., 1996; Arias
GonzAlez, 1998). Also with a reduction of piscivores in unprotected reefs in Belize there
has been an increase in certain prey species, such as smaller species of grunts and
parrotfishes (Sedbeny et al., 1996).
Abundance values for this reef (pN=80.60) are similar to values found in similar
studies, using the same technique in Belize (Sedbeny et al., 1996) (Table 17). However
cautibn should be used when making comparisons between different studies even with
the same technique, as in the Belize study, due to observer bias. A study in Saba
(Polunin and Roberts, 1993) used a modified point count technique for 15 minutes and a
radius of 5 m, while tlvs study used a point count for 5 minutes and a radius of 7.5 m.
For a heavily fished reef, Tres Cocos, barrier reef site, at the reef cut, abundance values
were 62.89 and for Glovers Atoll, at the forereef slope, a moderately fished reef, the
value was 118.72. Overall abundance at RPP would fall within the range between a
heavily fished and a moderately fished reef. For the different depth zones, values at RPP
ranged from 55.64 (patch), 72.42 (shallow), 83.56 (mid), and 1 10.78 (deep). Protected
reefs in Belize had greater abundance values, Lighthouse Reef, atoll, forereef slope had
245.33 and Hol Chan Marine Reserve, barrier reef, reef cut had 132.10.
The biomass value for this reef (p.B=41.74 gmA2) is similar to values found in
similar studies, using a similar technique in Belize and at Saba, Netherlands Antilles
(Polunin and Roberts, 1993) (Table 17). Biomass values at RPP range from 28.74-59.66
g n ~ - ~ , and on a lightly fished reef in Saba biomass was 27 gm-', while a protected,
unfished reef at Hol Chan Marine Reserve was 77-340 g n ~ - ~ . Values at RPP would seem
to be closer to a lightly fished reef. Removing the spawning aggregation of Acanthurus
coereleus from the mid reef zone, the biomass ranges from 28.74-48.05 grr~-~, which is
still in the same range.
A comparison of values for fish families reveals that the abundance of
epinephelins at RPP are less than half of that for fished and unfished reefs at Saba
(Roberts, 1995) (Table 18). The haemulids are slightly more abundant at RPP versus
fished and untished reefs of Saba. Fished reefs at Saba and Belize showed higher values
I
for haemulids, especially for smaller species (H. Jlavolineatum), possibly an increase in
abundance due to lack of fished predators. Larger haemulids such as H. sciurus and H.
plumieri exhibited low abundance values for fished reefs in Saba and Belize as well as at
RPP. Scarids were more abundant at RPP versus Saba for fished and unfished reefs.
Smaller scarid species were more abundant in fished versus unfished reefs in Belize
(Sedberry et al., 1996). Lutjanid abundance was slightly higher at RPP than for fished
and unfished reefs at Saba. Values were very low for fished reefs in Belize. With the
exception of 0. chrysurus, abundance for lutjanids was low at RPP. For acanthurids,
abundance and biomass was higher at RPP than at Saba and Belize. In Belize acanthurid
Table 18. Mean biomass and abundance for selected families observed at Rancho Pedro Paila (RPP), Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999, with a comparison to similar studies with varying degrees of fishing pressure. P= protected, SP= somewhat protected, UP= unprotected, F=fished, LF=lightly fished, MF-oderately fished, HF=heavily fished.
