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241 
Caribbean coral reefs of Panama: 
present status and future perspectives 
H6ctor M. Guzm~n 
Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Republic of Panama 
ABSTRACT: One of the goals of this work is to review the conservation status of the Panamanian 
Caribbean coral reefs and to demonstrate that there is sufficient and adequate information to support 
new legislation for their protection. Panama has 1,295 km of coastline in the Caribbean with coral reefs 
along almost all its coast, from Punta Cabo Tibur6n on the border with Colombia to Punta Boca del 
Drago, 27 km from the border with Costa Rica. The total coral reef surface area is estimated at 754 
km 2, mostly fringing reefs. Of these reefs, 81% are located in the eastern region and form part of the 
indigenous reservation of Kuna-Yala. In this region, there is also the highest diversity of corals, 
octocorals and sponges, with 97%, 100% and 93%, respectively, of the total number for the country. 
Live coral cover has changed dramatically in the last 20 years, with a reduction as high as 50-70% in 
several areas along the coast. At present the highest average cover ranges between 25% and 30%, to be 
found in the eastern and western regions. The reefs have been affected by natural disturbances such as 
seawater warming, associated with E1 Niflo events, and by diseases. However, from a historical point of 
view, anthropogenic disturbances have gone beyond our general comprehension. They started with 
massive coral mining more than 500 years ago, at the time of the Spanish conquest, and have continued 
up to the present day, during the construction of the Panama Canal and with the traditional coral mining 
and landfilling practices used by the Kuna people. Five marine-terrestrial protected areas along the 
Caribbean coast do not include the best reefs of Panama or protection is inadequately enforced. Reefs 
are at high risk due to coastal development particularly that associated with tourism; 48.5% of the 
country's tourist attractions are located along the coastal zone. This creates a pressing and unavoidable 
need to protect coral reefs as part of an integrated national coastal management and conservation plan. 
1. INTRODUCTION 
Recently, the global destruction of coral reefs has been observed with concern by 
scientists and authorities in charge of the management of natural resources. Among the 
main causes of degradation are overfishing, sedimentation and enrichment by nutrients 
(eutrophication) (Roberts 1993). The consequences Of these activities affect not only the 
structure and function of reef ecosystems, but also have a socio-economic impact on 
many countries of the Caribbean region. 
Coral reefs are highly productive and diverse ecosystems that play an important role 
in the maintenance of the coastal zone, bringing direct benefits to the surrounding 
human settlements and the economies of several nations (Hatcher 1988, 1990; Spurgeon 
1992). Among these benefits are the protection of the coastal zone against erosion, food 
production, drug production and recreational tourism. The appraisal of each of these 
Latin American Coral Reefs, Edited by Jorge Cort6s 
�9 2003 Elsevier Science B.V. All rights reserved. 
242 H.M. Guzmdn 
benefits is a very difficult task, but it is generally agreed that suitable protection will 
result in an overall healthier economy that will benefit the inhabitants of the coastal 
zone (Spurgeon 1992). 
At present, the lack of protection of these ecosystems is the main indirect cause of 
their degradation in the Republic of Panama. Inadequate management of the coastal 
zone and deforestation of surrounding areas have accelerated the destruction of most 
reefs locally at a time when the country is gearing up for an increase in tourism 
development that, among other plans, contemplates the recreational use of these 
habitats. Virtually all Panamanian coast containing reefs has been incorporated into the 
national master plan for tourism development (IPAT/OEA 1993) and one would expect 
that before its implementation, a general land-use plan for the country would be in place 
to guide the long-term management of these resources, with due concern given to the 
coastal communities. Nevertheless, the trend of accelerated degradation of both reefs 
and local economies implies quite the contrary. 
This work provides, for the first time, a general review of our knowledge of the 
Caribbean coral reefs of Panama and an assessment of their conservation status. It aims 
also to demonstrate that there is sufficient and adequate baseline information about 
these reefs and related resources for use by local authorities to support the new and 
sound legislation that is urgently required for the long-term sustainable conservation of 
this ecosystem. An integrated conservation and management plan for the coastal zone is 
a pressing and unavoidable need. 
1.1. Hydrography and meteorology of Panama's Caribbean coast 
Panama has 1,295 km of coastline on the Caribbean side, characterized by a narrow 
continental shelf with a variable width that ranges from 5.5 km to a maximum of 39 km 
and an approximate surface area of 6,000 klTl 2 (IGNTM 1988). Caribbean tides in 
Panama are semidiurnal or diurnal (mixed), not clearly predictable and with an ampli- 
tude below 0.5 m. Tides are higher than usual during the dry season due to the influence 
of northeasterly winds (Glynn 1972). North and northeast winds have a strong influence 
along the coastal zone throughout the year, mostly from January to April (DMA 1988; 
Cubit et al. 1989). The central coast is particularly affected by wind-driven waves and 
tides during the dry season, while they have a lesser impact on the leeward side of the 
eastern (Kuna-Yala) and western (Bocas del Toro) archipelagos where the largest reef 
tracts occur. The main coastal current, coming from northern Nicaragua and Costa Rica 
and nmning eastbound, affects Panama throughout the year with possibly a greater 
influence between the months of June and August when it rtms closer to the coast 
(DMA 1988; Greb et al. 1996). 
The Caribbean is not subject to seasonal upwelling but it is affected by strong runoff 
from several rivers and by high rainfall (Cubit et al. 1989). Temperature remains 
relatively stable throughout the year (< 3~ variation) with the highest temperature 
recorded during the rainy season (April-December) (Cubit et al. 1989). Nutrients, 
plankton, turbidity and chlorophyll are less abundant in the eastern region of the country 
where the oceanographic conditions are typically oligotrophir (D'Croz and Robertson 
1997; D'Croz et al. 1999). Nevertheless, this pattern does not seem to hold along the 
western region, around Bocas del Toro, where the fiver runoff and rainfall are higher 
(Guzmfin and Guevara 1998a; L. D'Croz per. com.). 
The Caribbean coral reefs of Panama 243 
1.2. Research history 
The Panamanian reefs have been the subject of several studies over more than a 
century. However, it is difficult to establish when research started, because the records 
of several expeditions carried out during the 19 th century, many of them associated with 
the search for a shorter route for the building of the Panama Canal, are not easily 
available or well documented. Nevertheless, it is true that research in Caribbean Panama 
from that period onwards, and possibly up to the mid 20 th century, paid special attention 
to hydrographic and oceanographic descriptions and included massive collection cam- 
paigns and the description of new species of corals and reef-related organisms. For 
example, in 1874 Thomas Selfridge remarks in his book "Reports and Explorations and 
Surveys of the Isthmus of Darien" that during his San Blas (Kuna-Yala) explorations he 
used navigation charts prepared by Commander Lull with detailed localization of coral 
reefs (Heckadon-Moreno 2002). The U.S. Fish Commission steamer Albatross dredged 
the coast off Col6n in 1884;one of the collected octocoral species, Nicella schmitti, was 
described later (Bayer 1961). During the zoological explorations of the Italian naturalist 
Enrico Festa in 1895, reef flats close to the city of Col6n were briefly described 
following an all but unsuccessful attempt to dredge the bottom but which ended up with 
the collection of organisms during the extreme low tide (Heckadon-Moreno 2001). 
Many such collections took place in shallow areas or by dredging and among some 
interesting records are those of Meek and Hildebrand (1923) who collected and identi- 
fied some 237 fish species along the "extensive reefs" from Col6n to Portobelo between 
1911 and 1912. Taylor (1929), collected and made observations on the macroalgae of 
reef and rocky areas and deLaubenfels (1936) collected sponges manually and by raking 
the intertidal zone. He reported 21 species of which 33% were hitherto unknown to 
science. Likewise, Rubinoff and Rubinoff (1962) produced new fish records for the 
Caribbean central coast. 
However, it was only shortly after the mid-20th century, and with the support of the 
Smithsonian Tropical Research Institute, established in the old Canal Zone, that the 
more detailed studies of the coral reefs along Caribbean Panama began. The first re- 
search work took place at Galeta Island, about 15 km east of the entrance of the Panama 
Canal and at Kuna-Yala (San Blas), in the eastern region of the country, where, in the 
early 1970s, the first two marine laboratories, or field stations, in Caribbean Panama, 
and possibly in all Central America, were built. In 1974, the Smithsonian Institution 
Environmental Sciences Program was established in Galeta, with the purpose of 
monitoring long-term climatic and hydrological changes (Meyer and Birkeland 1974); 
this work is still in progress today (Cubit et al. 1989) and it has been expanded to other 
regions of the country. About 350 contributions have been published from the marine 
and coastal environments nearby or from around the central region. There, research has 
focused on population dynamic studies and the behavior of plants and animals from reef 
flats and subtidal reef areas (Caldwell 1985; Cubit et al. 1986; Sebens 1982; Lasker 
1979, 1980). Herbivory processes in reefs (Hay 1981, 1984) and the impact of petro- 
leum on coral reproduction, reef structure and subsequent reef recovery have also been 
studied (Burns and Knap 1989; Jackson et al. 1989; Guzrrfin et al. 1991, 1994; Guzmfin 
and Holst 1994). Meanwhile, at the Kuna-Yala field station several research projects 
were initiated, with 150 scientific reports published up to its closure in 1997. These pro- 
jects brought an enormous contribution to the knowledge of the reproductive and evolu- 
tionary biology of octocorals (Brazeau and Lasker 1988, 1990, 1992; Coffroth et al. 
