Caribbean Coral Reef Ecosystems • Nat ional Museum of Natural His tory

CCRE Annual Report 2014

Smithsonian Institution

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Quamino Cays

Lagoon Cays

Channel Cay

Crawl Cay

Tarpum Cay

Coco Plum Cay

Man-o'-War Cay

Twin Cays

Carrie Bow Cay

South Water Cay

Manatee CayCat Cay

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Mosquito CayColumbus Cay

Sandfly CayHutson Cay Cross Cay

Garbutt Cay

Stewart Cay

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Wee Wee Cay

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Elbow Cays

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Bakers RendezvousGladden CaysRendezvous Cay

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CCRE ANNUAL REPORT

2014

National Museum of Natural HistorySmithsonian Marine Station at Fort PierceCaribbean Coral Reef Ecosystems Program

Fort Pierce, FL 34949

December 2014

Table of Contents

CCRE 2014.........................................................................................................................................................................1

Flashbacks...........................................................................................................................................................................2

Acknowledgements ........................................................................................................................................................5

Projects..................................................................................................................................................................................6

Tennenbaum Marine Observatory Network.................................................................................................6

Biodiversity Studies ................................................................................................................................................9

Ecology....................................................................................................................................................................19

Invertebrates............................................................................................................................................................32

Contributions 2014 .......................................................................................................................................................36

Participants 2014 ..........................................................................................................................................................37

Photograph & Art Credits ..........................................................................................................................................39

CCRE 2014 This year we witnessed the beginnings of new and exciting marine science endeavors as well as the continuation of the vital long-term environmental and biological observations that make Carrie Bow Cay a valued site for coral reef studies. Projects related to the Tennenbaum Marine Observatory Network (TMON) began in earnest: first with the Thalassia Experimental Network (p. 11) that lays the foundation for future seagrass ecosystem observations and later the Ocean-BITEMAP project (p. 9) to develop the protocols for globally-replicated fish feeding assays. The newly-launched Carrie Bow BioCode project (p. 14) aims to develop a library of genetic sequences for the entire marine biota in the habitats surrounding the station, and would be the first of its kind in the Atlantic basin. Projects like these are at the heart of the TMON mission to understand coastal marine life and its role in maintaining resilient ecosystems around the world.

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A nurse shark swims near Carrie Bow Cay.

Flashbacks1971 • National Museum of Natural History’s I.G. Macintyre (geology & sedimentology), W. Adey, P. Kier, T. Waller (paleobiology), A. Dahl (botany), A. Antonius (postdoctoral fellow, invertebrate zoology), M. Rice, and K. Ruet- zler (invertebrate zoology) found the program Investigations of Marine Shallow Water Ecosystems (IMSWE).1972 • IMSWE search party identifies Carrie Bow Cay on the barrier reef of Belize as ideally located and affordable site for long-term, collaborative field research on tropical coastal ecosystems • Establishment of principal reference transect across the Belize barrier reef just north of Carrie Bow Cay1974 • Hurricane Fifi destroys laboratory structures, uproots coconut trees, and reduces the surface area of Carrie Bow Cay by about one third, to 0.4 hectare. 1975 • EXXON Corporation provides grant for study of the coral reef ecosystem at Carrie Bow Cay. • Marine and terrestrial post-hurricane surveys.• Establishment of all-manual meteorological station. 1976 • Refinement and calibration of profiles and maps with the aid of vertical aerial photographs taken by Royal Sig- nals Detachment helicopter • Introduction of aerial photography by helium balloon for community mapping • Submersible tide recorder installed at Carrie Bow Cay concrete dock. 1977 • Field trip to Carrie Bow Cay by participants of the Third International Coral Reef Symposium.• Aerial and underwater surveys expanded to cover the entire barrier reef of Belize • Geology team drills first cores to determine reef history • EXXON’s The Lamp publishes article on company-sponsored research at Carrie Bow Cay (“Where seaworms glow.”).1978 • Hurricane Greta destroys Carrie Bow Cay field station. 1979 • Post-hurricane survey and rebuilding of laboratory with several improvements • Count of participating scien- tists and of published scientific contributions both pass the 50 mark; 23 scientific institutions are now collabora- ting with NMNH. 1980 • EXXON Corporation funds new initiative: comprehensive study of a western Atlantic mangrove swamp ecosys- tem, now known as SWAMP (Smithsonian Western Atlantic Mangrove Program) • Mapping of Twin Cays, principal site of SWAMP, by aerial photography and ground truthing.1981 • Initiation of Art in a SWAMP project where scientific illustrators and scientists collaborate in analysis and picto- rial rendition of mangrove communities in time and space • Employment of H. Edgerton underwater time-lapse camera with strobe light (on loan from the inventor) to record day-night activity in benthic communities • Vibracoring at Twin Cays to determine internal structure and development.1982 • Publication of The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, 1: Structure and Communities. Smithsonian Institution Press (K. Ruetzler & I.G. Macintyre, eds.). 1983 • New weather protected and enlarged seawater system for laboratory experiments installed on Carrie Bow Cay • Series of extremely low tides at noon time were observed to have catastrophic effects on reef and mangrove organisms. 1984 • First automated weather station installed at Twin Cays • Cooperation with Belize Government identifying coastal marine areas suitable for natural resource conservation • Busiest year since program start: 8 months con- tinuing laboratory operation for 45 research staff. 1985 • First year of operation of Caribbean Coral Reef Ecosystems (CCRE), a new program of the National Museum of Natural History. It replaces the old IMSWE project and supplements the ongoing SWAMP program which is supported by a renewed annual grant by the EXXON Corporation. 1986 • Renovations on Carrie Bow Cay to accommodate dry-laboratory space, added living quarters, and boat, diving, and laboratory equipment • Mangrove vegetation map for Twin Cays completed • Published scientific contribu- tions pass the number 200.1987 • Record visitation of Carrie Bow laboratory, 120 total: 90 scientists and assistants; others dignitaries, including the Prime Minister of Belize, Smithsonian administrators, and media people working on documentaries and news-related productions • Continued facility renovation, including addition of solar photovoltaic system, large seawater tank, two fiberglass whalers, fluorescence microscope, and time-lapse video recorder with underwater camcorder.

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1988 • Mangrove workshop for 37 EXXON-SWAMP scientists at Solomons, Maryland, entitled A Mangrove Ecosystem: Twin Cays, Belize. 1989 • Science as Art exhibit at the Smithsonian’s S. Dillon Ripley Center displays scientifically important and aesthetically pleasing products from SWAMP mangrove research, such as community drawings, paintings, photo- graphs, and sculpture-like epoxy casts of soft-bottom animal burrows • Vandalized and malfunctioning weather station reconditioned and relocated to the Carrie Bow field laboratory • Increasing problems with anthropogenic stresses at research sites, such as heavy tourist visitation, garbage dumping, and clear-cutting mangrove trees. 1990 • CCRE-SWAMP program represented at first Caribbean Coastal Marine Productivity workshop, Jamaica, CARICOMP is a program for Caribbean-wide monitoring of environmental quality in reefs, mangroves, and seagrass meadows. 1991 • Belize Forestry Department helps stopping disturbances to SWAMP research sites. Belize Department of Natural Resources reviews legislation with intention of declaring Carrie Bow Cay - Twin Cays area protected research site • CCRE-SWAMP program staff participates in developing Belize Tropical Forestry Action Plan and helps designing Institute for Ecology to be based in Belmopan. 1992 • CCRE-SWAMP researchers produce video documentary on mangrove swamp biology • Unprecedented, severe problem with hydrozoan stings to snorkelers and divers in the Carrie Bow area traced to microscopic siphonophorans • CCRE-SWAMP staff and Belize Fisheries Department and Agriculture representatives conduct first workshop for Belize high-school teachers entitled Mangrove Conservation through Education • CCRE-SWAMP lecture series started in Belize City, co-hosted by Belize Audubon Society • CCRE officially joins the CARICOMP network and initiates monitoring program.1993 • Belize Ministry of Natural Resources grants rights to Twin Cays for mangrove research • Launching of new 8 m (25 ft) research vessel Physalia, funded by a grant from the U. S. National Science Foundation, extends research radius over most of central and southern Belize • Ivan Goodbody pioneers surveys of Pelican Cays, a tunicate heaven at SSW of Carrie Bow.1994 • Start of collaborative surveys and experimental projects in the Pelican Cays • Pelican Cays workshop, co-hosted by Candy Feller (SERC), at Edgewater, Maryland.1995 • Finalized lease with the Villanuevas of Placentia to southern portion of Northeast Cay, Pelican group, to establish a field base for future studies • Malcolm Spaulding develops plans for new integrated environmental sensing system with radio- telemetry link to the University of Rhode Island’s COASTMAP network. 1996 • Installation by Tom Opishinski of self-contained Endeco-YSI-Campbell monitoring station of meteorological and oceanographic parameters and hookup to Internet • Visit of field party from 8th International Coral Reef Symposium, Panamá. 1997 • Celebration of the 25th birthday of the Carrie Bow Marine Field Station • New U. S. National Science Foundation grant allows purchase of a second 8-m (25 ft) boat to back up the heavily used Physalia (under construction) • International team of seven expert systematists conducts workshop at Carrie Bow Cay to quantify the unusually high sponge diversity of the Pelican Cays • Number 500 reached in CCRE scientific contributions • Carrie Bow Field Station, including laboratories, weather station, kitchen, and living quarters is consumed by an accidental electrical fire which was apparently sparked by a short in the wiring and aided by dry, termite-riddled lumber and strong northerly winds. Luckily, no-one was hurt. 1998 • Island clean-up and design for new field station completed. Construction work initiated but delayed by flooding and coastal erosion from hurricane Mitch • Completed editorial work on CD-ROM containing over 100 represen- tative CCRE scientific papers that resulted from research at Carrie Bow Cay • Cosponsored Smithsonian (STRI) exhibit Our Reefs –Caribbean Connections in Belize City. Contributed large poster describing 25 years of CCRE coral reef research in Belize • Serious coral bleaching and die-off on reefs off Carrie Bow and Pelican Cays observed, partly caused by hurricane Mitch. 1999 • Rededication ceremony for the new Carrie Bow Marine Field Station, in August • BBC team (Bristol, UK) films segments for its Blue Planet TV series, including (with E. Duffy) eusocial shrimps living in sponges. 2000 • Publication of Natural History of Pelican Cays, Belize, in Atoll Research Bulletin (Macintyre & Ruetzler, eds, 2000) • Replacement of environmental monitoring station lost in the 1997 fire • Initiation of Twin Cays Biocomplexity Study funded by an NSF grant (to I. Feller & colleagues).2001 • Completion of 3-room cottage over the eastern shore of Carrie Bow Cay • Hurricanes Michelle and Iris (October) barely miss Carrie Bow Cay, causing some damage to buildings and heavy beach erosion and devastate

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(Iris, in particular) large areas in southern Belize • Signing of MoU with Belize Fisheries Department officially acknowledging the Carrie Bow Marine Field Station as a nationally recognized laboratory • Publication of Golden (50-year anniversary) issue of Atoll Research Bulletin recognizing prominent coral reef scientists through their autobiographies, several of them participants in the CCRE Program.2002 • Founding of the Smithsonian Marine Science Network (MSN), incorporating the CCRE Program and the Carrie Bow Marine Field Station • Number 600 reached of CCRE scientific contributions • Ranger Station established on southeast Twin Cays by Belize Fisheries Department to oversee South Water Cay Marine Reserve.2003 • Cristián Samper, recently appointed director of the Smithsonian’s Natural History Museum, visits the Carrie Bow station in July, dives on the barrier reef, and snorkels in mangroves habitats • Hurricane Claudette threatens Carrie Bow (July) and necessitates temporary evacuation • Smithsonian Secretary Larry Small visits the Carrie Bow lab in December and dives on the reefs • Twin Cays Mangrove Biodiversity Conference is held at Ft. Pierce, Florida (December), convened by Klaus Ruetzler, Ilka Feller, and Ian Macintyre, and cosponsored by Valerie Paul of the Smithsonian Marine Station at Ft. Pierce.2004 • CCRE Postdoctoral Fellowship established • Hurricane Ivan causes substantial coastal erosion of Carrie Bow Cay • Atoll Research Bulletin volume dedicated to Twin Cays Mangrove Biodiversity goes to press • Number 700 reached of CCRE scientific contributions • Carla Dietrich takes over from Michelle Nestlerode as CCRE research assistant • Addendum to MoU with Belize Fisheries Department signed, clarifying intellectual property rights and issues of bioprospecting sponges in particular • CCRE Program Administrator Marsha Sitnik (recently, administrative advisor) retires.2005 • A total of 13 hurricanes formed this season. Three category five hurricanes (Katrina, Rita and Wilma) caused substantial coastal erosion and damage to the Carrie Bow facilities. The record number of 25 named storms in the Caribbean broke the previous record (from 1933) of 21 named storms • An external scientific review of the CCRE Program was conducted and resulted in a strong endorsement of the program’s mission and accom- plishments • Over 50 new CCRE scientific contributions were published.2006 • The first Belize National Marine Science Symposium, cosponsored by Belize Fisheries and Forestry departments and the Hugh Parkey Foundation, took place and CCRE was represented with 4 talks and 8 posters, including a review of 35 years of Smithsonian Marine Science in Belize • CCRE hosted the U. S. Ambassador and 35 Embas- sy staff for a picnic, including a tour of the Carrie Bow lab facilities • More than 130 Smithsonian Associates, North Carolina teachers, and members of the Sierra Club visited Carrie Bow for guided tours of facilties and ongo- ing projects • A film crew for a Discovery channel in The Netherlands worked at Carrie Bow to document Gordon Hendler’s work on newly discovered brittle-star light-sensing organs • The CCRE program and the Carrie Bow Marine Field Station, along with all other Smithsonian marine programs and facilities, took part in an external re- view ordered by the Smithsonian Undersecretary for Science; The efficiency and scientific productivity of the program and its field station received excellent marks.2007 • Hurricane Dean strikes Northern Belize and Yucatan, Mexico (August), Felix passed over Honduras south of Belize (September); both cause major beach erosion at Carrie Bow Cay but no damage to buildings. 2008 • The Belize Minister of Natural Resources and his staff visit our facilities and tour the Pelican Cays to view dam- age caused by mangrove clear-cutting in this part of the Southwater Cay Marine Reserve. 2009 • Ilka “Candy” Feller was again offered use of Light Hawk, a volunteer pilot-based organization at Lander, WY, to observe and photograph environmental damage to mangrove coast and cays. •Pro- ceedings of the first Smithsonian Marine Science Symposium highlight CCRE’s diverse contributions to knowledge of the biology and geology of the Mesoamerican Barrier Reef, Belize •Mike Carpen- ter retired after 25 years of service as CCRE Operations Manager and will build a new home in the woods of Georgia •Klaus Ruetzler resigned as CCRE Director after 25 years in this position (and a total 37 years as leader of the IMSWE, SWAMP, and CCRE programs). He will be followed by Valerie Paul of SMSFP.2010 • Director Valerie Paul and new staff at Fort Pierce assume responsibility for CCRE • Michael Carpenter, Zach Foltz, and Woody Lee spend three weeks on Carrie Bow Cay, for training and transition • A new CCRE website is launched: www.ccre.si.edu • U.S. Ambassador Vinai Thummalapally and five others from the U.S. Embassy in Belize visit Carrie Bow Cay on August 30, 2010 • Belize Fisheries establishes the South Water Caye Marine Reserve Conservation Zone, a no-take zone encompassing the area around Carrie Bow Cay (www.swcmr.org).

