!
Physis
Vol. VIII Fall 2010
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Journal of Marine Science
Physis !
! ! ! ! !
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Journal of Marine Science
CIEE Research Station Bonaire
Tropical Marine Ecology & Conservation Program Vol. VIII Fall 2010
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PHYSIS: !"#$%
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“We must all hang together, or assuredly, we shall all hang separately.”
-Benjamin Franklin
The definition of “Physis” is something not easily translated. Scientifically, Physis is referred to as “nature,” but the word conjures much more emotion than a simple definition can convey—Physis is life. Physis represents the natural course of all activity on Earth, with every species interconnected in the multifarious web of life, and subsequently death. This autonomous cycle has existed for billions of years, yet within a microsecond of geologic time, we as humans have threatened to upset the balance of our planet.
Newton’s third law states “for every action, there is an equal and opposite reaction.” Although originally written to describe motion, this law is applicable to any system with interconnected elements. As we advance through time, mankind is
constantly pushing the boundaries of knowledge, further unraveling the secrets of life.
However, instead of considering the “equal and opposite reactions,” we are focused on the “action.” We have become entranced with ideas of progress, while disregarding the steps we take to get there. Without a doubt, the aspect of nature that we are most careless with is the ocean. Through overfishing, pollution, habitat destruction, bioinvasion, disease, and climate change, humans have forever altered the ocean.
Despite its past resilience, the sea is on the verge of total collapse. We can no longer afford to believe that we transcend the laws of nature.
As humans, we are a part of nature, and therefore it is our responsibility and our duty to protect and conserve our environment, guaranteeing that future
generations of not only humans, but also all of nature are able to enjoy it. Only through understanding the interconnectivity of life and taking responsibility for our actions is this possible. We must learn from our past and study the ocean in order to devise conservation methods that are impervious to human ignorance.
We are a small band of aspiring scientists who are learning how to conserve arguably one of the most precious resources to man: the ocean. For 3 & months 16 undergraduate students came together in order to study the marine realm. Our efforts are focused in Bonaire, but what we are learning has worldwide implications. We have put theory into practice and taken our first steps in the marine conservation direction. This volume of Physis compiles the culmination of our mission here in Bonaire – to learn about the marine environment and how we can protect it to reach a state of Physis.
Ian Kroll
Osha Ann Rudduck Daniel Sternberg
CIEE Bonaire Fall 2010
Photo credits:
Front cover: Michael Hansen Back cover: Paul Patitsas
FOREWORD
The Council on International Educational Exchange (CIEE) is an American non-profit organization with over 100 study abroad programs in 41 countries around the world. Since 1947, CIEE has been guided by its mission:
“To help people gain understanding, acquire knowledge, and develop skills for living in a globally interdependent and culturally diverse world.”
The Tropical Marine Ecology and Conservation program in Bonaire is a one-of-a- kind program that is designed for upper level undergraduates majoring in Biology. The goal of the CIEE Research Station Bonaire is to provide a world-class learning experience in Marine Ecology and Conservation. The field-based science program is designed to prepare students for graduate programs in Marine Science or for jobs in Natural Resource Management and Conservation. Student participants enroll in six courses: Coral Reef Ecology, Marine Ecology Field Research Methods, Advanced Scuba, Tropical Marine Conservation Biology, Independent Research and Cultural & Environmental History of Bonaire. In addition to a full program of study, this program provides dive training that prepares students for certification with the American Academy of Underwater Scientists, a leader in the scientific dive industry.
The student research was conducted within the Bonaire National Marine Park with permission from the park and the Department of Environment and Nature, Bonaire, Dutch Caribbean. The focal point of the research this semester was Lac bay, a beautiful mangrove- lined bay with seagrasses, sandflats and coral reef habitats. The importance of the bay has been recognized at the international level through its designation as a Ramsar site. Students presented their findings in a public forum on the 23
rdand 24
thof November 2010 at the station for the general public.
The proceedings of this journal are the result of each student’s Independent Research project. The advisors for the projects published in this journal were Rita B.J. Peachey, PhD and Maria C. Uyarra, PhD. In addition to faculty advisors, each student had an Assistant Instructor and Interns that were directly involved in logistics, weekly meetings and editing student papers.
Photo credits (left to right): Quinn de la Concepcion, Marissa Paulling, Paul Patitsas, Amanda Brunner
CIEE FACULTY
Rita Peachey, Ph.D
Resident Director, Independent Research Advisor
Cultural & Environmental History of Bonaire Instructor
Caren Eckrich
Dive Safety Officer Instructor for Coral Reef Ecology, Marine Ecology Field Research Methods,
Advanced SCUBA
Maria C. Uyarra, Ph.D
Tropical Marine Conservation Biology Instructor, Independent ResearchAdvisor
Dr. Maria C. Uyarra is the Tropical Marine Conservation Bioloy faculty and Independent Research advisor for CIEE.
She holds a BSc in Biology from the Universidad de Navarra (Spain) and a MSc in Applied Ecology and
Conservation, and a PhD in Biology from the University of East Anglia (UK). Dr. Uyarra is a conservation ecologist with interest in interdisciplinary research. Her research focuses on the interaction between natural and
anthropogenic disturbances and coral reefs. She is also interested in the exploration on the impacts, adaptation and mitigation strategies of human populations to climate change.
Dr. Rita Peachey is the Resident Director in Bonaire.
She received her B.S. in Biology and M.S. in Zoology from the University of South Florida and her Ph.D. in Marine Sciences from the University of South Alabama. Dr. Peachey’s research focuses on ultraviolet radiation and its effects on marine invertebrate larvae and is particularly interested in issues of global change and conservation biology.
Rita teaches Independent Research and Cultural and Environmental History of Bonaire.
Professor Caren Eckrich is the Coral Reef Ecology Faculty and the Dive Safety Officer for CIEE. She holds a B.S. in Wildlife and Fisheries Management from Texas A&M University and a M.S. in Biological Oceanography from the University of Puerto Rico in Mayaguez. Caren is also the instructor for Marine Ecology Field Research Methods and Advanced SCUBA. Caren’s research interests include fish
behavior, seagrass and algal ecology, sea turtle ecology,
and coral disease and she is currently working on a
study of algae that over grow corals in Lac Bay.
CIEE FACULTY
Photo credit (left to right): Marissa Paulling, Michael Hansen, Michael Hansen, Amanda Brunner, Michael Hansen, Beth Tyrie
Amy Wilde
Administrative Assistant
Anouschka van de Ven
Assistant Resident Director
Amy Wilde is the Administrative Assistant for CIEE.
She holds a Bachelor of Science degree in Business Administration, as well as, a Masters of Science in Management Administrative Sciences in Organizational Behavior, from the University of Texas at Dallas. She has worked in call center management for the insurance industry and accounting for long term care while living in
Texas. Amy currently provides accounting and administrative support for staff and students at CIEE.
Anouschka van de Ven is the Assistant Resident Director for CIEE. She is a PADI Dive Instructor and underwater videographer. She assists with Advanced SCUBA and the Cultural and Environmental History of Bonaire courses.
