Bocas del Toro Research Station - Smithsonian Tropical Research ...

Bocas del Toro
Research Station
FY08-FY09
Biennial Report

Inaugurated in 2003, the laboratory building was designed to be environmentally
friendly. Its 10,000 square feet of lab and office space feature skylights that
provide natural lighting and a roof that doubles as a rainwater collector. The
6-hectare grounds include a small area of forest, a mangrove swamp and a lake
that is home to caimans and numerous species of water-birds.
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Bocas del Toro Research Station
The Bocas del Toro Research Station (BRS), a field station
of the Smithsonian Tropical Research Institute (STRI) on
Panama’s western Caribbean coast, is an ideal platform
for both marine and terrestrial research. The station hosts
a diverse group of Smithsonian staff scientists, visiting
scientists and students.
Activities at the station focus on the Smithsonian
Institution’s primary mission: the increase and diffusion
of knowledge. BRS visitors are engaged in research on
the biodiversity, ecology, paleontology, archaeology and
sustainable development of the Bocas del Toro region.
Educational and outreach activities include K-12 school
group visits to the station, development workshops for local
teachers and advanced training in biodiversity research for
graduate students.
Founded in 1998, the BRS is among the preeminent field
stations in the Caribbean. The campus provides field
accommodations, a fully operational research laboratory,
SCUBA facilities and a fleet of small boats to more
than 300 visiting scientists each year. A new dormitory
is planned to accommodate the growing number of
researchers.
Table of contents
Bocas del Toro Research Station 03
Research Highlights 11
Outreach, Education and Training 27
BRS Visitors and Usage 33
3 - Year Bibliography 49
Acknowledgments 56
Financial Contributions 57
Make a Donation 58

1970/1980’s
1990’s
1998
1999
2001
2002
2003
2004
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Charles Handley (1924-2000), curator of mammals at the Smithsonian’s National Museum of
Natural History, surveyed the mammal fauna of the Bocas del Toro region.
Anthony Coates, then STRI’s deputy director, coordinated the Panama Paleontology Project’s
work in Bocas del Toro to document the rise of the Isthmus of Panama.
STRI purchased six-hectares on Isla Colon for the development of a permanent research station
on the Caribbean. During the next several years, representatives from STRI met with Bocas
authorities and community representatives to present STRI’s mission and plans for the area and to
solicit feedback.
STRI joined CARICOMP (the Caribbean Coastal Marine Productivity Program) to monitor coral,
mangrove, and seagrass productivity near the BRS.
A dormitory to house 16 people was completed. The station began hosting visiting scientists as
well as STRI scientists and students.
STRI appointed Rachel Collin, a staff scientist
studying the evolution and life histories of marine
gastropods, as the first director of the BRS.
The state-of-the-art laboratory building was inaugurated in October. Then Smithsonian Secretary
Lawrence M. Small and Panama’s former president, Mireya Moscoso de Arias, were present. The
Mellon Foundation funded an Organization for Tropical Studies (OTS) course in Tropical Marine
Ecology. The first invertebrate biodiversity workshop, supported by the Smithsonian Institution’s
Marine Science Network and the Smithsonian Women’s Committee, documented exceptional
marine biodiversity. STRI appointed Gabriel Jacome as BRS station manager.
The seawater system was completed,
greatly increasing capability for
experimental research. A second
invertebrate workshop added to the
catalogue of local marine biodiversity.

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An issue of the Caribbean Journal
of Science featured the fauna and
environment of Bocas del Toro. The first
Training in Tropical Taxonomy course
instructed 14 international students in
taxonomy and systematics of sponges.
Casa Hoch and Casa Cofrin were completed, increasing the station’s capacity to 24 resident
researchers. The SI Board of Regents and SI’s Women’s Committee visited the BRS. The Bocas
del Toro biodiversity database reached 3,000 entries.
Donations from the Cofrin and Hoch families enabled the design of a new dormitory and common
area. The SI National Board visited the BRS. The original sawmill building was demolished and
the reception area was remodeled. BRS publications topped 200.
The Bocas del Toro biodiversity database exceeded 6,000 entries and 8,000 photographs. The BRS
hosted the first annual Bocas del Toro Feria Ambiental, bringing together over 20 local NGOs
and governmental agencies dedicated to environmental concerns.
SI Secretary G. Wayne Clough and SI Undersecretary for
Science Charles Alcock visited the station. Panama’s Marine
Environmental Authority (ARAP) declared the coastline
adjacent to the BRS as a special use area for research and
education.
The BRS Training in Tropical Taxonomy program received
support from the U.S. National Science Foundation (NSF)
for a PanAmerican Advanced Studies Institute on tropical
phycology.
STRI’s construction team finalized plans for new dormitory
and dining facilities and remodeled the laboratory to provide
more space for the increasing number of scientific visitors.
2005
2006
2007
2008
2009

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Bocas del Toro Research Station

Biologically rich marine and terrestrial habitats in the
province of Bocas del Toro, on the Western Caribbean
coast of Panama, provide BRS visitors with a wealth of
field opportunities. The complex geography of the region
contributes to its unique biodiversity and easy access to
many habitat types.
Mangrove-fringed mainland peninsulas and islands,
seagrass beds, and patch reefs delimit two distinct bays:
the Bahía Almirante and the Laguna de Chiriquí. Reefs
are best-developed in the Bahía Almirante. The Laguna de
Chiriquí, which receives most of the terrestrial run-off, is
dominated by seagrass and mangrove ecosystems.
The Bocas del Toro mainland is covered by montane and
lowland humid tropical forest and banana plantations.
There are large expanses of peat-swamp forest along the
coast of the Bahía Almirante. Deforestation is common on
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the islands of the archipelago, which are primarily covered
by secondary forest, pasture, and teak plantations. Despite
this, some old-growth forest still remains on Isla Colon and
Isla Bastimentos.
Bocas del Toro holds vast fossil deposits. Fossil reefs and
other outcrops from the last 18 million years are common
throughout the area. Fossil molluscs, foraminifera,
bryozoans, and corals are especially well studied.
Easy access from the BRS to these numerous habitats
and rich outcrops has enabled a diversity of studies aimed
at understanding the environmental changes associated
with the rise of the Isthmus of Panama, subsequent
environmental changes, and the resulting evolutionary
divergences between the Pacific and Caribbean biotas.
The BRS is located on a narrow isthmus between the town of Bocas del Toro and the majority of Isla Colon. The exposed coast of the island (bottom
left) is fringed by a sandy beach lined by sea-grape and sea-almond trees. The more protected Bahia Almirante side (top right) is fringed with red
mangroves and seagrass beds. The dock, automated data station for environmental monitoring and the seawater intake pile are on the bay side of the
station.

The region around the BRS receives about 3-5 meters of
rainfall annually. Temperature and rainfall do not show
the pronounced seasonal patterns typical of the Pacific
Coast of Panama. To allow comparisons with other
STRI sites around Panama, Bocas del Toro is the focus of
considerable environmental monitoring. Weekly and daily
measurements of rainfall and air and sea temperature are
taken at the research station.
Recent analysis of the BRS’s water quality monitoring data
by station director Rachel Collin has shown that despite the
continued deforestation on the mainland and development
in the islands, water quality has not shown any significant
decrease over the last 10 years. However, clarity and
chlorophyll concentration hover near the levels indicative
of eutrophication.
Seventy-one coral sites, 52 seagrass sites, and 5 mangrove
sites have been surveyed and there are several permanent
monitoring sites in each habitat. The BRS is also part
of CARICOMP, a Caribbean-wide program to monitor
the productivity of coral reefs, mangroves and seagrass.
Standardized CARICOMP monitoring protocols are
followed in all three habitats.
Casa Hoch and Casa Cofrin accomodate course professors and scientific
visitors who bring their families.
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Extensive mangroves, peat swamps, and other wet low-lying habitats
provide ideal environments for caimans (pictured) and American
crocodiles. Several caiman are year-round residents at the BRS,
delighting visiting researchers and school children alike.
The explosive success of the BRS and the rapidly
increasing number of scientific visitors is evident in the
statistics on station usage. This continued growth has
prompted plans for additional housing. With generous
donations from the Cofrin and Hoch families, a design is in
place for an additional dormitory and dining hall complex
that will increase the station’s capacity to house visitors by
nearly fifty percent.
The 2009 STRI master plan calls for the future construction
of facilities dedicated to outreach and education as well as
expanded dock and maintenance areas.

The BRS 7-boat fleet supports marine research and moves terrestrial
workers between remote islands.
Matumbal Bay Special-Use Zone
On December 22 2008, Dr. Reynaldo Pérez Guardia,
General Administrator of Panama’s Marine Resources
Authority (ARAP), signed Administrative Resolution
No. 10 establishing the Matumbal Reserve. Located
at Matumbal Bay in the Bahia Almirante, the reserve
is designated for research and education. STRI will
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SCUBA diving is an important tool even in shallow water. The BRS
accounts for more than 70 % of the dives at STRI’s facilities.
collaborate with ARAP to monitor and manage the
reserve. A socioeconomic analysis of its area of influence
and surveys of marine resource diversity and usage are
underway. These, in conjunction with environmental data
from other BRS biological surveys, will contribute to a
new management plan.
Bocas del Toro
Research Station
The Matumbal Bay reserve extends from the Saigon neighborhood to the edge of the Sunset Point development
project and includes 34.21 hectares of marine habitats, primarily reefs and mangroves.

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Working with frogs means getting wet: Andrea Rudh and Björn
Rogell, graduate students from Uppsala University in Sweden, study
the evolution and genetics of color variation in strawberry poisondart
frogs, Oophaga pumilio in the forest fragment on BRS property.

Research highlights
Research at the BRS ranges from deep time to real time, from tree-tops to the
ocean’s depth, and from ephemeral microbial communities to ancient corals.
Most projects at the BRS are conducted in a comparative
framework and center on the goals of the Smithsonian
Marine Science Network: understanding spatial and
temporal patterns of marine biological diversity and the
effects of human activities. A smaller group of studies
focus on physiology, development, behavior of a single
species or on applied topics.
Many of the projects at the BRS are linked by common
themes. The impacts of local development and global
anthropogenic change are important foci. Ana Spalding’s
work on re-making lives abroad takes an anthropological
perspective, while change in land-use and land reforms are
studied by Gayatri Thampy. Surveys to detect and identify
exotic or invasive species in marine fouling communities
are conducted by SERC’s Greg Ruiz and STRI’s Mark
Torchin and colleagues. Long-term anthropogenic changes
in reef community composition are being described by
Katie Cramer.
Major efforts to document the biodiversity of the southern
Caribbean are based at the BRS. An NSF-funded inventory
Fossilized shell deposits. Uplift pushed deep
sea sediments to the surface during 20 million
years, creating the Panama land bridge, which
linked two continents and separated the Pacific
from the Caribbean.
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A mantled howler monkey troupe often visits
the station. The BRS is only a distance from
several high biodiversity protected areas on
islands and on the mainland.
of Panamanian macroalgae is spearheaded by Brian Wysor,
Wilson Freshwater, Suzanne Fredericq and Jim Norris.
Extensive surveys of marine and freshwater shrimps by
Arthur Anker and Sammy de Grave show that Bocas del
Toro is one of the top-ten global sites for shrimp diversity.
Rosana Rocha has shown that Bocas del Toro has the
highest diversity of tunicates of any site in the Caribbean.
Finally, the NSF-funded Porifera Tree of Life project
headed by Bob Thacker is running a series of workshops on
sponge phylogeny.
In FY08-FY09 the BRS hosted 436 scientific visitors.
They, their projects, and their publications are listed at the
end of this report. Because the more than 100 ongoing
projects at the BRS are too numerous and diverse to
describe in full here, this section highlights the work
of students and post-doctoral researchers who received
fellowships from the Smithsonian Institution and STRI, and
projects of visiting researchers in two important thematic
areas: 1) Anthropogenic impacts and conservation in Bocas
del Toro and 2) Night Life in Bocas.
Fungal endophytes isolated from cacao
leaves—yet another source of cryptic
biodiversity in Bocas del Toro. These
fungi may play an important role in plant
protection.

SI/STRI fellows
Digging underwater is hard work.
Katie Cramer uses SCUBA as
she carefully removes layers of
sediment and coral skeletons
to determine the effects of the
history of land use on reef heath.
An oil-drum helps Cramer standardize the size and depth of the
excavation.
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KATIE CRAMER
STRI Short Term Fellow (2007), STRI Predoctoral
Fellow (2008-2009)
Historical changes in Panama’s Caribbean coral
communities: the land-use connection.
Sadly, Caribbean coral reefs are among the most degraded
reefs on the planet. Corals, the physical and ecological
foundation of reef ecosystems, have declined greatly in
abundance during the past fifty years. This decline is
due, in part, to the influx into reef waters of land-based
sediments and pollutants from activities such as agriculture
and urbanization. These materials smother the corals,
thus negatively impacting their feeding, growth and
reproduction.
To determine the state of coral communities before the
onset of major land-based human disturbances, Cramer
studies reefs in Bocas del Toro (western Panama) and Bahia
Las Minas/Costa Arriba (central Panama). Her sites span a
range of influence from land-based runoff.
Cramer digs through the sediments surrounding living
reefs, and collects calcium carbonate skeletons of dead
colonies from the past several centuries in order to
“reconstruct” coral communities by developing lists of their
former inhabitants.
Rubble from four distinct layers is classified into species
categories and radiocarbon dated to describe community
change over time. Chemical analyses of sediments within
each rubble layer link these changes with the reefs’ relative
exposure to land-based runoff. Cramer also surveys
contemporary reef communities for comparison.
Results from a preliminary study show a dramatic
decrease in coral diversity and in the relative abundance
of branching species, accompanied by an increase in
the relative abundance of weedy, low relief species and
disturbance-tolerant massive species during the past fifty
years. Influence from land-based runoff guaged by distance
from the mainland and river mouths partially explain the
changes in dominance which have occurred for the most
abundant species.