Family Study Region Protection1 Biomass Abundance Fishing (gm-*> Pressure
Epinephelinae This study, 1999 RPP SPILF 7.48 1.89 (1.16-34.28) (1.73-2.17)
Van Sant, 1998, pers. RPP SPLF observ.1data 1.03
Roberts, 1995 Saba SPLF 3.0-3.4
Roberts, 1995 Saba PLTF 3.7-5.1
Haemulidae This study, 1999 RPP SPLF 2.87 3.13 (2.20-2.78) (2.21-3.35)
Van Sant, 1998, pers. RPP SPILF observ.1data 2.07
Roberts, 1995 Saba UPIF 1.8-1.9
Roberts, 1995 Saba P L F 1.5-1.7
Scaridae This study, 1999 RPP SPILF 5.57 7.48 (4.30-7.00) (6.17-8.5 1)
Van Sant, 1998, pers. RPP SPLF observ./data 3.56
Roberts, 1995 Saba SPILF 4.7-8.7
t Roberts, 1995 Saba P/UF 3.8-9.5
Lutj anidae This study, 1999 RPP SPLF 3.42 3.22 (1.84-3.56) (2.19-3.87)
Van Sant, 1998, pers. RF'P SPLF observ.1data 3.3 1
Roberts, 1995 Saba SPLF 0.1-0.6
Roberts, 1995 Saba PNF
Acanthuridae This study, 1999 RPP SPILF 12.87 17.25 (1.71-37.02) (4.31-38.14)
Van Sant, 1998, pers. RPP SPILF observ.1data 3.56
Roberts, 1995 Saba SPILF 3.8-5.6
Roberts, 1995 Saba P/UF
abundance was greater at fished versus unfished reefs, and this was suspected due to lack
of fished predators; however, acanthurids were more abundant in unfished areas in Saba.
Greater abundance of acanthurids could be due to several factors at RPP. The spawning
aggregation contributed mostly to this surge in abundance when values are compared to
preliminary data from one year previous. Furthermore, the lack of predation from fishers
and piscivorous fishes could contribute significantly to the high abundance in A.
coeruleus and other herbivores such as smaller scarids. Normally, many species of
herbivores are less abundant in deeper habitats, such as the forereef slope, as in the mid
and deep reef zones at RPP. Foraging areas are usually in shallow reef zones such as the
reef crest, back and patch reef, such as in the patch and shallow reef zone at RPP.
Filamentous algae, sea grasses, and macro-algae are more abundant in these zones. The
deep reef had very low values for abundance of blue tang as would normally be true at
the mid reef. However, an estimated 2000 individuals dominated the mid reef for three
days, May 9- 1 1, 1999. Abundance values for herbivores were much higher at RPP
versis fished and unfished areas at Saba (Table 19). Fished and unfished areas at Saba
had very similar values for abundance. Biomass values for herbivores and piscivores
were different between 1998 and 1999 data at RPP. Biomass was higher for piscivores
during 1999 versus 1998. Biomass of epinephelins, scarids, and acanthurids were higher
during 1999 versus 1998 at RPP, while that for lutjanids and haemulids were very similar
(Table 18).
The reefs in the area of RPP, such as Boca Paila and Tampalam, are considered to
be semi-protected because certain forms of fishing are restricted. However, results of
Table 19. Mean values for biomass and abundance for trophic guilds observed at Rancho Pedro Paila (RPP), Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999, with a comparison to similar studies with varying degrees of fishing pressure. P= protected, SP= somewhat protected, LP= unprotected, F=fished, LF=lightly fished, MF=moderately fished, HF=heavily fished.
Trophic Guild Study Region Protectioll/ Biulllass Abundance Fishing Pressure (gm-7>
Herbivores This study, 1999 RPP SPILF 19.49 25.84 (7.14-42.20) (12.47-44.67)
Van Sant, 1998, RPP SPILF 3.56 pers. observ./data Roberts, 1995 Saba SPIF 8.5-14.3
Saba PAJF 8.5-16.3
Piscivores This study, 1999 RPP SPILF 8.18 3.94 (3.58-25.63) (2.70-4.73)
Van Sant, 1998, RPP SPILF 3.3 1 pers. observ./data Roberts, 1995 Saba SPILF 5.3-5.6
Saba PAJF 5.5-7.3
visual census data taken during May 1999 suggests that abundance and biomass values
for fikhes targeted for fishing are low. Few or no large schools of snappers (Lutjanidae),
grunts (Haemulidae), goatfishes (Mullidae), or large parrotfishes (Scaridae) were
observed. During 1998, large groupers (Epinephelinae) were absent and no individuals
of red grouper, Epinephelus morio, Nassau grouper, E. striatus, red hind, E. guttatus, or
tiger grouper, Mycteroperca tigris were observed. Out of over 250 point counts samples
taken during 1998 and 1999 only one Nassau grouper was observed. Abundance of
smaller scarids was higher (Sparisoma aurofrenatum and Scarus iseri) and acanthurids
were higher than other fished and unfished reefs. However, very little fishing activity or
obvious evidence of destructive fishing practices were observed, ie., abandoned gear,
such as traps, lines, lures, and trawls.