She has a B.A., First Class Honours Degree in
Communications Studies, from the London Metropolitan University and worked in television and advertising in Amsterdam before moving to Bonaire. Anouschka provides administrative support for the research station and is
responsible for the website and public relations. She is also
a volunteer operator at the hyperbaric chamber.
INTERNS
Mary Allen assisted Dr. Rita Peachey with the Independent Research and Cultural Environmental History of Bonaire classes.
Mary is originally from Tulsa, OK and has lived in Texas for the past 10 years. She holds a B.S. in Marine Biology and a M.S. in Marine Resources Management from Texas A&M University.
Alex served as a teaching assistant for Dr.
Maria C. Uyarra for the course Tropical Marine Conservation Biology. He also assisted advisees with their Independent Research projects. Originally from London, he holds a BSc in Freshwater and Marine Biology and an MSc in Applied Marine Science. He has been working in marine conservation for some years and is passionate about tropical marine ecosystem dynamics.
Mary Allen
Sarah Hogan
Alex Leman-Lawrie
Sarah Hogan assisted Dr. Rita Peachey’s advisees with their Independent Research projects. She was also an assistant for the Tropical Marine Conservation Biology course. She has a B.S. in Marine Biology from UCLA and has worked as a
research technician for a variety of
terrestrial and marine studies, including a
long-term coral health survey in Panama.
STUDENTS
AnnaRose Adams
BioResource Research Oregon State University
Ashland, Oregon
Alexandra Gulick
Marine Biology Oregon State University
Halfway, Oregon
Andrew Brendle
Integrative Physiology University of Colorado Boulder
Denver, Colorado
Amanda Brunner
Marine Biology Oregon State University
Kent, Washington
Quinn de la Concepcion
Biology Colorado College
Phoenix, Arizona
Michael Hansen
Global Studies University of Minnesota
Maplewood, Minnesota
Gina Jaber
Elementary Education Juniata College Johnstown, Pennsylvania
Ian Kroll
Biology Vassar College Calabasas, California
STUDENTS
Paul Patitsas
Marine Conservation Juniata College Huntingdon, Pennsylvania
Marissa Paulling
Biology University of Oregon
Fremont, California
Osha Rudduck
Biology DePaul University
Sydney, Australia
Katie Shoultz
Biological Science Arizona State University
Chandler, Arizona
Thomas Stabler
Biology Goucher College Keene, New Hampshire
Daniel Sternberg
Marine Biology University of Oregon
Portland, Oregon
Beth Tyrie
Biology Wofford College Glasgow, Kentucky
Rachael Vietheer
Biology Ursinus College Bethpage, New York
TABLE OF CONTENTS
Map of the Caribbean, including Bonaire and Lac Bay….….………...…..1
Community responses of site-specific fish assemblages to Ramicrusta sp. overgrowth of hydrocoral Millepora complanata
AnnaRose Adams ………2
Effects of the addition of Queen conch (Strombus gigas) shells as a refuge in reef, mangrove, and sand flat communities
Andrew Brendle ……….10
Are differences in Green sea turtle bites in a tropical bay affected by epiphytes on the turtle grass, Thalassia testudinum?
Amanda Brunner ………16
The relationship between the herbivore Diadema antillarum and a red crustose alga, Ramicrusta sp., in a shallow back reef community
Quinn de la Concepcion ………21
TABLE OF CONTENTS
Comparison of foraging tactics and striking
efficiency of adult Egretta tricolor within mangrove and seagrass beds of a tropical coral bay
Alex Gulick ………26
Ontogenetic differences in the use of the mangrove and coral reef habitats by the Rainbow parrotfish, Scarus guacamaia, in Lac Bay, Bonaire
Michael Hansen ………..…31
Phenotypic characteristics of the upside-down jellyfish, Cassiopea xamachana, within the mangrove environments of Lac Bay, Bonaire
Gina Jaber ………...36
Abundance and activity of the upside-down jellyfish, Cassiopea xamachana, a potential bioindicator of mangrove fitness in Lac Bay, Bonaire
Ian Kroll ……….42
TABLE OF CONTENTS
Abundance, density, and distribution of Queen conch (Strombus gigas) in Lac Bay, Bonaire, Dutch Caribbean
Paul Patitsas ………...………49
The feeding association between Queen conch, Strombus gigas, and epibiont assemblages on the turtle grass, Thalassia testudinum
Marissa Paulling ………56
How physical characteristics of the seagrass Thalassia testudinum affect current velocities in coastal ecosystems
Osha Rudduck ………61
Habitat preferences of the commercially important species Great barracuda, Sphyraena barracuda: a case study in Lac Bay, Bonaire
Katie Shoultz ………..65
TABLE OF CONTENTS
Color difference of a red crustose alga (Ramicrusta sp.) overgrowing corals: light sensitivity or response to Millepora complanata metabolites
Thomas Stabler ………...73
Does Lac Bay still lack an important benthic herbivore? Diadema antillarum population density, size distribution, and recruitment rates in Lac Bay, Bonaire, Dutch Caribbean
Dan Sternberg ………...77
Avoidance and preference of specific colors and substrates by the Peacock flounder, Bothus lunatus, through limited adaptive camouflage expression
Beth Tyrie ………...84
Interactions between root-fouling epibionts and the abundance and richness of fish in the mangroves of Lac Bay, Bonaire, Dutch Caribbean
Rachael Vietheer ………90
Photo credits (top to bottom): AnnaRose Adams, Beth Tyrie, Rachael Vietheer, Michael Hansen, Nick Kontonicolas, Michael Hansen, Gina Jaber, Rachael Vietheer, Paul Patitsas, Michael Hansen, Paul Patitsas, Michael Hansen, AnnaRose Adams, Paul Patitsas, Beth Tyrie, Rachael Vietheer
Bonaire
Klein Bonaire
North
5 km Caribbean Sea
Lac Bay
1
!"##$%&'() *+,-"%,+,) ".) ,&'+/,-+0&10) 1,2) 3,,+#4536+,) '") Ramicrusta sp.
"7+*6*"8'2)".)2(9*"0"*35)Millepora complanata
AnnaRose Adams Oregon State University
Corvallis, Oregon adamsan@onid.orst.edu
:4,'*30'
Human and natural disturbances to seascapes can alter benthic community structure, impacting resource
!"!#$!%#$#&'()*+(,#-,.+(&+*/,#0($.".$12(3#&.41/.0#50(51,(!11.6%$!-.1(!+.("7$8.+!%$.(&*(&,.1.(0,!8-.1(%.0!71.(*)(
their dependence upon particular habitats. The presence of a new alga Ramicrusta sp. in Lac Bay, Bonaire has shifted the benthos to a crustosecovered seascape, posing a potential threat to the community structure of its +..)12( 9,#1( 1&7:'( #8".1&#-!&.1( &,.( +.1/*81.( *)( 1#&.41/.0#50( 51,( !11.6%$!-.1( &*( Ramicrusta sp. overgrowth of
&,.(,':+*;*!8(5+.(0*+!$<(Millepora complanata. Visual surveys conducted by snorkel throughout the back reef
!11.11.:(51,(!%78:!80.(!8:(:#".+1#&'(=#&,(+.1/.0&(&*(!(-+!:#.8&(*)(Ramicrusta sp. overgrowth of M. complanata.
