Orchids of Paradise

Exploring the lower Talamanca Seashores of Costa Rica, the

“Coast of Plenty” of Columbus
by Franco Pupulin and diego Bogarín

846 Orchids november 2018   www.AOS.org

WHERE IT ALL BEGAN The Italian explor-
er and navigator Cristoforo Colombo
(Christopher Columbus, c. 1451–1506)
discovered the paradise during his fourth
voyage across the Atlantic Ocean, when
searching for a passage west to the Indian
Ocean and the Orient. He had sailed
from Cádiz (Spain) in May 1502, and
after landing on the island of Martinique
(Martinica), Santo Domingo and Jamaica,
he eventually reached Central America,
anchoring off the coast of Honduras at
the end of July. Two weeks later his ships
landed on the continental mainland, and
from here they began exploring the coast
to the south, arriving in Almirante Bay
in Panama on October 16, 1502. During
this cabotage navigation, on September
18, 1502 Colombo anchored off Quiribrí 2
Island (later known as Isla Uvita) in front
of what is now Limón, Costa Rica. Even
though he apparently never actually
came ashore, the natural beauty of the
coast, its exuberant vegetation and the
abundance of wildlife (and maybe the
gold ornaments worn by local dignitaries),
were material enough to convince him to
baptize the newly discovered land as Costa
Rica, the Coast of Plenty. On the shores
of the island, one can still find plants of
Brassavola nodosa, much as Columbus’s
crew could have seen them five centuries
ago.
Interestingly, the paradisiac Caribbean 3
coasts south of Limón had to wait a long
time to be really settled. During the Limón, is not mentioned before 1852, and
colonial period, the region remained it was not until 1867, when the plans of
sparsely populated, mostly because of the the construction of a railroad connecting
frequent raids by the Miskitos Zambos from the highlands to the sea began taking
Nicaragua and Honduras, who attacked shape, that Limón was chosen as the
Spanish-held territories and independent site of a major port to export the coffee
indigenous groups in the area, capturing produced in the Central Valley. The town
people to be traded under slavery to was officially founded in 1871.
British merchants. Spanish attempts to AFRO-CARIBBEAN COSTA RICA Since 4
reduce the region of the Cabécar and the beginning of 19th century, however, [1] The Caribbean coast of Costa Rica; inset:
Bribri tribes during the 16th and 17th fishermen of Afro-Caribbean origin, mostly
A specimen of Brassavola nodosa pho-
century never resulted in more than brief from Panama and the West Indies, began
tographed at Westfalia, a few kilometers
settlements or conquests. The “town” of to settle in the Caribbean coast of Costa
Badajoz (actual Sixaola) was founded by Rica. They stayed in temporary camps south of Isla Uvita, on the Caribbean
Hernán Sánchez de Badajoz in 1539 and during fishing seasons to plant coconuts, shores of Costa Rica. Photograph by
disappeared the next year; Santiago de cassava and yams. Starting in 1828, some Franco Pupulin.
Talamanca, founded in 1605 close to the of these temporary settlements became [2] The First Landing of Christopher Colum-
mouth of Río Sixaola, was destroyed in permanent; fishermen lived there with bus in America”, painted in 1862 by the
1610; San Bartolomé de Duqueiba, on the their families. Spanish Dióscoro Teófilo Puebla Tolín.
shores of the Telire river, was founded in The railway and the port at Limón, [3] Cristoforo Colombo’s name is remem-
1659 , and burned in 1662. The Caribbean on the Atlantic Coast, brought to Costa bered in the name of Costa Rican
coast was never effectively settled or Rica workers from Italy, China and the currency, the Colón (Spanish transla-
developed by the Spanish. On a map of Caribbean. On December 20, 1872, the
tion of Colombo). Here the navigator is
the isthmus prepared in 1774, the only first boat from Jamaica arrived at the port
portrayed on a coin of 1897.
settlement recorded on the Caribbean of Limón with 123 workers to work on the
coast of Costa Rica is Mohin (later Moín), railroad. The next year, Jamaican workers [4] The Uvita Island, Columbus’s first landing
north of the actual port of Limón. The in the port already numbered 1,000. With point in Costa Rica, off the coast from
same “capital” of the Caribbean province, the financial crisis of the railways in 1890, Limón. Photo by FotoAereasCR.