Based on conclusions drawn from the analysis of the data, recommendations for
management are suggested. An approach for reef fish management of the Florida Keys
National Marine Sanctuary has suggested comparing mean length as a biological
indicator for assessing the stock status or health of fishes (Ault et al., 1997), since fishing
selection tends to reduce mean length and thus egg production. Bohnsack and Harper
(1988) state that biomass data are important for studying and modeling ecosystem
structure, trophic relationships, population dynamics, species importance, stock
characteristics, and fisheries exploitation. No-take marine reserves have shown to be
more effective in protecting and restoring reef fish stocks than more traditional measures,
such as quotas, and size and bag limits (Beets and Friedlander, 1999; Bohnsack, 1999).
The American Fisheries Society (AFS) recognizes that reef fishes must be conservatively
managed to avoid rapid overfishing and stock collapse (Coleman et al., 2000). This is
due to the fact that reef fish communities comprise slow-growing, late maturing fishes
such as snappers and groupers. The Society recommends that fishing mortality should be
fi
maintained at or near natural mortality and cautions that an imbalance in the normal sex
ratio may occur rapidly during harvesting of many reef fishes. This could lead to stock
collapse because many reef fishes mature first as female but then become male later in
life; most of the older, larger individuals in the population are male. Conventional
management tools, such as Spawner Biomass Per Recruit may lead to to overly optimistic
conclusions and should be used cautiously. Since many reef fishes form predicatable,
localized, and seasonal spawning aggregations they are vulnerable to overharvesting,
AFS recommends their protection.
This study and sampling design could be a model to quantitatively monitor long-
term trends in community structure, diversity, and stocks as well as investigate factors
that are altering these populations. Also, fishing effort and other anthropogenic effects
need to be assessed and monitored. Fishing activities need to be monitored and more law
enforcement is needed. Spawning areas need to be documented and monitored, as in A.
coeruleus at RPP, and for E. striatus for Southern Quinta Roo (Anguilar-Perera and
Anguilar-Davila, 1996) and protected measures need to be in place to protect these
spawning populations. The spawning aggregation of E. striatus is protected directly from
spears and gillnets since 1993, but hook and line fishing is still allowed as well as gillnets
outside of the aggregation are depleting migrating grouper. Since 1999 the aggregation
of A. coeruleus has not been observed nor has any other spawning events been
documented for this region. These aggregation sites as well as others need to be
protected in a network of reserves to insure spawning and recruitment success.
Recruitment and spawning success needs to be evaluated for protected and unprotected
areas. Some areas have low abundance for certain groups in the absence of fishing, such
#
as grunts, snappers, and groupers (Polunin and Roberts, 1993). It is suspected that
recruitment is low because critical spawning areas or stocks are depleted, or possibly
nursery habitats are impacted in other areas. Target fish populations have been shown to
recover swiftly to protective measures and reductions in fishing pressure (Roberts and
Polunin, 1994; Roberts, 1995). Roberts (1995) has suggested that protection of
vulnerable species is only likely to be successful if networks are established throughout
species ranges to link larval supply and settlement areas. It will be important to continue
monitoring this site, especially with future changes in levels of protection or changes in
neighboring areas such as the E. striatus spawning aggregation site in southern Quintana
Roo. This study represents a valuable dataset for future comparisons following any
changes in reserve status, natural changes, or anthropogenic changes that occur. It is
recommended that sampling should be replicated in the near future at RPP and a
comparison made with the 1999 data. Also during the same time sampling could be done
at other reef sites outside of the reserve, with varying degrees of protection and fishing
intensity, and a comparison could be made with RPP.
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t
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Appendix I. Phylogenetic listing of all fish species (128) censused at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999, including frequency of occurrence, mean number (pN), and rank.
FAMILY/SPECIES RHINCODONTIDAE
Ginglymostoma cirratum MURAENIDAE
Enchelycore carychroa Muraena miliaris
SYNODONTIDAE Synodus intermedius Synodus sp.