>#)).+.80.1(%.&=..8(51,(!%78:!80.(!8:(:#".+1#&'(:#:(8*&(1#-8#50!8&$'(0,!8-.(=#&,(#80+.!1#8-(!$-!$(*".+-+*=&,<(
which is a likely product of little change in coral structural complexity. However, at maximum Ramicrusta sp.
*".+-+*=&,(:.0$#8.1(#8(51,(!%78:!80.(!8:()!6#$'(:#".+1#&'(=.+.(/+.1.8&<(#8:#0!-(&,!&(Ramicrusta sp. could be
!)).0-(+.1*7+0.1()*+(1#&.41/.0#50(51,2(?.17$&1(+.".!$("!+#!%$.(0*6678#&'(+.1/*81.1(%'(1#&.41/.0#50(51,(&,!&(
suggest effects of Ramicrusta sp. overgrowth on these assemblages require further investigation.
;%'*"9$0'&"%
The propensity for an organism to return to the 1!6.( $*0!&#*8( @#2.2( A1#&.( 5:.$#&'BC( 0!8( /$!'( !(
1#-8#50!8&( +*$.( #8( &,.( 17+"#"*+1,#/( *)( &.++.1&+#!$(
!8:( 6!+#8.( *+-!8#161( @3=#&;.+( DEEFC2( ?.6!#8#8-(
within the same locale can secure shelter from /+.'(@G*8.1(DEHIC<()**:(@J7+1!$$(DEKLM(N.+=.#O(!8:(
J!$-.$P.+P.8( QRRKC<( !8:( 8.1-( -+*78:( @S!+".'(
.&( !$2( DEKEM( >*6.#.+( !8:( T*$#8( DEEKC2( U,!&.".+(
%.8.5&(#1(+.0.#".:()+*6(+.6!#8#8-(1#&.41/.0#50<(#&(#1(
%.$#.".:( &,!&( 1#&.( 5:.$#&'( #80+.!1.1( =,.8( &,.( 0*1&1(
to survivorship increase with relocating (Switzer DEEFC2(
In the marine environment, benthic structural complexity can have a substantial impact on +.-7$!-(1#&.41/.0#50(51,(!11.6%$!-.12(3&+70&7+!$$'(
complex habitat, such as corals, provide a greater number of shelter sites for prey species (Hixon
!8:( V..&1( DEEFM( W+#.:$!8:.+( !8:( X!++#1,( DEEHC(
which can decrease predation rates and increase the 17+"#"*+1,#/(*)(1#&.41/.0#50(51,(@V.7P.+1(!8:(G*8.1(
DEEKM( Y$6!8'( QRRIC2( Z8( !::#&#*8<( /+#*+( +.1.!+0,(
has demonstrated that topographic complexity and diversity of benthic structures (corals, gorgonians, .&02C(!+.(/*1#&#".$'(0*++.$!&.:(=#&,(51,(!%78:!80.<(
+#0,8.11<(!8:(0*6/*1#&.(:#".+1#&'(@W+#.:$!8:.+(!8:(
X!++#1,(DEEHM([+!&=#0P.(!8:(3/.#-,&(QRR\C2((
U,.8( !( %.8&,#0( 0*6678#&'( #1( :#1&7+%.:<( #&(
can have cascading effects on the composition of
#&1( +.1#:.8&( 51,( !11.6%$!-.12( J!&7+!$( !8:( ,76!8(
disturbances to seascapes can reduce benthic 0*6/$.]#&'(!8:(:#".+1#&'<(0!71#8-(+.:70&#*81(#8(51,(
/*/7$!&#*81(@Y$"!+.;4W#$#/(.&(!$2(QRREM(X!::!0P(.&(
!$2(QRRE!C2(Z8(^!8*.,.(V!'<(_!,7<(,76!84#8:70.:(
eutrophication consequently produced an algal
dominated seascape of Dictyosphaeria cavernosa
@V!88.+(DEKFM(`!+!-*1(DEH\C2(9,#1(/,!1.41,#)&($.!:(
&*(!(1#-8#50!8&(:.0$#8.(#8(0*+!$(0*".+(@`!+!-*1(DEH\M(
3=*8(.&(!$2(QRRDC(!8:(!$&.+.:(1/.0#.1(!%78:!80.1(
within planktonic and nektonic communities
@V!88.+( DEKFC2( Y( 1#6#$!+( !$-!$4:*6#8!&.:( 1,#)&(
=!1( :*076.8&.:( #8( G!6!#0!( &,+*7-,*7&( &,.( DEHR1(
!8:( DEER1( !1( !( 0*6%#8.:( /+*:70&( *)( *".+51,#8-<(
Hurricane Allen, and the pathogeninduced die off of the herbivorous urchin, Diadema antillarum
@S7-,.1(DEHK<(DEEIC2(9,.(6!11#".($*11(*)(Acropora cervicornis and increased macroalgal cover throughout the area transformed the behavior and
!%78:!80.( *)( 51,( !11.6%$!-.1( @^!7)6!8( DEHFC2(
W*+(.]!6/$.<(!11.6%$!-.1(*)(10!+#:1(:#6#8#1,.:(#8(
size and did not resume normal feeding behavior 78&#$(&,.(&,+..1/*&(:!61.$51,(@Stegastes planifronsC(
+..1&!%$#1,.:(&.++#&*+#!$#&'(@^!7)6!8(DEHFC2(
3#&.41/.0#50( !11.6%$!-.1( !+.( /!+$!+$'(
vulnerable to seascape alterations because of their :./.8:.80.( 7/*8( !( 1/.0#50( $*0!$.( )*+( 17+"#"!$2( Z8(
+.1/*81.( &*( &,.( Z8:*4X!0#50( DEEH( 0*+!$( %$.!0,#8-(
.".8&<( `78:!'( @QRRIC( :.6*81&+!&.:( &,!&( -*%#.1(
which inhabited only one species of coral (i.e.
A1#&.( 1/.0#!$#1&1BC( .]/.+#.80.:( !( -+.!&.+( :.0$#8.( #8(
abundance than gobies which could inhabit multiple 0*+!$1( @#2.2( A1#&.( -.8.+!$#1&1BC2(Y( 1#6#$!+( :.0$#8.( #8(
abundance was documented in Osymonacanthus longirostris,(!(1#&.41/.0#!$#1&(5$.51,<()+*6(&,.(1!6.(
%$.!0,#8-( .".8&( @^*P#&!( !8:( J!P!;*8*( QRRDC2(
Q
Throughout the Caribbean, the decline of Acropora from Hurricane Allen induced a shift in the
!"#$%"%&'()*)()+,)&-*&%!)&."/0)120!&S. planifrons,
$+,()"0$+3& ."/0)120!& 3"(.)+0& -+& ,-("1& !)".0& -*&
Montastrea&45(),!%&)%&"16&7898:.
In Jamaica, a recently described crustose red alga, Ramicrusta textilius, has been documented -;)(3(-<$+3&,-("1&45=)0,!)1&"+.&>"=+.)(0&788?:6&@+&
Lac Bay, located on the Caribbean island of Bonaire,
"+& =+$.)+%$2).& 0'),$)0& -*& Ramicrusta has been found overgrowing 14 species of coral (Eckrich et
"16& $+& '()00:6& @%& $0& ,=(()+%1A& =+B+-<+& <!)%!)(& %!$0&
genus of algae is native or invasive to the Caribbean.