AMY FREESTONE
Smithsonian Institution Postdoctoral Fellow (2006),
Smithsonian Marine Science Network Postdoctoral
Fellow (2007-2009)
Does predation in a marine system differ between
temperate and tropical latitudes?
Eat or be eaten: predation is one of the major ways that
species interact. The idea that species interactions could
be more intense in the tropics than in temperate areas was
proposed to explain higher tropical biodiversity. Amy
Freestone compares how species interaction-strength varies
with latitude, working at Smithsonian marine stations along
the Atlantic coast from Maryland to Panama.
Freestone deploys plates-- artificial “habitats” which are
naturally colonized by tunicates, cnidarians (coral, sea
anemones, hydroids), sponges, bryozoans, tube-building
worms and other organisms--across 32° of latitude, from
the temperate zone to the tropics. Different levels of
predation are created by caging the plates to exclude
predators.
Freestone’s results indicate a clear shift in the impact
of predation on community diversity from temperate to
tropical sites. Tropical communities are nearly three times
more diverse in the absence of predators, indicating that
predators are a very strong selective force here. While
overall predation effects were strong and consistent in
the tropics, predation pressure often differed significantly
between habitats. In contrast, diversity of temperate
communities did not vary with predation pressure, although
relative abundances of certain taxa (e.g., tunicates) were
strongly affected.
Her results suggest that even if predation is an important
community driver at all latitudes, it has the greatest impact
on species diversity patterns in the tropics. In collaboration
with scientists both at STRI and at the Smithsonian’s
Environmental Research Center in Maryland, Freestone
will study the way biological processes affect the success or
failure of marine invasions.
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a b
Three-month-old communities that had developed in the absence of
predators were exposed to natural predators in the field. Diversity of
sessile invertebrates declined significantly after only one week. Photos
show the same community from Bocas del Toro before (a) and after (b)
exposure to predators.
Diverse sessile invertebrate communities developed on experimental
panels from which predators were excluded. This panel shows a
community from Bocas del Toro.
Experimental panel
with predator exclosure
deployed near Bocas del
Toro Research Station.

Aminur Rahman studies the evolutionary mechanisms that lead to
speciation and high tropical biodiversity.
Two very similar species of sea urchins live in the same habitat and
reproduce at roughly the same time. In fact, they can even interbreed,
producing hybrid offspring. So how do they maintain their identities as
individual species?
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AMINUR RAHMAN
Smithsonian Marine Network Postdoctoral Fellow
(2007-2008), Smithsonian National Board Fellowwith
funds from the Hoch Family (2008-2009)
How do two closely-related sea urchin species
maintain separate identities?
In 2009, scientists and friends celebrated Darwin´s 200th
birthday. Darwin proposed that small differences between
organisms make some more capable of surviving and
reproducing and result in the evolution of unique new
species. Two species of sea urchins that belong to the same
genus live in close proximity in the Caribbean.
From a genetic point of view, Lytechinus willamsi and
Lytechinus variegatus are quite similar. They produce
sperm and eggs at roughly the same time and they can
fertilize each other. However, a molecule called bindin
involved in sperm recognition is distinct in the two
species and is one of the characteristics researchers use
to distinguish them. What mechanisms maintain their
separate species identities? One possibility is that hybrids,
the offspring of pairings between the two species, do not
survive and reproduce as well as the offspring of each
parent species do.
Aminur Rahman visited Bocas del Toro to find out whether
the mechanisms resulting in the isolation of these two
species are at the mating stage, or later in the lives of the
urchins. He set up a series of experiments, rearing the
offspring of hybrid crosses to see how their survival and
reproduction compares to that of each parent species. He
succeeded in producing a huge number of larvae and
juveniles.
So far, it looks as if the offspring of the hybrid crosses do
not survive as well as the offspring of the crosses within
Lytechinus variagatus, but do better than the offspring of
crosses within L. williamsi. There is no answer yet, but
Rahman and other evolutionary biologists will continue to
work in Bocas to untangle explanations for the origins and
maintenance of tropical biodiversity.

KRISTIN HULTGREN
Smithsonian Marine Network Postdoctoral Fellow
(2007-2008)
Phylogenetically conserved niches and the
biodiversity of symbiotic shrimp.
How do so many different species originate and how do
they successfully coexist in diverse tropical ecosystems?
The diverse shrimp species coexisting within individual
sponges in coral reef ecosystems may provide important
clues. Kristin Hultgren tests two opposing hypotheses for
species coexistence in a diverse clade of sponge-dwelling
tropical snapping shrimps, Synalpheus.
“Habitat filtering” is an idea to explain why coexisting
species of shrimp should be closely related, with similar
adaptations that suit them to their common sponge habitat,
whereas “competitive exclusion,” posits that coexisting
species should be distantly related, because close relatives
are more likely to exclude one another within a sponge.
Shrimp community data from sponges collected at
Smithsonian field stations in Bocas del Toro and Belize as
well as in Barbados, Curaçao, and Jamaica strongly support
the habitat filtering hypothesis.
Synalpheus in sponges are more closely related than
expected. For at least one sponge, Lissodendoryx
colombiensis, within-sponge niche partitioning may
facilitate coexistence between closely related Synalpheus
species: larger species tend to live in larger holes near the
center of the sponge and prefer larger holes in habitatchoice
assays in the laboratory, and smaller species prefer
smaller holes that occur near the exterior of the sponge.
Although it has long been thought that this group speciated
mainly by host-related divergence driven by competitive
exclusion of close relatives, these results suggest that
alternate mechanisms may drive its rapid radiation.
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Collecting sponges at Punta Caracol in Bocas del Toro.
Synalpheus bocas, a new species of Caribbean sponge-dwelling snapping
shrimp in the gamberalloides group recently described from Bocas del
Toro by Anker and Toth (2008). Synalpheus shrimp have an enlarged
major chela used for fighting, and sponge-dwelling species have a
streamlined body shape well-suited to life in sponge canals.
Close-up of the sponge Lissodendoryx colombiensis, showing sponge
canals used by the shrimp.

How did striking differences in skin coloration of strawberry dart-poison
frogs come about? Do color patterns change based on mate choice
behavior? Do frog predators prefer one color pattern over another?
Corrine Richards raises frogs and their tadpoles in hundreds of small
terraria lining the walls of the lab building.
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CORRINE RICHARDS
Smithsonian Post-doctoral Fellow (2007-2009)
Why are frogs from different islands different
colors?
Strawberry dart-poison frogs, Oophaga pumilio, on the
islands of Bocas del Toro and the adjacent mainland exhibit
an amazing array of skin color patterns. How did such a
stunning example of biological diversity come about?
Previous researchers have suggested that the geographical
differences in frog coloration in Bocas del Toro probably
evolved since the archipelago took on its present form only
about 6,000 years ago. Given their recent origins, their
geographic proximity and their unusual mating and parental
care behaviors, these frogs provide a unique opportunity to
observe speciation in action.
As their name implies, dart-poison frogs have toxic skin.
Toxic animals use bright colors to warn predators that
eating them could be dangerous. Perhaps more brightly
colored frogs evolved on islands with more frog predators.
However, different color patterns also could have come
about as rivals compete for eye-catching mates.
Corrine Richards sets up breeding experiments among frogs
collected throughout Bocas del Toro. Are hybrid offspring
of differently colored frogs healthy? Can they reproduce?
How do they compare to the offspring of frogs from the
same place? Left to their own devices, do the frogs choose
mates showing different color patterns? Richards hopes to
find out whether a frog’s mate choices are learned or innate
by moving tadpoles to different parents.
Richards also keeps an eye out for the chytrid fungal
pathogen that probably caused severe frog die-offs in
upland areas throughout Central America.

COLIN QUINN
Smithsonian Short Term Fellow (2007-2008)
Arsenic accumulation: what does it signify for
plant function and ecology?
Some plants accumulate unusually large amounts of toxic
metals such as copper, zinc, nickel, selenium or arsenic,
a phenomenon known as metal hyperaccumulation.
The Dixie silverback fern, Pityrogramma calomelanos,
accumulates arsenic. Colin Quinn would like to know
more about the mechanisms of arsenic accumulation in
plants and how toxic compounds affect the fern’s ecology.
Ultimately--what is the role of arsenic hyperaccumulation
in the ecosystem?
Hyperaccumulated nickel, zinc, and selenium defend
plants against pathogens and herbivores. Plants may have
evolved to become metal hyperaccumulators because plants
containing higher levels of heavy metals survive attacks by
insects or browsers and by pathogens, whereas unprotected
plants succumb. Quinn will investigate the protective
effect of arsenic against herbivory.
Arsenic hyperaccumulation is facilitated by organic
phosphorous compounds, inositol phosphates, which are
secreted from the roots of the ferns. Phosphate compounds
may release arsenic from the soil making it more available
to the plant.
The origins of different chemical forms of inositol
phosphate in soils, which often account for a large fraction
of soil organic phosphorous, remain unknown. One goal
of Quinn’s research is to see whether the forms of inositol
phosphates secreted from Dixie silverback fern roots
contribute to important levels of this compound found in
the soil, a secondary effect of arsenic hyperaccumulation
which may be important on an ecosystem level.
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Colin Quinn studies the role of heavy metal accumulation in plants.
When fern roots secrete phosphate compounds, does that increase the
plant’s ability to take up arsenic?
The Dixie silverback fern, Pityrogramma calomelanos, is a perennial
plant native to the tropics of Central and South America, which has
spread to other tropical regions around the world. The undersides of the
fronds of the genus Pityrogramma characteristically have powdery white
or yellow ‘dust,’ making them easy to identify in the field.

Gayatri Thampy interviews heads of households on Isla Colon to
understand how land tenure is changing. She plans to extend her study
to neighboring islands.
Children playing among mangrove trees in one of the newly settled areas
on Isla Colon. How does the influx of foreign investment and tourism
influence the lives of the traditional residents of Bocas del Toro?
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GAYATRI THAMPY
STRI Short Term Fellow (2008-2009)
Land use and land tenure reforms in Bocas del
Toro, Panama.
Since the early 1990s Panama has enacted a series of
land reforms to encourage citizens to seek formal, private
land-tenure rights. The Bocas del Toro archipelago
has been targeted by the Panamanian government and
by international finance organizations for reforms that
encourage direct foreign investment and tourism. The
result, in many cases, has been increased land insecurity for
rural people. However, the effects of these policies vary
from household to household. Gayatri Thampy would like
to know why there is variation in the way people respond
to reforms and how their decisions articulate with macroeconomic
structures.
Currently, her research focus is Isla Colon, where she
interviews household heads from each locality and street.
She examines the effect of each household’s location,
land-tenure rights, social capital and ethnic affiliation on
decisions pertaining to changes in land-tenure decisions,
land-alienation decisions, and forms of resistance to land
reform and tourism development. She places her results
in a political-economy framework to understand whether
decisions concerning changes in land-tenure rights relate
to socio-economic levels and ethnicity, both of which may
mediate access to information and resources needed for
changing tenure rights.
As her study has progressed, she has become interested
in the process of colonization and establishment of tenure
claims in new localities by residents displaced by land
conflicts and by incoming migrant wage laborers from the
mainland. She plans to extend this study to the neighboring
islands of Bastimentos, Carenero, San Cristobal and Cayo
de Agua.

Research Focus 1: Human Impacts and Conservation
LAURA MAY-COLLADO
George Mason University, University of Puerto Rico
Biology of bottlenose dolphins.
Coastal dolphins live in an aquatic environment strongly
influenced by nearby activities on land.
Laura May-Collado studied dolphin communication for
her PhD thesis, comparing bottlenose dolphin populations
in Costa Rica’s Gandoca-Manzanillo Wildlife Refuge with
populations in Bocas del Toro. Each environment has a
unique “soundscape” or acoustic context. Dolphins in Costa
Rica experience loud wave action, a low-frequency sound,
whereas dolphins in Bocas, where tourism leads to heavy
boat traffic, experience high-frequency motor sounds.
Bottlenose dolphins whistle in several social contexts.
Whistles appear to play an important role in individual
recognition, bonding and communication among group
members. Dolphins in Costa Rica emit short, low
frequency whistles. Dolphins in Bocas tended to emit
longer, high-frequency whistles, particularly when several
tourist boats were present. May-Collado proposed that
variation in whistle structure shows how dolphins adapt
to local and changing habitat conditions, resulting in
differences between populations at different geographic
scales.
With colleagues at the University of Puerto Rico, she
will set up an Aquatic Automated Remote Biodiversity
Monitoring Network based on passive acoustic technology
which will make 24-hour monitoring of marine
biodiversity, ambient noise, dolphin presence and acoustic
behavior via ethernet possible.
Since 2004, May-Collado’s team has observed and photoidentified
nearly 100 dolphins. She will use social-network
theory to understand their complex behavior. Finally, she
hopes to predict future threats to dolphin populations and
to provide useful scientific information to marine managers
and conservation groups.
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Laura May-Collado records the sounds of bottlenose dolphins and the
ambient sounds, such as wave action or motorboat noise, that affect their
ability to communicate.
Bottlenose dolphins, Tursiops truncatus, will be monitored 24 hours
a day by a new Automated Remote Biodiversity Monitoring Network
designed to better understand their complex social behavior and their
responses to environmental hazards.

Anne and Peter Meylan processing blood serum samples from hawksbill
turtles captured in nets at the Zapatilla Cays. These samples will be used
to determine the concentration of circulating testosterone.
Peter Meylan using laparoscopy to examine the gonad of a live hawksbill
turtle. Laparoscopy allows the determination of gender, maturity and
reproductive status of mature animals. Local anesthesia is used before
the 5mm scope is introduced into the body cavity.
MY 505, a 73 cm adult male hawksbill turtle, Eretmochelys imbricata,
captured in a net at the Zapatilla Cays in 2003, 2004, 2006 and 2008.
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PETER and ANNE MEYLAN and field team
Florida Fish and Wildlife Conservation
Commission and Eckerd College
Ecology and migration of sea turtles of Bocas del
Toro Province, Panama.
Since 1979, this project has contributed to the conservation
of marine turtles in Bocas and the Caribbean by improving
understanding of the biology of green turtles, hawksbills,
loggerheads and leatherbacks. Early years of field work
were spent in aerial and ground reconnaissance of nesting
beaches and foraging areas throughout Bocas del Toro.
Every year since 1990, the Meylans have spent a month
sampling sea turtles in the waters of the Bastimentos Island
National Marine Park. Most sea turtle studies focus only
on females on the nesting beach. The “in-water” sampling
with traditional tangle nets used in this project allows
access to males, juveniles and mating pairs.
Netting studies suggest that each of the four species
of sea turtles uses park waters for a different set of life
history stages. Netting makes it possible to study the
biology of reproductive male and female green turtles en
route to the nesting beach at Tortuguero, Costa Rica, as
well as that of reproductive hawksbills that nest in the
park, itself. Laparoscopy and hormone assays reveal
the gender, maturity and reproductive condition of the
animals. Genetic studies along with satellite tracking and
flipper-tagging provide a better understanding of sea turtle
populations on a regional scale.
This project contributes to a larger effort coordinated by the
Caribbean Conservation Corporation to recover the nesting
population of hawksbill turtles that once flourished along
the Bocas coastline. From May to November, they monitor
sea turtle nesting on the three most important beaches in
the park: the two Zapatilla Cays and Playa Larga. Using
a standard method, CCC, under the direction of Cristina
Ordoñez, monitors hawksbill nesting at Chiriqui Beach,
Escudo de Veraguas and Playa Colorado. Standardized
monitoring of beaches across the province reveals that
hawksbill conservation efforts are paying off.