HOLOCENTRIDAE Holocentrus adscensionis Holocentrus bullisi Holocentrus marianus Holocentrus rufus Holocentrus sp. Holocentrus vexillarius Myripristis jacobus
AULOSTOMIDAE Aulostomus maculatus
SERRANIDAE Epinephelus cruentatus Epinephelus fulvus Epinephelus guftatus Epipephelus striatus Hypoplectrus guftavarius Hypoplectrus nigricans Hypoplectrus puella Hypoplectrus unicolor Liopropoma rubre Mycteroperca bonaci Mycteroperca tigris Serranus tabacarius Serranus tigrinus
GRAMMIDAE Gramma loreto
PRIACANTH IDAE Priacanthus cruentatus
APOGONIDAE Apogon sp.
BRACHIOSTEGIDAE Malacanthus ~lumieri
Common name Freq occ Carpet sharks Nurse shark 1 Moray eels Chestnut moray 1 Goldentail moray 2 Lizardfishes Sand diver 1 Lizardfish, unidentified 1 Squirrelfishes Squirrelfish 32 Deepwater squirrelfish 3 Longjaw squirrelfish 7 Longspine squirrelfish 19 Squirrelfish, unidentified 2 Dusky squirrelfish 3 Blackbar soldierfish 18 Trumpetfishes Trum petfis h 7 Seabasses and groupers Graysby 57 Coney 74 Red hind 16 Nassau grouper 1 Shy hamlet 9 Black hamlet 3 Barred hamlet 17 Butter hamlet 4 Peppermint bass 3 Black grouper 10 Tiger grouper 2 Tobaccof is h 4 Harlequin bass 47 Basslets Fairy basslet 56 Bigeyes Glasseye snapper 1 Cardinalfishes Cardinalfish, unidentified 1 1-ilefishes Sand tilefish 16
Rank
Appendix I. Continued.
FAMILYISPECIES CARANGIDAE
Common name Jacks Yellow jack Bar jack Scad, unidentified Perm it Snappers Mutton snapper Schoolmaster Gray snapper Dog snapper Mahogany snapper Yellowtail snapper Mojarras Silver jenny Grunts Black margate Porkfish Tomtate Caesar grunt French grunt Spanish grunt Sailors choice White grunt Bluestriped grunt Bonnetmouths Boga Porgies Saucereye porgy Drums Highhat Jacknife fish Spotted drum Reef croaker Goatfishes Yellow goatfish Spotted goatfish Chubs Bermudalyellow chub Butterflyfishes Longsnout butterflyfish Foureye butterflyfish Spotfin butterflyfish Reef butterflyfish Banded butterflyfish
-
Freq occ Rank
Caranx bartholomaei Caranx ruber Decapterus sp. Trachinotus falcatus
LUTJANIDAE Lutjanus analis Lutjanus apodus Lutjanus griseus Lutjanus jocu Lutjanus mahogoni Ocyurus chrysurus
GERREIDAE Gerres cinereus
HAEMULIDAE Anisotremus surinamensis Anisotremus virginicus Haemulon aurolineatum Haemulon carbonarium Haemulon flavolineatum Haemulom macrostomum Haemulon parra Haemulon plumieri Haemulon sciurus
INERMIIDAE lnermia vittata
SPARIDAE Calamus calamus
SClAENlDAE Pareques acuminatus Equetus lanceolatus Equetus punctatus Odontoscion dentex
MULLIDAE Mulloidichthys martinicus Pseudupeneus maculatus
KYPHOSIDAE Kyphosus sectatrix/incisor
CHAETODONTIDAE Chaetodon aculeatus Chaetodon capistratus Chaetodon ocellatus Chaetodon sedentarius Chaetodon striatus
Appendix I. Continued.