Ramicrusta nanhaiensis, a relative of R. textillius, -($3$+"%)0& *(-/& %!)& 5",$2,& C,)"+D& -')+$+3& %!)&
possibility that the Ramicrusta observed throughout
%!)&E"($##)"+&,-=1.&#)&"+&$+;"0$;)&0'),$)0&45=)0,!)1&
"+.&>"=+.)(0&788?:6 At this present time, no data are available on the extent, causes, and effects of this algal genus overgrowing Caribbean corals.
The ability of crustose algae to compete with coral for space is currently poorly understood.
F"%=("1$0%&.)0,($'%$-+0&#A&G$+,B!&49?8H:&"+.&;"+&.)&
I-)B& 49?J?:& '(-;$.)& );$.)+,)& %!"%& ,(=0%-0)& "13")&
can overgrow corals, inducing coral mortality. The presence of Ramicrusta overgrowth in Jamaica and Bonaire demonstrates a need to understand crustose algaecoral competition and its effects on the trophic communities which rely upon live coral for habitat.
Ramicrusta dominated seascapes could potentially have dire consequences to reef community structure, such as the alterations documented throughout K"/"$,"&"+.&L"+-)!)&M"A&4M"++)(&9?NOP&L"=*/"+&
9?QOP&R"("3-0&9?QSP&I=3!)0&9?QND&9??HP&>%$/0-+&
)%&"16&7889:6
The overall purpose of this study was to examine the impacts of Ramicrusta sp. overgrowth -*& ,-("1& -+& 0$%)T0'),$2,& ())*& 20!& "00)/#1"3)06&
U$;)+& %!"%& 0$%)& 0'),$"1$0%0& ,"+& #)& /-()& ;=1+)("#1)&
%-& !"#$%"%& "1%)("%$-+0& 4L-B$%"& "+.& F"B"V-+-& 7889P&
R=+."A&788HP&5(),!%&)%&"16&7898:D&%!)A&"()&"+&$.)"1&
study organism to investigate the trophic effects of Ramicrusta sp. overgrowth. To examine this ()1"%$-+0!$'D& 0$%)T0'),$2,& ())*& 20!& <)()& -#0)(;).&
over an overgrowth gradient of Ramicrusta sp. on
%!)& !A.(-V-"+& 2()& ,-("1& 4Millepora complanata) in Lac Bay. M. complanata is an important source -*&!"#$%"%&0%(=,%=()&*-(&/=1%$'1)&E"($##)"+&())*&20!&
0'),$)0P&/-0%&+-%"#1A)++$$.0&4F=(0"11&9?NJ:&"+.&
'-/",)+%($.0&4E1"(B)&9?NNP&W"1.+)(&"+.&X-#)(%0-+&
9?Q8:6&M. complanata is a structurally diverse coral 4E1"(B)& 9?NN:D& /"B$+3& $%& ,"'"#1)& -*& !-=0$+3& !$3!&
"#=+."+,)0& "+.& .$;)(0$%$)0& -*& 0$%)T0'),$2,& 20!&
"00)/#1"3)0&4G($).1"+.)(&"+.&5"(($0!&9??Q:6&
This study predicted that increases in Ramicrusta sp. overgrowth on M. complanata will have negative consequences on the abundance and .$;)(0$%A& -*& 0$%)T0'),$2,& ())*& 20!& "00)/#1"3)0& $+&
Lac Bay. Diversity and abundance were examined at the family level over a gradient of Ramicrusta sp. overgrowth of M. complanata. Diversity was measured on three levels: richness, evenness, and composite diversity. The following hypotheses were tested:
H1Y& Z!)& "#=+."+,)& -*& 0$%)T0'),$2,& 20!& <$11&
decrease as the percent Ramicrusta sp. overgrowth increases on M. complanata.
H7Y&Z!)&*"/$1A&($,!+)00&-*&0$%)T0'),$2,&20!&<$11&
decrease as the percent Ramicrusta sp. overgrowth increases on M. complanata.
H3: The family composite diversity of site
0'),$2,& 20!& <$11& .),()"0)& "0& %!)& Ramicrusta sp.
overgrowth increases on M. complanata.
H4Y&Z!)&*"/$1A&);)++)00&-*&0$%)T0'),$2,&20!&<$11&
decrease as the percent Ramicrusta sp. overgrowth increases on M. complanata.
!"#$%&"'()"*+),$#-.+()
/#0+1)(&#$)
[",&M"A&497\&J]&H69Q^FD&JQ\9O]&OO689^W:&$0&1-,"%).&
-+& %!)& <$+.<"(.& 4)"0%)(+:& 0$.)& -*& %!)& 0-=%!)(+&
Caribbean island of Bonaire. The back reef of the
#"A&,-+%"$+0&!$3!&"#=+."+,)0&-*&%!)&2()&,-("1D&M.
complanata, near its reef crest. Recently, Ramicrusta sp. has been observed overgrowing M. complanata
%!(-=3!-=%&%!)&#"A&4_,B($,!&)%&"16&$+&'()00:D&/"B$+3&
it the second area in the Caribbean to document the presence of the crustose alga.
A total of 13 sites were surveyed along the back ())*& -*& [",& M"A& 4G$36& 9:6& `0$+3& U--31)& _"(%!D& 78&
)a=$.$0%"+%&'-$+%0&4"''(-b$/"%)1A&J7&/&"'"(%:&<)()&
'1",).&'"("11)1&497S&/&+-(%!)"0%:&%-&,-+%-=(&-*&%!)&
reef crest. Seven points that would have been in sand patches were removed as a potential study sites.
234$%&,$*#"')+$(&5*
c=($+3& C,%-#)(& 7898D& 0=(;)A0& <)()& ,-+.=,%).& #A&
snorkel once at the 13 sites. During each survey,
3
6&57) 8 Image of 13 study sites (represented as
%!=/#%",B0:&<$%!$+&[",&M"AD&M-+"$()&4U--31)&_"(%!&
7898:6&Z!)&())*&,()0%&$0&.)0$3+"%).&#A&%!)&<!$%)&1$+)&
to the left of the study sites
!
observers followed a compass heading towards the crest at an angle perpendicular to the reef
!"##$%&'(")*+,- ./0o12- 34$5*,6- 7%4+8- *98- 7:*9- wave action was too rough to continue data collection
!"##$%&'(")*+,-.;-(-<$%(-$**<-=$*6)12->"98%(-=%$"+- heads of M. complanata 7*$*- 6"(#+*8- 7'):'9- "- ?- m belt in front of the observer. Randomization was 8*)*$('9*8- @,- "- (%8'A*8- 6,6)*(")'=- 6"(#+'9B- method. Because the entire population of M.
complanata in Lac Bay is unknown it was not possible to determine the sampling interval (k1-@"6*8- upon traditional systematic sampling methods.
Instead, the k value was predetermined by random 94(@*$-B*9*$")'%9-@*)7**9-"-A&*8-$"9B*-
!?- C- k- C/;1- #$'%$- )%- 64$5*,62- D'):'9- *"=:- 64$5*,E- every kth M. complanata head seen within the sample area was sampled.