ROSANA MOREIRA DA ROCHA
Universidade Federal do Paraná, Brazil
Tunicates as models for biological invasions.
Tunicates, or sea squirts, are surprisingly diverse in
Bocas del Toro. Rocha surveyed tunicates during the 1st
Invertebrate Taxonomy Workshop in 2003. Her list of 58
species, including many species new to science and unique
to the area, makes Bocas one of the five most diverse sites
for tunicates in the Caribbean. In 2008, Rocha returned for
a year’s sabbatical to collect more material and publish new
species descriptions.
Her observation that some species in Bocas are widely
distributed led her to new studies using tunicates as a model
for biological invasions. Bocas is growing quickly: more
artificial structures, which often support exotic species, are
being submerged. Passing ships carry tunicates and other
invertebrates on their hulls.
Experimental plates deployed at five sites revealed more
diversity and more colonial species near Bocas town.
Ascidia sydneiensis and Styela canopus, both likely to be
introduced species, were common at all sites. Species on
artificial substrates close to natural mangrove habitats were
not very different between sites, indicating that artificial
substrates could act as stepping stones for introduced
species that invade natural habitats.
Initial studies of salt tolerance of species in the Order
Phlebobranchia revealed that some tunicates could control
ion exchange and cellular volume in salinities as low
as 15 parts per thousand indicating that these species
have the potential to survive under different salinity
conditions. Additional experiments will reveal behavioral
and ecological differences between native and introduced
species.
In 2009, Rocha was an instructor in the BRS Taxonomy
and Biology of Tunicates course which contributed greatly
to the known biodiversity of tunicates in the region. She is
planning to publish a field guide to the tunicates of Bocas.
21
Rocha snorkels among mangrove roots and dock pilings to deploy her
settlement plates.
Sandwich of experimental plates deployed for two months shows
significant growth of tunicates inside the sandwich. Heavy sedimentation
usually prohibits tunicate growth on top of the sandwich.
Styela canopus is a globally distributed species probably introduced to
the West Atlantic by ship transport.

Eric Nyman and Sebastien Tilmans build a model treatment tank that
collects household wastewater and produces methane gas that can be
used for cooking.
Tilmans works closely with community residents and government
agencies to design affordable treatment systems that meet local needs.
22
SEBASTIEN TILMANS
Graduate Student, Stanford University
Fulbright Fellowship (2007-2008); STRI Research
Fellow (2009)
Building sustainable wastewater treatment systems.
Bocas del Toro is a nexus of human and ecological
diversity. Indigenous people, Europeans, banana growers,
Afro-Antilleans, tourists and US expatriates create an
eclectic mix of isolated communities whose major link is
the water around them. Recent explosive development
undermines environmental processes and threatens the
natural beauty that attracts people to the region.
Sebastien Tilmans works with local individuals and groups
to tailor low-impact, decentralized methods for watershed
conservation and restoration to the hydrologic and
ecological conditions of Bocas del Toro. A 2007-2008 pilot
project yielded a comprehensive USAID-funded project
to restore damaged watersheds in Old Bank, a community
on Bastimentos Island. The project includes installation
of wastewater treatment systems, ecosystem restoration
and reforestation. The project is part of a master plan for
watershed management elaborated in collaboration with
the community of Old Bank. Ongoing monitoring of the
project will document and evaluate the project’s benefits for
local ecosystems and public health.
Because Bocas is growing, treatment solutions can often
be incorporated into present development, preventing the
emergence of a pollution problem instead of reacting to it
with difficult retrofits later. The current project proposes
an example of appropriate conservation and ecosystem
protection, which could be adapted in other sensitive parts
of the archipelago.
Water scarcity and pollution are growing global challenges.
Tilmans believes that as the diversity of natural ecosystems
is the source of their resilience, the plethora of different
human cultures empowers us. Solutions to the future’s
complex environmental challenges must be multi-faceted
yet elegant, working in symbiosis with local culture and
biodiversity.

Research Focus 2: Night Life in Bocas
DON LEVITAN, NANCY KNOWLTON
Florida State University, Smithsonian National
Museum of Natural History
How do coral species maintain their identities
during reproduction?
During dramatic coral spawning events, eggs and
sperm from different coral species may mix in the
water. Research divers, primarily from the Smithsonian
Institution, Florida State University and Scripps Institute of
Oceanography, led by Don Levitan and Nancy Knowlton,
have monitored spawning yearly since 2000 at a permanent
site on a reef off of Solarte Island to understand how coral
species maintain their identities during this orgy.
In Panama, the Montastraea annularis coral species
complex -- three closely related coral species -- spawn
shortly after the full moon in September. Across the
Caribbean M. franksi spawns on average 100 minutes past
sunset while M. faveolata and M. annularis spawn 200
minutes past sunset. The latter two species produce eggs
and sperm that cannot cross-fertilize, but M. franksi, the
early spawning species, produces eggs and sperm that are
completely compatible with those of the late spawning
species M. annularis. Is this subtle difference enough to
maintain the distinct identities of these two species?
Divers mark corals that are preparing to spawn with red
chemical lights. Spawning brings the red lights to the
surface. A chase boat collects floating eggs and records its
position with a GPS. In the lab the team determines the
fraction of fertilized eggs.
Coral colonies appear to spawn within the same five-minute
window each year! Cues from lunar and sunset times,
probably modified by chemical signals released by nearby
corals, seem to affect the synchronicity of the release.
The team has conducted surveys at other sites around the
Caribbean to compare the behavior of the same species
under different conditions. Will climate change and local
human development influence coral health and identities
as coral reefs become fewer and farther apart? Continued
long-term monitoring will provide some of the answers.
23
Every year in early September coral researchers arrive to monitor
dramatic, synchronized coral spawning events that usually occur after
the first full moon. Back row: Maria Adreani, Hironobu Fukami, Don
Levitan, Nancy Knowlton, Pablo Munguia, Davey Kline and Neilan
Kuntz . Front row: Javier Jara, Jodi Grayson, Tamara McGovern and
Nicole Fogarty.
Coral colony releasing eggs. Three closely related coral species may
spawn within the same five-minute window each year. What cues
determine the timing of the release?

Nectar-drinking may have evolved several times among the leaf-nosed
bats. This nectar-feeding bat, Lonchophylla robusta, lives in caves in
Bocas del Toro.
The spear-nosed bat lives in harems in caves. Bat researchers discovered
that these bats produce the longest echolocation signal ever recorded.
This wide variety of signals may explain why the leaf-nosed bat group,
which this species belongs to, is so successful.
24
ELISABETH KALKO, MAURICE THOMAS and
students
STRI, Palm Beach Atlantic College, University of
Ulm, Odense University
Bat diversity and behavior.
The late Charles O. Handley, Jr., from the Smithsonian’s
National Museum of Natural History, first cataloged bat
diversity in Bocas del Toro in the 1950’s. Maurice Thomas
has continued to study bat distribution and conservation in
this changing Caribbean coastal landscape. In 2009, STRI
staff scientist Elisabeth Kalko, her students and colleagues,
joined Thomas for a month of bat research.
To complement previous work on the variability in
echolocation-call design in New World leaf-nosed bats, the
Phyllostomidae, they focused on the greater spear-nosed
bat, Phyllostomus hastatus, which forms large harems in
cave systems on Isla Colon. By recording echolocation
calls they discovered that P. hastatus emits species-specific
ultrasound signals. Signal duration is highly variable,
ranging from short calls when the animals leave a cave, to
the longest calls ever recorded for phyllostomid bats when
the bats fly high in the sky. Their observations confirm the
high behavioral plasticity in this family, which is likely to
be one of the main factors leading to its successful radiation
and ecological diversity.
Lonchophylla robusta, a specialized nectar-feeding bat,
belongs to a small group of bats characterized by distinct
tongue morphology. It may have a nectar-drinking
mechanism very different from the majority of Neotropical
nectar-feeding bats. The team used high-speed video to
analyze tongue movements during nectar extraction and
evaluated the amount of nectar bats drank per unit hovering
time in flight cage experiments.
Bat researchers will continue to study the ecology and
behavior of one of the most diverse groups of mammals in
the tropics in order to better understand how such diversity
arose and is maintained.

BRYSON VOIRIN
Max Planck Institute for Ornithology
Sleep in the three-toed sloth, Bradypus variegatus.
Why sleep? Even though sleepiness increases the time
it takes to respond to danger, every animal ever studied
spends at least some time in this non-responsive state. An
electroencephalogram (EEG) is a measure of brain wave
activity that tells us whether an animal is awake or asleep.
Until recently, the size of EEG devices made it impractical
to study sleep in wild animals. Almost all information
about sleep is based on studies of captive animals housed
individually under constant conditions with unlimited food,
water and protection from predators. New, portable EEG
recording devices are small enough for animals to carry in
the wild.
In 2008, Voirin, working with Niels Rattenborg and
colleagues, conducted the first sleep study of any animal
living in the wild--monitoring three-toed sloths, Bradypus
variegatus in their natural habitat at STRI’s research station
on Barro Colorado Island. They were surprised to find that
wild sloths sleep over six hours less than captive sloths do.
Sleep researchers previously thought that the amount of
sleep an animal requires is largely inflexible.
How does the risk of being eaten influence the amount,
timing, and intensity of sleep and the proportion of time
spent in different types of sleep (i.e., rapid eye movement
(REM) sleep and non-REM sleep? Voirin noticed that
three-toed sloths, Bradypus pygmaeus, on Isla Escudo de
Veraguas in Bocas del Toro, where no predators exist, are
often active during mid-day and relatively tame. Closely
related three-toed sloths, Bradypus variegatus, on the
nearby mainland, where natural predators and humans
exist, are less obviously active during the day and often flee
when approached. Now he is using the EEG recorder to
explore these surprising observations further.
25
Niels Rattenborg (left) designed the first study of sleep in a wild animal,
which revealed that wild sloths spend significanlty less time sleeping
than captive sloths do. Now he and Bryson Voirin (right) will compare
sloth sleep patterns in habitats with and without predators.
Three toed sloths on the
mainland in Bocas del
Toro seem to sleep less
than pygmy sloths on
the island of Escudo de
Veraguas where there
are no predators.
Voirin mounts small electroencephalogram devices on the sloths’ heads
to record sleep patterns.

Phycology researchers Brian Wysor and Wilson
Freshwater work with SI’s Amy Driskell to sort and
begin the process of obtaining DNA barcodes for
the seaweeds in Bocas del Toro. DNA barcoding is
done in collaboration with the Training in Tropical
Taxonomy program.
26

Outreach, education, training
Outreach and education at the Bocas del Toro Research
Station spans a range of programs targeting K-12 students,
university undergraduates, graduate students and young
professionals. K-12 education includes visits to local
schools, some of which are in remote mountainous
locations, and student visits to the BRS. The station also
offers an annual training workshop for local K-12 teachers
coordinated by STRI’s education specialist, Lidia Valencia.
More than 200 university undergraduates visited the
station as participants in courses hosted by the Bocas
Research Station (see page 46), and more than 40
participated as research assistants and interns in numerous
research projects. Our outreach specialists reach local
undergraduates by visiting the University of Panama
in Changuinola. Finally, graduate students and young
professionals obtain advanced training at the BRS where
27
they learn specialized skills for studying biodiversity from
the Training in Tropical Taxonomy program.
Activities at the station also touch the general public from
around the world. Bi-weekly station open houses received
close to 1500 visitors from 21 countries in FY08-09. The
BRS has an active web presence. The BRS Biodiversity
database, the most visited of all STRI web pages, received
40,000 unique hits during this two-year period. The virtual
mangrove walkway and real-time biological monitoring
data are also popular sites.
A self-guided interpretive nature walk and educational
kitchen garden are currently under development. This
expansion of the on-site public programs include the pond,
forest, and mangrove eco-systems that are present on the
BRS property.
Children visiting the BRS during the annual Feria Ambiental learn about the famous frogs of Bocas by making frog origami and playing the frog lifecycle
game. Entertainment for children during the fair also includes activities for making things out of discarded plastic bottles and turtle art.

Local school outreach
Display aquaria and touch tanks are an important part of the outreach
program. Visiting scientists use hands-on activities to illustrate aspects
of the biology of different marine invertebrates and kids can see corals in
action in the display tanks.
28
The BRS works closely with local schools. Marlon Smith,
the BRS outreach and education specialist organizes
6-8 class visits to the station every month, and travels to
schools that do not have easy access to the BRS.
The program consists of activities tailored to the
educational level and prior knowledge of each group.
Students compare ecosystems, use laminated photo sheets
and keys to identify organisms, test water quality and make
connections between land use and coral reef health. BRS
researchers often participate, sharing their expertise and
enthusiasm with the students.
Over 1600 students participated in this program in FY08-09.
The BRS hosts an annual workshop for local teachers,
organized in conjunction with Panama’s Ministry of
Education (MEDUCA). Local teachers spent time in
the field with outreach staff, listened to scientists present
their work and developed effective classroom exercises
to incorporate biological information into the standard
curriculum. In 2009 the BRS hosted an additional
workshop organized by USAID and Conservation
International to train teachers and government employees
in best practices for environmental tourism.
Sebastien Tilmans demonstrates how to make biodiesel from waste cooking oil from a local restaurant, during the
annual joint BRS and MEDUCA teachers training workshop.

Community outreach
The public programs at BRS extend past formal education
programs into a variety of community outreach activities,
such as monthly public presentations. The annual Earth
Day beach clean up is organized in collaboration with
ANAM, IPAT, Promar and local business. In 2009 the
2-day event attracted 230 volunteers who collected over 4
tons of garbage and materials for recycling.
Another popular community event is the Enviromental
Fair. Every year the BRS opens its doors to the public for
two days of presentations, forums and information booths
on environmental issues and events in the region. Recent
topics have included environmentally sensitive waste-water
treatment, recycling in Bocas del Toro, and installation
of a system of buoys to prevent anchoring in sensitive
coral reef areas. NGO’s, indigenous groups as well as
governmental organizations from the region participate in
these educational activities. Entertainment is provided by
local dance schools and features traditional dances of the
region. Children’s activities combine learning with fun.
In 2009, the BRS also hosted a photography course
organized by Proyecto Impacto, a non-profit organization
dedicated to the empowerment of indigenous youth through
hands-on training in photography and multimedia. Ten
kids of different ethnicities and nationalities attended the
month-long course. A photo exhibit at the end of the course
featured the students’ impressive work.
29
Nearly 4 tons of garbage were collected during the 2009 Earth Day
beach clean-up.
Kids combine art and science to understand color variation in local frogs
during the summer program at the BRS.
The Feria Ambiental attracts NGOs and governmental organizations working on environmental issues in the region. Public participation is
encouraged with food and entertainment. These dancers from an Almirante school demonstrate dances from the Afro-Caribbean tradition in Bocas
del Toro.