244 H.M. Guzmdn 
1992; Coma and Lasker 1997; Lasker 1985, 1990; Lasker et al. 1996, 1999), echino- 
derms (Lessios 1981, 1987, 1991, 1998; Lessios et al. 1984b)and reef fish (Cole 1988; 
Hoffman and Robertson 1983; Robertson 1984, 1992; Robertson et al. 1976, 1988, 
1993, 1999; Warner 1975, 1984, 1990; Warner and Hoffman 1980), to mention but a few. 
Among some of the pioneering work, stands out the concern about the imminent 
migration of marine organisms throughout the Panama Canal (Chickering 1930; 
Rubinoff 1965, 1968; Rubinoff and Rubinoff 1968), the impact of petroleum on the 
Col6n coastal reefs (Rtitzler and Sterrer 1970; Birkeland et al. 1976), coral mining in 
Kuna-Yala (Porter and Porter 1973), as well as the comparisons of the biota on both 
sides of the isthmus (Glynn 1972; Porter 1972a, b, 1974). These authors described for 
the first time the structure, diversity and distribution of coral reefs off the eastern and 
central regions of Caribbean Panama, from the mouth of the Chagres River to the 
Diablo River in Nargana, Kuna-Yala. 
It is worth mentioning some classical contributions, amongst which are a few 
controversial ones, which have set guidelines for further basic research in the Caribbean 
region. Included amongst these are: biological changes in marine ecosystems associated 
with the Panama Canal and the possible construction of a sea-level canal (Rubinoff 
1968, 1970; Topp 1969), sexual selection and sex change in reef fishes (Warner et al. 
1975), tests on the molecular clock in sea urchins (Lessios 1979), massive mortality of 
the long-spined sea urchin Diadema antillarum (Lessios et al. 1984a), settlement of 
coral larvae due to chemical stimuli from coralline algae (Morse et al. 1988), long-term 
population changes in reef fishes as a consequence of the Diadema die-off (Robertson 
1987, 1988, 1991), the impact of petroleum upon reefs and other coastal communities 
(Jackson et al. 1989) and the controversial definition of the Montastraea annularis sib- 
ling species complex (Knowlton et al. 1992). 
2. DESCRIPTION OF THE REEFS 
2.1. Geological origin 
Panama's origin is reflected mostly in the formation of the Central American isth- 
mus, the closure of which affected the circulation patterns of the oceans and the climate 
worldwide, with changes in the distribution of plants and animals (Coates 1997). In 
geological terms, the Central American region is of recent origin, starting with the 
formation of islands in what is currently eastern Panama and the subsequent formation 
of the southern half of the isthmus in the mid-Miocene, about 11 million years ago. In 
the following million years (Ma), the region became an archipelago and the circulation 
between the Caribbean and the Pacific Ocean was restricted, with the creation of new 
coastal-marine habitats (Coates 1997). At the end of the Miocene (7-6 Ma), the depth of 
the oceans surrounding Panama was about 150 m and by the end of the Pliocene (ca. 4 
Ma) they had become shallower, with depths less than 50 m. The isthmus was probably 
closed about a million years later. It is presumed that towards the end of the Pliocene 
there were still marine corridors that connected the two oceans until their final closure 
(Coates et al. 1992; Coates and Obando 1996; Coates 1997). The definitive separation 
of the two oceans took place in the southern region of Central America with the Isthmus 
of Panama being the last land to emerge (Coates et al. 1992; Collins and Coates 1999). 
After the closure of the isthmian barrier, a temporary breach may have occurred during 
The Caribbean coral reefs of Panama 245 
the Ice Age glaciation and associated interglacial warming and changes in sea level 
(Coates and Obando 1996; Coates 1997). It is estimated that over the past 20,000 years the 
sea level may have risen up to 135 m as modem glaciers have been melted (Coates 1997). 
Although small patch reefs appeared in Panama's Caribbean by the mid-Miocene, 
large-scale coral reefs and the associated coralline sediments started to appear mainly in 
the upper Pliocene to the Pleistocene, occurring at relatively deep levels of 80-100 m 
during this latter period (Collins et al. 1999). During the formation of the isthmus and 
particularly during the Pliocene and Pleistocene turnover periods (4-1 Ma), there were 
speciation and extinction episodes that dramatically transformed the Caribbean's coral- 
line fauna; approximately 80% of the more than 100 coral species of the Miocene- 
Pleistocene disappeared and 60% of the current species were originated. The diversity 
of corals in the Caribbean zone off Costa Rica and Panama during this period was 
greater than that in other sites in the Caribbean region, although the change off Panama 
started more recently (3-2.2 Ma), completing itself by about 1.6-1.2 Ma (Budd et al. 
1999). According to Budd et al. (1999), the diversity of coral species found in the fossil 
record is higher off Costa Rica (82 species) than off Panama (35 species) (Budd et al. 
1999). This is dissimilar to the record of present day species which shows that Panama 
almost doubles (>65 species) the diversity of Costa Rica(Guzmfin and Guevara 1998b, 
2001). Regarding fossil azooxanthellate corals, the highest diversity of the wider Carib- 
bean appears off the Dominican Republic and Panama, with 20 and 15 species respec- 
tively, although this possibly reflects a greater collection effort in these regions (Cairns 
1999). 
Even though not all the reef structures of the country have been dated in order to 
determine their origin, it is considered that recent reefs (Holocene) started to develop in 
Caribbean Panama about 7,000 years ago with the formation of fringing reefs typical of 
the Caribbean and dominated by Acropora palmata (see Macintyre and Glynn 1976). 
This species built reefs mainly in exposed areas and allowed the development of other 
massive coral species such as Diploria strigosa, Stephanocoenia michelinii, Porites 
astreoides and Montastraea annularis along back reef areas and upper and intermediate 
slopes about 5,000 years ago. At the same time, a lagoon environment developed and 
species such as P. furcata and A. cervicornis appeared (Macintyre and Glynn 1976). A. 
palmata kept pace with the rising and changing sea level that occurred at the end of the 
Holocene transgression when rapid reef growth took place. The accretion rate of these 
reefs, based mainly on this dominant species, fluctuated between 10.8 and 1.3 m/1000 
years, with an average of 3.9 m/1000 years (Macintyre and Glynn 1976). Within the 
3,000 to 2,000 year B.P. period, the dominance of A. palmata started to decrease. 
Vertical development of these reefs was subsequently limited by changes in the sea 
level and possible sedimentation during the last 2,000 years; corals reduced their active 
contribution to the construction of new reef framework, particularly in habitats exposed 
to high energy (Macintyre and Glynn 1976; Macintyre et al. 2001). 
2.2. Reef distribution and changes in coral cover 
Panama has coral reefs along almost all its Caribbean coast, from Punta Cabo Tibu- 
r6n on the border with Colombia (08~ - 77~ to Punta Boca del Drago 
(09~ 82~ 27 km from the border with Costa Rica (Fig. 1). The 
development and structure of these reefs varies dramatically with the degree of wave 
and wind exposure and their proximity to the mainland, being better developed on the 
246 H.M. Guzmrn 
Fig. 1. Map of the Republic of Panama showing the four political provinces on the Caribbean side (Kuna- 
Yala, Colrn, Veraguas and Bocas del Toro) and the three arbitrarily defined regions where the main reef 
structures have been described until now (eastern, central, western; indicated between diamond symbols). 
From east to west, the eastern region is divided into three sub-regions and limits are indicated (cross 
symbols); Tubuala, Ailigandi, Nargana. Details for the Bocas del Toro archipelago are shown in Figure 7. 
leeward side of side of insular systems. In particular, the best reef growth and 
development takes place in the Bocas del Toro and Kuna-Yala archipelagos, where 
there are hundreds of cays, islands and islets surrounded by fringing coral reefs. These 
islands are of calcareous or volcanic origin and their surfaces range between a few m 2 
and 61 km 2 (Isla Colrn, Bocas del Toro). 