FAMILYISPECIES Common name Freq occ POMACANTH IDAE Angelfishes
Holacanthus ciliaris Queen angelfish 4 Holacanthus tricolor Rock beauty 76 Pomacanthus arcustus Gray angelfish 21 Pomacanthus paru French angelfish 9
POMACENTRIDAE Damselfishes Abudefduf saxatilis Sergeant major 8 Chromis cyaneus Blue chromis 93 Chromis insolatus Sunshinefish 4 Chromis multilineatus Brown chromis 2 Microspathodon chrysurus Yellowtail damselfish 75 Stegastes diencaeus Longfin damselfish 34 Stegastes fuscus Dusky damselfish 40 Stegastes leucostictus Beuagregory 43 Stegastes partitus Bicolor damselfish 106 Stegastes planifrons Threespot damselfish 38 Stegastes sp. Damselfish, unidentified 1 Stegastes variabilis Cocoa damselfish 39
ClRRHlTlDAE Hawkfishes Ambylicirrhitus pinos Redspotted hawkfish 12
LABRIDAE Wrasses Bodianus rufus Spanish hogfish 36 Clepticus parrai Creole wrasse 43 Halichoeres bivittatus Slippery dick 58 Halichoeres garnoti Yellowhead wrasse 147 Halichoeres maculipinna Clown wrasse 27 Halichoeres pictus Rain bow wrasse 58 Halichoeres poeyi Blackear wrasse 10 Halichoeres radiatus , Slippery dick 26 Halichoeres sp. Wrasse, unidentified 3 Hemipteronotus splendens Green razorfish 1 Lachnolaimus maximus Hogfish 59 Thalassoma bifasciatum
SCARIDAE Cryptotomus roseus Scarus coelestinus Scarus iseri Scarus guacamaia Scarus sp.
Scarus taeniopterus Scarus vetula Sparisoma atomarium Sparisoma aurofrenatum Sparisoma chrysopterum
Bluehead wrasse Parrotfishes Bluelip parrotfish Midnight parrotfish Striped parrotfish Rainbow parrotfish Scarinae, unidentified juvenile Princess parrotfish Queen parrotfish Greenblotch parrotfish Redband parrotfish Redtail parrotfish
pN Rank
Appendix I. Continued.
FAM ILYISPECI ES Sparisoma rubripinne Sparisoma sp.
Sparisoma radians Sparisoma viride
SPHYRAENIDAE Sphyraena barracuda
OPISTOGNATHIDAE Opistognathus aurifrons
CLlNlDAE Malacoctenus triangulatus
BLENNllDAE Ophioblennius atlanticus
GOBllDAE Coryphopterus glaucofraenum Coryphopterus personatus/ hyalin us Gobiosoma evelynae Gobiosoma oceanops Ptereleotris calliurus
ACANTHURIDAE Acanthurus bahianus Acanthurus chirurgus Acanthurus coeruleus
SCOMBRIDAE Scomberomorus regalis
BALISTIDAE Balistes vetula Melichthyes niger Aluterus scriptus Cantherhines pullus Monacanthus tuckeri
OSTRACllDAE Lactophyrs triqueter
TETRAODONTIDAE Canthigaster rostrata Sphoeroides spengleri
Common name Freq occ Yellowtail parrotfish 27 Sparisominae, unidentified juvenile 2 Bucktooth parrotfish 6 Stoplight parrotfish 83 Barracudas Great barracuda 10 Jawfishes Yellowhead jawfish 3 Clinids Saddled blenny 19 Combtooth blennies Redlip blenny 7 Gobies Bridled blenny 4 Maskedlglass goby
6 Sharknose blenny 1 Neon goby 16 Blue goby 1 Surgeonfishes Ocean surgeonfish 144 Doctorfish 31 Blue tang 118 Mackerels and tunas Cero 9 Triggerfishes Queen triggerfish 4 Black durgon 1 Scrawled filefish 1 Orangespotted filefish 5 Slender filefish 1 Boxfishes Smooth trunkfish 9 Puffers Smooth puffer 59 Bandtail puffer 3
Rank 45
Appendix 11. Species grouped by family (Randall, 1967), including length-weight conversion formulae used for estimating biomass (Bohnsack and Harper, 1982), used in this study at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