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 franco pupulin
5

many Jamaicans did not return home, but

remained in the province of Limón on the
Caribbean coast of Costa Rica, working
in agriculture and later for the banana
industry. They did not speak Spanish and
retained Jamaican customs. They had
their own schools with teachers brought
from Jamaica, proud of being subjects of
the British crown. Caribbean Costa Rica
defined itself, in ethnical, cultural, and
linguistic terms, as an enclave of Afro-
Caribbean descendants, speaking a creole
language mostly derived from English,
but with influences from the Hispanic,
indigenous, Chinese and African cultures
(which academics define as Limonian-
creole and Mekatelyu). To a large extent, 6
this human and cultural landscape
continues today. The economy of the large [5] A small house close to the sea, among
banana monocrop industry, started in the coconut palms, in the vicinity of Cahuita.
Caribbean plains in the 1870s, and more Local houses were mostly built on stilts to
recently the activities connected to a reduce humidity in the living spaces and
subsistence tourism trade, have disrupted to avoid the frequent floods of the rivers.
the physical and cultural isolation of the [6] Antique map of Costa Rica and Nica-
Afro-descendant community of the
ragua from 1764 by Jacques Nicolas
Caribbean coast, but the seashore and
Bellin (1703–1772). No settlements are
the low mountain landscapes, with their
bright greens and light blues, are still recorded on the Caribbean coast of
reminiscent of the paradise that Colombo Costa Rica.
saw more than five centuries ago. [7] Painted by the Spanish Pedro Berru-
A DIVERSITY HOTSPOT The Carib- guete (1450–1540), this is probably the
bean slopes of the Cordillera de Talamanca, only portrait of Christopher Columbus
which descends steeply to the sea through for which he served as an actual model,
a narrow strip of lowland, are considered being still alive. Both the famous portraits
one of the richest hotspots in Costa Rican of Columbus by Sebastiano del Piombo
biodiversity (Elizondo et al. 1989, Zamora (1519) and Ridolfo del Ghirlandaio (1520)
2008, Kohlmann et al. 2010, Kohlmann
were painted posthumously.
2011). In botanical terms, however, it 7

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is one of the less explored regions of
the country. The original vegetation of
the coastal plains has been practically
extirpated to make room for banana
plantations, and the exuberant foothill
regions up to the intermediate elevations
of the Cordillera of Talamanca, for sure
among the richest in orchids on the entire
planet, are virtually inaccessible because
of the absence of roads and paths. The
historical impediment in access to these
regions is still reflected in the reduced
system of roads, most of which were built
only during the second half of the last
century. The Caribbean Talamanca region
is likely the last real, and hard-to-break
frontier to complete the inventory of the
Costa Rican orchid flora. Accessing, naming
and cataloguing the orchid diversity of this
region is also one of the more relevant
commitments of the scientists working
with the Lankester Botanical Garden at
the University of Costa Rica. In particular,
our interest was focused on surveying
the resilience of the orchid flora of the
Caribbean coastal region and to compare
it, in terms of diversity and composition,
with that found on the Pacific coast,
where a study was done 20 years ago. The
inventory of the orchid flora in a national 8
park of about 988 acres (400 ha) and its
immediate surroundings in the humid light yellow brain, elkhorn and blue [8]Map of Costa Rica, with a detail (inbox) of
regions of the central Pacific revealed staghorn corals, gorgonians and sea fans, Cahuita National Park. The pale orange
the existence of 55 species belonging to as well as more than 500 species of fish, area on the general map indicates the
32 genera (Pupulin 1998a, 1998b, 2001). are protected. It is also home to the only biographic region known as “lower Tala-
Was it true that we could find a richer example of coastal vegetation that has manca”.
orchid flora in a comparable region along remained undisturbed for more than 50
the Caribbean coast? Does the Caribbean years. Established in 1970, the national list of 85 orchid species in 48 genera
orchid flora include the same genera as park covers a marine area of 55,200 acres given by Estrada Chavarría (2014a) for
that of the Pacific? And, within each given (22,300 ha), with a 600-acre (242-ha) the lower Talamanca region, in fact, not
genus, are the species the same on both coralline reef, and 2,732 acres (1,106 ha) only includes plants from the foothills of
coasts? Is there a correspondence in the of land area, including lowland tropical the Matama and Talamanca chains, but
relative frequency of the species between wet forest, marshlands, mangroves and also from the Chimú and Muchilla peaks
the two regions? In 2012 we started the sand vegetation habitats. Around the (at 2,516 feet [767 m] and 2,887 feet [880
work to answer these questions. park, across the main road that connects m], respectively), which are obviously
CAHUITA NATIONAL PARK The coast- the town of Limón with the Panamanian outside the biogeographic circumscription
al plains between the Caribbean Sea border, there are a series of low hills of the region and far beyond the scope of
and the Talamanca range were severely known as Fila Carbón, not exceeding our research. In particular, it is probable
deforested during the first decades of the 656 feet (200 m) in elevation, that were that several of the recorded species in
20th century, at the times of the greatest promising spots to be also explored for Pleurothallis sensu latissimo, as well as
expansion of the banana plantations orchids. species of Camaridium (Maxillaria) and
under the control of the United Fruit The Baja Talamanca (lower Talamanca) Lycaste, do not form part of the regions’s
Company. The narrow strip of woods now region, varying in elevation from sea level orchid flora, and others were likely
protected by the Cahuita National Park, to about 1,300 feet (400 m), is one of the misidentified. In this sense, an orchid
and the mangroves of the Manzanillo 25 floristic regions proposed by Zamora flora of Cahuita National Park and its
Wildlife refuge a few miles (kilometers) and collaborators (2004) to interpret the surroundings (including the mangroves
south, are likely the only partial remains rich flora of Costa Rica. The tree flora, as of the Gandoca-Manzanillo Wildlife
of the original coastal vegetation. For well as the water plants and the bushes Refuge a few miles (kilometers) south
this reason, we decided to begin our of the national park, have been intensely along the coast) would constitute a close
inventory of the orchid flora of southern investigated by Sánchez Vindas (1983, approximation to the knowledge of the
Caribbean Costa Rica here. The Cahuita 2001), but the orchid composition of real orchid diversity of the whole “lower
National Park is probably best known for the native vegetation has not been dealt Talamanca,” extending from south of Río
its coral reefs close to the shore, where with in detail up to now. The excellent Matina to the Río Sixaola along the border