In order to simplify the presentation of this work in the following sections, the 
Caribbean coast has been arbitrarily divided into three large regions (Fig. 1), partly 
because the greatest reef development was found in them, but also because these were 
the areas where most reefs have been described to date and information is available. 
These areas, in turn, have been subdivided into smaller subregions, sometimes based on 
political units, which are suggested as potential areas of conservation or management. 
Likewise, this section shows existing trends in coral cover for the intervening regions, 
based on historical records and current monitoring programs. 
2.2.1. Eastern region. If we analyze the distribution of reefs by political unit or at 
provincial level, we fred that the most extensive and best-developed reefs off Panama 
are located at Kuna-Yala (San Bias), with approximately 610 km 2. So far, reef surface 
area has been estimated using only satellite images and aerial photographs for this 
region. These reefs are part of the indigenous reservation or "Comarca" of Kuna-Yala, 
an area entirely under the custody and management of the Kuna autonomous people and 
without interference from the government of Panama since 1938 (Tice 1995; Ventocilla 
et al. 1995; Howe 1998). The archipelago is made up of about 365 coral islands 
(Ventocilla et al. 1995). Along its length, hundreds of shallow reef patches cover more 
than 26 km2; most of them close to the mainland. Fringing coral reefs run along the 
The Caribbean coral reefs of Panama 247 
continental coast forming extensive reef flats bordered by gentle slopes down to 10-15 m. 
About 2-5 km offshore and around the islands and cays, large reef complexes develop 
with several habitats including internal enclosed lagoons with coral-dominated slopes, 
external barriers and deep reefs. Seaward reef fiats that are part of the external barriers 
are vast, dominated by spurs and grooves up to 15 m deep, and covered mainly by 
crustose coralline algae and several coral species that are resistant to wave action, e.g. 
Agaricia agaricites, Porites astreoides, Diploria spp. and Siderastrea siderea, among 
others. It is important to point out that these reef fiats were described at first as algal 
ridges, especially those of the Cayos Holand6s (Glynn 1973). Recently it was discov- 
ered that these structures had been formed about 2,000-2,800 years ago and were built 
with extensively lithified Agaricia and Millepora rubble, which is characteristic of storm 
deposits coated by cmstose coralline algae (Macintyre et al. 2001). Therefore, they may 
not be considered as true algal ridges (sensu Adey 1978). 
, Within this province or eastern region (Fig. 1), the largest reef tracts occur in the 
'corregimiento" (jurisdiction) of Nargana (ca. 365 km2), followed by Ailigandi (ca. 152 
km 2) and Tubuala (ca. 93 km2). Active vertical reef development can be observed to 
more than 45 m deep in the Nargana sub-region, particularly in areas that are farther 
away from the mainland, such as Cayos Holand6s and Limones. 
The reefs of Punta San Blas, Nargana, have been surveyed and monitored for more 
than 30 years (see Porter 1974). Changes in live coral cover show a gradual decrease 
from ca. 40% in 1983 (see Shulman and Robertson 1996) to ca. 15% in 1997, with a 
slight recovery of ca. 5% for 2001 (Fig. 2). 
Qx Kuna-Yala 
o~ 30 - 
L_ 
> 
O 
-- 20 - 
0 
> 
�9 n 1 0 - 
.... 
4 0 - 
Isla Grande 
Bahia Mina,, 
Bocas del Toro 
I " I I . . . . I I I " I " I ' 1 ' I ' ' ' 1 ' " I' I . . . . . . . 
83 85 86 88 90 92 93 97 98 99 00 01 02 
Year Survey 
Fig. 2. Long-term changes in live coral cover in four reef areas along the Caribbean coast of Panama. 
Analyses based on published (Guzmhn et al. 1991" Shulman and Robertson 1996) and unpublished 
(Smithsonian Institution Marine Environmental Science Program) data sets. 
248 H.M. Guzm6n 
2.2.2. Central region. This region includes the Province of Col6n, with an estimated 
reef surface area of 48 km 2, from the Kuna-Yala border to the Bel6n River, the western 
provincial border. The most extensively studied reefs in this region are located between 
Isla Grande and the mouth of the Chagres River, including Portobelo and Bahia Las 
Minas (Guzrn,hn et al. 1991; Guzrn~n and Hoist 1994). All these fringing reefs follow 
the mainland coastline throughout their extension with little vertical development - not 
more than 15 m deep. This is due mainly to the effect of the northeast trade winds and 
ocean waves that have a direct influence on this coastal region throughout the year. 
The best development occurs in the areas of Isla Grande and Bahia Las Minas (Fig. 1). 
In both regions, macroalgae cover comprises 80% in most reefs (Guzmhn et al.1991; 
Guzmhn and Hoist 1994). Live coral cover has changed remarkably in the reefs of Isla 
Grande-Portobelo since 1985, where there were reefs with an average cover of approxi- 
mately 25% that, following natural and anthropogenic disturbances (see Sections 3 and 
4), gradually decreased to levels close to 10% in 1992 (Fig. 2). As in the eastern region, 
a slight recovery of about 7% (Fig. 2) was observed in the late 1990's. The reefs of this 
region are composed mainly of Diploria clivosa, Agaricia tenuifolia, A. danai and 
Millepora complanata coral species in shallow areas of up to 2 rn deep, bordered by a 
relatively short slope dominated by D. strigosa, Colpophyllia spp. and Siderastrea 
siderea. Currently, small and isolated patches of Acropora palmata are found between 
Isla Grande and near Portobelo where they formerly made up vast fringing reefs parallel 
to the coast. The reefs of this subregion have shorter reef fiats and a greater number of 
shallow and deep reef patches. 
Towards the Bahia Las Minas area, live cover accounted for 22% in 1985 but 
decreased in 1986 due to a major oil spill (Jackson et al. 1989; Guzrn~n et al. 1991), 
reaching about 5% by the end of 1992 with a very slight recovery of ca. 5% towards 
1998 (Fig. 2). The composition of species and zonation is similar to those previously 
mentioned, but with a greater abundance and development of Siderastrea siderea 
colonies. This subregion has large reef fiats with a surface area of 4 to 24 ha (Guzrn~n 
and Hoist 1994) constantly exposed to high wave energy and quite short reef slopes of 
less than 10 m. The three largest and most representative reefs of this subregion, Isla 
Payardi, Isla Largo Remo and Isla Galeta, have areas of 48 ha, 44 ha and 35 ha, respec- 
tively (Guzrnhn and Hoist 1994). 
The reefs located west of the Panama Canal entrance were studied only in the early 
90s (Guzn~n and Hoist 1994). In general, they are not highly developed and have a 
quite homogeneous relief from the reef fiat towards the base. Siderastrea siderea, 
Diploria clivosa and Millepora complanata dominate the reef environments of this 
sector (Guzmhn and Hoist 1994). A total of seven reefs have been described west of 
Bahia Las Minas (Punta Galeta) as far as the mouth of the Chagres River, most of which 
are formed by vast reef fiats of up to 23 ha. The largest of these reefs located in Isla 
Media Palma has reef fiats of 16 ha and subtidal reefs of 18 ha (Guzrr~n and Hoist 
1994). The combined coral cover for this sector is low, ranging between 3% and 11% 
by 1992 (Guzmhn and Hoist 1994). It is important to point out that vast reefs existed 
within the Bahia Lim6n, the entrance to the Panama Canal. However, they were 
completely destroyed (see section 4) and, currently, there are only small patches of up 
to 4 m deep dominated by Millepora and Siderastrea corals located on the western 
margin of the bay and parallel to the old Fort Sherman. 
The Caribbean coral reefs of Panama 249 
2.2.3. Western region. There are reefs all along the coast from the Chagres River to the 
Calov6vora River where the western region begins (see Fig. 1). These reefs, which 
cover the western part of the Province of Col6n and the coast of Veraguas, have been 
visited but not documented to date. They might be described as extensive low relief reefs 
or subtidal reef platforms, composed mainly of macroalgae and the corals Siderastrea 
siderea and Diploria clivosa. Their vertical development is limited by the direct impact 
of waves and winds that may not allow the formation of intertidal reef fiats either. At 
the base of these reefs, small patches of Agaricia spp. and sponges are often found. 