Biodiversity Database and WWW Presence
Coordinated by BRS director, Rachel Collin, the Bocas
biodiversity database provides a list of plants and animals that
are known to occur in the Bocas del Toro province. Users can
search for a particular term or browse the database by group.
Descriptions, photographs, maps of known occurrences,
references and videos (when available) are included to
aid in species identification. Work continues to add more
information for the 6,500 species records as well as increase
the number of species included.
An example of one of the species pages shows how the database displays
morphological information, photographs and maps of the species
occurrences in the region.
30
The database is useful for scientists planning work at the
Bocas del Toro Research Station, field workers trying to
identify organisms they have seen in the field, and students
working in this part of Panama. The website receives more
than 1,600 unique visitors per month, with over 88,000
pageviews in the last year, and around 40,000 visitors over this
2-year period.
The data will soon be merged with the Encyclopedia of Life.
EOL is supporting a meiofauna workshop in 2010 to help add
data for these little-known groups.
In addition to the biodiversity database the BRS has an active
presence on Facebook. The page receives an average of
more than 500 hits a month and has recruited an average of
seventy fans a month since the initiation of this page in April
2009. The station’s Twitter feed and YouTube site also attract
attention from a diverse audience.
Average Visitors per Month
2000
1000
500
400
300
200
100
0
Bocas del Toro
Species
Database
Virtual and Actual visits
BRS
YouTube
BRS
Website
Station
Outreach
Visitors
FY09
FY08
BRS
FaceBook
Year
The number of public visits both to the station and to the internet site
continue to increase. The online biodiversity database receives four
times as many visits per months than any other outreach activity so far.

Training in Tropical Taxonomy
One of the major impediments to the documentation and
conservation of global biodiversity is the shortage of U.S.
and international scientists trained in taxonomy (finding,
describing and naming organisms). The BRS’s Training in
Tropical Taxonomy (TTT) program, is one of fewer than
a handful of programs worldwide aimed at addressing this
shortage.
The TTT runs a series of courses to train students and
biologists interested in biodiversity and taxonomy of
tropical marine animals. This training enables them to
efficiently collect and preserve material for subsequent
taxonomic work, sort the material and to use existing keys
or monographs to identify material to species. During the
course each student learns to identify the species in the
Bocas del Toro region.
Nearly half of the students are from the U.S. These courses
promote development of personal connections between
taxonomists in different countries and foster collaborations
between scientists in developed and developing countries
(see graph for breakdown of participants by country).
With the support of SENACYT, Panama’s governmental
office of science and technology, the SI’s Marine Science
Network and donor Paul Peck, courses on shrimp, tropical
algae, and tunicates were offered in FY08-09.
In 2009 Rachel Collin, Brian Wysor, and Suzanne
Fredericq obtained an NSF PanAmerican Advanced Studies
(PASI) grant to support an advanced workshop in Tropical
Phycology. Ten international experts worked with 20
graduate students and postdocs for 3 weeks. In addition
to initiating a number of international collaborations,
the PASI resulted in a bilingual phycology resource web
page that provides the academic community with lecture
materials on algae for a variety of educational levels as
well as laboratory protocols and a bilingual glossary of
phycological terms. The BRS aims to increase Panamanian
participation and urges Panamanians to apply for future
courses.
Thailand
Canada
Taiwan
New Zealand
France
Australia
Portugal
Slovenia
Ireland
Philippines
Martinique
Norway
Ecuador
British VI
31
West Indies
UK
TTT students painstakingly collect in the rocky intertidal.
Dissecting a sponge to find the tiny shrimp that live within its chambers.
El Salvador
Netherlands
Israel
Germany
Italy
Countries represented in TTT courses
Costa Rica
Cuba
Spain
Mexico
Puerto Rico
Venezuela
Brazil
Colombia
USA
50
40
30
20
10
0
Number of Student s

32
Phycologist Suzanne Fredericq from the University of
Lafayette in Louisiana examines each blade of seagrass
closely to find tiny epiphytic seaweeds that grow
there. Small algae like this make up much of the algal
diversity on coral reefs and in seagrass meadows.

BRS visitors and usage
Since its inauguration in 2003, the number of BRS scientific
visitors has increased five-fold from 75 in FY02 to 320 in
FY09 (see below). In addition there were 134 other visitors
including journalists and educators. Of the scientific visitors
39% were students in courses, 20% were graduate students,
undergrads and interns. BRS scientific visitors came from
over 35 countries. The intensity of usage as measured by
person days has likewise increased from 855 person days in
Number of Scientific Visitors
400
350
300
250
200
150
100
50
58
0
FY-01 FY-02 FY-03 FY-04 FY-05 FY-06 FY-07 FY-08 FY-09
Fiscal Year
Recent increases in the number of person days with little increase in
the number of visitors shows that visitors are now staying longer at the
station.
Training for teachers
and government
employees
Undergraduates,
Interns,
Volunteers
Post-docs,
Grad students
and STRI
Fellows
75
BRS Scientific Visitors
127
Visitors by Category
Scientific
Researchers
163
Undergraduate
courses
TTT courses
SI/STRI employees
Teachers and Panamanian government employees attending courses at
the BRS account for a significant proportion of overnight stays.
303
267
304
351
321
33
FY02 to over 7,000 in FY09. The average visitor stays at
the BRS for 22 days.
A comparison of the number of visitors at the BRS with
other facilities at STRI shows that the BRS has a similar
demand to Barro Colorado Island. Almost 30% of STRI
visitors do some work in Bocas del Toro.
Scientific Person Days
Average Persno Days
8000
7000
6000
5000
4000
3000
2000
BRS Scientific Usage Intensity
1949
2918 2918
3495
5295
6352
7156
1000
855
590
0
FY-01 FY-02 FY-03 FY-04 FY-05 FY-06 FY-07 FY-08 FY-09
Fiscal Year
The number of person days spent at the station has increased steadily
since its inauguration in 2003.
1000
800
600
400
200
Biannual Patterns in BRS Usage
Average FY02-03
Average FY04-05
Average FY06-07
Average FY08-09
0
Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sept
Months
There is a distinct seasonal pattern in the number of visitors with peaks
in July and August and a slow season in December and January.

Accommodations
Accommodations at the Bocas del Toro Research Station
include a dormitory with a common kitchen area that
houses 16 people, and two 2-bedroom houses (Casa Hoch
and Casa Cofrin) that can house 4 visitors each.
Occupancy of the houses in FY08-09 reached a peak
monthly occupancy of 99% in September 2008. The
average annual occupancy was 43% in FY09. A continued
increase is expected for FY10.
Person Days
Person Days
250
200
150
100
50
0
500
400
300
200
100
House Occupancy
FY06
FY07
FY08
FY09
“Ideal”
Oct Nov Dec Jan Feb Mar Apr May Jun Jul
Month
Dorm Occupancy
FY06
FY07
FY08
FY09
“Ideal”
Aug
Sep
0
Oct Nov Dec Jan Feb Mar Apr May Jue Jul Aug Sep
Month
Occupancy of both houses and dormitories is increasing steadily, but
monthly occupancy rates reflect the seasonal variation in station usage.
An Ideal occupancy rate of 60% gives time for maintenance and cleaning
between the numerous short visits. 60% occupancy is standard for hotels
in the US.
34
The dormitories consist of 4 rooms, each with 4 beds.
Occupancy was slightly higher than in the houses, with a
maximum monthly average of 95% in August, 2009. The
average annual occupancy was 40% in FY08 and FY09.
Overall, the dormitories are very crowded during the busy
season and pleasantly quiet in the off- season. 17% of the
station visitors live off-campus in remote locations or in the
town of Bocas del Toro.
Person Days
Number of Visitors
4000
3500
3000
2500
2000
1500
1000
500
0
250
200
150
100
50
0
FY06
FY06
Comparative Residency
FY07
FY07
Year
FY08
Comparative Residency
Year
FY08
FY09
FY09
Dormitory Town Houses
Long-term BRS residents stay in town, while short-term visitors, the vast
majority, stay at the station. The station occupancy rates have remained
stable. Accommodations are full during the busy season, whereas there is
space available in the off-season. Long-term visitors are staying longer.

Scientific Visitors
STRI Scientists
Marcy Balunas
Tropical disease drug discovery from marine and plant sources in
Panama (as part of the existing ICBG project).
Eldredge Bermingham
Molecular Ecology and Evolution in Hypoplectrus coral reef
fishes.
Carla Chizmar
Environmental leadership training initiative; biodiversity day activities.
Anthony Coates
Dissection of an extinction; Collection of fossil samples for the
Museum of Biodiversity in Panama.
Mary-Alice Coffroth, University of Buffalo - STRI Research
Associate.
The future of Caribbean reefs.
Rachel Collin, Director
Life history evolution in Calyptraeid gastropods.
Juan Del Rosario
Nutrient limitation of phytoplankton growth in the Bocas del Toro
Archipelago.
Kimberly Diver
Tropical disease drug discovery from marine and plant sources in
Panama (as part of the existing ICBG project).
Sergio Dos Santos
Monitoring of mangrove ecosystems in Bocas del Toro.
35
Amy Driskell
Development of a DNA barcode reference database of the fauna
and flora of Bocas del Toro.
Ilka Feller, SERC
Latitudinal variations in ecological stoichiometry in mangrove
communities.
Carmen Galdames
Inventory of Panama’s flora.
Edward Allen Herre
Figs and their associated organisms, sex ratio evolution, effects of
population structure, mycorrhizae, mutualism, parasitism, plantinsect
interactions.
Amalia Herrera
Morphological and molecular studies of neogene cupuladriids of
Tropical America.
Barbara Christine Hoekenga
Coral spawning in Montastraea annularis complex.
Javier Jara
Morphological and molecular studies of neogene cupuladriids
of Tropical America; Coral spawning in Montastraea annularis
complex.
Elisabeth Kalko
A study of reproductive patterns, community structure and
population fluctuations of bats in cave roosts on Isla Colon and Isla
Bastimentos, Bocas del Toro Province, Panama.
Karl Kaufmann
Monitoring of mangrove ecosystems in Bocas del Toro.
Nancy Knowlton
Coral spawning in Montastraea annularis complex.
Harilaos Lessios
Reproductive isolation between two recently diverged sympatric
species of Atlantic sea urchins (Genus Lytechinus): is it prezygotic
or postzygotic?
Juan Maté
Tropical Marine Ecology course instructor.
Javier Mateo-Vega
Environmental leadership training initiative.
Anne Meylan, Florida Fish and Wildlife Conservation Commission
- STRI Research Associate.
Ecology and migrations of sea turtles of Bocas del Toro, Panama.

Peter Meylan, Eckerd College - STRI Research Associate
Ecology and migrations of sea turtles in Bocas del Toro, Panama.
Robert Ricklefs, University of Missouri-St. Louis – STRI
Research Associate.
Molecular Ecology and Evolution in Hypoplectrus coral reef
fishes.
Rebecca Rissanen
BRS education and public programs development.
Felix Rodriguez
Morphological and molecular studies of neogene cupuladriids of
Tropical America.
Rosana Rocha, STRI Sabbatical Fellow, Universidade Federal do
Paraná.
Survey of Bocas del Toro biodiversity.
James Roper, STRI Sabbatical Fellow, Universidade Federal do
Paraná.
Genetic and behavoral evolution of island vertebrates since
isolation.
Gregory Ruiz, SERC
Shifts in predation pressure with latitude.
36
Carmen Schloeder
Parasites, trematodes, shifts in predation pressure with latitude.
Ashleigh Smythe, USNM Post-doc
Phylogeny of the marine nematode subclass Enoplia.
Maria Stapf
Inventory of Panama’s flora.
Ricardo Thompson
Monitoring of mangrove ecosystems in Bocas del Toro.
Mark Torchin
Parasites trematodes; shifts in predation pressure with latitude.
Noris Toribio
Inventory of Panama’s flora.
Francis Torres
Inventory of sponges and associated fungi.

STRI Fellows
Arthur Anker, STRI Post-doc Fellow.
Shrimp Taxonomy 2008, course instructor.
Gail Ashton, SERC Post-doc Fellow.
Shifts in predation pressure with latitude.
Augustin Cardona, STRI Post-doc Fellow.
Dissection of an extinction.
Anne Chamberlain, SERC Short-term Fellow.
Latitudinal variations in ecological stoichiometry in mangrove
communities.
Katie Cramer, STRI Pre-doc Fellow, University of California-
San Diego.
Historical changes in coral communities along a gradient of land
use in Panama.
Maya DeVries, STRI Short-term Fellow, University of California-
Berkeley.
Does morphological and functional specialization always go
hand-in-hand? Feeding morphology and ecology in mantis shrimp
(Stomatopoda).
John Douglass, STRI Post-doc Fellow.
Preliminary visit to plan a neurobiological research collaboration
on fiddler crab visual signalling.
David Farris, STRI Post-doc Fellow.
Collection of fossil samples for the Museum of Biodiversity in
Panama.
Amy Freestone, Smithsonian Marine Science Network
Postdoctoral Fellow.
Shifts in predation pressure with latitude.
Fernando Garcia, STRI Short-term Fellow.
The interaction of light and soil moisture availability in the
photosynthetic performance of tropical tree seedlings: implications
for seedling growth and survival in seasonal tropical forests.
Eva Garen, STRI Post-doc Fellow.
Environmental leadership training initiative.
Andres Gomez, Short-term Fellow.
Dissection of an extinction.
Keri Goodman, STRI Short-term Fellow.
Regulation of macroalgae communities via sea urchin grazing:
density-dependent factors and algal palatability.
37
Mary Hart, STRI Pre-doc Fellow, University of Kentucky.
Effect of density and risk on the mating behavior of an egg-trading
simultaneous hermaphrodite, Serranus tortugarum.
Alexandra Hiller, STRI Post-doc Fellow.
Reproductive isolation between two recently diverged sympatric
species of Atlantic sea urchins (Genus Lytechinus): is it prezygotic
or postzygotic?.
Kristin Hultgren, Smithsonian Marine Science Network
Postdoctoral Fellow.
Examining the processes regulating coexistence in Synalpheus in
a geographic context.
Santosh Jagadeeshan, STRI Post-doc Fellow.
Rapidly evolving sex and reproduction related genes and speciation
in sea urchins.
Omar Lopez, STRI Post-doc Fellow.
The interaction of light and soil moisture availability in the
photosynthetic performance of tropical tree seedlings: implications
for seedling growth and survival in seasonal tropical forests.
Osamu Miura, STRI Post-doc Fellow, Tohoku University.
Shifts in predation pressure with latitude.
Eric Nyman, STRI USAID Fellow.
Sustainable storm water and sanitation methods in Bocas del Toro.
Aaron O’Dea, STRI Post-doc Fellow.
Dissection of an extinction.
Laetitia Plaisance, NMNH-Fellow.
Coral spawning in Montastraea annularis complex.
Oscar Puebla, STRI Post-doc Fellow.
Molecular Ecology and Evolution in Hypoplectrus coral reef
fishes.
Colin Quinn, STRI Pre-doc Fellow, Colorado State University.
Plant arsenic hyperaccumulation: functional significance and
ecological impacts.
Aminur Rahman, STRI Post-doc Fellow.
Reproductive isolation between two recently diverged sympatric
species of Atlantic sea urchins (Genus Lytechinus): is it Prezygotic
or Postzygotic?
Corinne Richards, STRI Post-doc Fellow.
Selection and the rapid evolution of morphological variation among
strawberry poison-dart frogs of the Bocas del Toro archipelago.
Nikki Strong, STRI Post-doc Fellow.
Collection of fossil samples for the Museum of Biodiversity in
Panama.