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with Panama.
Walking along the beach within the
Park, or along the trail that flanks the
beach a few yards (meters) inside the
forest, is an unforgettable experience,
something really close to what one can
imagine as a paradise on earth. A few
yards (meters) more inland from the
coast, an impenetrable net of plants and
roots emerging from the salt waters of
large swamps make the area practically
inaccessible. This flooded forest near the
coast is locally known as yolillal, from the
vernacular name of the Raphia taedigera,
or yolillo, a palm that almost exclusively
dominates the vegetation. In other parts,
the yolillal is substituted by mangroves,
where Avicennia germinans, Conocarpus
erecta, Laguncularia racemose and
Rhizophora mangle, are the most common
species (Estrada Chavarría 2014b).
Further inland there are partially flooded
areas with a forest mostly dominated by
the cativo, Prioria copaifera (Fabaceae),
a tree up to 100–130 feet (30–40 m)
tall, often very rich in epiphytic flora,
as well as “sangrillo woods,” another
kind of seasonally flooded forest almost
exclusively composed by Pterocarpus
officinalis (sangrillo) (Cifuentes &
Masterson 1991, Estrada Chavarría
2014b). This tree gave origin to the Cahuita
name, from the indigenous miskita words
kawi (sangrillo) and ta, point, or sangrillo-
point. Fortunately, in 2016 rangers opened
an elevated walkway that runs inside the
flooded vegetation crossing the tidal
marsh to solid ground, allowing a closer
inspection of the large trees that emerge
from the water.
HOW MANY ORCHIDS ARE
THERE? Within the Park we established
133 (146) plots of approximately 1,076
square feet (100 sq m), where orchid
specimens were individually identified
and counted to assess their relative
frequency. Of these, 114 observation
plots were made in the gallery forest that
covers the main trail within the Park, some km) around the National Park. In only 19 Table 1. The list includes species recorded
33–39 feet (10–12 m) from the shore. The of the 170 plots of Cahuita National Park within the limits of the Park, as well as those
shrub vegetation growing on the beach and surroundings (=11%) were no orchids recorded from outside the Park, including the
was sampled with 25 plots, and another observed. coastal vegetation of Gandoca-Manzanillo
seven plots were established in the After six years of visits and explorations Wildlife Refuge (GMWR) and the hills of Fila
flooded area of mangroves. We also made of the park and its surroundings, as well Carbón, close to the Park. Species of which
24 plots in the immediate vicinities of the as searches in the national herbaria that we have not observed natural populations
Park (not more than 984 feet [300 m] host plants from lower Talamanca, we
within the park are marked with an asterisk.
from the northern and western borders), have now recorded 51 orchid species in
to survey orchids that could probably 33 genera, with 16 new taxa to be added situ after collecting them in the Park. A
be found within the park but cannot be to the general list by Estrada Chavarría few of the collected specimens have still
observed in the protected area due to the (2014a). Most of the species were refused to flower, so their identification
inaccessibility of its inward limit. Finally, fully documented with photographs, at the specific level is only tentative.
we searched on the low hills of the Fila botanical illustrations and Lankester Although some of the taxa were obviously
Carbón, in a radius of about 3.1 miles (5 Digital Composite Plates, often done in expected from the region (such as species