The area defined here as the western region comprises the whole Bocas del Toro 
archipelago (ca. 3,500 km2), formed by 6 large forested islands and hundreds of small 
mangrove cays (Fig. 1). Surrounding all these islands and along a large part of the 
mainland coast, fringing coral reefs up to 20 m deep and dozens of shallow reef patches 
have developed. The archipelago has an estimated reef surface area of 87 km 2, the 
second largest of the Caribbean of Panama. The reefs of the archipelago have been 
evaluated in detail since 1997 and live coral cover of up to 50% has been reported, 
depending on the depth, reaching up to 90% in shallow areas (Guzrmin and Guevara 
1998a, b, 1999, 2001). Leeward reefs are generally more developed than those off the 
exposed northern shore of the islands (Guznfin and Guevara 1998b), with a somewhat 
characteristic zonation with shallow areas built mainly by Porites furcata, bordered by 
Agaricia danae or A. tenuifolia on the upper slope. From the intermediate slope area 
towards the reef base, the reefs are composed of massive corals, mainly Colpophyllia 
spp., Montastraea franksi, M. cavernosa, Siderastrea siderea and Stephanocoenia 
intercepta. Typical deeper habitat coral species, Agaricia undata and A. lamarcki, are 
observed at 15-20 rn deep. In Bahia Almirante and parallel to the coast from Puerto 
Almirante to the northern side of Isla Crist6bal, there is an almost continuous fringing 
reef more than 30 km long, composed exclusively of Porites furcata in the shallower 
areas (Guzrrfin and Guevara 1998a, b). It is important to note that within the archipelago 
there are healthy and vast populations of Acropora palmata and A. cervicornis corals 
(Guzrmin and Guevara 1998a, 1999, 2001), now considered endangered throughout the 
Caribbean (Precht et al. 2002). Over the past four years, unlike the rest of the country, a 
gradual decrease of up to 10% in the live coral cover has been observed (Fig. 2). 
2.3. Reef conservation status 
Caribbean reefs have been visited relatively recently and information about live coral 
cover includes data from 1998 to date. Almost all the reefs along Caribbean Panama 
have been affected by natural or man-made disturbances in one way or another (see 
sections 3 and 4), and in some locations these temporal changes have been recorded as 
part of monitoring programs in combination with historical records (Ogden and Ogden 
1994; Shulman and Roberson 1996; Guzrrfin et al. 2003) (Fig. 2). 
Figure 3 summarizes our current conservation status for most Caribbean reef areas in 
Panama. We observe that none of the subregions has an average live coral cover higher 
than 30%; the reefs of the Nargana sector in Kuna-Yala are those in the best condition 
but under high risk (Guzn~n et al. 2003). The worst conserved reefs are those located in 
the central region of the country, from the Chagres River to Isla Grande, with live coral 
cover lower than 15% (Fig. 3). It is important to point out that these cover values 
represent a wide spatial scale and that they are the result of a combination of observa- 
tions from several reefs described individually within each subregion, with a minority of 
250 H.M. Guzm6n 
30 
L _ 
o 20 
0 
0 0 10 
~ 
_.1 
0 
14 
6 
. - - - - 15 
Tubuala Ailigandi Nargana 
29 42 
Isla Bahia Bocas Bocas 
Grande Minas Mainland Islands 
Fig. 3. Summary of most recently updated percentage live coral cover in seven sub-regions along the 
Caribbean coast of Panama. Survey years are 2001 for Tubuala-Nargana, 1998 for Isla Grande-Bahia 
Las Minas and 1998-2000 for Bocas del Toro. Number of reefs is indicated at the top of each bar. 
reefs showing cover reaching 50%. Nevertheless, we believe that the statistical data 
from the more than 100 reefs quantitatively described, truly represent the spatial scale 
that is necessary to update the country's status because both highly degraded and 
relatively pristine reefs have been included in the analysis. 
2.4. Diversity 
The diversity of hermatypic scleractinian corals in Caribbean Panama was estimated 
at 49 species in the early 1970s, excluding four species of Millepora (Porter 1972a). 
This diversity, all found in the Kuna-Yala region, seemed impressive and was similar to 
the diversity observed for other reefs across theCaribbean, such as those off Jamaica 
(Porter 1972a). The absence of the species Dendrogyra cylindrus was considered curious 
because this coral was abundant off all the Caribbean islands. Likewise, the author 
questioned the presence of Solenastrea off Panama. 
Almost two decades later, the presence of 61 species of hermatypic corals is reported 
off Panama, including three species of Millepora (Hoist and GuzmAn 1993). This list 
includes for the first time, the presence of D. cylindrus off the central region of the 
country, confrere the presence of two species of Solenastrea, includes a new, recently- 
described species, Porites colonensis, off Col6n and includes separately the three sibling 
species of the now controversial Montastraea complex (sensu Knowlton et al. 1992). In 
this chapter, the list of species for Caribbean Panama is updated, including the distribu- 
tion of coral species at sub-regional level, in addition to octocorals and sponges. The 
diversity of corals increased to a total number of 70 species, including four species of 
Millepora (Table 1). Among the new records is the species Goreaugyra memoriales, 
which has only been found in Kuna-Yala, the country's highest diversity area (97%). 
The status of this species has been recently questioned (see Veron 2000). The central 
and western regions contain 77% and 87%, respectively, of the country's diversity. A 
lower number of species is found in the Bahia Las Minas area, with 51 species, while 
the highest diversity is found in Nargana, with 66 species (Table 1). 
The Caribbean coral reefs of Panama 251 
TABLE 1 
List of scleractinian and hydrocoral (Millepora) coral species from the Caribben coast of Panama by 
study regions. 1, Tubuala; 2, Ailigandi; 3, Nargana; 4, Portobelo-Isla Grande; 5, Rio Chagres-Bahia Las 
Minas; 6, continental areas; and 7, insular areas. 
Species Eastern Region Central Region Western Region 
1 2 3 4 5 6 7 
A cropora cervicornis X X X X X X X 
A cropora palmata X X X X X X X 
Acropora prolifera X X 
Agaricia agaricites X X X X X X X 
Agaricia carinata X X X X 
Agaricia crassa X X 
Agaricia danai X X X X X X X 
Agaricia fragilis X X X X X X X 
Agaricia grahamae X X X X 
Agaricia humilis X X X X X X X 
Agaricia lamarcki X X X X X X X 
A garicia purpurea X X X X X X X 
Agaricia pusilla X X X X X X X 
Agaricia sp. X X 
Agaricia tenuifolia X X X X X X X 
Agaricia undata X X X X 
Colpophyllia amaranthus X X X 
Colpophyllia breviserialis X X X X X X X 
Colpophyllia natans X X X X X X X 
Goreaugyra memorialis X 
Dendrogyra cylindrus X 
Dichocoenia stellaris X X X X X 
Dichocoenia stokesi X X X X X X X 
Diploria clivosa X X X X X X X 
Diploria labyrinthif ormis X X X X X X X 
Diploria strigosa X X X X X X X 
Eusmilia fastigiata X X X X X X X 
Favia fragum X X X X X X X 
Isophyllastrea rigida X X X X X X X 
Isophyllia sinuosa X X X X X X X 
Leptoseris cucullata X X X X X X X 
Madracis decactis X X X X X X X 
Madracis formosa X X X 
Madracis luciphila X X X X X X 
Madracis mirabilis X X X X X 
Madracis pharensis X X X X X X X 
Madracis senaria X 
Manicina areolata X X X X X X X 
Manicina mayori X X X X X 
Meandrina brasiliensis X X X X X 
252 H.M. Guzm6n 
Table 1 cont. 
Species Eastern Region Central Region Western Region 
1 2 3 4 5 6 7 
Meandrina danae X X X 
Meandrina meandrites X X X X X X X 
Millepora alcicornis X X X X X X X 
Millepora complanata X X X X X X X 
Millepora squarrosa X 
Millepora striata X X X X X X X 
Montastraea annularis X X X X X X 
Montastraea cavernosa X X X X X X X 
Montastraea faveolata X X X X X X X 
Montastraea franksi X X X X X X X 
Mussa angulosa X X X X X X X 
Mycetophyllia aliciae X X X X X X X 
Mycetophyllia danaana X X X X X X X 
Mycetophyllia ferox X X X X X X X 
Mycetophyllia lamarckiana X X X X X X X 
Mycetophyllia reesi X X X X 
Oculina difusa X X 
Porites astreoides X X X X X X X 
Porites branneri X X 
Porites colonensis X X X X X X X 
Porites divaricata X X X X X X X 
Porites furcata X X X X X X X 
Porites porites X X X X X X X 
Scolymia cubensis X X X X X X X 
Scolymia lacera X X X X X X X 
Siderastrea radians X X X X X X 
Siderastrea siderea X X X X X X X 
Solenastrea bournoni X X X X X �9 
Solenastrea hyades X X X X X X X 
Stephanocoenia intersepta X X X X X X X 
Total 70 56 61 66 54 51 57 59 
Soft corals or octocorals represent 38 species, with the highest diversity in Kuna- 
Yala (Table 2). The only previous list containing this information concerned Kuna-Yala 
(Clifton et al. 1997). It reported 26 species without including the genera Iciligorgia and 
Nicella. N. schmitti was previously described for Panama (Bayer 1961) and, according 
to the report, was widely distributed in all the regions. Two possible new species for 
Kuna-Yala are included in the current list (Table 2). One hundred percent of the 
country's species are present in the eastern region, whereas only 60.5% and 82% are 
found in the central and western regions, respectively. It is worth mentioning that only 
24 species were known for the eastern coast of Mexico and Central America (Bayer 
1961). 