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2011 • Carrie Bow Cay recieved major infractructure improvements, including 10 kw diesel generator, improved photovoltaic system, air compressor for SCUBA, and composting toilets • CCRE contibutions list reached #900 • Randi Rotjan, Peter Gawne, Jay Dimond, Scott Jones, and Zach Foltz initiated 24 permanent reef transects, establishing a long-term reef monitoring program to assess the effects of the SCWMR no-take zone • CCRE director, Valerie Paul, along with Raphael Ritson-Williams, Scott Jones, and Nicole Fogarty receive an award from the Smithsonian Endowment Program for “Population dynamics of threatened Caribbean acroporid corals at Carrie Bow Cay, Belize.” • Tropical Storm Harvey forces evacuation and makes landfall in Belize; no damage was caused to the station.2012 • The CCRE program celebrated its 40-year anniversary since Drs. Klaus Rutlzer and Arnfried Antonious

happened upon Carrie Bow Cay and secured an agreement to use it for field operations • Hurricane Ernesto tracked near Belize in early August, theatening to force evacuation of the island; the storm tracked north and caused no damage to the island. • Sherry Reed assumed responsibilities as Dive Safety Officer for the CCRE program • Over 30% of dives logged at the Smithsonian were completed in the waters surrounding Carrie Bow Cay, a total of 875 dives.

2013 • In October 2012, the CCRE program welcomed members of the Summit Foundation and Wildlife Conservation Society (WCS) to Carrie Bow Cay for a tour of the facilities and a visit to the Acropora-dominated reef adjacent to the island. Participants included Summit Foundation Chairman Roger Sant and WCS president Cristián Samper. • Several photographers and videographers from National Geographic, the BBC, and Public Television used the island as a venue for their work • On-site tours of CBC continued to reach a large public audience, with 1556 people touring the lab in FY2013. • The Smithsonian Institution launched the Tennenbaum Marine Observatory Network, a world wide network of coastal ecological field sites, and Carrie Bow Cay was named one of the first five participat-ing sites.

2014 • Smithsonian Secretary Wayne Clough visited the station with newly-appointed Tennenbaum Marine Observatory Network (TMON) director Emmet Duffy and CCRE director, Valerie Paul • The first experiments under the aus-pices of MarineGEO and the TMON were launched, starting with the Thalassia Experimental Network (a multi-site seagrass experiment, see p. 11) and later Ocean BITEMAP (fish feeding assays, see p. 9) • Over 100 individuals visited Carrie Bow Cay, and nearly 1600 individuals participated in station tours. • CCRE scientific contributions exceed 950 papers.

Acknowledgements

Our research is hosted by the Belize Fisheries Department and we thank Ms. Beverly Wade and Mr. James Azueta and staff for collaboration and issuing permits. The owners and dedicated staff of Pelican Beach Resort in Dangriga provided logistical support for our fieldwork.; Earl David and his staff provided boat transportation, as well as invaluable advice and support. Numerous volunteer managers helped run the field station and assisted in research activities; we greatly ap-preciate their many efforts: Reg Horning, Craig Sherwood, Greg & Joann Dramer, Gary Peresta, Jim Taylor, Tanya Ruetzler, Joel & Linda Moore, Jerry & Sandy Alanko, Ed James, Keith & Shirley Parsons, Nicky & Marilyn Haren, and Daniel Gouge. In Fort Pierce, we sincerely thank Joan Kaminski for administrative advice and assistance with many fund management tasks. CCRE Dive Officer Sherry Reed was very helpful in assisting dive operations at Carrie Bow. Many thanks to Laura Diederick for her editorial eye, sharing her expertise in science communication, and lending valuable advice. In Washington, Klaus Ruetzler and Mike Carpenter are always willing to share wisdom stemming from their many years of experience in Belize. A number of people at NMNH are always willing to answer questions: Charmone Williams, Marty Joynt, Mike McCarthy, Carol Youmans, and JoAnna Mullins among many others. We also thank the Smithsonian offices of the Undersecretary for Science and the Director of the National Museum of Natural History for continued support. The CCRE program is supported by Federal funding complemented by the Hunterdon Oceanographic Re-search Fund.

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MarineGEO & The Tennenbaum Marine Observatories Network

On the Road in Belize: The Secretary's Travel Journal

Secretary Wayne Clough

The following is an excerpt from Secretary Clough's travel journal blog entry following his visit to Carrie Bow Cay. The original story can be found here: http://www.e-torch.org/2014/05/on-the-road-in-belize-i-the-secretarys-trav-el-journal/

What is the connection between Mauna Kea in Hawaii; the South Pole; Mount Hopkins in Arizona; Saint Lawrence Island in the Bering Strait; the Las Campanas Observatory in Chile; Mpala, Kenya; tiny Worland, Wyoming; the Sa-cred Valley in Peru; Port Au Prince, Haiti; and Barro Colo-rado Island in Panama? The obvious answer is the Smith-sonian−these are just a handful of the many places where scholars from the Smithsonian do field research that asks and answers fundamental questions about our planet. The less obvious answer is me. These are all places I have vis-ited as Secretary to learn more about how the Smithsonian

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really works. But there is one last site that I need to visit in my last year as Secretary because of the role it will play in in a project I helped launch.

Carrie Bow Cay appears as a dot on a map off the southern coast of Belize, and is the only place where the Smith-sonian has year-round physical facilities that I have not visited as Secretary. Although Carrie Bow Cay itself may be small, it is surrounded by one of the largest, most ac-cessible, richest and least disturbed marine habitats in the world, a resource for our ocean scientists and hundreds of others from the United States and around the world. It will become even more important since it is slated to be one of the first five sites for our Tennenbaum Marine Observato-ries Network (TMON), a new form of global ocean obser-vation platform that will allow scientists and policy mak-ers to see with clarity the changes occurring in the ocean and how this happens over time. The Network will start with Smithsonian-controlled locations, including Carrie Bow Cay. Carrie Bow Cay is an ideal site for TMON be-cause of its natural setting and because researchers work-ing there over time have already established baselines as to species diversity and the overall marine environment. As the project moves forward, we will be working with partners such as the National Oceanic and Atmospheric Administration, Woods Hole Oceanographic Institution, universities and cooperating countries to develop other locations for the Network. The agenda for our brief trip is packed in order for me to see as much of the science and preliminary TMON work going on at Carrie Bow

Cay as possible. I am fortunate to be joined by Emmett Duffy, director of TMON, and Valerie Paul, director of Carrie Bow Cay and our Fort Pierce Marine Sta-tion in Florida as well as a key participant in TMON. They can provide valuable insights into the project’s future plans and they are bringing along several post-doctoral students who are already working on different investiga-tions that will contribute to the observations.

Just a spit of land, Carrie Bow Cay is roughly 350 feet long by 120 feet wide and is graced by 60 or so coconut palms. You can see other islands nearby, some of which have been developed as

Figure 1- From left, Justin Campbell, Valerie Paul, Craig Sherwood, Caitie Kuempel, Wayne Clough, Martha Nichols, Zach Foltz, Solomon Chak, and Emmett Duffy.

low-key resorts. The developments presage change in this previously undisturbed area; our observational platform here will be crucial in documenting its impact.

The waters of the Caribbean are an artist’s palette of col-ors−green-blues dappled by cloud shadows are interrupted by patches of bright azure blue signaling the presence of shallow coral reefs. The water is in some places so shallow the reefs rise just above its surface, but less than a half mile out toward the ocean they drop precipitously to over 100 feet in depth. At Carrie Bow, the ocean-side reef, which is part of the great Mesoamerican Barrier Reef, rises to the surface about 200 yards offshore, creating a breakwater. The shallow calm waters behind it are patrolled by the pel-icans and several small lemon sharks looking for snacks. This setting with its readily accessible shallow reefs, deep reefs, mangroves and sea grasses attracts both scientists and tourists alike.

We will begin our trip by visiting a shallow reef site where Emmett and his graduate student Solomon Chak can find specimens of snapping shrimp, a remarkable tiny shrimp

that has a large claw that can be rapidly closed to make a very audible snap. These shrimp are social creatures like ants or wolves in that they create family groups that work together to survive. They are found in colonies living in sponges. Very unusually for invertebrates, shrimp pairs raise families together and are possibly mated for life.

After being fueled by a hearty breakfasts, we load our gear on the boat in order to go out to a reef that Emmett had ex-plored years before that will likely have snapping shrimp populations. In addition to Val, Emmett, and Solomon, we are joined by Johnny Gibbons (Smithsonian press officer for science), Zach Foltz (CBC station manager), Caitie Kuempel (STRI), and Justin Campbell (SI post-doc) (Fig-ure 1). Emmett checks several sites before finding the snapping shrimp he seeks. Several of us put on snorkeling gear while others don SCUBA equipment in order to go deeper and collect specimens.

Below the surface a forest of staghorn coral appears, some areas lightly populated and others forming large colonies that create hotels for the tropical fish that appear in abun-

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Figure 2- Secretary Clough takes a closer look a snapping shrimp under Solomon’s supervision.

dance. As soon as I get my head underwater I hear a slight snapping noise. The tiny snapping shrimp below are let-ting us know they know we are here. It is hard to believe that such a small organisms could possibly make such loudnoises, but it happens because there are so many of them.

Fish are everywhere, some flashing by in schools, others guarding holes and crevices in the coral where they make their homes. They are spectacularly diverse in coloring, shape and size and I remember that some species vary in color and shape depending on age and sex. I realize that it is going to be a challenge to name them all, but at Carrie Bow I can call on some of the world’s experts to help me. In total, I compiled a list of the 43 species I saw—a small percentage of the species that live in the seas around Car-rie Bow. Sadly, Val tells me that in spite of seeming vital-ity of marine life, the reefs are only moderately healthy as they are declining due to the effects of ocean warming and acidification, contamination of the water by humans, and for other reasons not fully understood yet.

Back on shore I spend some time with Justin, Caitie and

Solomon talking about their research interests and plans for future experiments. Their enthusiasm reminds me of my time as a doctoral student at UC Berkeley and the excitement that comes in the search for new knowledge. Solomon caps our discussions off when he offers us an up-close-and-personal encounter with one of the snapping shrimp specimens he collected earlier in the day. Tiny when viewed by eye, under the microscope these shrimp look formidable waving their big claw and unintimidated by any threat we giants might make (Figure 2). So it is that from big sharks and rays to tiny crustaceans, we have in one day experienced the wonders of our ocean, courtesy of a dot of land called Carrie Bow Cay (Figure 3).

It has been a short but entirely effective visit as I have been able to develop an appreciation for the importance of Carrie Bow Cay as a resource for Smithsonian research-ers and hundreds of other colleagues who have visited. I am also more than ever encouraged by the Tennebaum Marine Observatory Network concept and its value to our understanding of our ocean that is facing ever increasing pressures from climate change, development and global marine traffic.

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Figure 3- Secretary Clough takes a look at what's beneath the waves near Carrie Bow Cay.