FAMILIES LUTJANIDAE (Snap~ers) Ocyurus chrysurus Lutjanus apodus Lutjanus griseus Lutjanus mahogani Lutjanus jocu Lutjanus analis EPINEPHELINAE (Groupers) Epinephelus fulvus Epinephelus striatus Epinephelus cruentatus Mycteroperca bonaci Epinephelus guttatus Mycteroperca tigris HAEMULIDAE (~runts) Haemulon sciurus Haemulon plumieri Haemulon flavolineatum Haemulon parra Anisotremus virginicus Haemulon carbonarium Anisotremus surinamensis Haemulom macrostomum Haemulon aurolineatum ACANTHURIDAE (Sur~eonfishes) A canthurus coeruleus A canthurus bahianus Acanfhurus chiurgus SCARIDAE (Parrot-fishes) Scarus iseri Sparisoma aurofrenatum Sparisoma viride Scarus taeniopterus Sparisoma chrysopterum Sparisoma rubripinne Scarus vetula Sparisoma atomarium Sparisoma radians Scarus coelestinus Scarus guacamaia Scarus sp. Sparisoma sp. Cryptotomus roseus POMACENTRIDAE (Damselfishes) Abudefduf saxatilis Microspathodon chrysurus Stegastes fuscus Stegastes variabilis Chromis cvaneus
Length -Weight Conversion Formulae
Appendix 11. Continued
FAMILIES Chromis insolatus Chromis multilineatus Stegastes sp. Stegastes diencaeus Stegastes partitus Stegastes planifrons Stegastes leucostictus MULL1 DAE (Goatfishes) Mulloidichthys martinicus Pseudupeneus maculatus
Length-Weight Conversion Formulae W=1.282035*1 O-~(XL*I 0) '.lSIY
W=1.282035.1 O-~(XL-I 0) '.1519
W=4.48849 01 O-~(XL*I 0) 2.8956
W=4.48849 -1 O-~(XL-I 0) W=1.282035-1 O'~(XL-I 0) W=5.26987 -1 ~"(xL-1 0) 2.8569 W=3.92916 -1 O'~(XL-I 0) 2.8868
LABRIDAE (Wrasses) Hemipteronotus splendens Thalassoma bifasciatum Bodianus rufus Halichoeres bivittatus Halichoeres garnoti Halichoeres maculipinna Halichoeres pictus Halichoeres poeyi Halichoeres radiatus Halichoeres sp. Lachnolaimus maximus Clepticus parrai *CHAETODONTIDAE (Butterflvfishes) Chaetodon aculeatus Chaetodon capistratus Chaetodon ocellatus Chaetodon sedentarius Chaetodon striatus *POMACANTHIDAE (Angelfishes1 Hola~anthus ciliaris Holacanthus tricolor Pomacanthus arcuatus Pomacanthus paru
* Both Chaetodontidae and Pomacanthidae are merged together during analyses by family.
Appendix 111. Species grouped by trophic guild (Randall, 1967), including length-weight conversion formulae used for estimating biomass (Bohnsack and Harper, 1982), used in this study at Rancho Pedro Paila, Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico, May 1999.
TROPHIC GROUPS - . . . . - - . - - . -
HERBIVORES Acanthuridae Scaridae Kyphosus sectatrix/incisor Ophioblennius atlanticus Coryphopterus glaucofraenum Melichthyes niger Microspathodon chrysurus Stegastes fuscus Stegastes variabilis SESSILE ANIMAL FEEDERS Chaetodontidae Pomacanthidae Abudefduf saxatilis Aluterus scriptus OMNIVORES Malacoctenus triangulatus Cantherhines pullus Stegastes sp. Stegastes diencaeus Stegastes partitus Stegastes planifrons Stegastes leucostictus BENTHlVORES Haemulidae Mullidae Gerres cinereus Balistes vetula Calahus calamus Canthigaster rostrata Enchelycore carychroa Equetus lanceolatus Equetus punctatus Holocentrus adscensionis Holocentrus bullisi Holocentrus marianus Holocentrus rufus Holocentrus sp. Holocentrus vexillarius Hypoplectrus guttavarius Hypoplectrus nigricans Hypoplectrus puella Hypoplectrus unicolor Lactophyrs triqueter Liopropoma rubre Malacanthus plumieri Muraena miliaris Serranus tabacarius Serranus tigrinus Sphoeroides spengleri
Lenath-Weiaht Conversion Formulae
Appendix 111. Continued.