-D:*9-"9-%@6*$5*$-="(*-7'):'9-/-(-%<-"-6"(#+*- M. complanata, a 1 min period would begin to allow A6:- )%- $*=%5*$- <$%(- "9,- %@6*$5*$- 8'6)4$@"9=*62- During this time period, volumetric dimensions of ):*-=%$"+-!=(1-"98-#*$=*9)-Ramicrusta overgrowth 7*$*- *6)'(")*82- F*$=*9)- "+B"+- %5*$B$%7):- 7"6- categorized as follows:
;-G-H%-I5*$B$%7):-!;J1
/-G-K'9'("+-I5*$B$%7):-!/L?0J1
?-G-F"$)'"+-I5*$B$%7):-!?ML0;J1 .-G-K%8*$")*-I5*$B$%7):-!0/LN0J1 O-G-K"&'(4(-I5*$B$%7):-!NML/;;J1
After the 1 min recovery period, the number of A6:-'9-"66%='")'%9-7'):-M. complanata was recorded
"==%$8'9B- )%- <"('+,2- P:'6- 6)48,- "664(*8- ):")- A6:- 7'):-:'B:-6')*-A8*+'),-)%-M. complanata would live 7'):'9-=+%6*-#$%&'('),-%<-):*-=%$"+2-Q'6:-"66%='")'%9- 7"6-=%96*R4*9)+,-8*A9*8-"6-'98'5'84"+6-7'):'9-;-)%- O;-=(-%<-M. complananta. The number and family
%<-6')*L6#*='A=-A6:-7*$*-$*=%$8*8-"6-):*-9%$):7"$8- facing side of each coral head was visually scanned from left to right.
!"#"$"%"&'()(
Fish abundance vs. percent algal overgrowth Q'6:-"@498"9=*-7"6-8*)*$('9*8-<%$-*"=:-%@6*$5*8-M.
complanata and averaged within algal overgrowth
=")*B%$'*62-S-9%9#"$"(*)$'=-T$46U"+-V-D"++'6-%9*L way analysis was employed to test for differences in (*"9-A6:-"@498"9=*-"(%9B-%5*$B$%7):-=")*B%$'*6-
!W-G-;2;012-
Fish richness vs. percent algal overgrowth
Richness was measured at the family level,
$*#$*6*9)'9B-):*-)%)"+-94(@*$-%<-<"('+'*6-'8*9)'A*8- at each M. complananta head. Average richness was tested across algal overgrowth categories by a one
7",-9%9#"$"(*)$'=-T$46U"+-V-D"++'6-)*6)-!WG;2;012- Composite diversity vs. percent algal overgrowth
Composite diversity was calculated using the 3:"99%9LD'*9*$-X'5*$6'),-Y98*&-!F'*+%4-/ZMM1[-
H\-G-]-#i ln(pi1
where pi is the proportion of individuals observed 7'):'9-*"=:-<"('+,2-K*"9-=%(#%6')*-8'5*$6'),-7"6-- calculated for each algal overgrowth category and
=%(#"$*8- 46'9B- "- 9%9#"$"(*)$'=- T$46U"+- V-D"++'6- )*6)-!WG;2;012-
Fish evenness vs. percent algal overgrowth
Evenness was determined through calculating the 3:"99%9LD'*9*$-^5*99*66-Y98*&-!F'*+%4-/ZMM1[
^-G-_\`-+9!31
7:*$*- _\- '6- ):*- 3:"99%9LD'*9*$- X'5*$6'),- Y98*&-
"98- 3- '6- <"('+,- $'=:9*662- K*"9- *5*99*66- %<- *"=:- algal overgrowth categories was compared by a 9%9#"$"(*)$'=-T$46U"+-V-D"++'6-)*6)-!WG;2;012-
H%$("+'),- 7"6- )*6)*8- <%$- "++- <%4$- 5"$'"@+*6- (abundance, richness, composite diversity, and
*5*99*661- 46'9B- S98*$6%9LX"$+'9B- )*6)62- X")"-
"9"+,6'6-7"6-=%(#+*)*8-46'9B-K'9')"@-!a2/M12-
*+(,&#($
S- )%)"+- %<- /N;- =%$"+- :*"86- %<- M. complanta were observed across the spectrum of algal overgrowth
=")*B%$'*6-$"9B'9B-<$%(-;-)%-/;;J-Ramicrusta sp.
cover. Of all M. complanata %@6*$5*8E-bNJ-*&:'@')*8- some degree of Ramicrusta sp. overgrowth. The highest proportion of M. complanata was found to be ("&'("++,-%5*$B$%79-!NM-V-/;;J1-@,-Ramicrusta 6#2E- 7:'=:- "==%49)*8- <%$- ./J- %<- "++- 6"(#+*62- P:*- lowest number of M. complanata was observed within the no overgrowth category, indicating that /0J-%<-"++-6"(#+*6-7*$*-"@6*9)-%<-Ramicrusta sp.
!Q'B2-?12-
AndersonDarling normality tests showed that abundance, richness, composite diversity,
*5*99*66- 7*$*- 9%)- 9%$("+- !#c;2;;012- P:*$*<%$*E-
4
-)./$0$Comparison of the total number of Millepora complanata observed in each algal overgrowth
=")*B%$,-!9G/N;1
0 10 20 30 40 50 60
0% 1 - 25% 26 - 50% 51 - 75% 76 - 100%
Number of coral heads observed
Algal Overgrowth Category
!"!#$%$&'(%)*+,%-./$0+1+2$00).+('.(.+3'%'+'�"4'5+
("+ *"&#$%'+ 6.7+ $..'&80$9'+ *7$%$*('%).()*.+
(abundance, richness, composite diversity, and ':'!!'..;+ 8'(3''!+ Ramicrusta sp. overgrowth categories.