The Caribbean coral reefs of Panama 253 
TABLE 2 
List of octocorals species from the Caribben coast of Panama by study regions. 1, Tubuala; 2, Ailigandi; 
3, Nargana; 4, Portobelo-Isla Grande; 5, Rio Chagres-Bahia Las Minas; 6, continental areas; and 7, 
insular areas. 
Species Eastern Region Central Region Western Region 
1 2 3 4 5 6 7 
Briareum asbestinum 
Erythropodium caribaeorum X X 
Ellisella barbadensis X X 
Ellisella elongata X X 
Ellisella sp. X 
Eunicea calyculata X X 
Eunicea fusca X X 
Eunicea laxispica X X 
Eunicea mammosa X X 
Eunicea succinea X X 
Eunicea tourneforti X X 
Gorgonia flabellum X X 
Gorgonia mariae X X 
Gorgonia ventalina X X 
Iciligorgia schrammi X X 
Muriceopsis flavida X X 
Muriceopsis sulphurea X X 
Muricea elongata X X 
Muricea laxa X X 
Muricea muricata X X 
Muricea atlantica X 
Muricea pinnata X X 
Nicella schmitti X X 
Nicella sp. 1 X 
Mi ra l I n ~n 9 ~)( 
= 
Plexaura kuna X X 
Plexaura flexuosa X X 
Plexaura homomalla X X 
Plexaurella nutans X X 
Plexaurella sp. 
Pseudoplexaura porosa X X 
Pseudopterogorgia acerosa 
Pseudopterogorgia americana X X 
Pseudopterogorgia bipinnata X X 
Pseudopterogorgia kallos 
Pterogorgia anceps X X 
Pterogorgia citrina X X 
Pterogorgia guadalupensis X X 
Total 38 30 34 
X X X X X 
X X X X X 
X X X X 
X 
X X X X X 
X X X X X 
X X X X X 
X X X X X 
X X X X X 
X X X X X 
X X X X X 
X X 
X X X X X 
X 
X X X X X 
X X X X 
X X 
X X 
X X X X X 
X X X 
X X X 
X X X X 
X X X 
X X X X X 
X X X X X 
X X X 
X X X 
X X X X X 
X X 
X X X X X 
X X X X 
X 
X X X X X 
X X X X 
X X 
33 23 23 25 29 
254 H.M. Guzmdn 
TABLE 3 
List of sponges from the Caribben coast of Panama by study regions. 1, Tubuala; 2, Ailigandi; 3, 
Nargana; 4, Portobelo-Isla Grande; 5, Rio Chagres-Bahia Las Minas; 6, continental areas; and 7, insular 
areas. 
Easto'n Region 
1 2 3 
Central Region Western Region 
4 5 6 7 
Acarnus souriei X 
Adocia implexiformis X 
Agelas clathrodes X 
Agelas confiera X 
Agelas dispar X 
Agelas flabelliformis X 
Agelas sceptrum X 
Agelas schmidti 
Agelas wiedenmyeri 
Amphimedon compressa X 
Amphimedon erina X 
Amphimedon rubens 
Anthosigmella varians X 
Artemisina melana 
Apysilla gracilis X 
Aplysina archeri X 
Aplysina aurea X 
Aplysina cauliformis X 
Aplysina fistularis X 
Aplysina fulva 
Aplysina lacunosa 
Callyspongia armigera 
Callyspongia longissima 
Callyspongia vaginalis X 
Calyx podatypa 
Ceratoporella nicholsoni 
Chelonaplysilla erecta X 
Chondrilla nucula 
Cinachyra alloclada 
Cinachyra sp. X 
Clathria aspera X 
Cliona aprica 
Cliona delitrix 
Cliona lampa X 
Cliona langae X 
Cribrochalina vasculum X 
Desmapsamma anchorata X 
Dictyodendrilla nux 
Diplastrella megastellata X 
Diplastrella ministrella 
X X X 
X X X X 
X X X X X 
X X X X X X 
X X X 
X X 
X X X 
X 
X 
X X X X X X 
X X X X X X 
X 
X X X X X X 
X X 
X X X X X X 
X X X X X 
X X X X X X 
X X X X X X 
X X X X X X 
X X X X X X 
X X X X X 
XX X X X X X 
X X 
X 
X 
X X X X X 
X X 
X X X X X X 
X X X X X X 
X X X X X 
X X X X X 
X X 
X X X X X X 
X X X X X 
X X X X 
X X 
X X X X 
X X 
The Caribbean coral reefs o f Panama 255 
Table 3 cont. 
Eastern Region Central Region Western Region 
1 2 3 4 5 6 7 
Discodermia dissoluta X 
Ectyoplasia ferox X X X X X X X 
Geodia neptuni X 
Haliclona hogarthi X X X X X X 
Halisarca sp. X X X X X X X 
Hyattella intestinales X 
Iotrochota birotulata X X X X X X X 
Ircinia campana X X X X X X X 
Ircinia felix X X X X X X X 
Ircinia strobilina X X X X X X X 
Ircinia variabilis X X X 
Leucandra aspera X X X X X X 
Monanchora barbadensis X X X X X X X 
Monanchora unguifera X X X X X X X 
Mycale americana X X X X 
Mycale arndti X X X X X X X 
Mycale laevis X X X X X X X 
Mycale laxissima X X X X X X X 
Neofibularia nolitangere X X 
Niphates amorpha X X X X X X X 
Niphates digitalis X 
Niphates erecta X X X X X X X 
Niphates ramosa X X X 
Oceanapia bartschi X X X X X 
Pachypellina podatypa X X 
Petrosia pellasarca X X 
Phorbas amaranthus X X X X X 
Plakortis angulospiculatus X X X X X X X 
Plakortis haHchondroides X X X X X X X 
Pseudoceratina crassa X X X X X X X 
Pseudaxinella lunaecharta X X X X X X X 
Ptilocaulis walpersi X X X X X X X 
Sigmadocia caerulea X X X 
Sigmadocia piscaderaensis X X 
Siphonodictyon coralliphagum X X 
Spheciospongia vesparium X 
Spirastrella coccinea X X X X X X X 
Spirastrella mollis X 
Tedania ignis X 
Teichaxinella burtoni X X 
Tethya crypta X 
Topsentia amorpha X X 
Ulosa ruetzteri X X X X X X X 
256 H.M. Guzmrn 
Table 3 cont. 
Eastern Region 
1 2 3 
Central Region Western Region 
4 5 6 7 
Verongula reiswigi X 
Verongula rigida X X X X X X X 
Xetospongia muta X X X X X X X 
Xetospongia proxima X X 
Xetospongia rosariensis X X X X X X X 
Total 88 54 63 73 52 47 54 56 
Sponges are an important sessile group in reefs, of which we recognize 88 species 
for the Caribbean coast (Table 3). Of these species, 93% are found in Kuna-Yala, 
followed by 72% for Bocas del Toro (63 species) and about 59% and 53% for the areas 
of Isla Grande and Bahia Las Minas, respectively. Previously, 58 species of sponges 
were reported for Kuna-Yala, including species that inhabit seagrass meadows and 
mangrove roots (Clifton et al. 1997). I agree with these authors that the diversity of 
sponges off Panama has been underestimated and that more efforts are needed to 
identify those that occur. More than 640 sponges have been reported for the wider 
Caribbean (van Soest 1984). 
Figure 4 summarizes the diversity of the three groups of sessile organisms previously 
presented and includes figures for each region along Caribbean Panama. Generally, we 
may say that the country's highest diversity is found off Kuna-Yala. 
Fig. 4. Total number of species for major sessile taxa (scleractinian corals, octocorals, sponges), in 
three regions of the Caribbean coast of Panama. 
3. NATURAL DISTURBANCES 
Panama has been affected mainly by sea warming and diseases in corals and other 
reef-related organisms (Ogden and Ogden 1994; Shulman and Robertson 1996). In the 
early 1980s, the massive mortality of Diadema antillarum, which apparently began near 
Panama, started to have a negative impact on reefs all over the Caribbean because of the 
The Caribbean coral reefs of Panama 257 
loss of an important herbivore that controlled macroalgae growth (Lessios et al. 1984a; 
Hughes 1994; Lessios 1998; Hughes and Tanner 2000). Sea urchin densities, although 
they show some recovery in Jamaica's reefs (Edmunds and Carpenter 2001), have not 
reached levels comparable to those reported off Panama before the die-off, especially in 
Kuna-Yala (Guzrnfin et al. 2003), and similar to those in other regions such as Florida 
(Chiappone et al. 2002). 