TMON Field Expedition: Ocean BITEMAP

Scott Jones, Emmett Duffy, Justin Campbell, Zach Foltz, Michelle Weber, Michael Goodison, Angelo Sparado

By creating the first global network of biological observa-tories for the ocean, the Smithsonian has set out to uncover vital information about the state and functions of marine ecosystems on a scale never before attempted. With a core network of research stations throughout the Atlantic al-ready in place, the newly-established Tennenbaum Marine Observatory Network (TMON), led by Dr. Emmett Duffy, is uniquely positioned to ask and answer important ques-tions through its distinctive approach to collecting data. “Traditionally, oceanographic observatories have focused on physical measurements, like temperature, salinity, pH, and dissolved oxygen, leaving the biological observations behind,” said Duffy. While those physical measurements are a critical part of characterizing an ecosystem, replicat-ed biological experiments on a global scale could move knowledge about the ocean in a new direction. Calling

upon the wealth of expertise throughout the Smithsonian, and under the direction of Duffy, TMON is now actively developing the methods it will soon launch worldwide.

In September 2014, Duffy assembled a team to visit Car-rie Bow Cay with the focus on developing the methods for observing two key ecological functions: predation and herbivory in marine fishes. In other words, what method would best measure the activities of “fish as predators” and “fish as grazers” over time and between locations, whether on a tropical coral reef or a temperate saltwater marsh? Differences in herbivory and predation are impor-tant questions in ecology, as they are the very processes that structure biological communities and food webs. Predators can have ripple effects throughout the food web from the top down, while herbivory can do the same from the bottom up. In coral reefs, the intensity of grazing ac-tivity plays a key role in how a reef functions. Under in-tense grazing, corals can dominate a reef, supporting a rich assemblage of fish and invertebrates that associate with them. Remove the herbivores and algae can quickly, per-haps even permanently, dominate a reef, greatly reducing diversity and function of the system.

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Figure 4- Assembling "squid pops" at night to be deployed the next day. Michele Weber (left) attaches disks of dried squid to garden stakes with monofilament line while Michael Goodison (right) punches out the disks.

To target the predators, Duffy and his team deployed “squid pops” in a variety of habitats surrounding Carrie Bow. Squid pops are intentionally simple: a small disk of dried squid tethered to a garden stake with fishing line (Figures 4 & 5). “If these things are going to be replicat-ed all over the world,” explained Duffy, “they have to be very simple.” The method for gathering data is simple, too. Squid pops are deployed and observed after the first hour and again after 24 hours. Presence and absence is measured so that the results are a simple 1 or 0. “It’s very satisfying. During what other biological experiment can you exit the water with your data in hand? It’s never hap-pened for me!,” Duffy enthused (Figure 6).

Collecting data on herbivores proved a bit more challeng-ing. With the success of the squid pops, the logical step was to design an herbivore-friendly equivalent. Emmett’s team had come armed with two types of dried algae, purchased from the same Asian markets that had supplied the dried squid. The algae could be easily punched into disks and tethered to stakes, and initial results were promising. In the end, though, the “kelp pops” made from store-bought algae proved overwhelmingly unappealing to herbivores. “It’s no wonder nobody has tried this [method] before,” la-mented a disappointed Duffy. Other options were quickly explored then dismissed. “Sea lettuce”, a widespread type of algae, could not be used because of regional diversity – the same genus of sea lettuce may be found in each obser-vatory location, but the same species would likely not be. And even the same species of algae could have differences in palatability from one region to the next. Several fish species responded to the iceberg lettuce appropriated from

Carrie Bow’s kitchen, but there were durability issues with that option, and iceberg would likely be difficult to obtain in the Network’s most remote locations.

Employing precisely the same methods at each research site is the cornerstone of the Network’s initiative and no small task. Visually observing fish is relatively easy in tropical locations like Belize. The water is usually warm and clear, so a combination of divers and cameras can be used to watch the fish come and go. In other regions, the colder, cloudier Chesapeake Bay, for example, reduced visibility is a major obstacle. Enter the DIDSON (Dual-frequency Identification Sonar), an imaging sonar tech-nology that records fish as they pass in and out of view. This cutting edge technology can gather information on fish size and abundance, even in turbid conditions.

The team returned to the U.S. having learned a lot about how best to measure fish feeding intensity and armed with new ideas on how to best replicate “kelp pops”. If suc-cessful, the project could produce some of the first high-resolution data on patterns of marine predation and graz-ing intensity across space and time.

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Figure 6- TMON Director, Emmett Duffy supervises a squid pop deployment near Carie Bow.

Figure 5- Squid pops ready to be packaged up and deployed.

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Thalassia Experimental Network: Underwa-ter Meadows and Resilient Seas

Justin Campbell, Emmett Duffy, Valerie Paul, Jennifer McMillan

Note: This article appears on the MarineGEO website, marinegeo.si.edu

If not for the flash of an octopus as it propels through the water or the swish of sediment as a manatee lumbers through an expanse of emerald green fronds, seagrass meadows might be mistaken for a lush and unruly lawn, maybe even the tall grasses of a mountain meadow. To scientists, however, they are more like tropical rainforests. Home to an amazing diversity of life, seagrass beds are important indicators of the overall health of coastal eco-systems—and they can be just as sensitive as their terres-trial counterparts to changes in the environment (Figure 7).

Seagrasses are angiosperms—plants that produce flowers and seeds—that have adapted to life in the sea. With long narrow blades, they grow in extensive meadows through-out the world’s shallow coastal regions, both temperate

and tropical, providing important ecosystem services. Seagrasses help stabilize sediment, recycle nutrients, and protect our shorelines. They also serve as juvenile nurser-ies for fishes and shellfish, homes for endangered animals like manatees and sea turtles, and as habitats for commer-cial and recreational fisheries. But these complex ecosystems are in danger, says Justin Campbell, a Smithsonian postdoctoral fellow who devel-oped an interest in the effects of climate change on sea-grasses during his graduate work at Florida International University in Miami. “In addition to climate change, there are a number of local and regional factors harming sea-grasses, such as shifts in water quality, overfishing, and shoreline modification. We need rigorous, powerful ex-perimentation to begin to understand how these stressors interact and ultimately influence seagrass beds.”

As a Smithsonian postdoctoral fellow, Dr. Campbell has continued his work with seagrasses by helping launch the Thalassia Experimental Network (TEN). A MarineGEO initiative, TEN has established monitoring and parallel experiments in beds of turtlegrass (Thalassia testudinum), the dominant seagrass found throughout south Florida and the Caribbean. Based at the Smithsonian Marine Station

Figure 7- Seagrass ecosystems host a diverse array of fish and invertebrates.

(SMS) at Ft. Pierce, Florida, TEN is replicating these ex-periments at a Smithsonian field site in the Florida Keys as well as at the Smithsonian Tropical Research Institute (STRI) at Bocas del Toro, Panama, and the Smithsonian’s Caribbean Coral Reef Ecosystems Program (CCRE) at Carrie Bow Cay, Belize (Figure 8).

The goal of TEN is to understand vulnerability of sea-grasses to nutrient pollution (eutrophication) and decline of grazing animals, and to investigate how those mecha-nisms vary among locations. This issue is important for coastal systems worldwide, where fertilizer and septic runoff are common contributors to altered ecosystems. Results from this project will provide scientists with a wealth of data on how human impacts affect vegetated near-shore ecosystems and, by extension, the species that depend on them for shelter and sustenance.

Just like any other plant, seagrasses need sunlight to thrive and reproduce. Excess nutrients can increase the growth of algae, which cloud the water and overgrow the seagrasses, reducing the amount of sunlight reaching the plants. Al-though such algal blooms represent a major cause of sea-grass decline around the world, the complex interactions among nutrients, plants, and grazers that influence them remain poorly understood.

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The TEN experiments mimic these stressors, says Camp-bell, thus examining how seagrasses at different locations respond to nutrient loading and fishing pressure. Each site consists of 60 small field plots of turtlegrass in shallow wa-ters that receive similar amounts of light. Half the patches are treated with the same amount of slow-release nutri-ent fertilizer. The other 30 seagrass plots are left untreated. These beds are being monitored over time to describe and compare the structure and functioning of the seagrasses within the plots. “We monitor various aspects of seagrass health, such as growth rates and shoot density,” says Campbell. “It is vitally important that we do exactly the same thing in exactly the same way across all four sites. That way we ensure that differential responses between sites are not due to distinctions in our experimental design or methods.” This standardization of methodology among sites is a key theme of MarineGEO, and TEN represents an important first set of coordinated experiments across the sites.

Full analysis of the data gathered by the TEN experiment will start in fall 2014, but early findings are compelling. “What seems to be emerging initially is that nutrients influence vulnerability of seagrasses to grazing, but that there are differences in how seagrass beds respond,” says Campbell. Attention now is shifting to what parameters vary across geographic location that might explain these

Figure 8 - Justin Campbell (right) and Caitie Kuempel (left) pound PVC pipes into the bottom as they construct their experimental plots at the Carie Bow Cay site in the Thalassia Experimental Network.

differences. The coordinated experiments thus open up a host of opportunities for further research.

Campbell and the other researchers are also interested in the role that larger herbivores such as turtles and ur-chins play in determining the health of tropical seagrasses. “These animals can consume substantial quantities of sea-grass vegetation, and in some locations drastically change the structure of the seagrass bed,” Campbell explains. “While seagrasses are sensitive to ‘bottom-up’ influenc-es of excessive nutrients in certain regions of the world, other areas may be increasingly influenced by changes in these herbivore populations.” The TEN experiment thus is also assessing changes in herbivory. Of the 60 plots at each site, half are grown in cages to exclude these herbi-vores, while the other half are left exposed.

Finally, increased acidification of seawater is another fac-tor impacting seagrass health. Oceanic CO2 levels have been increasing over the past 200 years, and the Smithso-nian has recently begun to develop a baseline for measur-ing CO2 concentrations in their marine monitoring sites at SMS in Ft. Pierce, at SERC in Maryland, and at STRI’s

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Bocas del Toro facility in the Caribbean. These measure-ments are tracking variations in pH and acidity but—im-portantly—also assisting scientists to identify reasons for those variations.

For instance, monitoring has revealed extreme variability in pH at local levels. Campbell explains: “Just like ter-restrial plants, seagrasses use CO2 during the day in order to produce carbohydrates, which are essentially their fuel. During the day CO2 levels decline, due to photosynthesis. At night they increase, due to respiration.”

And while excess CO2 has proven generally to slow growth of coral reefs, Campbell’s research shows that cer-tain seagrass beds actually thrive in waters with high acid-ity. Those studies could prove invaluable to understanding how certain marine ecosystems can resist, or even thrive under, increased levels of CO2.

Campbell is excited to be part of research that meshes so closely with his own long-term goals. “I’m broadly interested in how coastal environments respond to en-vironmental change. Many of these changes stem from

Figure 9- Smithsonian research diver Caitie Kuempel transports PVC pipes for a seagrass plot.

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human activity, and my research examines the ultimate consequences of our actions. Seagrass decline can result from multiple factors, thus we have a complex interplay of local and regional stressors such as eutrophication, and broader global stressors such as elevated temperatures and CO2 from climate change. My work aims to address these factors, both singly and combined, across a variety of spatial and temporal scales.”

Tracking and understanding these causes and effects is a rigorous and long-term challenge. “These systems are amazingly complex and comprised of a diverse array of organisms, and they all show slightly different responses. Which means, unfortunately, there is no general blanket statement that we can make regarding how a particular organism will respond to some sort of stress, whether nu-trient enrichment, CO2, or even elevated temperature.”

The Smithsonian’s MarineGEO initiatives make it pos-sible for the kind of in-depth, broadly based, long-term re-search necessary to understand this complexity—and the role human beings play in changing the coastal environ-ments of which seagrass beds are such a vital part. “The

work that we can do across the network is increasingly powerful,” says Campbell, “because now we can paint a picture of how these systems respond not only in one loca-tion but across larger geographic scales, such as across the Caribbean.”

Biodiversity StudiesCarrie Bow Cay BioCode: Next-Generation Natural History

Scott Jones, Seabird McKeon, Michele Weber, Amanda Windsor, T.J. Yokachonis, Stephen Harris, J.T. Boehm

In the 40 years that Carrie Bow Cay has been a field sta-tion, an impressive array of marine life has been cataloged at the Smithsonian’s National Museum of Natural History (NMNH): more than 3000 species, spanning 24 plant, algae and animal phyla, containing 670 distinct families and 1518 genera. While this may seem comprehensive, scientists estimate that we have only discovered about 1/3

Figure 10 - A BioCube in mangrove habitat.

of the species that live in the ocean, so there is still a long way to go before we have an accurate picture of marine biodiversity. Given the rate that human beings are driving the loss of species, it is more important than ever that we concentrate efforts on better understanding biodiversity. In an attempt to capitalize on new technologies and utilize the resources that have been put into documenting the di-versity of the region, scientists at the Smithsonian have set out to generate a genetic library, or a “biocode”, of the major marine ecosystems around Carrie Bow Cay. A biocode is a comprehensive record of the taxonomic and phylogenetic diversity of a given place and time. It can be thought of as a library for our knowledge of biodiversity. When paired with traditional museum-based collections and images, the biocode becomes a powerful resource to understand patterns and address questions about ecologi-cal and evolutionary processes for a specific location.