!"#$%&'()*&)+,%
Across the gradient of algal overgrowth, site
.#'*)6*+ 6.7+ $8-!5$!*'+ )!*%'$.'5+ <%"&+ =+ ("+
>?@+ Ramicrusta *":'%$9'+ AB)9C+ D;C+ E(+ &$F)&$0+
":'%9%"3(7+ A>G+ 1+ H==@;I+ 6.7+ $8-!5$!*'+ 5'*0)!'5+
("+:$0-'.+$##%"F)&$('+("+!"+":'%9%"3(7+A=@;+AB)9C+
D;C+ J)<<'%'!*'.+ )!+ $8-!5$!*'+ 3'%'+ !"(+ 5'('*($80'+
8'(3''!+ $09$0+ ":'%9%"3(7+ *$('9"%)'.+ A,%-./$00+ 1+
2$00).K+LMNHDCMOI+!NH>=I+#N=C=H=;C+P)97+:$%)$8)0)(4+
about mean abundances indicates more sampling is
%'Q-)%'5+("+5'('*(+)<+.)9!)6*$!(+5)<<'%'!*'.+'F).(C+
!"#$%-&."/0%1"+$),##
B).7+ <$&)04+ %)*7!'..+ 5'&"!.(%$('5+ $+ (%'!5+ .)&)0$%+
("+ .)('R.#'*)6*+ 6.7+ $8-!5$!*'C+ S)*7!'..+ )!*%'$.'5+
$*%"..+=+("+>?+@+Ramicrusta overgrowth, followed 84+ $+ 5'*0)!'+ $(+ &$F)&$0+ ":'%9%"3(7+ A>GRH==@;+
AB)9C+ D;C+ P"3':'%I+ (7'+ 5'*0)!'+ .''!+ )!+ %)*7!'..+
is less pronounced, equating to levels similar to
#$%()$0+ ":'%9%"3(7+ AMGR?=@;+ AB)9C+ D;C+ T)9!)6*$!(+
differences in richness were not detected between
":'%9%"3(7+*$('9"%)'.+A,%-./$00+1+2$00).K+LMN+HMCHOI+
!NH>=I+#N=C=HG;C+
!"#$%-&."/0%+2.32#"4,%*"5,1#"40%&)*%,5,)),##
Composite diversity and evenness followed a .)&)0$%+%).'+A=+1+>?@;+$!5+<$00+A>G+1+H==@;+.''!+)!+
%)*7!'..+ AB)9C+ D;C+ U"(7+ *7$%$*('%).()*.+ 5'&"!.(%$('+
5).()!*(+ )!*%'$.'.+ <%"&+ !"+ ":'%9%"3(7+ A=@;+ ("+
&)!)&$0+":'%9%"3(7+AH+1+M?@I+*$('9"%4+H;I+37)*7+
3$.+ !"(+ #%'.'!(+ <"%+ $8-!5$!*'+ $!5+ %)*7!'..+ AB)9C+
D;C+ J'*0)!'.+ )!+ *"&#".)('+ 5):'%.)(4+ $!5+ ':'!!'..+
$(+&$F)&$0+":'%9%"3(7+A>?+1+H==@;+<'00+("+:$0-'.+
.)&)0$%+("+#$%()$0+":'%9%"3(7+AMG+1+?=@;C+V"&#".)('+
5):'%.)(4+5)5+!"(+.)9!)6*$!(04+5)<<'%+8'(3''!+0':'0.+
of Ramicrusta+ .#C+ *":'%$9'+ A,%-./$00+ 1+ 2$00).K+
LMNGCD=I+ !NH>=I+ #N=CH>O;C+ B$&)04+ ':'!!'..+
'F7)8)('5+.)&)0$%+,%-./$00+12$00).+%'.-0(.+ALMN+WCDDI+
!NH>=I+#N=CDGD;C+
Discussion
X7'+ #%'5)*('5+ 5'*%'$.'.+ )!+ 6.7+ $8-!5$!*'+ $!5+
diversity (family richness, evenness, and composite 5):'%.)(4;+3)(7+)!*%'$.)!9+Ramicrusta sp. overgrowth were not evident in this study. High variability was apparent among groups, indicating that further sampling is required to detect if differences exist.
Alternatively, additional factors not assessed in this .(-54+ &$4+ 8'+ )!:"0:'5+ )!+ %'9-0$()!9+ .)('R.#'*)6*+
6.7+ $..'&80$9'.+ .-%%"-!5)!9+ M. complanata. A potential confounding factor, not accounted for in this study, is the role of coral volume and complexity
!"67% 8 Comparisons of Ramicrusta sp. overgrowth
*$('9"%)'.+$!5+&'$!+$;+E8-!5$!*'I+8;+S)*7!'..I+*;+
V"&#".)('+5):'%.)(4I+$!5+5;+Y:'!!'..+AZ+TJ;+"<+6.7+
associated with Millepora complanata in Lac Bay, Bonaire
"!+(7'+!-&8'%+"<+6.7+$!5+6.7+<$&)0)'.C
M. complanata was sampled regardless of size, or complexity. This is problematic because it
*"!<"-!5.+(7'+$8)0)(4+("+*"&#$%'+6.7+$..'&80$9'.+
between coral heads of different physical characteristics. Corals that are larger in volume are .)9!)6*$!(04+&"%'+.(%-*(-%$004+*"�'F+AP"08%""/+'(+
$0C+M==M;C+[!*%'$.'5+.(%-*(-%$0+*"�'F)(4+7$.+8''!+
*"%%'0$('5+3)(7+7)97'%+6.7+$8-!5$!*'+$!5+%)*7!'..+
AB%)'50$!5'%+ $!5+ \$%%).7+ H]]O^+ P"08%""/+ '(+ $0C+
M==M;I+37)*7+).+0)/'04+5-'+("+(7'+)!*%'$.'5+!-&8'%+
"<+7"0'.+$:$)0$80'+<"%+.7'0('%+AP)F"!+$!5+U''(.+H]]D^+
B%)'50$!5'%+$!5+\$%)%).7+H]]O;C+_).-$0+'.()&$('.+"<+
M. complanata volume completed throughout this study demonstrate a positive correlation to both
$8-!5$!*'+ $!5+ %)*7!'..+ AB)9C+ W;I+ 37)*7+ )!5)*$('.+
that size of M. complanata may have )!`-'!*'5+
(7'+ *"&&-!)(4+ *"&#".)()"!+ "<+ .)('R.#'*)6*+ 6.7+
-2.0 -1.0 0.0 1.0 2.0 3.0 4.0 5.0 6.0
0% 1 - 25% 26 - 50% 51 - 75% 76 - 100%
Mean number of individuals ± SD
-1.0 -0.5 0.0 0.5 1.0 1.5 2.0 2.5 3.0
0% 1 - 25% 26 - 50% 51 - 75% 76 - 100%
Mean number of families ± SD
-0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1.0 1.2
0% 1 - 25% 26 - 50% 51 - 75% 76 - 100%
Mean Composite Diversity ± SD
-0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1.0 1.2
0% 1 - 25% 26 - 50% 51 - 75% 76 - 100%
Mean Evenness ± SD
Algal Overgrowth Category
$;
8;
*;
5;
?
assemblages. An additional confounding factor was that M. complanata heads that were not overgrown with Ramicrusta sp. tended to be small in volume, which could potentially account for the
!"#$ %&'()%(*+$ %()$ ,-*.(+//$ "&/+,0+)$ -($ 1.+$ 23$
"0+,4,"#1.$*%1+4",5$67-48$9:8$
The structural complexity did not appear in
;+!)$ "&/+,0%1-"(/$ 1"$ )-<<+,$ &+1#++($ *"!"(-+/$ "<$
similar size at different levels of overgrowth. In ,+/="(/+$1"$*.%(4+/$-($/+%/*%=+>$/-4(-;*%(1$)+*!-(+/$
-($ /-1+?/=+*-;*$ ;/.$ %//+@&!%4+/$ .%0+$ &++($ !%,4+!5$
dependent on the reduction of live coral cover and /1,'*1',%!$*"@=!+A-15$6B,%1*.+11$+1$%!8$C22DE$F,%.%@$
+1$%!8$C22GE$B,%1*.+11$+1$%!$C22HE$I-!/"($+1$%!8$C22HE$
B%))%*J$ +1$ %!8$ C22K%:8$ L"#+0+,>$ -($ ,+/="(/+$ 1"$
MKHC?MKH9$N!$O-("$*",%!$&!+%*.-(4>$I+!!-(41"($%()$
P-*1",$ 6MKHQ:$ <"'()$ 1.%1$ ="@%*+(1,-)$ %&'()%(*+$
was unaffected by declines in live coral cover.