Ana Spalding, STRI Shot-term Fellow, University of California-
Santa Cruz.
Re-making lives abroad: lifestyle migration and socioenvironmental
change in Bocas del Toro.
Gayatri Thampy, STRI Short-term Fellow, Ohio State
University.
Land use and land tenure reforms in Bocas del Toro.
Simon Tierney, STRI Post-doc Fellow.
Evolution of the nocturnal sweat bee genus Megalopta (Halictidae):
integrating molecular and behavioural evolution.
Sebastien Tilmans, STRI USAID Fellow.
Sustainable stormwater and sanitation methods in Bocas del Toro.
Eva Toth, STRI Post-doc Fellow.
Behavioral and genetic analysis of the principal of sociality in
sponge dwelling shrimp.
Dagoberto Venera, STRI Short-term Fellow.
Effects of macroalgae on the recovery of bleached corals.
38
John Wilk, STRI Short-term Fellow, University of Illinois-
Chicago.
Evolution of developmental patterns in Isognomon (Bivalvia:
Pterioidea): using cross-isthmus geminate species pairs to expose
changes in ontogenetic vectors and ecophenotypic responses.
Dan Lee Warren, STRI Pre-doc Fellow, University of California-
Davis.
Gamete plasticity in the bluehead wrasse.

Scientific Researchers
Catherine Caballero, INDICASAT.
Inventory of sponges and associated fungi.
José Cruz, INDICASAT.
Inventory of sponges and associated fungi.
Molly Cummings, University of Texas.
Poison or passion: warning and attraction in a color polymorphic
frog.
Sammy De Grave, Oxford University.
Shrimp Taxonomy 2008 course instructor.
Cristina Diaz, Museo Marino de Margarita.
The Porifera Tree of Life (PorToL) working group.
Humberto Diaz, Duke University.
Tropical Marine Ecology 2007 course instructor.
Emmet Duffy, College of William and Mary.
Examining the processes regulating coexistence in Synalpheus in
a geographic context.
Shelley Etnier, Butler University.
Butler University course preparation.
Philip Fearnside, Instituto Nacional de Pesquisas Da Amazônia.
Global changes and Amazonian development.
Suzanne Fredericq, University of Louisiana at Lafayette.
NSF-PASI Advanced Tropical Field Phycology 2009/ Tropical
Field Phycology 2008 course instructor.
39
Wilson Freshwater, University of North Carolina Wilmington.
Marine algal diversity of southern Central America.
Neusa Hamada, Instituto Nacional de Pesquisas Da Amazônia.
Global changes and Amazonian development.
Mark Hay, Georgia Institute of Technology.
Experiments in aquatic chemical signaling.
April Hill, University of Richmond.
The Porifera Tree of Life (PorToL) working group.
Cecile Jolly, University of Bergen.
Shallow-water Astrophorida (Porifera: Demospongiae) of Bocas
del Toro.
Cynthia Kicklighter, Goucher College.
Within-species variation in prey escape and deterrence strategies.
James Klaus, University of Miami.
Geology of Isla Colon - Chronostratigraphy of fossil coral bearing
units.
David Kline, University of Queensland.
The ecological genetics of reef building corals on both sides of the
isthmus.
Dennis Lavrov, Iowa State University.
The Porifera Tree of Life (PorToL) working group.
Don Levitan, Florida State University.
Coral spawning in Montastraea annularis complex.
Stan Lindsteat, Northern Arizona University.
Plasticity of metabolic capacity in gray catbirds, Dumetella
carolinensis.
Roger Linington, University of California Santa Cruz.
Tropical disease drug discovery from marine and plant sources in
Panama (as part of the existing ICBG project).
Jose Lopez, Nova Southeastern University.
The Porifera Tree of Life (PorToL) working group.
Derek Madden, California State University.
Thorns and Insects as features associated with the daily ranging
patterns of tamarins and agoutis in a tropical moist forest.
Donald McNeill, University of Miami.
Geology of Isla Colon - chronostratigraphy of fossil coral bearing
units.
Todd Oakley, University of California - Santa Barbara.
Evolution of eyes and vision in marine invertebrates.

Maria Cristina Ordoñez Espinosa, Caribbean Conservation
Corporation.
Hawksbill turtle population recovery and research in the Comarca
Ngobe-Bugle Chiriquí Beach: Escudo de Veraguas and the
Bastimentos Island National Marine Park.
Niels Rattenborg, Max-Plank-Institute for Ornithology.
Sleep in the three toed sloth Bradypus variegatus.
Niamh Redmond, Smithsonian National Museum of Natural
History.
The Porifera Tree of Life (PorToL) working group.
Gregory Rouse, Scripps Institution of Oceanography.
Annelid Collection of Panama.
Carmen Salsbury, Butler University.
Butler University course preparation.
40
Paul Schaeffer, Miami University.
Plasticity of metabolic capacity in gray catbirds (Dumetella
carolinensis).
Adolphe Schluder, University of British Columbia.
The historical biogeography of Mesoamerica.
Davis Stephen, Texas A&M University.
Quantifying relationships between resource heterogeneity and plant
community structure in a coastal freshwater swamp of Panama.
Robert Thacker, University of Alabama at Birmingham.
Ecology and evolution of sponge-cyanobacteria symbioses.
Maurice Thomas, Palm Beach Atlantic University.
A study of reproductive patterns, community structure and
population fluctuations of bats in cave roosts on Isla Colon and
Isla Bastimentos.
Tiffany Troxler, Florida International University.
Quantifying relationships between resource heterogeneity and plant
community structure in a coastal freshwater swamp of Panama.
Marco Tschapka, University of Ulm.
Reproductive patterns, community structure and population
fluctuations of bats in cave roots on Isla Colon and Isla
Bastimentos.
Engel Vic, National Park Service.
Quantifying relationships between resource heterogeneity and plant
community structure in a coastal freshwater swamp of Panama.
Steve Vollmer, Northeastern University.
The ecological genetics of reef building corals on both sides of the
isthmus.
Gert Woerheide.
The Porifera Tree of Life (PorToL) working group.
Janie Wulff, Florida State University.
Influence of competition, mutualism, and physical environment on
patterns of sponge diversity and co-occurrence on Caribbean coral
reefs and mangroves.
Brian Wysor, Roger Williams University.
Marine algal diversity of southern Central America.
Steven Zeichner, George Washington University, Medical Center.
Figs and their associated organism, focused on: sex ratio evolution,
effects of population structure, mycorrhizae, mutualism, parasitism,
plant-insect interactions.

Post-doctoral Researchers
Maaike Bader, University of Oldenburg.
Changes in plant carbon balance of epiphytes along an altitudinal
gradient.
Elizabeth Borda, Scripps Institution of Oceanography.
Annelid collection of Panama.
Alexander Hayward, Oxford University.
Antipredator behaviour in the Caribbean reef squid (Sepioteuthis
sepioidea).
Laura May-Collado, George Mason University.
Status of the bottlenose dolphins from Bocas del Toro, Panama:
Implications for future local conservation and management plans.
David Luther, University of Maryland.
Latitudinal variations in ecological stoichiometry in mangrove
communities.
Sofie Sjogersten, University of Notthingham.
From tree species characteristics to population and community
dynamics – investigations with the individual-based forest model
FORMIND.
Christian Voolstra, University of California – Merced.
Coral reef health.
41
Graduate Students
Rebecca Albright, University of Miami.
The molecular ecology of photoreception in coral larvae.
Christopher Angioletti, Scripps Institution of Oceanography.
Historical changes in coral communities along a gradient of land
use in Panama.
Deanna Ashline, University of Buffalo.
The futures of Caribbean reefs.
Flavia Barbosa, University of Missouri.
Rapid, environmentally-induced color change in a polymorphic
poison frog.
Samir Bhatt, Oxford University.
Antipredator behaviour in the Caribbean reef squid, Sepioteuthis
sepioidea.
Brendan Biggs, Florida State University.
Influence of competition, mutualism, and physical environment on
patterns of sponge diversity and co-occurrence on Caribbean coral
reefs and mangroves.
Christina Blewett, University of Washington.
Soil nutrient dynamics.
David Bloom, University of Toronto.
Phylogenetics and evolution of anchovies.
Signe Brinklov, University of Southern Denmark.
Reproductive patterns, community structure and population
fluctuations of bats in cave roots on Isla Colon and Isla
Bastimentos.
Martin Breed, Uppsala University.
Population differentiation in strawberry poison frogs.
Rodrigo Carballo, Universidad de El Salvador.
Reproductive isolation between two recently diverged sympatric
species of Atlantic sea urchins, Genus Lytechinus: Is it Prezygotic
or Postzygotic?
Paco Cardenas, University of Bergen.
Shallow-water Astrophorida (Porifera: Demospongiae) of Bocas
del Toro, Panama.
Leandro Castaño, Organization for Tropical Studies.
Environmental leadership training initiative.
Andia Chavez-Fonnegra, Nova Southeastern University.
The Porifera Tree of Life (PorToL) working group.

Alexander Cheesman, University of Florida.
Soil nutrient dynamics.
Laura Crothers, University of Texas.
Poison or passion: warning and attraction in a colour polymorphic
frog.
Emily Dangremand, University of California –Berkeley.
Latitudinal variations in ecological stoichiometry in mangrove
communities.
Danyelle Dehner, University of Alabama at Birmingham.
Experimental manipulation of light and its impact on spongecyanobacteria
symbioses.
Amanda Fenner, University of Iowa.
Tropical disease drug discovery from marine and plant sources in
Panama (as part of the existing ICBG project).
Nicole Fogarty, Florida State University.
Coral spawning in Montastraea annularis complex.
Chris Freeman, University of Alabama at Birmingham.
The Porifera Tree of Life (PorToL) working group.
Tania Gonzalez, University of Ulm.
A study of reproductive patterns, community structure and
population fluctuations of bats in cave roosts on Isla Colon and
Isla Bastimentos, Bocas del Toro Province, Panama.
Zbigniew Grabowski, University of Connecticut.
Carbon estimation for avoided deforestation / NEO STUDENT.
Melanie Heckman, Georgia Institute of Technology.
Regulation of macroalgae communities via sea urchin grazing:
density-dependent factors and algal palatability.
Robert Hegna, Florida International University.
A study of aposematism in the strawberry poison dart frog
(Oophaga pumilio): The effect of frog color on predation.
Elizabeth Hemond, Northeastern University.
The ecological genetics of reef building corals on both sides of the
isthmus.
Zeehan Jaafar, National University of Singapore.
Sponge associations and interactions at Bocas del Toro.
Ehsan Kayal, Iowa State University.
The Porifera Tree of Life (PorToL) working group.
Marieke Keller, Leiden University.
Effects of density and risk on the mating behavior of an egg-trading
simultaneous hermaphrodite, Serranus tortugarum.
42
Dustin Kemp, University of Georgia.
Regulation of macroalgae communities via sea urchin grazing:
density-dependent factors and algal palatability.
Justin P. Lawrence, Michigan State University.
Conservation of a Polymorphic Frog, Dendrobatidae: Oophaga
pumilio, in Western Panama.
Amanda Lea, University of Texas at Austin.
Rapid, environmentally-induced color change in a polymorphic
poison frog.
Eric Lewallen, University of Toronto.
Phylogenetics and evolution of Anchovies.
Silvia Libro, Northeastern University.
The ecological genetics of reef building corals on both sides of the
isthmus.
Tse-Lyn Loh, University of North Carolina.
Sponge associations and interactions in Bocas del Toro.

Jose Loiza, McGill University.
Demographic history of Anopheles puntinacula in Panama.
Christian Luneburg, Babson College.
Study of the sleep in the three toed sloth (Bradypus variegatus).
Megan Lupek, University of Liverpool.
The role of odours for mate choice and social structure in Noctilio
albiventris, the lesser bulldog-bat.
Benjamin Mason, University of Miami.
The molecular ecology of photoreception in coral larvae.
Franklin Marek.
Rapid, environmentally-induced color change in a polymorphic
poison frog.
Carmel Norman, Northeastern University.
The ecological genetics of reef building corals on both sides of the
isthmus.
Kylee Pawluk, University of Victoria.
Coral spawning in Montastraea annularis complex.
Johanne Pelletier, McGill University.
Carbon estimation for avoided deforestation.
43
Jorge Pinzon, Penn State University.
Pocillopora spp and Acropora palmata as case studies in the coevolution
of coral-algal symbioses from Panamian Pacific and
Caribbean coral reef ecosystems.
Michael Reichert, University of Missouri.
Rapid, environmentally-induced color change in a polymorphic
poison frog.
Lisa Rodriguez, Eckerd College.
Ecology and migrations of marine turtles of Bocas del Toro.
Björn Rogell, Uppsala University.
Population differentiation in strawberry poison frogs.
Jose Rovira, Instituto Conmemorativo Gorgas.
Demographic history of Anopheles puntinacula in Panama.
Ewelina Rubin, Nova Southeastern University.
The Porifera Tree of Life (PorToL) working group.
Andrea Rudh, Uppsala University.
Population differentiation in strawberry poison frogs.
Madeleine Scriba, Max-Plank-Institute for Ornithology.
Sleep in the three toed sloth Bradypus variegatus .