To get started, the team needed a sampling strategy that captured the representative diversity of major habitats sur-

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rounding Carrie Bow Cay, but with a manageable amount of effort. The method they chose was the “one cubic foot” approach; that is, collect all the living organisms from one cubic foot sections (or “BioCubes”) of mangrove roots, a sea grass bed, and coral reef (figure 10). When forced to capture all living organisms within a constrained space, remarkable diversity emerges. For instance, more than 200 researchers were given five years to inventory all species on a reef in French Polynesia. A single BioCube from the reef produced more than 335 species (~10% of the total documented organisms) and included at least 17 species not found by previous collectors. Many of these inhabitants are tiny crustaceans like amphipods and juve-nile crabs that are often overlooked with other sampling methods.

In July 2014, a team of scientists led by Smithsonian Ma-rine Station ecologist Seabird McKeon traveled to Carrie Bow with an agenda of sampling three BioCubes. They started in the mangroves at Twin Cays, where they had to deal with stinging creatures of all types and visibility that could go from bad to worse with the kick of a fin. They then moved on to a seagrass bed and finally the back-reef lagoon. Once a site was picked, the team anchored a BioCube, set up a camera to film all the fish swimming through the cube, and finally extracted the area enclosed by the cube frame. That’s when the fun really got started. Each sample was brought back to the wet lab to be picked and pulled apart to find every last living organism large enough to see with the naked eye (Figure 11). Every plant

Figure 11 - Seabird McKeon (center) looks on as the group picks out all the creatures from a BioCube in Carrie Bow's lab.

Figure 12 - Sea anemones, crabs, shrimp. amd sponges are sepa-rated into cups before processing.

and animal was sorted and assigned a number. Each crea-ture was sorted and their numbers tallied to get an idea of the relative abundance of each organism and then selected three to be photographed and processed for DNA extrac-tion at NMNH (Figure 12). The specimens that got the extra attention were the voucher specimens that would be-come the beginning of the biocode library of Carrie Bow Cay. In total, the effort netted 680 specimens, 660 DNA vouchers, and 108 genomic tissue samples.

Additionally, McKeon and his colleagues have found ex-citing ways to use the project as a platform for outreach. In partnership with local non-profit the Biodiversity Center of Belize (BioBelize), they have already used the biocode project to help augment BioBelize’s efforts to support training of the next-generation of Belizean scientists to bet-ter understand and use the country's biological resources. After leaving the island, BioBelize utilized the specimen data collected to conduct a DNA barcoding workshop with five Belizean high school students utilizing predatory fish species collected from the BioCube effort. Each student received one fish, removed the gut, extracted the contents, performed DNA extraction, and gene amplification. Pre-liminary results indicate high quality extractions and high success rate in generating useful DNA sequences.

McKeon hopes the project will continue to grow; an in-ternship will start this winter dedicated to compiling the historical records of specimens, existing DNA sequences, and photos of samples taken from Carrie Bow Cay, in or-der to highlight ‘gaps,’ and appropriate taxa and ecosys-tem targets for future field expeditions. If he’s successful, the Carre Bow Cay Biocode will be the first of its kind in the Atlantic Ocean and will serve as template for future initiatives at other marine sites.

Decapod Biodiversity of Carrie Bow Cay

Darryl Felder

When friends hear that one of my projects is based at a small tropical research lab off the coast of Belize, they as-sume that I am out snorkeling and diving most of the day, taking flashy color photographs of plate-sized crustaceans for coffee-table books, and otherwise pretty much there to enjoy the tropical sun and sea breeze. While the work in Carrie Bow is without question enjoyable and rewarding to me, my collaborators, graduate students, and techni-cians, it is a far cry from what most envision.

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For a little history, my work in Belize began in the early 1980s, as a collaboration with two now-deceased Smithso-nian colleagues who were based at the National Museum of Natural History (NMNH). We initially set out to publish a faunal checklist based on what we thought could be a somewhat thorough collection of accessible intertidal and shallow subtidal decapod crustaceans (crabs, shrimp, and lobsters). I was included both for my interest in comparing diversity in Belize to that in tropical reaches of the Gulf of Mexico, and because I was interested in documenting coloration and studying its usefulness in defining species and populations. Early on, we made new discoveries and formally named new species, often utilizing coloration to distinguish them, but quickly realized that the task of fully documenting the decapod fauna would be a massive one, both to fully collect representatives and to deal with ex-tremely complex issues in identification of the organisms

collected. As in most tropical settings, the real diversity and abundance of decapods was composed of species that attain only thumbnail or smaller sizes. These may con-struct burrows in mud and sand or otherwise hide in small snail shells, crawl among the cracks and cavities of rocks and dead corals, or live as associates on or in cavities of other living marine organisms like sponges, anemones, corals, clams, snails, sea squirts, sea stars, sea cucumbers, seaweeds, sea grasses, and mangroves. To credibly rep-resent this diverse assemblage in the area around Carrie Bow Cay would require years of collecting effort, a vari-ety of technical approaches in collecting materials, and the emergence of new collaborators.

Figure 13 - The hermit crab Dardanus venosus.

After a long hiatus from the initial efforts, the task was taken up anew about ten years ago with another primary investigator and close friend from NMNH, Rafael Lemai-tre. Recently taking in yet another collaborator, Sammy De Grave from Oxford University, we are working toward completion of our coverage, but not without almost start-ing over! During the long period that the project was set aside, technology changed; methods for the identification of animals and the studying of their origins and relation-ships turned to gene sequencing. With an infusion of grant support from the Department of Energy and the National Science Foundation, my own home lab became heav-ily dependent upon what we call molecular phylogenetic techniques, by which we can use gene sequences to con-firm identities, closest relatives, and lineages of evolution-ary origin. Thus, we now can tell whether variant color patterns represent more than one species or just variations within just one species. Once this is known, the value of color as a recognition feature is greatly enhanced. One need only know the range of variation and fully document it for both sexes, in different habitats, and at different mat-urational stages. However, the early collections from Car-rie Bow were preserved in a way that did not allow speci-

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mens, even those previously photographed, to be used for gene sequencing. So the collecting and photography effort more or less had to be started over- this time with help of graduate students and technicians to expedite work back at the gene-sequencing laboratory. This involvement of a young cohort of future investigators is an important com-ponent of our project, as the ranks of more senior taxo-nomic specialists, such as myself, continue to dwindle.

Starting over was a tall order, but thankfully we had his-torical records of “what lives where” in the Carrie Bow vicinity. The single most productive return in terms of di-versity comes from breaking open old, eroded dead conch shells and coralline rubble. However, this requires hours of carefully chipping away with a hammer and chisel to remove intact little crabs, shrimp, and associated fauna. The targeted specimens are narcotized for color photogra-phy, followed by proper preservation for later genetic and morphological studies at the home laboratory. Sponges can also harbor diverse decapod assemblages, particularly the sponge masses growing on mangrove roots dangling in the water. Picking for hours through

Figure 14 - The shame-face crab, Calappa ocellata.

sometimes noxious tissues and toxic spicules of sponges, along with stinging hydroids, is a required but not envied duty. Likewise, using extraction pumps (called “yabby” pumps in Australia) in the swampy mud around the edges of mangrove-lined islands can be exhausting exercise on a hot calm day when the insects are hungry and where one can sometimes become too deeply mired to move. It guarantees a backache for the coming night, but the di-versity of highly adapted decapods to be extracted from that mud is remarkable. Then there is grassbed snorkeling, with face against bottom when the tide is right. This al-ways seems the time when those abundant little stingrays want to investigate the same small sand patch openings, the occasional box jelly drifts into contact with you, or nasty little parasitic isopods (fish lice related to pill bugs, but bigger and with serious jaws) manage to clasp onto skin underneath clothing.

So when the time comes for an evening break, it is usu-

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ally very welcomed…though all too short. Most evenings are spent hunched over the microscope for morphological studies or the camera stands for color macrophotography, with endless reworking of lens combinations and light-ing, cleaning and posing of subjects, writing field notes, and specimen logging until the lower back fully gives out (Figures 13-15). Then, well after dark is the best time for push-netting in the grassbeds. This is when nocturnal spe-cies of all kinds abound where they were not to be found a few hours previous. Getting a good photograph of these animals only pushes the nightly duties later, but it makes that eventual appointment with a bed washed by a steady sea breeze that much better. Sleep comes easily while waves crash onto the reef in the distance and occasional lightening from a thunderstorm glimmers on the horizon. To most it is not luxury on a tropical island, but it is one of the most ideal possible field settings for the kind of work I do.

Figure 15 - The snapping shrimp, Alpheus floridanus.

The seriousness of what we are doing at Carrie Bow is not lost across the years we have spent in these surround-ings, but instead is amplified by the perspective this gives. Over the last 35 years, we have not only seen the passing of two valued specialist colleagues involved in this study, we have also witnessed dwindling of the landmass and vegetation on this island that serves as the base of our proj-ect. At the same time we have documented the appearance of invasive species in the surrounding waters and declines in the health of corals and other reef organisms. Coastal ocean change poses increasing threats and we are now po-sitioned to document and better understand their impacts. Each year, we discover new species at Carrie Bow, and may not yet know what is already under threat. Some spe-cies that were not uncommon at Carrie Bow in the early 1980s we can no longer seem to find. It is sobering, and adds urgency to our efforts.

Ocean Sampling Day

Scott Jones

On the summer solstice (June 21, 2014), Carrie Bow Cay took part in Ocean Sampling Day (OSD), the larg-est ocean sampling event ever to take place on a single day. In total, 185 science teams all around the world took part in this massive event aimed at characteriz-

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ing the microbial communities in ocean water (Figure 16). Why orchestrate such a massive effort to collect things that are invisible to the naked eye? Well, it turns out that microbes (micro-organisms like bacteria and fungi) are a powerful way of assessing the health of the ocean because they drive most of the processes vital to ocean and human well-being. Over 5 million microbes can be found in one teaspoon of seawater and they com-prise 98% of the biomass of the world’s ocean, supply more than half of the world’s oxygen, and are the major processors of the world’s greenhouse gases. Scientists are only just beginning to understand their diversity, distribution, and function on a global scale.

Although many are visible with the use of a micro-scope, these microbes weren’t identified by human eyes. New DNA sequencing technologies allow us to not only to identify which microbes are in the sample, but also what function the microbes fill. These sequenc-ing technologies have only very recently been afford-able enough to reproduce on this scale. The process is simple: take a water sample, filter the contents through a very, very small filter, and then send the filters (which were immediately frozen) off to be analyzed.

The project was coordinated through the European-lead Micro B3 program (Biodiversity, Bioinformatics and Biotechnology, www.microb3.eu), and the Smithsonian

Figure 16 - A map of the 185 sites around the world participating in Ocean Sampling Day.

played a key role, by storing duplicate samples in its brand new BioRepository (http://www.mnh.si.edu/rc/biorepository/). Given that new technologies are devel-oping rapidly, having those samples in the deep freeze will allow future researchers to go back in time to sam-ple past ocean conditions in ways we can only imagine today. (OSD website link: http://www.microb3.eu/osd)

Environmental MonitoringCCRE Meterological and Oceanographic Monitoring Program

Tom Opishinsksi

The meteorological and oceanographic monitoring pro-gram initially established on Carrie Bow Cay in 1996 continued operation through 2014. A combination of oceanographic (temperature, salinity, turbidity, water lev-el, pH, and dissolved oxygen) and meteorological sensors (air temperature, wind speed/direction, relative humidity, barometric pressure, rainfall, and solar radiation - Figure 18) operate continuously to provide measurements of environmental conditions every 10 minutes. A datalog-ger records and transmits the measurements by radio fre-quency in real time to a base station located in Dangriga on the Belize mainland. The data is processed at the base station and assembled to form continuous time series data sets that are then uploaded via the Internet where it is im-mediately available for visualization and analysis on our web portal.

Due to a historical lack of monitoring over the entire Me-soamerican Barrier Reef, the environmental monitoring system (EMS) realizes a vision of Klaus Ruetzler, the founder of the CCRE program, to monitor conditions ex-isting on the reef to serve not only as a baseline for com-parisons of future conditions but also as an immediate re-source for researchers, scientists, environmental managers and the public. The EMS is completely automated and re-quires no human interaction except quarterly maintenance and service visits to calibrate sensors, perform repairs, and implement upgrades.

Data collected by the system continues to be analyzed to expand the products available to users of the web por-tal (http://nmnhmp.riocean.com). One such data prod-uct, introduced in 2011 and updated regularly as new data becomes available is the presentation of monthly wind rose charts (http://nmnhmp.riocean.com/windrose.

php?siteIndex=0). A wind rose is a graphic chart used to give a succinct view of how wind speed and direction are distributed at a particular location and for a specific time period. Viewed successively in chronological order the wind rose charts clearly illustrate seasonal patterns of the trade winds for the areas. In general the wind blows from a northeasterly to easterly direction from March through September and then transitions to northwest to north-northwesterly direction from October to February.

Following the tradition initiated by Klaus Ruetzler to es-tablish the first operational oceanographic and meteoro-logical monitoring system on the Mesoamerican Barrier Reef, we have initiated activities to expand our monitor-ing program with a new platform for sea level monitor-ing. The new system is designed to meet requirements established by the international Global Sea Level Observ-ing System (GLOSS) program for sea surface water level measurements to monitor sea level rise. As part of this ef-fort we performed an analysis of 10-plus year record of water level data (>450,000 samples) to correct for water density and atmospheric pressure changes. A two-year time series of the corrected data set has been submitted

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Figure 18- Meterological sensor tower at Carrie Bow Cay.