R.-/$ =.+("@+("($ #%/$ %11,-&'1+)$ 1"$ -(/-4(-;*%(1$
differences between the topographic complexity of dead and live coral, implying that pomacentrids will use coral as habitat regardless if it is living
",$ ("1$ 6I+!!-(41"($ %()$ P-*1",$ MKHQ:8$ F-0+($ 1.%1$
pomacentrids were the most abundant family
"&/+,0+)$ 6(SMKK:>$ -1$ -/$ ="//-&!+$ 1.%1$ -(/-4(-;*%(1$
)-<<+,+(*+/$ #+,+$ )+1+*1+)$ -($ ;/.$ %//+@&!%4+$
characteristics because structural complexity was similar between M. complanata with high and low Ramicrusta sp. overgrowth.
It is important to note the curved trend that was
"&/+,0+)$ <",$ %!!$ ;/.$ %//+@&!%4+$ *.%,%*1+,-/1-*/8$
Abundance, richness, composite diversity, and evenness all demonstrated a peak at moderate
"0+,4,"#1.$ 6QMTUQ3:>$ <"!!"#+)$ &5$ %$ )+*!-(+$
%1$ @%A-@%!$ "0+,4,"#1.$ 6UGTM223:$ 67-48$ 9:8$
Aside from the potential effects of differences in structural complexity that were not controlled
<",$ -($ 1.-/$ /1')5E$ <"")$ %0%-!%&-!-15$ @%5$ =!%5$ %()$
%))-1-0+$ ,"!+$ -($ ,+4'!%1-(4$ ;/.$ %//+@&!%4+/8$
Millepora provide habitat for many invertebrate species, most notably barnacles, crustaceans, and
="!5*.%+1+/$ 6V+#-/$ MKKC>$ MKKH>$ C22GE$ F%,*-%$ +1$
%!8$C22H:8$W<$1.+/+$/=+*-+/>$/"@+$%,+$+@&+))+)$-($
the tissue of Millepora spp. 6V+#-/$ MKKC>$ MKKH:8$
High percent cover of Ramicrusta sp. could mean that invertebrate habitat is overgrown, which could decrease their abundance on M. complanata. A
@%X",-15$ "<$ 1.+$ <%@-!-+/$ "&/+,0+)$ 1.,"'4."'1$ 1.-/$
/1')5$ %,+$ =,-@%,-!5$ -(0+,1-0",+/$ 6B%))%*J$ +1$ %!8$
C22K&:8$ R.+$ )+*!-(+$ "<$ -(0+,1+&,%1+$ %&'()%(*+$ "($
M. complanta would mean that less food would be
%0%-!%&!+$<",$/-1+?/=+*-;*$;/.>$<",*-(4$1.+@$1"$;()$
alternate resources on other corals.
In addition to overgrowth by Ramicrusta sp., there was a noteable prevalence of bleaching and overgrowth by turf and macroalgae on M.
complanata. Over half of all M. complanata covered to varying extents in Ramicrusta sp. were
&!+%*.+)$67-48$Q:8$Y($%))-1-"(>$"($%$!%,4+$=",1-"($"<$
M. complanata, Ramicrusta sp. is being overgrown by turf algae and occasionally macroalage Halimeda and Dictyota. I.-!+$&!+%*.-(4$#%/$'(-0+,/%!$%*,"//$
algal overgrowth categories, turf algae and turf algae with bleaching increased with increasing Ramicrusta /=8$*"0+,$67-48$Q:8$Z()+,$<%0",%&!+$*"()-1-"(/>$/'*.$
as high nutrients, turf algae can outcompete crustose
%!4%+$<",$/=%*+$6V-11!+,$%()$V-11!+,$MKHD:8$R.+$1,+()$
between turf algae and bleaching with greater Ramicrusta sp. overgrowth could signify that stress to M. complanata, or the presence of Ramicrusta sp., is giving turf algae some form of a competitive advantage.
R.+$ !%*J$ "<$ /1%1-/1-*%!$ /-4(-;*%(*+$ &+1#++($
/-1+?/=+*-;*$ ;/.$ %//+@&!%4+/$ %()$ Ramicrusta sp.
overgrowth of M. complanta does not negate its
="1+(1-%!$ +*"!"4-*%!$ /-4(-;*%(*+8$ Y(-1-%!$ +0-)+(*+$
for the minimal impact of Ramicrusta sp. on structural complexity further supports that changes -($ *"@=!+A-15$ ,+4'!%1+$ ;/.$ %//+@&!%4+$ ,+/="(/+$
to seascape alterations. Declines observed in
%!!$ ;/.$ %//+@&!%4+$ *.%,%*1+,-/1-*/$ %1$ @%A-@%!$
overgrowth also points towards potential negative community effects at high levels of Ramicrusta sp.
"0+,4,"#1.8$ 7-(%!!5>$ 1.+$ =,+0%!+(*+$ "<$ &!+%*.-(4$
and turf overgrowth on Ramicrusta covered M.
complanata demonstrates a peculiar coralturf
crustose competition dynamic. The combination of these results and additional observations provided by this study gives weight to the potential impacts of Ramicrusta sp. on coral reef ecosystems. The presence of this new alga deserves the attention of 1.+$ /*-+(1-;*$ %()$ @%(%4+@+(1$ *"@@'(-158$ ['*.$
!"#$%& Scatter plot demonstrating the relationship between Millepora complanata$0"!'@+$%()$%:$
\&'()%(*+$%()$&:$7%@-!5$,-*.(+//$"<$;/.$%//"*-
ated with M. complanata
%:
&:
G
is still unknown about Ramicrusta sp. and to fully understand its impact on the ecosystems it alters, further investigation into its ability to overgrow corals and resulting indirect effects on coral reef communities should be undertaken.
!"#$%&'()*(+($,-.