During the 1982/83 E1 Nifio, Kuna-Yala reefs were affected by the anomalous sea 
warming, with bleaching of 32% of the live coral cover that affected mainly species such 
as Agaricia spp. and Montastraea annularis, and caused a mortality of 53% in the 
Agaricia cover (Lasker et al. 1984). Reef bleaching was also observed in several regions 
of the country during 1995 and 1997/8, apparently related to the same phenomenon but 
without further implications (Clifton et al. 1997; Guzrnfin pers. obs.). The Kuna-Yala 
time-series showed remarkable changes in the coral cover during the 1980s, which 
gradually extended during the 1990s (Fig. 2). Nevertheless, it must be pointed out that 
during 1987, coral mortality was reported in the central region, especially in the Isla 
Grande-Portobelo area (Fig. 2), the cause of which was not determined. It is suspected 
that the mortality may have been related to the brief warming that took place during that 
year or to white-band disease (sensu Gladfelter 1982) as the population of Acropora 
palmata was the most seriously affected (Guzrnfin et al. 1991). 
Regarding coral diseases, the massive mortality that virtually destroyed the entire 
population of Gorgonia flabellum octocoral off Costa Rica in the mid-1980s (Guzmfin 
and Cort6s 1984) and affected the whole of Panama (J. Cubit per. com.; H.M. GuzmAn 
per. obs.) must be mentioned. From the start, it was suggested that some kind of disease 
was involved and the probable agent was subsequently found to be a fungus, 
Aspergillus (Smith et al. 1996). Likewise, since the end of the 80s, white-band and 
black-band diseases (sensu Antonius 1981) as well as other recently-described diseases 
(Richardson 1998) have been regularly observed in reefs all along the Caribbean coast, 
although never in epidemic proportions or causing generalized mortality. One report from 
1996-97 suggested a major outbreak of yellow-blotch/yellow-band disease off Panama 
at Kuna-Yala, affecting mostly Montastraeafaveolata (McCarty and Peters 2000). 
Finally, Panama is generally included in the area of the Caribbean region not 
affected by hurricanes (Cubit et al. 1989). Nevertheless, Hurricane Joan, the last 
hurricane to pass near this region in the last 100 years, had a devastating effect on the 
shallow-water sponge populations in Kuna-Yala (Wulff 1995). 
4. ANTHROPOGENIC IMPACT 
Without doubt, the reefs of the central region of Panama have been the most affected 
by man-made adventures. The impact of civilization, as it is now known, on these reefs 
goes back to the times of the Spanish conquest. The history is quite similar for most 
Caribbean countries, as coral, for many centuries, was the main raw material used in the 
construction of fortresses and settlements in the whole region. With regard to Panama, it 
is worth mentioning the construction of the City of Portobelo, located in the central 
region of the country. By the late 16 th century the first massive extraction of coral from 
nearby reefs had been carried out and continued for about 200 years (Fig. 5). In the 
Archivo General de Indias (AGI) coral mining activities are relatively well reported and 
provide great details on the periods of construction, demolitions due to redesign and the 
258 H.M. Guzmdn 
Fig. 5. Historical coral mining in Caribbean Panama. Seventeenth century Spanish fortification built 
with Montastraea, Colpophyllia and Siderastrea coral species in Portobelo, Panama (A); and detail 
of a coral wall (B). 
reconstruction of several military fortresses and public and religious buildings that 
protected and made up Portobelo (Webster 1970; Castillero 1990). Corals or "sea stones" 
were extracted inside and around Portobelo cove and in pits located 45 km (8 leagues) 
west of the city, between Bahia Las Minas and Bahia Lim6n (formerly Puerto Naos) 
The Caribbean coral reefs of Panama 259 
(AGI 1595 in Castillero 1990). By the mid-18 th century, coral was still being mined 
from these sites, which seemed "inexhaustible" (AGI 1756 in Castillero 1990). Based 
on several documents from the AGI, where the quantity of materials used for several 
constructions is described in detail, over 70,000 m e of coral was minedin a short period 
of time (Castillero 1990). The buildings and structures or ruins of Portobelo, which have 
been included in the UNESCO World Heritage since 1980, were measured on site 
(Table 4), yielding a total volume of 15,387 m 3, of which coral accounted for some 
12,923 m 3 (Jim6nez and Guzn~n in prep.). 
TABLE 4 
Estimated volume of existing Spaniard ruins constructed from the late 1500's to mid 1700's in Portobelo, 
Panama. Volume was measured in situ for each of the standing walls and coral volume was based on a 90 
• 3.6% mean coral cover obtained from transects ran across the walls (Jim6nez and Guzm~in in prep.). 
Name of Building Construction Period Building Volume Coral Volume (m 3) 
(~) 
Fort Santiago de la Gloria 1600-1760 7,420 6,554 
Town 1596-1744 191 I42 
Iglesia San Juan de Dios 1629-? 238 144 
Aduana --1630-1638 1,944 1,749 
Fort San Jer6nimo 1659-1758 1,320 1,188 
Fort San Pedro (San Carlos) 1682-1686 981 588 
Iglesia San Felipe 1600-1814 1,072 643 
Convento Las Mercedes ?-1736 51 33 
Fort San Fernando 1753-1758 2,170 1,882 
Estimated Total Volumes 15,387 12,923 
By the end of the 19 th century, the French had initiated the first excavations for the 
construction of the Panama Canal, which the Americans would continue after 1904 
(McCullough 1977). The entrance to the Canal from the Caribbean was located at Bahia 
Lim6n, where the extraction and dredging of all the coastal reefs started in order to fill 
the mangrove swamps over which military bases, airports and the City of Crist6bal were 
to be built (Comber 1916; Castillero 1962). Some records conform the extraction of 
approximately 20 million m 3 of reef coral and other material by the French Company 
from 1882 to 1885 and 33 million m s by the American Company from 1904 to 1907 
(Comber 1916; Rousseau 1916; Noriega 1986). Moreover, the American Company built 
two 3.3 km-long breakwaters at the entrance of Bahia Lim6n between 1910 and 1916. 
These were destroyed a number of times by storms and rebuilt every time using new 
material dredged from the reefs off Coco Solo island, located at the eastern edge of the 
Bay. According to the records, about 1.5 million m 3 of reef-excavated material was 
needed to build the western breakwater and about 400,000 m 3 (part of the record lost) 
for the eastern one (Rousseau 1916). Concurrent to the construction of the Panama 
Canal and especially after 191 I, numerous American military bases (e.g. Fort Sherman, 
260 H.M. Guzmim 
Fort DeLesseps and Fort Randolph) were built around the coastal area of the Canal, 
from the mouth of the Chagres River to the Isla de Largo Remo in Bahia Las Minas. 
The same pattem of filling the mangrove swamps with reef coral was always followed 
and continued through the two World Wars until 1944. During this period, new US 
Navy and Air Force military bases were build or enlarged on the islands of Margarita, 
Coco Solo, Largo Remo and Galeta that became peninsulas by means of landfill 
(Copeland 1964; Gardner and Carpenter 1965). 
In recent times, between 1958 and 1974, more than 5 million m s of reefs were dredged 
to fill 80 ha of mangrove swamps and marshes in Bahia Las Minas during the construc- 
tion of a refinery (see publicity leaflet of Refmeda Panamfi, S.A.). From the day the 
refinery started operating (1961-62) and continuing to this day, the area has been 
consistently polluted by hydrocarbons (Guzmfin and Jarvis 1996). Several oil spills have 
also affected and destroyed coral reefs in this area, particularly in 1968 and 1986 
(Jackson et al. 1989; Guzmfin et al. 1991, 1994). To this, we must add the generalized 
pollution of coral reefs with different metals from multiple activities that take place both 
near to and at a distance from the coastal zone. These include port, industrial and fam3ing 
activities (Guzmfin and Jim6nez 1992), which are responsible for the emission of high 
levels of extremely toxic elements such as mercury (Guzrr~n and Garcia 2002). 
Equally important is the destruction of reefs due to the traditional coral mining and 
landfilling practices of the Kuna people in Kuna-Yala. The Kuna modify and expand 
their populated islands from the shores to the edges of the shallow reef flats, marking 
the perimeter with coral walls for landfill (Fig. 6a-b). This activity started 150 years ago 
when the Kuna living in the forests between Colombia and Panama decided to settle 
along the archipelago on the coastal zone (Tice 1995; Ventocilla et al. 1995; Howe 
1998). The activity was first acknowledged in the early 1970s and the impact on reef 
habitats was brought to the attention of the authorities (Porter and Porter 1973). 