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to NOAA for a comprehensive analysis that precludes analysis of the entire data record. The data set is also being utilized by Smithsonian scientist Marguerite Toscano to benchmark peat cores taken on Twin Cays with the goal of advancing our understanding of historical mangrove re-sponse to sea level change.

This exciting expansion of monitoring capabilities, sup-ported by Smithsonian’s MarineGEO program, will include construction of a new offshore platform in early 2015. The platform will host multiple state-of-the-art vented water level sensors and will be equipped with a satellite transmit-

ter to send data in real time to NOAA and to our web portal. Measurement sampling will be increased from the present 10-minute rate to 6 minutes. In addition, the system design will meet protocols for tsunami warning systems, providing a crucial need for the Central American region.

EcologyAn Integrative Investigation of Population Connectivity Using the Coral Reef Fish Elacatinus lori

Peter Buston

Understanding the patterns, causes and consequences of larval dispersal is a major goal of 21st century marine ecology. Patterns of dispersal determine the rates of lar-val exchange, or connectivity, between populations. Both physical factors (e.g., water movement) and biological factors (e.g., larval behavior) cause variation in popula-tion connectivity. Population connectivity, in turn, has major consequences for all aspects of an organism’s biol-ogy, from individual behavior to metapopulation dynam-ics, and from evolution within metapopulations to the origin and extinction of species. Further, understanding population connectivity is critical for the design of effec-tive networks of marine reserves – vital tools in the devel-

Figure 20 - Boston University research divers collectiong neon gobies.

Figure 19 - The neon goby, Elacatinus lori, inhabits tube sponges.

opment of sustainable fisheries.

Over the last decade, three methods, each of which tells us something slightly different, have emerged as the leading contenders to provide the greatest insights into population connectivity. First, coupled biophysical models make as-sumptions regarding water flow, larval behavior and ecol-ogy, to predict population connectivity. Second, indirect genetic methods use spatial distributions of allele frequen-cies to infer population connectivity. Third, direct genetic methods use parentage analyses, tracing recruits to spe-cific adults, to measure population connectivity. Despite advances, lack of integration means that we do not know the predictive skill of biophysical models, or the extent to which patterns of dispersal predict spatial genetic struc-ture.

The overall objective of our work is to conduct an integrat-ed investigation of population connectivity, using all three methods, in one tractable system: the neon goby, Elaca-tinus lori (Figure 19), on the Belizean Barrier Reef. In 2011, we stayed at Wee Wee Cay and conducted studies of larval dispersal over short distances (1 km) on the reef ad-jacent to Carrie Bow, showing that we could trace recruits to parents using parentage analyses. In 2012, we used Car-rie Bow as a base to conduct studies of population genetic structure, showing that oceanic breaks between patches of reef were associated with strong genetic differentiation between sites (Figure 20). These preliminary studies laid the foundation for us to take things to a whole new level.

In 2013, we stayed at the Institute for Zoological Expedi-tions on Southwater Cay and conducted studies of larval dispersal over much longer distances (30 km) on the bar-

rier reef to the north and south of Carrie Bow. We hope that the findings of that work will emerge in 2015. This year, we used Carrie Bow as the starting point for an ex-pedition to conduct studies of population genetic structure at a much finer scale all over Belize. Six of us stayed at Carrie Bow: myself, a postdoc from University of Miami (David Lindo), a graduate student from Boston University (Cassidy D’Aloia) and three undergraduates from the lab (Diana Acosta, Philip Schmiege and Derek Scolaro). Car-rie Bow was the perfect launch pad and we are thankful to the staff there for a wonderful stay.

You can find out more about the whole trip by visiting Di-ana Acosta’s blog (http://bustonlab.wordpress.com/).

Can Seaweed-Coral Competition Make Sea-weeds More Palatable?

Guilherme Longo and Mark Hay

Seaweed-coral competition is increasingly common on modern coral reefs, but the detailed mechanisms and con-sequences of these interactions are still to be unraveled. The outcomes of seaweed-coral competition may vary with coral and seaweed species, resulting in variable im-pacts to reef structure and dynamics. Seaweeds can use different strategies to compete with corals, including shad-ing, abrasion, and producing chemical compounds that are toxic to corals. However, if the same chemical compounds also serve as anti-herbivore defenses, seaweeds compet-ing with corals could be even more herbivore resistant and less likely to be consumed by herbivorous fishes and sea

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Figure 21 - Interaction betwen Agaricia lettuce coral and seaweed Halimeda, resulting in coral bleaching.

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urchins. We video recorded approximately 500 coral colo-nies at Curlew Bank, near Carrie Bow, Belize to inves-tigate the frequency of coral-seaweed contacts, the sea-weeds and corals involved, and whether corals appeared to be damaged by these contacts. For the most common pair of interactors, we evaluated whether contact with coral made the seaweed more palatable to the sea urchin Diadema antillarum. Coral-seaweed contacts were partic-ularly frequent between Agaricia corals (mostly the thin leaf lettuce coral, A. tenuifolia) and the seaweed Halim-eda opuntia, with this interaction being associated with coral bleaching in 95% of contacts (Figure 21). Pooling across all coral species, H. opuntia was the seaweed most commonly contacting corals and most frequently associ-ated with localized bleaching at the point of contact. The common corals Agaricia and Porites (mustard hill cor-als) bleached with similar frequency when contacted by H. opuntia, but Agaricia experienced more damage than Porites when contacted by articulated coralline algae. When paired samples of H. opuntia that had been in con-tact with Agaricia and not in contact with any coral were offered to sea urchins (Diadema antillarum), urchins con-sumed about 150% more of thalli that had been compet-ing with Agaricia (Figure 22). Contact and non-contact algae did not differ in nutritional value, suggesting that anti-herbivore chemical defenses of Halimeda may have been compromised by coral-algal contact. If competition with corals commonly enhances seaweed palatability, then the dynamics and nuances of small-scale seaweed-coral-herbivore interactions at coral edges is deserving of greater attention in that such interactions could scale-up to have important consequences for coral resilience, the persistence of reef structure and function.

A 1000-year Record of Change in Caribbean Coral Reef Communities and Environments

Katie Cramer, Aaron O’Dea, Richard Norris

Caribbean reefs have declined dramatically since reef monitoring studies began in the 1980s, losing 50-80% of their living coral cover during this time. This decline is

typically attributed to outbreaks of coral bleaching and disease that first appeared in the 1980s and are linked to anthropogenic climate change. Although human-caused disturbances such as land-clearing and overfishing were negatively impacting Caribbean reefs for centuries or more, the lack of baseline data on reef communities and environments prior to the 1980s has made it difficult to assess the role of historical anthropogenic activities, hin-dering efforts to pinpoint the precise mechanisms of reef deterioration.

To help resolve this issue, our team traveled to Carrie Bow Cay Field Station to reconstruct a timeline of histori-

Figure 22- Diver operating the push corer.

Figure 23 - Section of sediment core.

cal change in Caribbean reefs by sampling material from the reefs of the past. Using a diver-operated push coring system (Figure 22), we collected sediment cores up to 5.5m below modern reefs at six different reef sites within the central Belizean lagoon (Figure 23 & 25). Along the length of each core, we are tracking pre-historical and his-torical change in: (a) reef environmental conditions from analysis of chemistry and grain size distribution of sedi-ments, and (b) composition and abundance of major reef organisms including corals, reef fish, urchins, algae, and sponges from skeletons, teeth, spines, plates, and spic-

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ules that are preserved in the sediments (Figures 24). This study, coupled with reef sediment cores we collected from Caribbean Panama in 2012, will provide a more accurate ecological baseline from which to assess the true timing, extent, and causes of the recent demise of Caribbean reefs. Understanding the natural state of reefs before large-scale human disturbance and pinpointing the ultimate drivers of recent reef degradation are essential for effective reef management and conservation.

Patterns of Insect Biodiversity in Pristine and Degraded Mangrove Habitats

Alexander Forde

Habitat loss and fragmentation, along with climate change and invasive species, continue to cause worldwide de-clines in biodiversity. Because diversity is linked to eco-system stability, productivity, and services, increasing our understanding of how natural and anthropogenic factors shape diversity is a critical pursuit. While the plight of in-land tropical forests is well-known, coastal tropical forests comprised of mangrove trees are similarly threatened by deforestation and over-exploitation. Approximately 30% of the world’s mangroves have been lost in the past three decades, and a further 1% is lost annually. The use of secondary forests (forests in the process of regenerating after deforestation) and agricultural areas in conservation plans has recently become a topic of great interest, yet data on levels of diversity across disturbance gradients in tropical forests is often lacking. In order to address this knowledge gap for mangrove forests, I surveyed insect

Figure 26- Insect traps in dwarf mangrove stand.

Figure 24- Various types of fish teeth found in the cores.

Figure 25- Research team with cores in the boat ready to transport.

communities in mangrove habitats that differed in their history of disturbance.

I conducted the insect survey during November 2013 on Twin Cays, a 92-ha mangrove island, 12km off the coast of Belize. Vegetation on the island primarily consists of red mangroves (Rhizophora mangle), which form tall stands (4-6m) in bands along the exterior of the island. These “fringe” forests grade into short (1m) stands in the island interior where low nutrient conditions cause stunt-ed growth. Portions of Twin Cays have been clear-cut in the past, and some of these areas were covered in mate-rial dredged from the nearby seafloor. For my study I fo-cused on five distinct habitat types: 1) Tall pristine stands of red mangroves (fringe), 2) short pristine stands of red mangroves (dwarf, Figure 26), 3) areas clear-cut in 2003 but not otherwise altered (cut), 4) areas cut and filled with dredge in 1995 and cut again in 2003 (cut and filled, Fig-ure 27), and 5) natural interior ponds lacking any vegeta-tion. The interior ponds serve as a reference for habitat to-tally unsuitable for sustaining local populations of insects.

To sample insects, I deployed traps consisting of 10x18cm cards covered on both sides by water-proof adhesive, sus-pended 1.25m above ground level by a PVC pipe. Traps

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were at least 8m from a habitat edge or from other traps, were set up in 10 separate locations on the island, and were collected after 6 days. An average of 18 traps were set in each habitat type. I identified insects caught by the traps to various taxonomic levels ranging from Order to Family, and assessed levels of biodiversity with the Shan-non index. The most common groups of insects collected were true flies, whiteflies, wasps, true bugs, and barklice.

Overall, pristine mangrove stands had 38% higher insect diversity compared to regenerat-ing areas, yet the abundance of insects on traps did not differ between pristine and cut areas (mean=47 per trap). The most prominent dif-ference between insect communities in pristine vs. cut areas was the greater predominance of true flies (Diptera) in the latter (79% of all in-dividuals vs. 48%). Differences in diversity be-tween all 5 habitat types are displayed in Fig-ure 28. Surprisingly, areas cut 10 years ago had similar levels of insect diversity compared with pristine dwarf stands, while areas cut and filled exhibited levels of diversity as low as that found above open water. These results highlight the fact that different types of disturbance can have drastically different outcomes, which can range from a return to somewhat “natural” levels of animal diversity after a decade vs. remaining completely unsuitable habitat.Fringe Dwarf Cut Cut&Filled Pond

Landscape type

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Figure 27- Insect traps in clear-cut and filled mangrove stand.

Figure 28- Diversity of insects in pristine and regenerating habitats on a mangrove

island (±SE). Means labeled with different letters are significantly different.

Marine Chemical Ecology and Invertebrate Taxonomy

Natassia Patin, Paul Jensen, Joseph Pawlik, Lindsay Deignan, Greg Rouse, and Fredrik Pleijel

The coastal town of Dangriga, Belize, is a small seaside town with no high-rise buildings, no dive resorts, no bars catering to expatriates. Many of the streets are unpaved, and chickens run with abandon between the houses on stilts. These houses come in all shapes and sizes, often painted in lively shades of violet and pink. The people there are unfailingly friendly, carefully adjusting their native Creole dialect to speak proper English with white foreigners. “We are recreating amongst ourselves,” de-clared a lady delightfully when asked about her card game. It’s easy to forget that this small community is just a short boat ride away from some of the world’s most stunningly beautiful coral reefs dotted with tropical cays. A quick reminder is the Pelican Beach Resort, which

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serves as the jumping-off point to Carrie Bow Cay, our destination for the upcoming week.

Carrie Bow, as it is affectionately known, has served as a venue for tropical marine research for over 40 years. The island is run by the Smithsonian Institution to fa-cilitate easy access to Mesoamerican Barrier Reef, the second largest barrier reef in the world (and largest in the northern hemisphere). Having survived hurricanes, erosion, and a devastating fire, Carrie Bow remains an unparalleled setting for studying coral reef and man-grove ecosystems. It provides excellent diving and small boating facilities, along with both wet and dry lab space for sample processing and experiments. The setting and facilities offered at Carrie Bow are the reason our group chose to spend a week here in September 2014. For sev-en days, we lived on an island that can be circumnavi-gated on foot in two minutes and used the small boats to explore the surrounding reef and nearby mangroves. Snorkeling and scuba diving allowed us to make obser-vations, collect samples, and conduct experiments in a way that would never be possible at our host institutions.