I would like to thank the staff of the CIEE Bonaire
!"#"$%&'()*$*+,-(.,%(/,0+#*+&$/(#122,%*3(4$-1#&%+2*(
%"5+"6(6$#(2%,5+7"7(89(!+*$(:"$&'"9;(4$%<(=+>,-;(
:$1/( :$*+*#$#;( $-7( ?$-( @%,//3( A$*$( &,//"&*+,-( 6$#(
assisted by Thomas Stalber. A special thanks to the
*'"#+#(&,BB+**""(,.(*'+#(#*179(C4$%<(=+>,-;()"/+-$(
="22"//;( $-7( )&,**( ="22"//D( .,%( *'"+%( &,-*+-1$/(
support. This research was supported in part by the Oregon State University ER Jackman Internship )122,%*( :%,0%$B3( E'$-<( 9,1( *,( )E?FG:G( .,%(
granting permission to conduct research within Lac H$9($-7(*'"(H,-$+%"(F$*+,-$/(4$%+-"(:$%<3
References
G/B$-9( I!( CJKKLD( A,"#( +-&%"$#"7( '$8+*$*(
complexity reduce predation and competition in
&,%$/(%"".(M#'($##"B8/$0"#N(O+<,#(PKQRJSTUJVL G/5$%"WUX+/+2( Y;( A1/59( F@;( I+//;( ZG;( [\*]( ?4;
^$*<+-#,-(G!(CJKK_D(X/$**"-+-0(,.([$%+88"$- /0*1. 2( :',*,#( ,.( `( &$*"0,%+"#( ,.( Ramicrusta( #23( ,5"%0%,6*'( &'$%$&*"%+#*+&#R( $D( H/"$&'+-0( CKa( ,5"%0%,6-D;( 8D(
Turf overgrowth of Ramicrusta #23(CLKa(,5"%0%,6-D;(&D(H/"$&'+-0($-7(*1%.(,5"%0%,6*'(,.(Ramicrusta #23(CVTa(
,5"%0%,6-D3( 7D( :"%&"-*$0"( ,.( Millepora complanata coral heads demonstrating these characteristics by algal overgrowth category
coral reefs: regionwide declines in architectural
&,B2/">+*93(:%,&(!(),&(H(JSQR`KP_b`KJT H$--"%( G=( CP_SLD( @$-",'"( H$9;( =$6$++R( 1%8$-
2,//1*+,-( $-7( $( &,%$/( %"".( "&,#9#*"B3( :%,&( Jnd
?-*([,%$/(!"".()9B(JRQVTUSKJ
H"1<"%#( Z);( Z,-"#( I:( CP__SD( =$8+*$*( &,B2/">+*9 B,7+M"#(*'"(+B2$&*(,.(2+#&+5,%"#(,-($(&,%$/(%"".(
M#'(2,21/$*+,-3(O"&,/,0+$(PPLRTKUT_
[/$%<"( !A( CP_SSD( =$8+*$*( 7"#*%1&*+,-( $-7( #2"&+"#
diversity of chaetodontid and pomacentrid M#'"#(-"$%(H+B+-+;(H$'$B$#3(4$%(H+,/(LKRJSSU JV_(
A,B"+"%( 4Y;( [,/+-( :Y( CP__SD( E%,2+&$/( %"".( M#'
S
34 54
"4 )4
!"#$%&%'(#'')*'#+&,%!-(.*/%*.(#%.()*0&*$*.1(
2344(5#)(67&(89-8:;<=>8
?7@)&7A( BC( ?%'*4( 56C( D*#7A*E( F2GH( I&%( ")*!!J1 Crustose, calcareous algal bloom (Ramicrusta
!"1J( ,0*)'),$&%'( !74*)#7+&%&#% corals, gorgonians, a hydrocoral, sponges, and other
#4'#*(&%(K#7(2#EC(2,%#&)*C(L*+A*)4#%.!(M%+&44*!1(
Coral Reefs
N&%7@A( M?( IO:9PJ( 2&,4,'E( ,Q( +A*( )**Q( Q,)R&%'(
,)'#%&!R!( #+( N3%#Q3+&( M+,441( S%-( TA*( M+,44( ,Q(
N3%#Q3+&C(U,)&%'!(&%+,(#(7,)#4()**Q(#%.(+A*()*!34+!1(
TA*( F,E#4( 6,7&*+E( ,Q( K,%.,%C( ""( O>V<OV9(
N)&*.4#%.*)( M5C( D#))&!A( GH( IO::;J( W#U&+#+
7A#)#7+*)&!+&7!( #QQ*7+&%'( /!A( #!!*RU4#'*!( ,%(
#( W#$#&&#%( 7,)#4( )**Q1( G( ?X"( 5#)( 2&,4( ?7,4(
>>P-O<Y9
Z#)7&#( T5C( 5#++A*$!<B#!7,%( WC( N)#%@4&%<G3%&,)(
[( I>99;J( 5#7),Q#3%#( #!!,7&#+*.( $&+A(
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mortality of a coralinhibiting barnacle Megalanus stultus( IH#)$&%J( 3",%( +A*(
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and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata.
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9
Effects of the addition of Queen conch (Strombus gigas) shells as a refuge in
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Andrew Brendle
University of Colorado, Boulder Boulder, Colorado andrew.brendle@colorado.edu
Abstract
Marine organisms depend on complex habitats for protection from predators as well as safe nesting grounds.
There is a positive correlation between habitat complexity and species richness and abundance. The purpose of this study was to compare species richness and abundance of organisms in and around shells in three habitats within Lac Bay, Bonaire. In order to do this, the types of organisms inhabiting the shells were compared and changes in the types of organisms inhabiting the shells were observed over a fourweek period. This study utilized
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(Strombus gigas) shells as a type of natural refuge. Each week, sites in three habitats were observed to see what organisms were found around the shells. Three shells were then haphazardly collected and brought to shore where the shells and the water collected with them were examined. This study found the highest species richness and
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type and size of refuge.
Introduction
Predation rates can impact habitat choice and /;&.4."8%&"2'/%-(%*"&4+'%-&,"+4/*/<%A-&"8%&''(%>/5%
depend on complex habitats for increased protection and survival as well as for safe nesting sites (Wilson et al. 2005; Gratwicke and Speight 2005b). The most important aspects of complexity are the amounts of rugosity, hard substrata and refuge holes (Wilson et al. 2005; Gratwicke and Speight 2005a, 2005b).
Increases in any of these factors can result in higher
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(Wilson et al. 2005; Gratwicke and Speight 2005a, BCCD!E<%F*"88%!-04'0%/7'#4'/%"+0%G;.'+48'%>/5%"&'%
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they are more susceptible to predation (Wilson et al.
2005).
Although increased amounts of complexity, hard substrata and hiding holes should theoretically increase species richness and abundance in an area (Gratwicke and Speight 2005a; 2005b), there are large differences among habitats such as coral reefs, /"+0%1"2/%"+0%*"+,&-.'/<%A-&"8%&''(/%"&'%'92&'*'86%
complex habitats providing a large abundance of different shelters (Wilson et al. 2005; Gratwicke and Speight 2005b). Similarly, mangroves are complex 5"!42"2/)% 7&-.404+,% /5'82'&% (-&% *"+6% 267'/% -(% >/5%
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2005). They are also used as nursery grounds for G;.'+48'% >/5% -(% +;*'&-;/% /7'#4'/% HI-!'&2/-+% "+0%
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characterized by a large open matrix of shelter sites within the prop
roots (Wilson et al. 2005). In contrast to coral reefs
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relatively low complexity and small amounts of hard substrata (Wilson et al. 2005; Gratwicke and Speight 2005a).
Population distribution and reproductive success of an individual within a certain population depends on the ability of an individual to monopolize a resource. The defense of any resource has been predicted to evolve when monopolization 4/% '#-+-*4#)% 25"2% 4/)% 35'+% 25'% !'+'>2/% -(%
monopolization exceed the cost of defense (Baeza et
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refuges is usually more energetically costly, whereas small refuges are less energetically costly. Thus, small refuges are expected to harbor single or small groups of the same species, while large refuges should shelter large groups or aggregations because guarding is more energetically expensive (Baeza et
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habitat requirements, such as particular coral species, various echinoderms, and organisms with hole dimension requirements (Wilson et al. 2005).
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