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of Brassavola, Catasetum, Dimerandra,
Epidendrum, etc., or the ubiquitous
Ionopsis utricularioides and Rodriguezia
compacta, others represented, for us,
quite a surprise. Most remarkable were
the finding of Psychopsis krameriana,
a severely endangered species in Costa
Rica, growing in a dense wood at no
more than 656 feet (200 m) from the
seashore, and that of Kefersteinia alata
(new for the flora of Costa Rica) close to
a lateral access to the Park. Also, it was
extraordinary finding a large specimen of
Coryanthes horichiana, happily growing
on a tree planted on the sidewalk outside
a renowned restaurant in the village of
Puerto Viejo, a few miles (kilometers)
south of the Park. A few months after we
collected one of the small specimens of
franco pupulin

franco pupulin
Crths. horichiana, the tree was cut and
the small population of Coryanthes was
lost. Noteworthy was the finding of very 9 10
large populations of Vanilla planifolia,
which we documented both in the dense
woods in the inner parts of the Park
and along the shore, where the species
forms populations of literally a hundred
individuals in the white sands a few
yards (meters) from the sea. Among the
vanillas, we also found large specimens
of Vanilla cf. mexicana, a new record for
the flora of Costa Rica (voucher: Bogarín
9586). Also new for the Costa Rican flora
was Kefersteinia alata (voucher: Pupulin
8642), a species previously known only
from the Caribbean watershed of the
Chiriquí mountains in Panama.
The most frequent species within the
franco pupulin

Park was Catasetum maculatum, of which

184 individual plants were observed in 71
of the 170 plots, with a frequency of 42% 11
of the observed sites. The second most
frequent species at Cahuita was Vanilla
planifolia, recorded in 55 plots (=32%),
with 131 individuals. These two orchid
species may be considered fairly common
within the study area. Frequent species
include Epidendrum angustisegmentum,
recorded in 15 plots (=9%), and
Brassavola cf. grandiflora in 9 plots (5%).
Species observed in less than 3% of the
plots were Caularthron bilamellatum,
Ionopsis utricularioides (both in four
franco pupulin

plots), Cohniella (now Trichocentrum)

ascendens, and Nidema boothii (three
plots). Although they were recorded in
fewer plots, Inps. utricularioides and Trt. 12
ascendens form very large populations. [9] Punta Cahuita as seen from the hills close to the coast.
Epidendrum nocturnum, Mormolyca [10] The large Raphia palms dominating the yolillal.
now Maxillaria) hedwigae, Polystachya [11] Two researchers of the Lankester staff, Melania Fernéndez (left) and Christina Marie
masayensis, Rodriguezia compacta and
Smith (bright) checking a tree along the beach for epiphytic orchids.
Vanilla cf. mexicana were recorded in
[12] The brackish waters of the marsh inside the National Park.
two plots each (ca. 1%). All the remaining