Figure 29 - Lettuce sea slug, Elysia crispata

The trip was organized and led by Paul Jensen, a re-search microbiologist at Scripps Institution of Oceanog-raphy at UC San Diego. Our group studies a group of marine bacteria that produce bioactive chemical com-pounds, many of which have therapeutic potential. These bacteria, called Salinispora, inhabit tropical marine sedi-ments worldwide, and we had previously isolated strains from the Bahamas and off the Yucatán peninsula. We hoped to find more strains at Carrie Bow to better our understanding of their biogeography. In addition, we at-tempted ambitious projects to test potential ecological functions of their compounds, or secondary metabolites (“secondary metabolites” refers to compounds that are not necessary for the short-term survival or an organism, unlike primary metabolites like amino acids or sugars). As a fourth-year graduate student in Paul’s lab, my thesis work has focused on the effects of these secondary me-tabolites on other members of the bacterial community. At Carrie Bow, Paul and I combined field experiments with molecular microbiology to gauge how Salinispora compounds affect the richness and diversity of their sur-rounding community (Figure 31). Although the samples have yet to be processed, we hypothesize that exposure to these potent molecules will produce a noticeable shift in community structure as determined by high-through-put DNA sequencing. In addition to these experiments, we tested a separate hypothesis that certain secondary metabolites deter grazing by higher-level predators like polychaete worms and benthic crabs. Having immediate access to the native habitat of our organisms provided an incredible opportunity to run these experiments with minimal delay and processing time.

Two other members of the group joined us from the University of North Carolina Wilmington to study the chemical ecology of a very different group of organ-

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isms. Professor Joe Pawlik has studied the chemical defenses of Caribbean sponges for over 20 years, pro-ducing groundbreaking studies on the role of feeding deterrence in sponge ecology. At Carrie Bow, Joe and his student Lindsey Deignan investigated predation on chemically defended sponges by molluscs. Previous observations of grazing scars on defended sponges, such as the large yellow tube sponge Aplysina fistu-laris, had been attributed to fish or turtle predation. During their night dives, Joe and Lindsey found that these scars are actually the result of nocturnal mollus-can grazing, mostly by cowries. They photographed cowrie predation on sponges and discovered that mul-tiple cowries may inhabit individual sponge tubes, and that their crawl-away juveniles feed on the interior of the sponge. Theirs will be the first report of these basic ecology and natural history features and was made pos-sible by the fantastic diving facilities on Carrie Bow Cay.

The remaining two members of our group were inver-tebrate taxonomists and underwater photographers Pro-fessor Greg Rouse, from Scripps Institution of Ocean-ography, and Professor Fredrik Pleijel, from Göteborg University in Sweden. Together, Greg and Fred are a vast resource of information on benthic invertebrates. From the historical details of taxonomic reshuffling to the feeding strategies of obscure microscopic worms, dinnertime conversation was never dull and offered a unique opportunity for microbiologists to trade ideas with sponge and invertebrate biologists. Greg and Fred collected an astonishing number of small invertebrates from the sand around the island, surrounding reefs, and

Figure 31- Research divers collecting sediment.

Figure 30- Polychaete fan worm, Sabellidae.

nearby mangroves. These animals resulted in wonder-ful photographs, some of which are included here, il-luminating in stunning detail the intricate beauty and diversity of the benthic macrofauna (Figures 29 & 30). As someone who does not get to visually appreciate my study organisms, it was remarkable to see how basic morphological features can provide insight into an ani-mal’s ecological role.

Up to six scientists are allowed on Carrie Bow Cay at any given time, so we were at full occupancy during our week’s stay. Despite the island’s tiny size it never felt crowded; the laboratories and bedrooms are spacious, with room for plenty of porch hammocks. The Smith-sonian also provides a cook to prepare all the meals and a station manager to handle the filling of scuba tanks, generator maintenance, and any broken equipment. We were lucky to have Martha, an exceptional cook who provided us with delicious, satisfying meals including plenty of vegetarian options. Keith, the station manag-er, has been volunteering for over twenty years at Car-rie Bow and was a wealth of knowledge on the island’s history, weather patterns, and local fishing. They both

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made our stay on the island extremely comfortable and gave us the freedom to spend as much time on our work as possible. Of course, that still left a few minutes each day for cocktail hour and hammock time – the perfect balance on a true tropical paradise island. With luck and continued funding, many others will be able to take ad-vantage of Carrie Bow Cay in the future and continue the scientific tradition that has flourished here for over four decades.

Coral Larval Ecology

Valerie Paul and Jennifer Sneed

Coral reef degradation is occurring rapidly on a global scale. In the Caribbean, depressed herbivory rates due to overfishing and the massive die-off of the sea urchin Di-adema antillarum have increased macroalgal abundance and the frequency of coral-algal competitive interactions. The proliferation of macroalgae on these coral-depauper-ate reefs threatens the health of corals and the resiliency of reef ecosystems. It has been proposed that in the ab-

Figure 32- Larval chambers are used to test the interactions between coral larvae and macroalgae.

sence of herbivorous pressure, macroalgae can become abundant on reefs and their presence puts into motion a positive feedback system that leads to an alternate stable state in which macroalgae are dominant. The production of potent allelopathic compounds by many macroalgae contributes to their ability to maintain a dominant state on reefs by negatively impacting coral health and inhibiting the recruitment of juvenile corals.

This past summer at Carrie Bow Cay we examined the effects of macroalgae on the settlement behavior of Acro-pora palmata (Elkhorn coral) larvae (Figure 32). We tested the response of A. palmata larvae to two species of macroalgae that are commonly found on reefs in the Caribbean, Halimeda opuntia and Dictyota pulchella. We had previously found that H. opuntia does not impact the overall settlement response of another coral, Porites astreoides. Contrary to what we expected, P. astreoides larvae were not repelled by the algae and in fact ~ 10% of the larvae settled on the algae itself. Similarly, settlement of A. palmata larvae was not reduced in the presence of the either H. opuntia or D. pulchella. However, unlike P. astreoides, A. palmata larvae did not settle on the sur-face of H. opuntia. This study emphasizes the differences in settlement behavior among different coral species and contributes to our knowledge of coral-algal interactions, which are becoming increasingly common on Caribbean reefs.

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Mesophotic Reefs

Michael Studivan and Josh Voss

The Voss lab traveled to the Smithsonian’s Carrie Bow Cay Field Station with two objectives in mind: (1) to scout out reef locations for PhD student Michael Studivan’s cor-al reciprocal transplant experiment, and (2) to determine if Carrie Bow is a suitable site for future Voss lab projects. Michael’s dissertation research aims to explore the differ-ences between shallow and mesophotic (30-80 meters) coral reef. His proposed reciprocal transplant will specifi-cally determine how the great star coral Montastraea cav-ernosa modifies its skeleton and gene expression to adapt to varying environmental conditions. This experiment re-quires relatively healthy populations of the species at both shallow (20 meters) and mesophotic depths (50 meters), and the ability to easily manipulate colonies and sample them over time. Over the next three years, the Voss lab plans to return to CBC to monitor and sample the same coral colonies and see how they change over time.

Figure 34- A colony of Monstastrea cavernosa tagged for future use in reciprocal transplant experiment.

Figure 33- Researcher Josh Voss tagging a coral colony,

During their recent trip in March of 2014, Dr. Joshua Voss and Michael Studivan surveyed several reefs and selected three sites for the transplant experiment, named Raph’s Wall, Tobacco Reef, and South Reef. These three sites have an outer reef crest at 20 meters depth along the bar-rier reef, with a wall-like slope with small ledges at 40 and 50 meters. The deeper reefs are dominated by sponges and whip coral species, but still have apparently healthy hard coral populations, including M. cavernosa and sev-eral species of the lettuce coral, Agaricia (Figures 33- 34).

Since mesophotic reefs are found below the limit for rec-reational SCUBA divers, Dr. Voss and Michael completed technical diver training that allows divers to dive deeper and longer safely. Using their advanced training, they deployed duplicate temperature/light data loggers at the shallow crest and deep ledge sites in order to characterize some of the environmental differences between depths. They also tagged and sampled (4 sq cm each) fifteen shal-low colonies from each site. The preliminary sampling will allow an initial description of coral population struc-ture using microsatellite analysis. Knowledge of structure and connectivity within a population is extremely impor-tant for an understanding of intraspecies dynamics and may provide necessary information for conservation and protection strategies of coral species.

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Additionally, the samples from their recent trip will be used to determine the basal gene expression profiles of the shallow corals. By looking at gene expression, they are able to see a snapshot of how a coral is reacting to environ-mental stimuli, as expression of a certain gene often leads to a protein used under those conditions. Advanced molec-ular tools like these are now available to provide scientists with more information about a coral species than can be observed by analyzing its structure and appearance alone. In order to determine how the coral changes over time dur-ing the transplant experiment, they need to compare it to normal levels of gene expression, such as those exhibited by the unmanipulated colonies sampled in March.

With the first data from this project, the Voss lab will be able to return to CBC as soon as June 2014 to move colonies and start the transplant experiment. They will be sampling each of the 45 total colonies at the outset of the experiment, and will return in October 2014 for short term responses, and then each following May until 2017 for long term responses. During their first trip in March, they accomplished their research objectives, enjoyed the company of the station managers and cook, and managed to find some time to relax on the island! They invite you to see our webpage (http://vosslab.weebly.com/news.html) for blog posts about our trip and lab Flickr (https://www.

Figure 35 - Carrie Bow Cay station managers taking bi-weekly secchi disk readings in front of the station.

flickr.com/photos/hboi_voss/sets/72157642012263954/) for photos. Finally, once logistics around technical diving are sorted out, they are excited to return to the island for Michael’s project and future Voss lab research!

CARICOMP: 21 Years of Data Collection Contributing to Important Publications

Karen Koltes

Monitoring surveys and data collection were conducted in July 2014 under the Caribbean Coastal Marine Productiv-ity Program (CARICOMP). CARICOMP was launched at Carrie Bow Cay (CBC) in 1993 as part of a collabora-tive regional scientific effort among marine laboratories to study land-sea interaction processes, to monitor for change on local and regional scales and distinguish an-thropogenic change from natural variation, and to provide appropriate and reliable scientific information for natural resource management.

Standardized, synoptic measurements are made in the three primary Caribbean coastal ecosystems of coral

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reefs, seagrass beds and island mangroves, together with relevant and simple oceanographic and meteorological measurements. In addition to the standard CARICOMP measurements taken at CBC (Figures 35 - 36), enhance-ments to the protocols continue with photo-documenta-tion of coral reef transects, octocoral surveys to improve observations of population dynamics, in situ light intensity measurements and current flow monitoring in shallow and deep sites.

In March of this year, the CARICOMP network of collab-orators published the first regional synthesis of the nearly two-decades long record of semi-annual measurements on growth rates and biomass of the dominant Caribbean sea-grass, Thalassia testudinum (see van Tussenbroek, et al 2014, p. 38). Wide variations in community total biomass (285 to >2000 g dry m−2) and annual foliar productivity (<200 and >2000 g dry m−2) were found among sites. So-lar-cycle related intra-annual variations in T. testudinum leaf productivity were detected at latitudes > 16°N. Hur-ricanes had little to no long-term effects on these well-de-veloped seagrass communities at all but one station where the vegetation was lost by burial below ~1 m sand. At two

Figure 36 - Horizontal secchi disk readings near Twin Cays measures water clarity in the lagoon.

sites the seagrass beds collapsed due to excessive grazing by turtles or sea urchins (the latter in combination with human impact and storms). The low-cost methods of this regional-scale monitoring program were sufficient to de-tect long-term shifts in the communities, and fifteen (43%) out of 35 long-term monitoring stations showed trends in seagrass communities consistent with expected changes under environmental deterioration.

Data collected at Carrie Bow Cay under CARICOMP was also incoprorated in a landmark study, Status and Trends of Caribbean Coral Reefs: 1970-2012 (link). This report is the most detailed and comprehensive study of its kind published to date and is the result of a three-year joint effort of the International Coral Reef Initiative’s (ICRI) Global Coral Reef Monitoring Network (GCRMN), the Interna-tional Union for the Conservation of Nature (IUCN), and the United Nations Environmental Programme (UNEP). Long-term datasets such as those collected at Carrie Bow were an important part of generating the report’s conclu-sion that important herbivores, such as parrortfish, will play a large role in the preservation and recovery of Carib-bean coral reefs.

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CCRE South Water Caye Marine Reserve Reef Assessment Program

Scott Jones, Randi Rotjan, Zach Foltz, Peter Gawne, and James Dimond

This year marks the fourth full year that CCRE staff and collaborators from the New England Aquarium have implemented the South Water Caye Marine Re-serve Reef Assessment Program. This project is di-rected at assessing effects of the no-take conservation zone around Carrie Bow Cay on recovery of fish and coral populations. Permanent transects were estab-lished in June 2011 inside (12 transects) and outside (12 transects) the area’s boundary (Figure 37).