TROPHIC GROUPS Length-Weight Conversion Formulae Trachinotus falcatus W=3.19669 -1 o-"(xL*~ 0) L.YuuS
Bodianus rufus W=1.277615*1 O'~(XL-I 0) 3.0532
Halichoeres bivittatus W=1.54276 -1 O'~(XL*I 0) 3.2017 Halichoeres garnoti W=2.1923 -1 ~"(xL-1 0) 3.3747
Halichoeres rnaculipinna W=5.5924 -1 o - ~ ( x L * ~ 0) 3.6932
Halichoeres pictus W=1.54276 -1 O-~(XL*I o ) ~ . ~ ~ ~ ~ Halichoeres poeyi W=1.54276 -1 o - ~ ( x L * ~ 0) 3.2017
Halichoeres radiatus W=1.19646 -1 ~"(xL-1 0) 3.0382 Halichoeres sp. W=1.54276 *IO"(XL-IO)~~~~~~ Lachnolairnus maxirnus W=3.98382 -1 o '~ (xL*~ 0) 2.8828
PlSClVORES Epinephelinae Lutjanidae Aulostomus maculatus W=5.387658*1 o -~ (xL*~ 0) 2.8657 Synodus interrnedius W=9.954054*1 o"(xL*~ 0) 2.9988
Synodus sp. W=9.954054*1 o"(XL.1 0) 2.9988
Ginglyrnostorna cirraturn W=1.35487 -1 o - ~ ( x L * ~ 0) 2.8918
PELAGIC PlSClVORES Scornberornorus regalis W=8.83487 -1 o"(xL*~ 0) 2.9731 Sphyraena barracuda W=4.10677 - ~ o " ( x L * ~ o ) ~ ~ ~ ~ ~ ~ Caranx bartholornaei W=3.19669 *1 ~"(xL-1 0) Caranx ruber W=2.13599 *1 ~"(xL-1 0) 2.9545
PLANKTIVORES Thalassorna bifasciaturn W=1.29867 01 ~"(xL-1 0) 2.9162
Ambylicirrhitus pinos ~=9.618337*1 O-~(XL-I 0) 3.4266 Apogon sp. W=2.28402 01 O-~(XL-I 0) 2.9434
Chrornis cyaneus W=1.282035*1 o -~ (xL -~ 0) 3.1519
Chrornis insolatus W=1.282035*1 o '~ (xL-~ 0) 3.1519
Chrornis rnultilineatus W=1.282035*1 ~"(xL-1 0) 3.1519 Hern$teronotus splendens W=9.97241 01 ~"(xL-1 0) 2.9995 Clepticus parrai W=1.2218 - ~ o ~ ( x L , ~ o ) ~ . ~ ~ ~ ~ Coryphopterus personatus/hyalinus W=l .I41 6 -1 o-~(xL-I 0) 2.9674
Decapterus sp. W=1.3674 -1 O-~(XL-I 0) 2.9600
Pareques acuminatus W=5.47016 -1 ~"(xL-1 0) 3.2017
Gobiosorna evelynae W=5.8331 -1 ~"(xL-1 0) 3.1370
Gobiosoma oceanops W=5.8331 -1 ~"(xL-1 0) 3.1370
Grarnma loreto W=1.29957 -1 0 5 ( ~ L - l 0) 3.0475
lnerrnia vittata W=1.3674 01 o - ~ ( x L - ~ 0) 2.9600
Pterelotris calliurus W=1.29867 -1 o -~ (xL -~ 0) 2.9162
Monacanthus tuckeri W=1.20226 -1 04(xL-1 0) 2.6178 Myripristis jacobus W=4.78189 -1 04(xL-1 0) 2.4280
Odontoscion dentex W=1.03419 -1 ~"(xL-1 0) 3.0073 Opistognathus aurifrons W=9.52796 -1 ~"(xL-1 0) 2.9895
Priacanthus cruentatus W=2.19432 -1 o - ~ ( x L - ~ 0)
Appendix IV. Frequency of occurrence, mean number, rank abundance (N), and overall rank for all species (128) censused at Rancho Pedro Paila, Sian Ka'an Biosphere Resene, Quintana Roo, Mexico, May 1999. Listed in order by rank abundance in descending order.