Erik Sperling, Yale University.
The Porifera Tree of Life (PorToL) working group.
Shinichi Sunagawa, University of California- Merced.
Coral reef health.
Tracy Stetzinger, National University of Singapore.
Selection and the rapid evolution of morphological variation among
strawberry poison-dart frogs of the Bocas del Toro archipelago.
Anna Strimaitis, Florida State University.
Coral spawning in Montastraea annularis complex.
Anne-Marie Surlykke, University of Southern Denmark.
Reproductive patterns, community structure and population
fluctuations of bats in cave roots on Isla Colon and Isla
Bastimentos.
Casey Terhorst, Florida State University.
Coral spawning in Montastraea annularis complex.
44
Justin Yeager, East Carolina University.
Poison or passion: warning and attraction in a color polymorphic
frog.
James Bryson Voirin, Max Planck Institute for Ornithology-
Seewiesen.
Study of the sleep in the three toed sloth, Bradypus variegatus.
Kristine White, University of Southern Mississippi.
Diversity and ecology of leucothoid amphipods in tunicate host of
Panama.
Emma Wright, University of Nottingham.
Soil nutrient dynamics carbon and nutrient cycling in a tropical
wetland.

Volunteers, Undergraduates, Assistants and
Interns
Rafael Aizprua, Texas A&M University
Jessie Alden, Roger Williams University
Dalia Barragan, Universidad de Bogotá Jorge Tadeo Lozano
Narissa Bax, University of Hawaii
Mario Baylon, STRI
Jose Benchetrit, McGill University
Jessica Bolaños, INDICASAT
Ricardo Cossio, Universidad de Panamá
Marie-Claude Côté-Laurin, McGill University
Natalia Decastro, Universidad de Medellin
Luis Elizondo
Keith Erickson
Craig Erickson
Angie Carole Estrada, Universidad de Panamá
Ramon Fernandez, Universidad de Valencia
Catlin Fong, University of California - Santa Barbara
Deyvis Gonzalez, Colegio Jose Ibarra, Bocas del Toro
Molly Goodier, Eckerd College
Kasey Grace, Florida State University
Christine Gruman, University of Alberta
Heidi Guenther, Florida State University
Richard Harr, Suncoast High School
Sarah Holloway, University of Texas
Simon Hultby, Uppsala University
Silke Janssen, University of Hamburg
Guy Kieckhefer, Eckerd College
Erin Klamper
Ashley Lamb, University of Texas
Cara Lawrence, University of Kentucky
Amanda Lea, University of Texas
Mara McGrew, University of California
Jessica Miller, Duke University
Andrea Montalvo, School of International Training
Myriam Monroe
Sara Moron, Universidad Nacional de Colombia
Jaclyn Nora, Duke University
Sergio Nuñes
José Palacio, Fundación Vieto
Johana Parra, Universidad de los Andes
Lillian Perkins-High, School for International Training
Nico Reinhold, Technical University of Munich
Elizabeth Sargen, Roger Williams University
Isaac Standish, Michigan State University
Cristina Wong, University of Rochester
Apanie Wood, University of New South Wales
Christopher Wright
Alexander Zaharoff, University of California - Santa Barbara
Geoff Zawacki
45
Science Communicators
Asociación de Comunicadores de Bocas del Toro
Station visit
Mary Albion, Icon Films
Weird Creatures with Nick Baker - “Tamandua” & “Pygmy sloth”
Nick Baker, Icon Films
Weird Creatures with Nick Baker - “Tamandua” & “Pygmy sloth”
John Barrat, Smithsonian Institution Office of Public Affairs
Latitudinal variations in ecological stoichiometry in mangrove communities
James Chapman, Icon Films
Weird Creatures with Nick Baker - “Tamandua” & “Pygmy sloth”
Megan Gambino, Smithsonian Magazine
Coral spawning in Montastraea annularis complex
John Gibbons, Smithsonian Institution Office of Public Affairs
Latitudinal variations in ecological stoichiometry in mangrove communities
Marcos Guerra, STRI
Biodiversity day activities
Karin Hammerberg, National Geographic Explorer
Science of Sloth Sleep
Anne Kazel, Gold Coast Communications
Article for travel magazine
Beth King, STRI
Biodiversity day activities
Jan Sapp, York University, Toronto, Canada – STRI Research Associate
Research for book proposal
Ted Tuel, National Geographic Explorer
Science of Sloth Sleep
Simon Wagen, Icon Films
Weird Creatures with Nick Baker - “Tamandua” & “Pygmy sloth”
Kennedy Warne
Latitudinal variations in ecological stoichiometry in mangrove communities
Stefan Weisen, National Geographic Explorer
Science of Sloth Sleep
Solange Welch, Icon Films
Weird Creatures with Nick Baker -”Tamandua” & “Pygmy sloth”
Christian Ziegler, freelance photographer
Coral spawning in Montastraea annularis complex

Course participants –
Visiting Courses
Universidad de los Andes, Colombia
Instructor: Juan Sánchez
Ecologia Marina
September 26 – October 6, 2007 (22 students)
September 28 – October 4, 2008 (15 students)
Duke University
Instructor: Humberto Diaz, Richard Forward
Experimental Tropical Marine Ecology
October 6 – 13, 2007 (12 students)
October 11 – 18, 2008 (12 students)
School for International Training, Panama
Instructor: Alyson Dagang, Juan Mate, Michael Smalls (TA)
Tropical Marine Ecology
November 7 – 12, 2007 (8 students)
April 25 – 30, 2008 (18 students)
Conservation and Development 2008
November 1 – 6, 2008 (12 students)
Tropical Marine Ecology
May 3 – 8, 2009 (8 students)
University of Wisconsin-Green Bay
Instructor: Robert Howe, Amy Wolf, Anindo Choudhury, Mike
Draney, Vicki Medland
Tropical Biology
January 2 – 7, 2008 (10 students)
January 5 – 8, 2009 (19 students)
46
Butler University
Instructor: Shelley Etnier, Carmen Salsbury
Tropical Field Biology
March 9 – 14, 2008 (6 students)
Princeton University
Instructor: Stephen Pacala, Ryan Chisholm (TA)
Tropical Marine Ecology Course
February 3 – 13, 2008 (18 students)
February 23 – March 4, 2009 (16 students)
Radford Universtity
Instructor: Judy Guinan, Robert Sheehy
Environmental Biology in Panama
May 19 – 24, 2008 (9 students)
May 25 – 29, 2009 (9 students)
Georgia Institute of Technology
Instructor: Mark Hay
Experiments in Aquatic Chemical Signaling 2008
May 31 – June 30, 2008 (8 graduate students)
Harvard University
Instructor: Gonzalo Giribet, Casandra Extravour, Sebastian
Velez (TA), Vanessa Gonzalez (TA)
Organismic and Evolutionary Biology
March 22 – 29, 2009 (15 students)

Course participants –
STRI-TTT Courses
Tropical Field Phycology
July 9 - 23, 2008
Instructors: Suzanne Fredericq, D. Wilson Freshwater, Brian
Wysor
Students: Cindy Fernandez (Costa Rica), Martha Diaz
(Colombia), Olga Camacho (Colombia), Margarita Rosa Albis
Salas (Colombia), Andrea Planas Orellana (El Salvador), Ryan
Fikes (USA), Kevin Miklasz (USA), Samantha Schmitt (USA),
Thomas Sauvage (USA), Anna Fricke (Germany), Jimena
Samper Villareal (Costa Rica).
Shrimp Taxonomy (Caridea, Dendrobranchiata and
Stenopodidea)
August 4 - 16, 2008
Instructors: Sammy de Grave, Arthur Anker, Michel Hendrickx
Students: Betel Martinez (Mexico), Chris Ashelby (UK), Juan
Felipe Lazarus Agudelo (Colombia), Jure Jugovic (Slovenia),
Laura Anderson (USA), Lucas Simon Torati (Brazil), Nuno
Simoes (Portugal), Timothy Page (Australia), Leslie Harris
(USA), Carolina Rodrigues Tavares (Brazil), Patricio Hernaez
(Costa Rica), Javier Luque (Colombia), Nicola C. Dobson (UK).
47
Taxonomy and Biology of Tunicates
June 4-18, 2009
Instructors: Rosana Rocha, Gretchen Lambert, Charles Lambert
Students: Andrea Frey (USA), Christian Sardet (France),
Christina Simkanin (USA), Marc Rius (Spain), Stephan Bullard
(USA), Mike Page (New Zealand), Mari Carmen Pineda Torres
(Spain), Lisa Draughon (USA), Nadia Bonnet (Brazil), Gil
Koplovitz (Israel), Su Shih-Wei (Taiwan), Ronaldo Ruy (Brazil),
Ashley Callahan (Canada), Aida Hernández Zanuy (Cuba),
Betzabé Moreno Dávila (Mexico), Johanna Cannon (USA).
NSF Pan-American Advanced Studies Institute
Advanced Methods in Tropical Phycology
Aug. 14 - Sept. 4, 2009
Instructors: Juan M. Lopez-Bautista, Brian Wysor, Suzanne
Fredericq, D. Wilson Freshwater, Rafael Riosmena-Rodriguez,
Nadine Schubert, Guillermo Diaz-Pulido, Amy Driskell, Steven
Paton, Bernardo A.P. da Gama, Rachel Collin
Students: Craig Aumack (USA), Kyle Glenn (USA), Eddie
Parish (US Virgin Islands), Anchana Prathep (Thailand), Lizette
Irene Quan Young (Mexico), Dagoberto Venera (Colombia),
Cindy Fernández (Costa Rica), Nahid Osman (Sudan), Enrique
Peña (Colombia), Daniela Milstein (Brazil), Rebecca Guenther
(USA), Simona Augyte (USA), Rachel Kennison (USA),
Elizabeth Lacey (USA), Jasmine Ruvalcaba (USA), Ruben
Cabrera (Cuba), Neidy Cetz-Navarro (Mexico), Olga Tejada (El
Salvador), Haj Allali (USA), William Schmidt (USA).

48
Staining transverse sections of red algae (Rhodophyta) with methylene blue
makes the cellular structure easily visible. Cutting paper-thin sections and
staining them was one of the methods that students practiced in the NSF-
PASI Advanced Methods in Tropical Phycology workshop.

3 - year bibliography
In press
Baeza, J.A. Protandric Simultaneous Hermaphroditism Is
a Conserved Trait in Lysmata (Caridea: Lysmatidae):
Implications for the Evolution of Hermaphroditism in the
Genus. Smithsonian Contributions to Marine Sciences 38. In
press.
Collin, R., L. D’Croz, P. Gondola, J.B. Del Rosario and K.W.
Kauffman. Climate and Hydrological Factors Affecting
Variation in Chlorophyll Concentration and Water Clarity in
the Bahia Almirante, Panama. Smithsonian Contributions to
Marine Sciences 38. In press.
Diaz, M.C. and K. Rützler. Biodiversity and Abundance
of Sponges in Caribbean Mangrove: Indicators of
Environmental Quality. Smithsonian Contributions to Marine
Sciences 38. In press.
Feller, I.C., C.E. Lovelock and C. Piou. Growth and nutrient
conservation in Rhizophora mangle in response to
fertilization along latitudinal and tidal gradients. Smithsonian
Contributions to Marine Sciences 38. In press.
Bocas del Toro Research Station
Publications by Topic
Marine
Paleontology
(34)
Environmental
Biology
(14)
Anthropology
(4)
Marine Biology
(196)
Terrestrial
Biology
(60)
49
Li, C. and R. Collin. Imposex in One of the World’s Busiest
Shipping Zones. Smithsonian Contributions to Marine
Sciences 38. In press.
O’Dea A. Relation of form to life habit in free-living bryozoans.
Marine Ecology Progress Series.
Ostrovsky, A.N. and A. O’Dea. Comparative anatomy of the
internalincubational sacs in the genera Discoporella and
Cupuladria (Bryozoa,Gymnolaemata), and evolution
of brooding in the free-living Cheilostomata. Journal of
Morphology.
Puebla, O., E. Bermingham and F. Guichard. Estimating dispersal
from genetic isolation by distance in a coral reef fish
(Hypoplecturs puella). Ecology. In press.
Puebla, O., E. Bermingham and F. Guichard. Genetic signature
of local dispersal in a coral reef fish: demography and scale
matters. In revision.
Wulff, J. Sponge Community Dynamics on Caribbean Mangrove
Roots: Significance of Species Characteristics. Smithsonian
Contributions to Marine Sciences 38. In press.
Number of Publications
60
50
40
30
20
10
0
Bocas del Toro Research Station
Publications
13
2001
15
2002
17
2003
22
2004
35
2005
Year
28
2006
56
2007
47
2008
15
2009

50
2009 2008
Anker, A. C. Hurt and N. Knowlton. 2009. Description of cryptic
taxa within the Alpheus bouvieri A. Milne-Edwards, 1878
and A. hebes Kim and Abele, 1988 species complexes
(Crustacea: Decapoda: Alpheidae). Zootaxa 2153: 1- 23.
Barros, R.C., R.M. Rocha and M.R. Pie. 2009. Human-mediated
global dispersion of Styela plicata (Tunicata, Ascidiacea).
Aquatic Invasions 4: 45-57.
Granek, E.F., J.E. Compton and D.I. Phillips. 2009. Mangrove-
Exported Nutrient Incorporation by Sessile Coral Reef
Invertebrates. Ecosystems. Online.
Loh, T.L. and J.R. Pawlik. 2009. Bitten down to size: Fish
predation determines growth form of the Caribbean coral
reef sponge Mycale laevis. Journal of Experimental Marine
Biology and Ecology 374: 45-50.
Lovelock, C.E., M.C. Ball, K.C. Martin and I.C. Feller. 2009.
Nutrient Enrichment Increases Mortality of Mangroves.
PLoS ONE 4: e5600. doi:10.1371/journal.pone.0005600
Smith J.T., Jackson J.B.C. 2009. Ecology of extreme faunal
turnover of tropical American scallops. Paleobiology 35:
77-93.
Zotz, G. and N. Reuter. 2009. The effect of exposure to sea
water on germination and vegetative growth of an epiphytic
bromeliad. Journal of Tropical Ecology 25: 311-319.
Anker, A., C. Hurt and N. Knowlton. 2008. Revision of the
Alpheus formosus Gibbes, 1850 complex, with redescription
of A. formosus and description of a new species from the
tropical western Atlantic (Crustacea: Decapoda: Alpheidae).
Zootaxa 2008: 1-22.
Anker, A., C. Hurt and N. Knowlton. 2008. Revision of
the Alpheus websteri Kingsley, 1880 species complex
(Crustacea: Decapoda: Alpheidae), with revalidation of A.
arenensis (Chace, 1937). Zootaxa 1694: 51-68.
Anker, A., C. Hurt and N. Knowlton. 2008. Revision of the
Alpheus cristulifrons species complex (Crustacea: Decapoda:
Alpheidae) with description of a new species from the
tropical eastern Atlantic. Journal of the Marine Biological
Association of the United Kingdom 88: 543-562.
Anker, A. and E. Toth. 2008. A preliminary revision of the
Synalpheus paraneptunus Coutière, 1909 species complex
(Crustacea: Decapoda: Alpheidae). Zootaxa 1915: 1-28.
Baeza, J.A. 2008. Protandric simultaneous hermaphroditism
in the shrimps Lysmata bahia and Lysmata intermedia.
Invertebrate Biology 127: 181-188.
Baeza, J.A. and A. Anker. 2008. Lysmata hochi n. sp., a new
hermaphroditic shrimp from the southwestern Caribbean Sea
(Crustacea: Caridea: Hippolytidae). Journal of Crustacean
Biology 28: 148-155.
Baeza, J.A., J.M. Reitz and R. Collin. 2008. Protandric
simultaneous hermaphroditism and sex ratio in Lysmata
nayaritensis Wicksten, 2000 (Decapoda: Caridea). Journal of
Natural History 41: 2843-2850.
Carlon, D.B. and C. Lippe. 2008. Fifteen new microsatellite
markers for the reef coral Favia fragum and a new
Symbiodinium microsatellite. Molecular Ecology Resources
8: 870-873.
Collin, R. 2008. Development of Tylodina fungina Gabb, 1865
(Gastropoda: Notaspidea) from the Pacific Coast of Panama.
Veliger 50: 51-56.
Collin, R. and M. Miglietta. 2008. Reversing opinions on Dollo’s
Law. Trends in Ecology and Evolution. 23: 602-609.