Most monitoring plans measure the diversity and abundance of key reef organisms; some of the best programs also assess biomass of benthic reef builders and fishes. We have designed our program to assess similar ecological metrics, so as to be cross-compati-ble with historical and simultaneous efforts elsewhere

Figure 37 - Fish and coral populations are monitored bi-annually near Carrie Bow Cay.

in Belize and the western Atlantic Ocean. However, we have also added some innovative and critically im-portant assessments that yield information about key ecological rates and states that are thought to contrib-ute to reef resistance, resilience, and recovery in the face of negative impacts. Rates include the grazing rates of herbivorous parrotfishes and surgeonfishes. Benthic states include scleractinian coral health status as well as recruitment and growth dynamics. This plan will enable more comprehensive ecological monitor-ing, and inform models of reef dynamics that will be used to generate new insights into reef community structure in response to different reserve management regimes. This study is designed to take advantage of the strengths and capabilities of the Carrie Bow Cay Field Station and produce important information that will be applied to habitat management in the newly formed SWCMR no-take area.

To date, 24 permanent transects have been surveyed eight times, for a total of 192 transects, and results to date show that:

• Hard coral cover is holding steady at ~10%, supported by data collected via 2 methods: transects and photo-quadrat analysis

• As of yet, there are no obvious reserve-level effects on benthic structure or fish populations.

InvertebratesInvertebrate Vision

Michael Bok, John Kirwan, Erin Cummings, and Laura Bagge

When most people think about eyes or vision, worms aren't usually the first animals that come to mind. In a way, this is precisely why we are interested in this (somewhat ob-scure) topic: the eyes of worms – how they work and why they evolved. Likewise, when most people think of excit-ing scientific fieldwork doing marine biology in far-flung tropical islands, they probably don't conjure up images of diving and hauling rubble for strange kinds of worms, but that's exactly why we came to Carrie Bow Cay, off the coast of Belize.

Half a billion years ago, before animals with very good eyes appeared, the seas swarmed with a multitude of peculiar

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Figure 37 - Fan worm, notice paired eyepsots on crown.

Figure 38 - Photoreceptor on the crown of a fan worm.

soft-bodied animals, with very different body shapes, sizes and lifestyles, but many of which we could broadly lump together as 'worms'. Although there has been a tremendous amount of evolutionary change since then, this title could still apply to most of the groups of complex animals that are around today. In the meantime, a few groups of animals have cast aside their wormish origins and taken over the earth, seas and skies, especially the back-boned animals (vertebrates) and the arthropods (insects, crustaceans etc.). Many different factors make these sophisticated animals different, but undoubtedly one key aspect of their success has been the emergence of fast and acute vision.

Almost every group of animals can detect light in some way – but this ability (photoreception) is mostly slow and is used for checking the time of day or finding if you are in shad-ow. True 'vision' means forming an image from the light information around you in real-time and this is very chal-lenging. It is also extremely useful; the first animals with true vision and speed may have forced their neighbours to evolve fast or face extinction! So how did this happen and

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what circumstances make this valuable but very demanding trait necessary? To answer this, it is necessary to understand the anatomy and visual ecology of animals with increasing complexities of light detection, and especially those with the most basic image-forming vision.

We searched through the mangroves, seagrass beds, open water and reefs for worms that could help us explore these questions. Mike was mostly interested in the 'fan-worms' (Sabellida), strangely beautiful segmented worms that sport a startling array of different photoreceptor (visual cell) types among the many species (Figures 37 - 39). John was mostly interested in the errant polychaetes, more active seg-mented worms that run the gamut from odd and beautiful to monstrous and mean-spirited (they have jaws!) and come equipped with everything from the simplest photoreceptors to fully-fledged camera-type eyes. We also looked for other strange beasts such as flatworms, ribbon worms and arrow worms with all manner of simple eye arrangements.

We were assisted by our excellent scientific divers, Erin

Figure 39 - Fan worms are often colorful and can be found all over coral reefs.

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and Laura, and our top-notch station manager, Danny. We collected many fireworms (a type of colorful segmented worm with a nasty sting) from the reef as well as flatworms, bloodworms and giant-eyed ragworms. We collected fan worms of all stripes and eye types, some of which were tiny and some over a foot in length (Figure 39). Using a custom-made arena we checked if the more active worms would move towards a dark pattern, indicating that they can see a simple image and it appears that many of the worms can, even without sophisticated eyes. Being out in the field in places like Carrie Bow Cay is not only the best part of the job, it's also an essential component of natural science – ex-ploring the diversity of living things in their natural context helps to make sense of fundamental questions.

Super Size me! Bacteria Break Cell Division Size Limit

Silvia Bulgheresi, Nika Pende, and Nikolaus Leisch

The life of a cell is pretty straightforward: it doubles in size, divides in the middle and gives rise to two identical daughter cells. Then these start the cycle all over again. Therefore, it has been long assumed that cells belonging to the same population are similar in size and that binary fission cannot work over long distances. However, cell bi-ology traditionally focused on a few cultivable organisms, thereby neglecting how unicellular organisms multiply out in the wild.

In the past year, we showed that the bacteria coating two species of marine roundworms - despite growing so long

Figure 40- A) Scanning electron microscopy image of a Eubostrichus fertilis nematode and its crescent-shaped bacterial partners arranged on its surface like the layers of an onion (right). B) Eubostrichus dianeae nematode and its fur made

of filamentous bacterial cells, each attached with one end to the surface of the worm. C) Plot showing average FtsZ and DNA fluorescence along the E. fertilis symbiont cell length (in percentage) emitted by 77 dividing cells. D) Plot showing average FtsZ and DNA fluorescence along the E. dianeae symbiont cell length (in percentage) emitted by 54

dividing cells

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to be perceivable by the naked eye - divide regularly, as model bacteria do. The crescent-shaped bacteria coating the roundworm Eubostrichus fertilis and the filaments attached to E. dianeae can indeed be up to 45 µm and 120 µm-long, respectively. By applying cutting-edge im-aging and automated analysis of thousands of symbiont cells, we proved that (1) division is mediated by the tu-bulin homologue FtsZ and that (2) DNA is symmetrically segregated among the daughter cells (see Figure 40). Ad-ditionally, on the E. fertilis worm, we observed up to ten-fold length variation between short, proximal crescents and long, distal ones. This unprecedented size variability arises because, surprisingly, the E. fertilis symbiont may divide at every length comprised between 3 and 45 µm.

What supersizes the bacteria associated to the two Eubost-richus worms? Why are they differently arranged on their respective animal partners? One possible explanation is that each bacterial arrangement evolved to make the most out of its faithful partnership with the worm.

The major implication of this study is that yet unknown factors may regulate bacterial cell division both spatially and temporally. Given that these could be animal-derived and that microorganisms make up to 1 kg of our body weight, learning how animals may regulate their prolif-eration might come in handy.

The Evolution of Mating Displays in Ostracods

Emily Ellis, Todd Oakley, Nicholai Hensley

When we arrived at Carrie Bow Cay, we hit the ground running. The facilities are great for field work and the station manager on duty was extremely helpful in terms of getting us situated on the island. We immediately set out to find the creatures we came to study: ostracods! Ostracods are small crustaceans that inhabit most bodies of water, including the wonderful waters around Carrie Bow. For some species, we simply scooped and sieved sand from the water and then searched through the filtered samples for the little arthropods. But at night, we collect-ed a unique group of ostracods found solely in Caribbean waters. Right after sunset, when the moon isn't shining, the males of these ostracods come out and produce a vivid mating display to attract females. They make a series of bioluminescent pulses in the water, and each species cre-ates a unique signal pattern to attract the right mate. Hence their Japanese name, “umihotaru”, or sea firefly. These

displays are fantastic, like delicate streams of underwa-ter light, where dozens of individuals swimming over the patch reef and sand beds produced them simultaneously. On different nights, we repeated our sampling over differ-ent areas of the reef, as well as over some of the grass beds to capture different, but closely related species (Figure 41).

There are multiple reasons we are studying these amaz-ing performers. As an evolutionary biology lab at the Uni-versity of California, Santa Barbara, we are interested in the evolution of this complex mating system and the un-derlying genetic mechanisms that regulate the signal. We are also interested in how these species are related to each other, as there are roughly 64 species in the Caribbean and many of them are undescribed. During our time at Carrie Bow, we collected specimens and performed small experi-ments aimed at providing answers to these broad ques-tions.We also got to share some of our knowledge and show offour hard work with a visiting group of students from Cal-

Figure 41 - Belize ostracods.

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Figure 42 - Stomatopod aggressive display, meral spots are the small purple spots on each arm.

vin College who toured the facilities at Carrie Bow Cay. We showed videos and explained our research to a group of about 15 students. They seemed intrigued to look at theanimals and eagerly asked questions about what we were doing.

The Secret Signals in Stompatopod Battles

Amanda Franklin

Stomatopods (otherwise known as “mantis shrimp”) are highly aggressive marine crustaceans with one of the mostcomplex visual systems known in the animal kingdom. While there are numerous studies investigating how and what they see, there are surprisingly few studies address-ing what they use their vision for. Behavioral observa-tions have shown that stomatopods display coloured body patches during interactions with each other, suggesting a role for visual signals in communication. The meral spread is a stance commonly performed during combative encounters and displays the “meral spot” (Figure 42). Thispatch reflects UV light that is invisible to human eyes, butin the spectrum that mantis shrimp can see. I hypothesizethat ultraviolet reflectance of the meral spot is a signal offighting ability. Furthermore, chemical signals could play a role in the outcome of stomatopod interactions. To ad-dress this, I studied the stomatopod, Neogonodactylus oerstedii at Carrie Bow Cay, and experimentally manipu-lated the UV reflectance of the meral spot as well as chem-ical signal detection. Under natural lighting in the field, I filmed conflicts between a resident stomatopod in a bur-row (control vs. reduced UV reflectance) and an intruder (no modification to reflectance). Stomatopods are aggres-sive creatures. They have deadly weapons that can punch as fast as a bullet and they do not hesitate to use them. In fact, I’m quite glad I work with a small species, otherwise I would have needed several stitches by now. They do not only direct their aggression at me, but frequently it is di-rected towards another stomatopod. However, they don’t tend to go straight to the punching, they do some fierce displays first. Like many other animals, they try to make themselves bigger to appear more fearsome. They pull their raptorial appendages (the punching arms) out to the side and lift their body up. If neither agressor nor intruder runs away after this display, then things get more physical. One will move towards the other one, and, if it doesn’t lose its nerve, will try to punch its opponent! In many cas-es one will punch, then quickly curl up into a defensive ball, in case the other one retaliates. Sometimes they will exchange several punches before one admits defeat and runs away. Probably a good move since they are known

to sometimes kill one another in these fights. These inter-actions are over very quickly and there’s always a clear winner. In Belize I spent most of my days watching sto-matopods behaving aggressively which never got boring.

At Carrie Bow, I collected stomatopods from all around the island. I found them by probing small holes in coral, rock or conch shells, and if one was in there they would usually punch the probe. During the three weeks I caught 130 stomatopods – about 70 females and 60 males (Fig-ure 43). It’s pretty common to catch more females than males because males will leave their burrows in search of females. After mating with a female in her burrow she will kick him out and he goes on to search for another burrow. All this searching leaves him vulnerable to predation. Theresult? Males are eaten more often than females.

I take the rubble back into the lab, crack it open and extractthe shrimp. I then record body length, weight, color and take measurements of the meral spot with a spectropho-tometer. After that, all stomatopods are allocated into size-matched pairs. One of these will be the ‘resident’ and one the ‘intruder’. Half the residents have the UV reflec-tance of their meral spot hidden with paint (other half are controls) and half the intruders have their ability to detect chemical cues removed by dipping their antennae in fresh water (other half are controls). These manipulations affect

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Figure 43 - The sea water lab at Carie Bow Cay houses 130 stomatopods used in experimental trials.

the information that the intruder receives (i.e. the intruder will or will not have access to chemical cues and/or UV reflectance cues). In my experiments, I am looking at the behaviors of the intruder. I then put two shrimp together in a tub outside (so lighting conditions are natural), the resident has a burrow to defend (a plastic tube) and the intruder will usually fight the resident over ownership of the burrow. I film all these fights and record the behaviors to see whether the intruder is using UV reflectance or chemical cues in these fights. A variety of behavioral variables are recorded including time to approach, distance of approach, duration of encounter, number of punches and winner of the burrow. Afterwards, all the shrimp are returned to the reef. Hopefully, the results will indicate how important UV reflectance is to stomatopod conflicts.

CCRE Contributions Fiscal Year 2014Open access articles are indicated with clickable link

Bright, M., S. Espada-Hinojosa, I. Lagkouvardos, and J. Volland. 2014. The giant ciliate Zoothamnium niveum and its thiotrophic epibiont Candidatus Thiobios zoothamnicoli: a model system to study interspecies cooperation. Frontiers in Microbiology 5 (145): 1-13. link

Bucher, K.E., D. L. Ballantine, C. Lozada-Troche, and J. N. Norris. 2014. Wrangelia gordoniae, a new species of Rho-dophyta (Ceramiales, Wrangeliaceae) from the Tropical Western Atlantic. Botanica Marina 57 (4): 265–280.

Conway, K.W., C. Baldwin, M.D. White. 2014. Cryptic diversity and venom glands in western Atlantic clinngfishes of the genus Acyrtus (Teleostei: Gobiesocidae). PLoS ONE 9(5): e97664. doi:10.1371/journal.pone.0097664 link

DeBose, J.L., V.J. Paul. 2014. Chemical signatures of multi-species foraging aggregations are attractive to fish. Marine Ecology Progress Series 498: 243–248.