Recently, the impact of coral mining and landfilling was quantitatively assessed by 
means of direct measurement of the coral volume in the walls as well as the changes in 
the island surfaces, comparing photographic records from the 1960s with those of 2001 
(Guzmfin et al. 2003). While the authors assessed only 50% of the inhabited islands, 20 
km of walls with a coral volume estimated at 16,000 m 3 were measured in addition to an 
increased surface area of 6.23 ha due to coral landfilling. The growing tourist industry 
has further encouraged the Kuna to extract corals and other reef-related organisms for 
their sale as souvenirs (Fig. 6c). 
It is important to mention the impact of sedimentation on the reefs off the central and 
western regions of the country, a process that has resulted from indiscriminate defores- 
tation along the coastal zone and fiver basins. This has contributed in part to the gener- 
alized degradation of the landscape observed throughout the country for decades. This 
problem has worsened during the last decade in the Isla Grande-Portobelo sector and in 
the last 5 years in Bocas del Toro (GuzmAn and Guevara 1999, 2001), both associated 
with tourist development and the mismanagement of forests in the river basins which 
has increased the runoff into the Caribbean. It has been observed, moreover, that a 
substantial portion of the vegetation and tree trunks produced during the process of 
deforestation is carried by the rivers down to the sea, where, with unexpected fre- 
quency, physically destroy the reef shallow habitats (Kilar and Norris 1988). 
The Caribbean coral reefs of Panama 261 
Fig. 6. Traditional coral mining in Kuna-Yala, Panama. Coral Cay enlarged by coral landfilling (A); coral 
wall at the edge of island (B); and souvenirs made of small coral colonies (notice fresh coral tissue), 
conchs, etc. to be sold by Kuna people to tourists (C). Photos by M. Guerra (A), and C.E. Jim6nez (C). 
262 H.M. Guzm6n 
Finally, unregulated exploitation of reef-related fishing resources is decimating the 
natural populations and, consequently, is drastically reducing the income of coastal zone 
populations. Lobster exploitation has already resulted in a situation of overfishing in 
Kuna-Yala (Castillo and Lessios 2001) and in Bocas del Toro (GuzroAn and Guevara 
unpubl, data). Moreover, among other non-traditional resources, three species of sea 
cucumber have been briefly exploited in Caribbean Panama, reaching overfishing levels 
in at least two of the species, Isostichopus badionotus and Astichopus multifidus 
(Guzrnfin and Guevara 2002). It has been suggested that the existing low densities may 
not allow their reproduction and recovery in the mid-term (Guzmfin et al. 2003). 
5. PROTECTION, LEGISLATION AND MANAGEMENT 
5.1 Protected Areas 
Panama has four protected areas with reefs on the Caribbean side and under different 
management categories. The creation of a new protected area along the old Fort Sherman, 
from the mouth of the Panama Canal to the Chagres River, is at present under revision. 
Most areas have a rather weak legal definition as they were established by Directive 
Board Resolutions (Resoluciones de Junta Directiva; R.J.D.) or Ministry Decrees 
(Decretos de Gabinete) and not by national laws (see Navarro 1998). Moreover, at 
presentthere is a management plan only for Parque Nacional Portobelo, though plans 
are apparently being prepared for other areas and are expected by the year 2003. The 
protected areas are distributed throughout the three Caribbean reef regions previously 
defmed (Fig. 1), namely: 
(1) In the eastern region (Kuna-Yala), the Area Silvestre de Nargana with circa 
99,000 ha managed by the Kuna people and established in 1994 (R.J.D. 22-94). The 
extent of the maritime-coastal zone under protection is not clear, but it is indeed within 
the "corregimiento" and includes the best reefs in the country. Most reefs close to the 
mainland and near settlements have been undeniably altered by traditional coral mining 
(see Fig. 6 and Guzrnfin et al. 2003). These authors have suggested eight priority 
conservation areas and the need for the creation of a network of protected areas to serve 
as a marine corridor within the indigenous reservation, a proposal that is now under 
consideration by the local authorities. 
(2) In the central region, there are two existing protected areas and one being created. 
The Parque Nacional Portobelo, with 13,226 ha, was created in 1976 (Decree 91) and 
includes the reef areas from Portobelo to Isla Grande in a strip bounded by about 70 km 
of coast and extending 2,000 m seaward. Most of these reefs are affected by 
sedimentation and runoff associated with coastal tourist development. The second area, 
Area Silvestre Protegida of Isla Galeta established in 1997 (Act 21), includes a coastal 
area of no more than 25 km and reefs in an advanced state of deterioration, directly as a 
result of the 1986 oil spill (Guzmfin et al. 1991). The third (possibly) protected area of 
San Lorenzo includes underdeveloped, low-diversity reefs (GuzmAn and Hoist 1994). 
This area is in fact designated for the conservation of one of the few pristine forest areas 
in the country's central region, a move that would indirectly benefit the reefs. 
(3) In the western region, the Parque Nacional Marino Isla Bastimento was estab- 
lished in 1988 (R.J.D. 22-88) with 13,226 ha, of which 11,596 ha includes protected 
marine environments. Recently, it was reported that the highest diversity or reef- 
associated organisms and best reef development occur outside the protected area 
The Caribbean coral reefs of Panama 263 
(Guzrrfin and Guevara 1998b, 1999). This park is under enormous pressure due to the 
high level of sedimentation caused by deforestation on the neighboring islands and 
increasing tourism. At present, a management plan is being elaborated for this park. 
5.2. Legislation and Management 
In the previous sections, we have identified the present conservation status of many 
coral reefs on the Caribbean coast of Panama. We can confidently state that the diversity 
of scleractinian corals and other sessile organisms found here is representative of the 
wider Caribbean. On the one hand, we have observed that the percentage of live coral 
cover indicates that the reefs' health is not excellent. They have been gradually affected 
over the past 30 years by natural or anthropogenic disturbances. In some cases, the agent 
causing this deterioration has not been easy to identify, but in others, as in Bahia Las 
Minas where coral mortality above 80% has been reported, the problem can be clearly 
attributed to human disturbances (GuzmAn et al. 1991). On the other hand, it is 
important to point out that in most reefs where changes in coral cover, composition and 
diversity are monitored at present, a small recovery has been observed. Even though this 
improvement is lower than 10% and is not particularly significant, it indicates that the 
system has self-restorative capabilities. 
Both processes, the deterioration and the potential recovery, force us to recognize 
the inexorable need of exemplary legislation and better scientific management of the 
resources for the conservation of the country's reefs. The first reef conservation and 
environmental policy efforts started in 1977 when Panama became a signatory of 
CITES (Convention on International Trade in Endangered Species of Wild Fauna and 
Flora). Legislation developed fin'ther during the early 1990s when two resolutions 
(R.J.D.) prohibited the extraction of corals (INRENARE R.J.D. N ~ 033-93, September 
28, 1993) and reef fishes (Executive Decree N ~ 29, June 24, 1994) throughout the 
country. Nevertheless, because these two initiatives are mere "resolutions" and not 
national laws, in practice they have been documents that have never fulfilled their goal. 
The extraction of scleractinian corals, octocorals, anemones, sponges and reef fishes 
continues indiscriminately in several sectors of the country, and even worse, within 
protected areas. Moreover, Panama has signed Act 2 from 1995, which approves the 
Convention on Biological Diversity, and Act 9 of the same year, which regulates the 
Protection of Priority Protected Areas of Central America. More importantly for Panama, 
the Ley General del Ambiente N ~ 41 (Environment General Law) was established in 
1998, which, up to now (October 2002) has not been clearly regulated, creating thus a 
legal vacuum that does not allow the adequate protection and management of coral reefs 
and their related resources. 