DeSalvo, M.K, C.R. Voolstra, S. Sunagawa, J.A. Schwartz, J.H.
Stillman, M.A. Coffroth, A.M. Szmant and M. Medina.
2008. Differential gene expression during thermal stress
and bleaching in the Caribbean coral Monastraea faveolata.
Molecular Ecology 17: 3952–3971.
Darnell, M. Z., M.B. Ogburn and H. Diaz. 2008. A novel running
wheel apparatus to monitor locomotor rhythms in land crabs.
Marine and Freshwater Behaviour and Physiology 41: 205-
210.
Erwin, P.M. and R.W. Thacker. 2008. Cryptic diversity of the
symbiotic cyanobacterium Synechococcus spongiarum
among sponge hosts. Molecular Ecology 17: 2937-2947.
Erwin, P.M. and R.W. Thacker. 2008. Phototrophic nutrition and
symbiont diversity of two Caribbean sponge-cyanobacteria
symbioses. Marine Ecology Progress Series 362: 139-147.
Granek, E. and B.I. Ruttenberg. 2008. Changes in biotic and
abiotic processes following mangrove clearing. Estuarine,
Coastal and Shelf Science 80: 555 – 562.
Herrera-Cubilla, A., M.H. Dick, J. Sanner and J.B.C. Jackson.
2008. Neogene cupuladriidae of tropical America. II:
Taxonomy of recent Discoporella from opposite sides of the
Isthmus of Panama. Journal of Paleontology 82: 279-298.
Hooge, M.D. and S. Tyler. 2008. Acoela (Acoelomorpha) from
Bocas del Toro, Panama. Zootaxa 1719: 1-40.
Johnson, K.G., J.B.C. Jackson and A.F. Budd. 2008. Caribbean
reef development was independent of coral diversity over 28
million years. Science 319: 1521 – 1523.
Knowlton, N. and J.B.C. Jackson. 2008. Shifting baselines, local
impacts, and global change on coral reefs. PLOS Biology 6: e54.
Knowlton N. 2008. Coral reefs. Current Biology 18: R18-R21.
Lessios, H.A. 2008. The Great American Schism: Divergence
of marine organisms after the rise of the Central American
Isthmus. Annual Review of Ecology Evolution and
Systematics. 39: 63-91.
Loaiza, J.R., E. Bermingham, M.E. Scott, J.R. Rovira and J.E.
Conn. 2008. Species composition and distribution of adult
Anopheles (Diptera: Culicidae) in Panama. Journal of
Medical Entomology 45: 841 – 851.
51
López-Duarte, P.C. 2008. Selective tidal-stream transport
behavior of fiddler crab (Uca spp.) larva: comparisons
among species and different tidal regimes. Florida Institute
of Technology, Melbourne (FL).173 pages.
Lovelock, C.E. 2008. Soil respiration and belowground carbon
allocation in mangrove forests. Ecosystems 11: 342 – 354.
Maan, M.E. and M.E. Cummings. 2008. Female preferences for
aposematic signal components in a polymorphic poison frog.
Evolution 62: 2334-2345.
MacDonald, J.A. 2008. Variation among mangrove forests as
fish habitat: The role of prop-root epibionts, edge effects and
behavior in neotropical mangroves. Rutgers University, New
Brunswick, NJ. 153 pages.
MacDonald, J.A., T. Glover, and J.S. Weis. 2008.The impact of
mangrove prop-root epibionts on juvenile reef fishes. A field
experiment using artificial roots and epifauna. Estuaries and
Coasts 31: 981-993.
Mathews, L.M. and A. Anker. 2008. Molecular phylogeny reveals
extensive ancient and ongoing radiations in a snapping
shrimp species complex (Crustacea, Alpheidae, Alpheus
armillatus). Molecular Phylogenetics and Evolution 50: 268-
281.
May-Collado, L. and D. Wartzok. 2008. A comparison of
bottlenose dolphin whistles in the Atlantic Ocean: factors
promoting whistle variation. Journal of Mammalogy 89:
1229-1240.
Mejia, L.C., E.I. Rojas, Z. Maynard, S.Van Bael, A.E. Arnold,
P. Hebbar, G.J. Samuels, N. Robbins and E.A. Herre. 2008.
Endophytic fungi as biocontrol agents of Theobroma cacao
pathogens. Biological Control 46: 4-14.
Miglietta, M-P. and H.A. Lessios. 2008. A silent invasion.
Biological Invasions. Online.
Miglietta, M-P., M. Rossi and R. Collin. 2008. Hydromedusa
blooms and upwelling events in the Bay of Panama, Tropical
East Pacific. Journal of Plankton Research 30: 783-793.
O’Dea, A., J.B.C. Jackson, P.D. Taylor and F. Rodriguez. 2008.
Modes of reproduction in Recent and fossil cupuladriid
bryozoans. Palaeontology 51: 847-864.

Perez-Miles, F., R. Gabriel, L. Miglio, A. Bonaldo, R. Gallon,
J.J. Jimenez and R. Bertani. 2008. Ami, a new Theraphosid
genus from Central and South America, with the description
of six new species (Araneae: Mygalomorphae). Zootaxa
1915: 54-68.
Puebla O., E. Bermingham and F. Guichard. 2008. Population
genetic analyses of Hypoplectrus coral reef fishes provide
evidence that local processes are operating during the early
stages of marine adaptive radiations. Molecular Ecology 17:
1405-1415.
Rawlinson, K.A. 2008. Biodiversity of coastal polyclad flatworm
assemblages in the wider Caribbean. Marine Biology 153:
769-778.
Richards, C., A.J. Zellmer and L.M. Martens. 2008.
Batracochytrium dendrobatidis not detected in Oophaga
pumilio on Bastimentos Island, Panama. Herpetological
Review 29: 200-202.
Rodriguez, S., A. Croquer, H.M. Guzman and C. Bastidas. 2008.
A mechanism of transmission and factors affecting coral
susceptibility to Halofolliculina sp. infection. Coral Reefs
28: 67-77.
Schloeder, C. and H.M. Guzman. 2008. Reproductive patterns of
the Caribbean coral Porites furcata (Anthozoa, Scleractinia,
Poritidae) in Panama. Bulletin of Marine Science 82: 107-
117.
Summers, K., T.W. Cronin and T. Kennedy. 2008. Variation in
spectral reflectance among populations of Dendrobates
pumilio, the strawberry poison frog, in the Bocas del Toro
Archipelago, Panama. Journal of Biogeography 30: 35 – 53.
52
Taylor, A.S., J.L. Haynes and G. Holzman. 2008. Taxonomical,
nomenclatural and biogeographical revelations in the
Zamia skinneri complex of Central America (Cycadales:
Zamiaceae). Botanical Journal of the Linnean Society 158:
399-429.
Toth, E. and J.E. Duffy. 2008. Influence of sociality on allometric
growth and morphological differentiation in sponge-dwelling
alpheid shrimp. Biological Journal of the Linnean Society
94: 527-540.
Toth, E. and R.T. Bauer. 2008. Synalphus paraneptunus
(Crustacea: Decapoda: Caridea) populations with intersex
gonopores: a sexual enigma among sponge-dwelling
snapping shrimps. Invertebrate Reproduction and
Development 51: 49–59.
Vollmer, S.V. and D.I. Kline. 2008. Natural Disease Resistance in
Threatened Staghorn Corals. PLoS ONE 3: e3718.
Wulff, J.L. 2008. Collaboration among sponge species increases
sponge diversity and abundance in a seagrass meadow.
Marine Ecology 29: 193 – 204.
Wulff, J.L. 2008. Life-history differences among coral reef
sponges promote mutualism or exploitation of mutualism by
influencing partner fidelity feedback. American Naturalist
171: 597–609.
Zigler, K.S., H.A. Lessios and R.A. Raff. 2008. Egg energetics,
fertilization kinetics, and population structure in echinoids
with facultatively feeding larvae. Biological Bulletin 215:
191-199.

2007
Anker, A., C. Hurt and N. Knowlton. 2007. Revision of the
Alpheus nuttingi (Schmitt) species complex, with description
of a new species from the tropical eastern Pacific (Crustacea:
Decapoda: Alpheidae). Zootaxa 1577: 41-60.
Anker, A., C. Hurt and N. Knowlton. 2007. Three transisthmian
snapping shrimps (Crustacea: Decapoda: Alpheidae) associated
with innkeeper worms (Echiura: Thalassematidae) in Panama.
Zootaxa 1626: 1-23.
Arnold, E. and B.M.J. Engelbrecht. 2007. Fungal endophytes
nearly double minimum leaf conductance in seedlings of a
neotropical tree species. Journal of Tropical Ecology 23: 369-
372.
Baeza J.A., Reitz JM, Collin R. 2007. Protandric simultaneous
hermaphroditism and sex ratio in Lysmata nayaritensis
Wicksten, 2000 (Decapoda: Caridea). Journal of Natural
History 41: 2843-50.
Baeza, J.A. 2007. Sex allocation in a simultaneously
hermaphoditic marine shrimp. Evolution 61-10: 2360–2373.
Baeza, J.A. 2007. Male mating opportunities affect sex allocation
in a protrandric-simultaneous hermaphroditic shrimp.
Behavioral Ecology and Sociobiology 61: 365-370.
Baeza J.A. 2007. No effect of group size on sex allocation in a
protandric-simultaneous hermaphroditic shrimp. Journal of
the Marine Biological Association 87: 1169-74.
Baeza J.A. 2007. The origins of symbiosis as a lifestyle in marine
crabs (genus Petrolisthes) from the eastern Pacific: does
interspecific competition play a role? Revista de Biologia
Marina y Oceanografia 42: 7-21.
Bolaños, M.D., S.Y. Quiroga and M.K. Litvaitis. 2007. Five
species of cotylean flatworms (Platyhelminthes: Polycladida)
from the wider Caribbean. Zootaxa 1650: 1-23.
Browne, W.E., S.H.D. Haddock and M.Q. Martindale. 2007.
Phylogenetic analysis of lineage relationships among
hyperiid amphipods as revealed by examination of the
mitochondrial gene, cytochrome oxidase I (COI). Integrative
and Comparative Biology 47: 815-830.
53
Collin R, Chaparro OR, Winkler F, Veliz D. 2007. Molecular
phylogenetic and embryological evidence that feeding larvae
have been reacquired in a marine gastropod. In, pp. 83-92
D’Croz L, O’Dea A. 2007. Variability in upwelling along the
Pacific shelf of Panama and implications for the distribution
of nutrients and chlorophyll. Estuarine, Coastal and Shelf
Science 73: 325-40
Crowley, P.H. and M.K. Hart. 2007. Evolutionary stability of
egg trading and parceling in simultaneous hermaphrodites:
The chalk bass revisited. Journal of Theoretical Biology 246:
420–429.
De Grave, S. 2007. A new species of Pseudocoutierea Holthuis
from the Caribbean coast of Panama (Crustacea, Decapoda,
Palaemonidae), with a key to the genus. Zootaxa 1397: 29-37.
Diaz, M.C., R.W. Thacker and K. Rützler. 2007. Haliclona
(Soestella) walentinae (Chalinidae, Haplosclerida) and
Xestospongia bocatorensis (Petrosiidae, Haplosclerida),
two new sponge species from Caribbean Panamá with
filamentous cyanobacterial symbionts. In: Custódio, M. R.,
G. Lôbo-Hajdu, E. Hajdu, and G. Muricy (eds.). Porifera
Research: Biodiversity, Innovation, and Sustainability, Série
Livros 28. Museu Nacional, Rio de Janeiro: 31-36.
Diaz, M.C., R.W. Thacker, K. Rützler and C. Piantoni. 2007.
Two new haplosclerid sponges from Caribbean Panama
with symbiotic filamentous cyanobacteria, and an overview
of sponge-cyanobacteria associations. In: Custódio MR,
Lôbo-Hajdu G, Hajdu E, Muricy G (eds). Porifera research:
biodiversity, innovation and sustainability. Série Livros 28.
Museu Nacional, Rio de Janeiro: 31-39.
Erwin, P.M. and R.W. Thacker. 2007. Incidence and identity of
photosynthetic symbionts in Caribbean coral reef sponge
communities. Journal of the Marine Biological Association
of the United Kingdom 87: 1683-1692.
Erwin, P.M. and R.W. Thacker. 2007. Phylogenetic analyses of
marine sponges within the order Verongida: a comparison
of morphological and molecular data. Invertebrate Biology
126(3): 220-234.
Erwin, P.M. and R.W. Thacker. 2007. Incidence and identity of
photosynthetic symbionts in Caribbean coral reef sponge
communities. Journal of the Marine Biological Association
of the United Kingdom 87: 1683-1692.