Dimond, J.L., R.R. Pineda, Z. Ramos-Ascherl, and B. L. Bingham. 2013. Relationships between host and symbiont cell cycles in sea anemones and their symbiotic dinoflagellates. Biological Bulletin 225(2): 102-112.

Engene, N., T. Byrum, A. Thor, M.H. Ellisman, W.H. Gerwick, V.J. Paul. 2013. Five chemically rich species of tropi-cal marine cyanobacteria of the genus Okeania gen. nov. (Oscillatoriales, Cyanobacteria). Journal of Phycology 49 (6): 1095–1106.

Kirk, N.L., R. Ritson-Williams, M.A. Coffroth, M.W. Miller, N.S. Fogarty, S.R. Santos. 2013. Tracking transmission of Apicomplexan symbionts in diverse Caribbean corals. PLoS ONE 8(11): e80618. doi:10.1371/journal.pone.0080618 link

Leasi, F., J.L. Norenburg. 2014. The necessity of DNA taxonomy to reveal cryptic diversity and spatial distribution of Meiofauna, with a focus on Nemertea. PLoS ONE 9(8): e104385. doi:10.1371/journal.pone.0104385 link

Mathis, W. 2014. A Revision of the new world species of Paralimna Loew (Diptera: Ephydridae). Smithsonian Contribu-tions to Zoology Number 643.

Mathis, W.N., T. Zatwarnicki. 2013. A revision of the shore-fly genus Hydrochasma Hendel (Diptera, Ephydridae). ZooK-eys 363: 1–161. doi: 10.3897/zookeys.363.6482 link

Meyer, J.L., V.J. Paul, and M. Teplitski. 2014. Community shifts in the surface microbiomes of the coral Porites astre-oides with unusual lesions. PLoS ONE 9(6): e100316. doi:10.1371/journal.pone.0100316 link

Ott, J.A., H. R. Gruber-Vodicka, N. Leisch and J. Zimmermann. 2014. Phylogenetic confirmation of the genus Robbea (Nematoda: Desmodoridae, Stilbonematinae) with the description of three new species. Systematics and Biodiversity 12 (4): 434-455.

Ott, J., N. Leisch, H. R. Gruber-Vodicka. 2014. Eubostrichus fertilis sp. n., a new marine nematode (Desmodoridae: Stil-bonematinae) with an extraordinary reproductive potential from Belize, Central America. Nematology 16: 777-787. link

Pende, N, N. Leisch, H. Gruber-Vodicka, N. Heindl, J. Ott, T. den Blaauwen, S. Bulgheresi. 2014. Size-independent sym-metric division in extraordinarily long cells. Nature Communications 5: 1-10. DOI: 10.1038/ncomms5803 link

Ritson-Williams, R., S. N. Arnold, V. J. Paul, and R. S. Steneck. 2014. Larval settlement preferences of Acropora palmata and Montastraea faveolata in response to diverse red algae. Coral Reefs 33 (1): 59-66.

Ruetzler, K., C. Piantoni, R.W.M. van Soest, and M.C. Diaz. 2014. Diversity of sponges (Porifera) from cryptic habitats on the Belize barrier reef near Carrie Bow Cay. Zootaxa 3805(1): 001–129. link

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Sneed, J.M., K. Sharp, K. Ritchie, V.J. Paul. 2014. The chemical cue tetrabromopyrrole from a biofilm bacterium induces settlement of multiple Caribbean corals. Proceedings of the Royal Society - B 281 (1784), doi: 10.1098/rspb.2013.3086.

Steneck, R. S., S. N. Arnold, and P. J.Mumby. 2014. Experiment mimics fishing on parrotfish: insights on coral reef recov-ery and alternative attractors. Marine Ecology Progress Series 506: 115-127. link

van Tussenbroek B.I., J. Cortés, R. Collin, A.C. Fonseca, P.M.H. Gayle, et al. 2014. Caribbean-wide, long-term study of seagrass beds reveals local variations, shifts in community structure and occasional collapse. PLoS ONE 9(3): e90600. doi:10.1371/journal.pone.0090600 link

Villamizar, E., M.C. Diaz, K. Rutlzer, and R. De Nobrega. 2013. Biodiversity, ecological structure, and change in the sponge community of different geomorphological zones of the barrier fore reef at Carrie Bow Cay, Belize. Marine Ecol-ogy 35(4): 425-435. link

Windsor, A, and D. Felder. 2014. Molecular phylogenetics and taxonomic reanalysis of the family Mithracidae (Decapo-da: Brachyura: Majoidea). Invertebrate Systematics 28: 145–173.

2014 Participants* served as station manager

Acosta, Diana, Boston University, Boston, MA Alanko, Jerry & Sandy, Tilghman, MD*Altieri, Andrew, Smithsonian Tropical Research Institute, PanamaAlvarez, Marcos, Smithsonian Tropical Research Institute, PanamaAngioletti, Christopher, University of California, San Diego, CAAshbert, Miranda, Wildlife Conservation Society, BelizeBagge, Laura, Duke University, Durham, NCBoehm, J.T., University of New York, New York, NYBodart, Jake, University of Maryland, College Park, MDBok, Michael, Lund University, Lund, SwedenBurns, Virginia, Wildlife Conservation Society, Belize Buston, Peter, Boston University, Boston, MA Campbell, Justin, Smithsonian Marine Station, Fort Pierce, FL Canty, Steve, International Center for Marine Ecology, HondurasChak, Solomon, Virginia Institute of Marine Science, VAChernoff, Barry, Wesleyan University, Middletown, CTClough, Wayne, Smithsonian Institution, Washington, D.C.Craig, Catherine, University of Lousiana at Lafayette, LACramer, Katie, University of California, San Diego, CACummings, Erin, Duke University, Durham, NCD' Aloia, Cassidy, Boston University, Boston, MADavid, Alben, Dangriga, BelizeDeignan, Lindsey, University of North Carolina at Wilmington, Wilmington, NC De Gracia Taylor, Brigida, Smithsonian Tropical Research Institute, PanamaDixson, Danielle Georgia Institute of Technology, Atlanta, GADramer, Greg & Joann, Kalispell, MT* Duffy, Emmett, Smithsonian Institution, Washington, D.C.Ellis, Emily, University of California, Santa Barbara, CAEvans, Jonathan, Sewanee: The University of the South, Sewanee, TN Felder, Darryl, University of Lousiana at Lafayette, Lafayette, LA

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Felder, Jennifer, University of Lousiana at Lafayette, Lafayette, LAFelder, Richard, University of Lousiana at Lafayette, Lafayette, LAFieseler, Clare, University of North Carolina at Chapel Hill, Chapel Hill, NCFogarty, Nicole, Nova Southeastern University, Dania Beach, FLFoltz, Zachary, Smithsonian Marine Station, Fort Pierce, FLForde, Alexander, University of Maryland, College Park, MDFranklin, Amanda,Tufts Unviversity, Medford, MAGawne, Peter, New England Aquarium, Boston, MA Gibbons, Johnny, Smithsonian Institution, Washington, D.C.Goodison, Michael, Smithsonian Environmental Research Center, Edgewater, MD Gouge, Daniel, Williston, FL* Haren, Nick & Marylin, Anacortes, WA*Harris, Stephen, City University of New York, New York, NY Hay, Mark, Georgia Institute of Technology, Atlanta, GAHenley, Michael, Smithsonian National Zoological Park, Washington, D.C.Hensley, Nicholai, University of California, Santa Barbara, CAHorning, Reginald, Traverse City, MI*James, Edwin & Bonnie, Tilgman, MD*Jensen, Paul, Scripps Institution of Oceanography, La Jolla, CAJohnston, Lane, Smithsonian Marine Station, Fort Pierce, FL Jones, Scott, Smithsonian Marine Station, Fort Pierce, FLKirwan, John, Lund University, Lund, Sweden Koltes, Karen, U.S. Department of the Interior, Washington, D.C.Kuempel, Caitie, Smithsonian Tropical Research Institute, PanamaLeisch, Niko, University of Vienna, AustriaLeray, Matthieu, National Museum of Natural History, Washington, D.C.Lewis, Sara, Tufts Unviversity, Medford, MALindo, David, The University of Miami Rosenstiel School of Marine and Atmospheric Science, Miami, FLLittle, Mark, National Museum of Natural History, Washington, D.C.Longo, Guilherme, Georgia Institute of Technology, Atlanta, GAMasi, Joe, New England Aquarium, Boston, MAMcKeon, Seabird, Smithsonian Marine Station, Fort Pierce, FLMeyer, Julie, University of Florida, Gainesville, FLMoore, Joel & Linda, Shingle Springs, CA*Nash, Chloe, Wesleyan University, Middletown, CT Noren, Hunter, Nova Southeastern University, Dania Beach, FLNorris, Richard, University of California, San Diego, CAO'Dea, Aaron, Smithsonian Tropical Research Institute, PanamaOakley, Todd, University of California, Santa Barbara, CAOpishinski, Tom, Interactive Oceanographics, East Greenwich, RIPalacios-Theil, Emma, University of Lousiana at Lafayette, LAParsons, Keith & Shirley, Atlanta, GA*Paul, Valerie, Smithsonian Marine Station, Fort Pierce, FL Patin, Nastassia, Scripps Institute of Oceanography, La Jolla, CA Pawlik, Joseph, University of North Carolina at Wilmington, Wilmington, NCPende, Nika, University of Vienna, Austria Penland, Laurie, National Museum of Natural History, Washington, D.C.Peresta, Gary, Smithsonian Environmental Research Center, Edgewater, MDPleijel, Frederik, Gothenburg University, Gothenburg, SwedenPlaisance, Laetitia, National Museum of Natural History, Washington, D.C.Reichert, Mary, Georgia Institute of Technology, Atlanta, GA Riley, Megan, University of South Carolina, Columbia, SCRosebraugh, Sydney, National Museum of Natural History, Washington, D.C.Rotjan, Randi, New England Aquarium, Boston, MA

CCRE Staff:Valerie Paul, DirectorZach Foltz, Station ManagerScott Jones, Program Coordinator

Smithsonian Marine StationCaribbean Coral Reef Ecosystems ProgramFort Pierce, FL · Carrie Bow Cay, Belize

www.ccre.si.eduwww.sms.si.edu

Rouse, Greg, Scripps Institute of Oceanography, La Jolla, CA Safrit, Glen, University of North Carolina at Chapel Hill, Chapel Hill, NC Schmiege, Phillip, Boston University, Boston, MAScolaro, Derek, Boston University, Boston, MASherwood, Craig, Deale, MD*Simpson, Lorae', Smithsonian Environmental Research Center, Edgewater, MDSpadaro, Angelo, Old Dominion University, Norfolk, VASpathias, Hanae, Interamerican University of Puerto Rico, San Juan, PRSpiers, John, Fort Pierce, FL Stewart, Hannah USAStrindberg, Samantha, Wildlife Conservation Society, Belize Studivan, Michael, Harbor Branch Oceanographic Institute, Florida Atlantic University, Fort Pierce, FLTaylor, Jim & Tanya, Oxford, MS*Taylor, Sarah, New England Aquarium, Boston, MATeplitski, Max, University of Florida, Gainesville, FLTewfik, Alex, Wildlife Conservation Society, BelizeThacker, Cheryl, University of Florida, Gainesville, FLTipton, Michelle, Wesleyan University, Middletown, CT Tschirky, John, American Bird Conservancy, The Plains, VA Voss, Joshua, Harbor Branch Oceanographic Institute, Florida Atlantic University, Fort Pierce, FL Vilchis, Nacho, University of California, San Diego, CAVu, Ivana, University of North Carolina at Chapel Hill, Chapel Hill, NCWeber Michelle, National Museum of Natural History, Washington, D.C.Wesby, Dale, Wildlife Conservation Society, BelizeWindsor, Amanda, National Museum of Natural History, Washington, D.C.Wood, Abigail, Smithsonian National Zoological Park, Washington, D.C.Yockachonis, T.J., San Francisco State University, San Francisco, CA

Photograph & Art Credits: Front Cover: S. Jones, p.1 A. Wood, p.6 J. Gibbons, p.7 J. Gibbons, p.8 J. Gibbons, p.9 Z. Foltz, p.10 Z. Foltz (both), p. 11 S. Jones, p. 12 Z. Foltz, p.13 Z. Foltz, p.14 S. Jones, p. 15 S. Jones (both), p. 16 D. Felder, p.17 D. Felder, p. 18 D. Felder, p. 19 OSD Consortium, p. 20 T. Opishinksi, p. 21 P. Buston (both), p.22 G. Longo, p. 23 K. Cramer, p.24 K. Cramer, top left, bottom left, A. Forder, right, p. 25 A. Forde (both), p. 26 F. Pleijel, p. 27 F. Pleijel (both), p. 28 R. Ritson-Williams, p. 29 M. Studivan (both), p. 30 S. Jones, p. 31 S. Jones, p. 32 Z. Foltz, p. 33 M. Bok, p. 34 M. Bok, p. 35 S. Bulgheresi, p. 36 N. Hensley, p. 37 A. Franklin, p. 38 A. Franklin, Back Cover: S. Jones.

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