Species Acanthurus coeruleus Clepticus parrai Chromis cyaneus Thalassoma bifasciatum Halichoeres garnoti Stegastes partitus Acanthurus bahianus Sparisoma aurofrenatum lnermia vittata Caranx ruber Scarus iseri Ocyurus chrysurus Halichoeres bivittatus Sparisoma viride Halichoeres pictus Gramma loreto Chaetodon capistratus Holacanthus tricolor Pseudupeneus maculatus Caranx bartholomaei Microspathodon chrysurus Epinephelus fulvus Coryl~hopterus personatus/hyalinus Haemulon flavolineatum Kyphosus sectatrix/incisor Scarus taeniopterus Stegastes diencaeus Lachnolaimus maximus Stegastes planifrons Anisotremus virginicus Haemulon plumieri Canthigaster rostrata Stegastes fuscus Decapterus sp. Lutjanus mahogoni Bodianus rufus Haemulon sciurus Stegastes leucostictus Serranus tigrinus Epinephelus cruentatus Chaetodon striatus Halichoeres maculipinna
Freq occ 118 43 93
121 147 106 1 44 153
8 50 83 9 1 58 83 58 56 66 76 72 4
75 74 6
74 13 48 34 59 38 64 71 59 40 2
34 36 48 43 47 57 36 27
Rank Abund 2328 1282 1275 1274 975 685 538 52 1 408 336 28 1 253 25 1 209 197 180 151 140 131 125 122 116 115 1 06 101 99 92 91 88 84 84 83 77 75 73 73 70 69 66 64 62 59
Rank 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 2 1 22 23 24 25 26 27 28 29 30 30 31 32 33 34 34 35 36 37 38 39 40
Appendix IV. Continued.
Species Freq occ Stegastes variabilis 39 Mulloidichthys martinicus 15 Acanthurus chiurgus 31 Haemulon aurolineatum 33 Sparisoma atomarium Sparisoma rubripinne Sparisoma chrysopterum Myripristis jacobus Pomacanthus arcuatus Holocentrus adscensionis Halichoeres radiatus Abudefduf saxatilis Malacoctenus triangulatus Scarus sp. Lutjanus apodus Chromis insolatus Malacanthus plumieri Haemulon carbonarium Scomberomorus regalis Gobiosoma oceanops Holocentrus rufus Hypoplectrus puella Chaetodon ocellatus Epinephelus guttatus Lutjanus griseus Halichoeres poeyi Hemipteronotus splendens Ambylicirrhitus pinos Pomacanthus paru Ophioblennius atlanticus Mycteroperca bonaci 10 Sparisoma radians 6 Hypoplectrus guttavarius 9 Sphyraena barracuda 10 Holocentrus marianus 7 Calamus calamus 9 Lactophyrs triqueter 9 Opistognathus aurifrons 3 Chaetodon aculeatus 8 Halichoeres sp. 3 Serranus tabacarius 4 Aulostomus maculatus 7 Liopropoma rubre 3 Coryphopterus glaucofraenum 4 Holacanthus cikaris 4
Rank Abund 54 51 49 44 42 42 38 38 38 33 32 3 1 31 30 30 29 26 26 25 23 23 2 1 18 18 17 17 15 15 13 12 12 1 1 1 1 10 9 9 9 8 8 7 7 7 5 5 5
Rank 4 1 42 43 44 45 45 46 46 46 47 48 49 49 50 50 5 1 52 52 53 54 54 55 56 56 57 57 58 58 59 60 60 61 6 1 62 63 63 63 64 64 65 65 65 66 66 66
Appendix IV. Continued.
Species Freq occ p N Rank Abund Rank Hypoplectrus unicolor 4 1.25 5 66 Gtherhines pullus Apogon sp. Scarus coelestinus Chaetodon sedentarius Holocentrus bullisi Holocentrus vexillarius Lutjanus jocu Balistes vetula Scarus vetula Haemulon parra Gerres cinereus Holocentrus sp. Mycteroperca tigris Sparisoma sp. Hypoplectrus nigricans Lutjanus analis Sphoeroides spengleri Anisotremus surinamensis Monacanthus tuckeri Pt erelotris calliurus Chromis multilineatus Equetus punctatus Muraena miliaris Pareques acuminatus Aluterus scriptus Cryptofomus roseus Enchelycore carychroa Epinephelus striatus Equetus lanceolatus Ginglymostoma cirratum Gobiosoma evelynae Haemulom macrostomum Melichthyes niger Odontoscion dentex Priacanthus cruentatus Scarus guacamaia Stegastes sp. Synodus intermedius Synodus sp. Trachinotus falcatus