Gochfeld, D.J., C. Schloeder and R.W. Thacker. 2007. Effects of
anthropogenic disturbance on sponge community structure
and disease prevalence in Bocas del Toro, Panamá. In:
Custódio, M. R., G. Lôbo-Hajdu, E. Hajdu, and G. Muricy
(eds.). Porifera Research: Biodiversity, Innovation, and
Sustainability, Série Livros 28. Museu Nacional, Rio de
Janeiro. pp. 335-343.
Granek, E.F. and K. Frasier. 2007. The impacts of Red Mangrove
(Rhizophora mangle) deforestation on zooplankton
communities in Bocas del Toro, Panama. Bulletin of Marine
Science 80: 905-914.
Hochberg, R. 2007. Gastrotricha of Bocas del Toro, Panama: A
preliminary report. Meiofauna Marina 16: 101-107.
Hooge, M., A. Wallberg, C. Todt, A. Maloy, U. Jondelius and S.
Tyler. 2007. A revision of the systematics of panther worms
(Hofstenia spp., Acoela), with notes on color variation and
genetic variation within the genus. Hydrobiologia 592:
439–454.
Johnson K.G., J.A. Todd, and J.B.C. Jackson. 2007. Coral reef
development drives molluscan diversity increase at local and
regional scales in the late Neogene and Quaternary of the
southwestern Caribbean. Paleobiology 33: 24.
Kicklighter, C.E. and M.E. Hay. 2007. To avoid or deter:
interactions among defensive and escape strategies in
sabellid worms. Oecologia 151: 161–173.
Lessios, H.A. 2007. Reproductive isolation between species of
sea urchins. Bulletin of Marine Science 81: 191-208.
Lovelock, C.E., I.C. Feller, M.C. Ball, J. Ellis, A.M. Schwarz and
B. Sorrell. 2007. Growth Rate Hypothesis vs Geochemical
Hypothesis for variation in plant nutrients over latitude: a
test using mangroves. Ecology Letters 10: 1154-1163.
McKee, K.L, D. Cahoon and I.C. Feller. 2007. Caribbean
mangroves adjust to rising sea-level through biotic controls
on soil elevation change. Global Ecology and Biogeography
16: 545-556.
McPhail K.L, J. Correa, R.G. Linington, J. Gonzalez, E. Ortega-
Barrıa, T.L.Capson and W.H. Gerwick. 2007. Antimalarial
Linear Lipopeptides from a Panamanian Strain of the Marine
Cyanobacterium Lyngbya majuscula. Journal of Natural
Products 70: 984-988.
54
Neigel, J., A. Domingo and J. Stake. 2007. DNA barcoding as a
tool for coral reef conservation. Coral Reefs 26: 487-499.
O’Dea, A., J.B.C. Jackson, H. Fortunato, J.T. Smith, L. D’Croz,
K.G. Johnson and J.A. Todd. 2007. Environmental
change preceded Caribbean extinction by 2 million years.
Proceedings of the National Academy of Sciences 104:
5501–5506.
O’Dea, A., F. Rodriguez, C. DeGracia and A.G. Coates. 2007. La
paleontología marina en el Istmo de Panamá. Canto Rodado
2: 149-179.
O’Dea, A., F. Rodriguez and T. Romero. 2007. Response of
zooid size in Cupuladria exfragminis (Bryozoa) to simulated
upwelling temperature. Marine Ecology 28: 1-9.
Ordoñez, C., S.Troëng, A. Meylan, P. Meylan and A.Ruiz. 2007.
Chiriqui Beach, Panama, the most important leatherback
nesting beach in Central America. Chelonian Conservation
and Biology 6: 122-126.
Pearse, V.B. and O.Voigt. 2007. Field biology of placozoans
(Trichoplax): distribution, diversity, biotic interactions.
Integrative and Comparative Biology: 1-16.
Puebla, O., E. Bermingham, F. Guichard and E. Whiteman.
2007. Colour pattern as a single trait driving speciation in
Hypoplectrus coral reef fishes? Proceedings of the Royal
Society 274: 1265-1271.
Reynolds, R.G. and B.M. Fitzpatrick. 2007. Assortative mating
in poison-dart frogs based on an ecologically important trait.
Evolution 61: 2253-2259.
Rudh, A., B. Rogell and J. Hoglund. 2007. Non-gradual variation
in colour morphs of the strawberry poison frog Dendrobates
pumilio: genetic and geographical isolation suggest a role for
selection in maintaining polymorphism. Molecular Ecology Online.
Sanchez, J.A., C. Aguilar, D. Dorado and N. Manrique. 2007.
Phenotypic plasticity and morphological integration in a
marine modular invertebrate. BMC Evolutionary Biology
7:122. Online.
Saporito, R.A., M.A. Donnelly, R.A. Norton, H.M. Garraffo,
T.F. Spande and J.W. Daly. 2007. Oribatid mites as a major
dietary source for alkaloids in poison frogs. Proceedings of
the National Academy of Sciences 104: 8885-8890.

Saporito, R.A., M.A. Donnelly, P. Jain, H.M. Garraffo, T.F.
Spande and J.W. Daly. 2007. Spatial and temporal patterns
of alkaloid variation in the poison frog Oophaga pumilio in
Costa Rica and Panama over 30 years. Toxicon 50: 757-778.
Smith, M., K.S. Zigler and R.A. Raff. 2007. Evolution of directdeveloping
larvae: selection vs loss. BioEssays 29: 566–571.
Swain, T.D. and J.L. Wulff. 2007. Diversity and specificity of
Caribbean sponge-zoanthid symbioses: a foundation for
understanding the adaptive significance of symbioses and
generating hypotheses about higher-order systematics.
Biological Journal of the Linnean Society 92: 695-711.
Taylor, M.W., R.W. Thacker and U. Hentschel. 2007.
Evolutionary insights from sponges. Science 316: 1854-
1855.
Taylor, M.W., R.T. Hill, J. Piel, R.W. Thacker and U. Hentschel.
2007. Soaking it up: the complex lives of marine sponges
and their microbial associates. The International Society for
Microbial Ecology Journal 1(3): 187-190.
Tewfik, A., J.B. Rasmussen and K.S. McCann. 2007.
Simplification of seagrass food webs across a gradient of
nutrient enrichment. Canadian Journal of Fisheries and
Aquatic Sciences 64: 956-957.
Thacker, R.W., M.C. Diaz, K. Rützler, P.M. Erwin, S.J.A.
Kimble, M.J. Pierce and S.L. Dillard. 2007. Phylogenetic
relationships among the filamentous cyanobacterial
symbionts of Caribbean sponges and a comparison of
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55
Tomaiuolo, M., T.F. Hansen and D.R. Levitan. 2007. A
theoretical investigation of sympatric evolution of temporal
reproductive isolation as illustrated by marine broadcast
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Toth, E. and R.T. Bauer. 2007. Gonopore sexing technique
allows determination of sex ratios and helper composition in
eusocial shrimps. Marine Biology 151:1875–1886.
Troxler, T.G. 2007. Patterns of phosphorus, nitrogen and 15N
along a peat development gradient in a coastal mire, Panama.
Journal of Tropical Ecology 23: 683-691.
Uy, J.A.C. and A.C. Stein. 2007. Variable visual habitats may
influence the spread of colourful plumage across an avian
hybrid zone. Journal of Evolutionary Biology 20: 1847-1858.
Van Bael, S.A., P. Bichier, I. Ochoa and R. Greenberg. 2007. Bird
diversity in cacao farms and forest fragments of western
Panama. Biodiversity and Conservation 16: 2245-2256.
Vollmer, S.V. and S.R. Palumbi. 2007. Restricted gene flow in the
Caribbean staghorn coral Acropora cervicornis : Implications
for the recovery of endangered reefs. Journal of Heredity 98:
40-50.
Zamzow, J.P. 2007. Ultraviolet-absorbing compounds in the
mucus of shallow-dwelling tropical reef fishes correlate with
environmental water clarity. Marine Ecology Progress Series
343: 263-271.
Zeh, J.A. and D.W. Zeh. 2007. Mate choice by non-virgin
females contributes to reproductive isolation between
populations of the Harlequin beetle-riding pseudoscorpion.
Ethology 113: 1202-1211.

56
Acknowledgments
Rachel Collin Director
Gabriel Jacome Scientific Coordinator
Plinio Gondola Research Assistant
Dennis Allen Maintenance Supervisor
Urania Gonzalez Secretary
Scientific visitors to the BRS rely heavily on the station’s
staff, whose jobs include creative problem solving and
long hours as they welcome visitors and keep the station
running smoothly. Additional support for the BRS comes
from STRI’s headquarters in Panama City. Boat mechanics
Reynaldo Tapia and Pierre Fuentes and STRI’s Dive Safety
Officer, Edgardo Ochoa provide daily support and visit the
station monthly. Juan Mate and Orelis Arosemena work
closely with the station visitors to assist their applications
for collection and export permits.
During FY06-07, Director Eldredge Bermingham and
his staff, with the Development Office and the Special
Events Office, have worked to find supporters and to bring
them to Bocas where they experience research efforts first
hand. The Smithsonian Institution’s Office of Facilities,
Engineering and Operations (OFEO) has worked closely
with station staff to design and build new facilities and to
keep existing facilities operating well. Staff from STRI’s
Arcadio Castillo Research Technician
Doroteo Machado Maintenance
Gilberto Murray Mechanic Small Craft
Marlon Smith Public Program
Deselvia Mercado Janitor
Keyla Serrano Janitor
Mauricio Pineda Grounds Keeper
Office of Information Technology have kept the BRS
online and in communication with the rest of the world.
The Visitors Office in Panama City helps process the
visitors’ applications and Nelida Gomez and Nilka Tejada
coordinate visits from university classes. Procurement and
Accounting Offices keep supplies coming. The Human
Resources office has endeavored to understand the needs of
the staff.
None of this would be possible without the support of
the government of Panama and STRI’s External Affairs
Office under the direction of Advisor, Elena Lombardo.
The aquatic resources (ARAP) and environmental
(ANAM) authorities graciously grant collection and export
permits for research projects. The Ministry of Education
(MEDUCA) works closely with BRS teacher training. The
Ministry of Foreign Relations helps with visas, whereas
safety and security are coordinated with the National Police
and the Ministry of Health.

Financial Contributions
THE BRS GRATEFULLY ACKNOWLEDGES
DONATIONS TO BOCAS DEL TORO
MARINE STATION DURING FY2008-FY2009
Dr. and Mrs. David A. Cofrin have been deeply involved
over the past several years in Bocas del Toro’s expansion,
contributing most recently to the purchase of a glassbottom
boat and to the new dormitory, the construction of
which is imminent.
The Hoch Family (Hoch Charitable Trust) has
generously supported Bocas del Toro for almost a decade,
contributing most recently to senior scientist residences
and the new dormitory. In addition, the Hochs have funded
postdoctoral two- year fellowships for research at the BRS.
The Hunterdon Fund and the Johnson Fund through
the Smithsonian Marine Science Network, have provided
funding for the operational costs and salaries needed to
maintain Bocas del Toro, support for post-doctoral fellows
and contributions to the Training in Tropical Taxonomy
program.
Mr. Paul Peck’s gift to Bocas del Toro has made it possible
to extend the Training in Tropical Taxonomy Program into
FY09.
SENACYT , the Republic of Panama’s office of science
and technology, NSF, the United States National Science
Foundation, and the Encyclopedia of Life have provided
grants to support the Training in Tropical Taxonomy
Program.
The Smithsonian Institution Women’s Committee
has contributed generously over the past 5 years to the
documentation of biodiversity in Bocas del Toro as well as
to the Training in Tropical Taxonomy Program.
USAID and Chemonics provided support for the
watershed restoration project in Old Bank on the island of
Bastimentos.
STRI mourns the loss of our dear friends and
enthusiastic supporters:
Frank W. Hoch (1921-2007)
David A. Cofrin (1923-2009)
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Cofrin visit to Bocas del Toro. From left to right: Maria Inez Jaramillo,
Lisa Barnett, Edith Cofrin, Mary Ann Cofrin (mother), Mary Ann Cofrin
(daughter), Dr. David Cofrin, Rachel Collin, Tony Coates, Gregory Kiss,
David Dilcher. Carlos and Camilo Jaramillo were also present.
Lisina, Steven and Jane Hoch, in front of Hoch House.
U.S. Ambassador to Panama, Barbara J. Stephenson visited the station
with USAID representatives in August 2009. Shown here with Alejandro
Arze, STRI Protection and Security Manager (left) and Gabriel Jacome,
Bocas Research Station Manager (right).

Help a K-12 student visit the BRS: The low income of
residents in the Bocas del Toro region means that school
groups often do not have the funds available to reach the
BRS and participate in educational activities. We send our
public programs coordinator to visit the schools, but there
is no substitute for the hands-on experiences on-site at the
BRS. Only $5/student can help bring a class from a local
school for a visit.
Help an undergraduate or graduate student: The
Training in Tropical Taxonomy program is a one-of-a-kind
program that aims to counter the growing crisis in the loss
of expertise in taxonomy and the study of biodiversity. We
bring experts and students from around the world together
for intensive training workshops. Donations to this fund
will help provide fellowships to help a needy student
participate in this competitive program.
Director’s Discretionary Fund: Let station Director Dr.
Rachel Collin decide where your donation can do the most
to help outreach, education, public programs or science at
the BRS. Contributions to this fund are used to purchase
upgrades to microscopes and other scientific equipment
as well as to provide fellowships to excellent graduate
students working at the station. They can also be used to
supplement the outreach and education programs that the
BRS offers.
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Make a Donation
With your help we can do more…
To donate please see our web site at:
www.stri.org/BocasDonations
Contact Lisa Barnett at the Smithsonian Tropical Research
Institute’s Development Office for gifts of stock, planned
giving, or bequests.
Lisa Barnett: E-mail: barnettl@si.edu
Phone: 202-633-4014
Smithsonian Tropical Research Institute
P.O. Box 37012, MRC 705
Washington, DC 20013-7012

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The seagrass beds in Bocas del Toro are particularly rich in biodiversity.
The cushion stars, Oreaster reticulatus, are an attraction at Starfish
Beach. The feeding activity of the sea cucumber, Holothuria mexicana,
helps to keep the sediment clean.
We thank the following people for their contributions to this report:
Rachel Collin, Beth King, Ricardo Chong, Rebecca Rissanen, Lina Gonzalez, Annette Aiello, Javier Luque, Maria Fernanda Vinasco, Marcos Guerra,
Christian Ziegler, Steve Paton, Edgardo Ochoa, Arthur Anker, Chip Clark, Bryson Voirin, Rosana Rocha, Peter and Anne Meylan, Laura May-Collado,
Gayatri Thampy, Corrine Richards, Kristin Hultgren, Amy Freestone, Katie Cramer, Aaron O’Dea, Christian Sardet, Nancy Knowlton, Elisabeth
Kalko, TTT-students.

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To learn more about the BRS visit www.stri.org/bocas
e-mail: collinr@si.edu
BRS: (country code 507) 212-8550
STRI Panama: (country code 507) 212-8000
Washington D.C.: (202) 633-4014