The following three examples elaborate the above statements and highlight the lack 
of legal attention bestowed upon reefs. At the same time, they reveal how the available 
scientific information has not been used appropriately in the implementation of 
protection measures for these ecosystems. Firstly, we have pointed out before that the 
best reef development and the largest reef extension in Panama is in Kuna-Yala under 
the autonomous political regime of the Kuna people. Since the early 1970s, there has 
been scientific information that, although not quantitative, has reported the traditional 
practices of coral mining and landfiUing, rather than the building of new communities on 
mainland, in order to keep pace with population growth (Porter and Porter 1973). Both 
researchers made repeated calls to the Kuna authorities and the central government, 
264 H.M. Guzmfn 
suggesting that coral extraction be controlled, and that the reef be sustainably managed, 
for the benefit of the Kuna people. Both parties have ignored the warnings and 
consequently reef destruction has reached unprecedented levels. What is worse, there is 
now an increase in the extraction of materials from other shallow habitats such as seagrass 
meadows and beaches (Guzmfin et al. 2003). Until 2001 reef deterioration as a result of 
this practice was monitored and was found to be limited to the western region or Punta 
San Bias (Shulman and Robertson 1996) while very little was known about the general 
state of reefs in the remaining area. However, the state of the reefs and some aspects of 
their diversity throughout the entire indigenous reservation are now well-known (Guzrnfin 
et al. 2003). Secondly, in the early 1990s the government gave permission to Spanish 
architects to extract coral from the Portobelo area reefs to be used as raw material in the 
restoration of some buildings erected by the Spaniards 500 years ago (see Fig. 5). Their 
intention, unconceivable to scientists now, was mainly to restore the Customs House 
(Aduana) using the same material with which it was originally built, which was coral. 
Some of the Portobelo National Park reefs had been monitored since 1985 (Guzmfin et al. 
1991) and in 1992 it was reported that live coral cover within this area was lower than 
10% (Fig. 2). Both the authorities and the Spanish technicians that in 1992 elaborated the 
first management plan for the protected area were fully aware of the existing scientific 
information. Years later, in 1997, which was declared International Year of the Reef, live 
coral extraction continued in order to complete the restoration work and, ironically, the 
local media reported that with this work "technical assistance wasoffered to train more 
than 30 Portobelo residents in restoration work with coral" (Santamaria-Loo 1997). 
Thirdly and f'mally, studies carried out in Bocas del Toro in the late 1990s identified 
sensitive and priority areas for conservation throughout the archipelago (Fig. 7), 
suggesting that the currently protected marine area did not include some of the 
important resources that needed to be preserved (GuzroAn and Guevara 1998b, 2000). 
These areas were selected on the basis of the presence of the greatest diversity 
("hotspots") of corals and other sessile organisms within the archipelago (sensu Briggs 
2002) and the existence of important populations of species considered to be 
endangered, such as Acropora palmata and A. cervicornis (Precht et al. 2002). In spite 
of public disclosure of the results and direct lobbying before the authorities as well as 
national and international non-governmental organizations since 1998, these areas 
remain unprotected to date, although we acknowledge that efforts are being made to 
change the current park limits. Time is important as a gradual deterioration of the reefs 
has been observed within these areas over the past three years (see histograms in Fig. 7). 
6. CONCLUSION AND RECOMMENDATIONS 
The examples above refer to the best reef areas of Panama, which, without doubt, are 
the ones that should be protected. In short, analyzing the future prospects for the country's 
Caribbean reefs and rating them according to their current risk level due to environmental 
pressures (sensu Bryant et al. 1998), we can say that more than 80% of the coral reefs are 
at risk (Fig. 8). Indeed, 50% of the areas have a high-risk level. When we analyze in detail 
these levels for each Province, we observe that in Col6n 100% of the reefs are high-risk, 
followed by 75% and 50% for Bocas del Toro and Kuna-Yala, respectively (Table 5). 
These risk levels go hand in hand with the increase in the coastal population estimated at 
between 60% and 290% for the last 40 years (see Table 5; Stmglia and Winter 2002). 
The Caribbean coral reefs of Panama 265 
Fig. 7. Priority conservation area of Bocas del Toro, western Caribbean Panama, and changes in live 
coral cover from 1999 to 2002 at two reef-monitoring sites within the critical areas. Area containing 
the highest diversity of coral, octocorals and sponges or "hot spot" within the archipelago (light 
gray), and areas of relatively abundant endangered coral species Acropora spp. (dark gray). Notice 
the polygon that limits the existing marine protected area of PNMIB (modified from Guzm/m and 
Guevara 1999). 
TABLE 5 
Estimated reef area, percentage of reefs at risk, and demographic growth between 1960 and 2000, for 
geographical provinces along the Caribbean coast of Panama (see Fig. 1). Percent reef areas at risk 
(medium-high) were estimated combining four threat factors: coastal development, marine pollution, 
overexploitation, and inland pollution and erosion (Bryant et al. 1998). Population data include only those 
"Corregimientos" (smallest administrative political units) close to the coastal zone (CGR 2001). 
Province Reef Area High Risk Population Population Change 
(km 2) (%) (1960) (2000) (%) 
Kuna-Yala 610 50 19,509 31,619 62.1 
Col6n 48 100 76,845 123,198 60.3 
Veraguas 8.2 25 616 2,340 289.9 
Bocas del Toro 87.4 75 27,238 103,307 279.3 
266 H.M. Guzm6n 
.~ 6O 
t ~ ~ 
n, 50 
O0 
~ 40 
~ ao n,, 
= 20 
r 
L _ 
�9 10 13. 
0 m 
Low Medium High 
Threat Category 
Fig. 8. Percentage of reefs at risk along the Caribbean coast of Panama, according to threat categories 
(sensu Bryant et al. 1998). 
One of the most important risk factors is the development of the coastal zone and the 
resultant overpopulation (Bryant et al. 1998). Tourism may be considered as the most 
important activity because 48.5% of the 1,398 tourist attractions included in the Tourist 
Development Master Plan (Plan Maestro de Desarrollo Turistico) of Panama (IPAT 
/OEA 1993) are in the coastal zone (JICA/IPAT 1995). Prospectively, 90.3%, 59%, 
45.7% and 40.4% of these coastal tourist attractions, in comparison with non-coastal 
ones, are located in the Provinces of Kuna-Yala, Col6n, Veraguas and Bocas del Toro, 
respectively. Doubtlessly, reefs in these areas are an important tourist attraction but the 
reefs also fulfill a very important role in the productivity and functioning of a very 
complex coastal ecosystem. It is obvious that the maintenance of these high levels of 
tourist attraction will depend exclusively on the integral conservation of these coastal 
areas and their reefs and some kind of control on demographic growth (see Stmglia and 
Winter 2002). 
There is nothing new about all of the above. However, I would like to close this 
chapter with some personal comments on aspects that may be of help in the 
conservation of our reefs. It is well-established that in recent years coral reefs have 
become very popular on the environmental agenda of some governments and, especially 
non-governmental organizations (NGOs). On the one hand, I acknowledge the impor- 
tant work carried out by some of these NGOs in the diffusion of the magnitude of reef 
deterioration and status worldwide and of the long-term effect this degradation will 
have on the coastal zone in general if this trend is not reversed. On the other hand, it is 
to be noted that after all these multi-million dollar campaigns, very little change has 
been observed. Deterioration continues unchallenged in most countries, especially in the 
Neotropics. With a few localized exceptions that do not represent the spatial scale of the 
problem, it was possible to create protected areas that work, instead of the so-called 
"paper parks", and the problem has been halted. Considering the above and after partici- 
pating personally in several planning workshops (technical and communal) for reef 
sustainable conservation, I can state with no doubt that something is amiss. Although 
scientists have taken the position of helping in the conservation process and are 
constantly supplying the information requested by the NGOs and resource managers, no 
The Caribbean coral reefs of Panama 267 
real solutions to the local and regional problems are emerging from this process. Part of 
the answer may lie in the lack of empowerment and organization of the civil society, but 
it is to be noted that the excessive financial resources available for conservation have 
been invested almost exclusively in planning workshops. We seem to have totally 
forgotten that, for the implementation and functioning of these plans, a very important 
ingredient is needed: the will of the political class. In short, we have common immedi- 
ate conservation goals, but they must be better focused on the professional informing 
and lobbying of legislators and government officers who are the only ones that have the 
power to decide the future and management of resources in our countries. 
ACKNOWLEDGMENTS 
I wish to thank the government of the Republic of Panama, the National 
Environmental Authority (ANAM), the Panama Maritime Authority (AMP) and the 
Congreso General Kuna for giving me collection permits and access to the reefs 
throughout the country for many years. I thank Carlos A. Guevara for all the support 
given to the acquisition of field information; without his help, it would have been 
impossible to describe the reefs of Panama. Thanks to I. Herngndez, for preparing the 
database, K. Kaufman for processing the data and L. Gonzhlez for photograph prepara- 
tion. I wish to thank the Editor, J. Cort6s, for his constant support and patience. The 
information contained in this chapter has been obtained with the financial support of the 
Smithsonian Tropical Research Institute and the Smithsonian Institution Environmental 
Sciences Program, as well as with the partial support of The Nature Conservancy, 
Fundaci6n Natura de Panam,5, Fundaci6n PROMAR de Panamfi and AEK-PEMASKY. 
This chapter is dedicatedto Irene, Adriana and Carolina, pillars of my existence. 
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