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 7DEOH&RPSDULVRQEHWZHHQWKHRUFKLGÁRUDVRI/RZHU7DODPDQFDDQG&HQWUDO3DFLÀFLQ&RVWD5LFD
Lower Talamanca (Caribbean) &HQWUDO3DFLÀF
Acianthera
A. ellipsophylla (Bogarín 9573)
A. hondurensis (Karremans 5865) *
Anathallis
A. lewisae (Pupulin 360)
Aspasia
A. epidendroides (Pupulin 311)
Brassia
B. caudata (Bogarín 11243)
Brassavola
B. cf.JUDQGLÁRUD (Pupulin 8275) B. nodosa (Pupulin 331)
Bulbophyllum
B. oerstedii (Pupulin 453)
Camaridium
C. neglectum (Pupulin 363)
Campylocentrum
C. fasciola (Smith 422)
C. micranthum (Pupulin 332)
C. panamense (Pupulin 305)
C. tyrridion (Pupulin et al. 1029) *
Catasetum
C. maculatum (Pupulin 8330) C. maculatum (Pupulin 327)
Caularthron
C. bilamellatum (Bogarín 9572) C. bilamellatum (Pupulin 355)
Cohniella
C. ascendens (Pupulin 8278) C. ascendens (Pupulin 322)
Coeliopsis
C. hyacynthosma (Pupulin 1153) *
Coryanthes
C. horichiana (Bogarín 11231)
Dimerandra
D. emarginata (Bogarín 9930) D. emarginata (Pupulin 325)
Encyclia
E. alata* (herbarium record)
Epidendrum
E. anceps (herbarium record)
E. angustisegmentum (Bogarin 9574)
E. congestoides (Castelranco s.n.) *
E. congestum (Pupulin 296)
E. eburneum (Bogarín 11254)
(ÁH[XRVXP(Bogarin 11232)
E. fulfordianum (Bogarín 11228)
E. isomerum (Bogarin 11237) E. isomerum (Pupulin 1658) *
E. montis-narae (Pupulin et al. 1140) *
E. nocturnum (Bogarin 9933) E. nocturnum (Pupulin 335)
E. cf. schomburgkii (Pupulin 8638) *
E. sculptum (Pupulin 373)
E. stamfordianum (Pupulin 370)
E. strobiliferum (Karremans 5864) E. strobiliferum (Castelranco s.n.) *
E. vulgoamparoanum (Pupulin 328)
Epidendrum sp.* (herbarium record)
Epistephium
E. ellipticum* (herbarium record)
Galeottia
*JUDQGLÁRUD(Bogarín 11246)
Gongora
Gongora sp. (Bogarín 11234)
Gongora sp. (Bogarín 11229)
Gongora sp. (Castelranco s.n.) *
Heterotaxis
H. crassifolia (Pupulin 312)
Ionopsis
I. satyrioides (not vouchered) I. satyrioides (Pupulin 309)
I. utricularioides (Pupulin 8337)
Kefersteinia
K. alata (Pupulin 8642)
K. costaricensis (Bogarín 11252)
K. orbicularis (Pupulin et al. 1170)
species marked with an asterisk were collected outside the physical boundaries of the protected areas.
852 orchids november 2018 www.AOS.Org
Lower Talamanca (Caribbean) &HQWUDO3DFLÀF
Leochilus
L. labiatus (Bogarin 10414)
L. scriptus (Pupulin 374)
Lockhartia
L. micrantha (Pupulin 313)
Macradenia
M. brassavolae (Pupulin 8632)
Maxillaria
M. uncata (Pupulin 8650)
Maxillariella
M. oreocharis (Pupulin s.n.)
M. cf. poneranta (Pupulin 345)
Mormolyca
M. hedwigae (Bogarín 9934) M. hedwigae (Pupulin 1840)
Nidema
N. boothii (Bogarín 9584)
N. ottonis (Pupulin 318)
Notylia
N. cf. lankesteri (Pupulin 8332)
N. pittieri (Pupulin 328)
Notylia sp. (Karremans 5863)
Oncidium
O. polycladium (Pupulin 292)
O. stenobulbon (Pupulin 303)
O. stenotis (expected)
Palmorchis
Palmorchis sp. (Pupulin 8643)
Polystachya
P. foliosa (Bogarin 11250) P. foliosa (Pupulin 468)
P. masayensis (Pupulin 8290)
Prosthechea
P. abbreviata (Mena s.n.)
P. chacaoensis (Pupulin 1034)
3WDUGLÁRUD (Pupulin 1431) *
Psychopsis
P. krameriana* (Bogarín 10418)
Psygmorchis
P. pusilla (Fernandez 988)
Rhetinantha
R. aciantha (Bogarín 11711)
R. scorpioidea (Merli s.n.) *
Rodriguezia
R. compacta (Pupulin 8296)
Sacoila
S. lanceolata (Pupulin 356)
Scaphyglottis
S. bidentata (Pupulin 2194) *
S. boliviensis (Pupulin 1854) *
S. micrantha (Pupulin 377)
6PLQXWLÁRUD(Pupulin 8627)
S. prolifera (Pupulin 471)
S. stellata (Pupulin 300)
Sobralia
S. fenzliana (Pupulin 320)
S. macrophylla (Castelranco s.n.) *
Stanhopea
S. cirrhata (Pupulin 1872) *
S. ecornuta (Bogarín 11258)
Stelis
Stelis sp. (Bogarín 11265)
Stelis sp. (Pupulin 8640)
Trigonidium
T. egertonianum (Bogarin 11259) T. egertonianum (Pupulin 368)
Trizeuxis
T. falcata (Pupulin 473)
Vanilla
V. aff. PH[LFDQD(Bogarín 9586)
V. planifolia (Pupulin 8274) V. planifolia (Pupulin 1429) *
Xylobium
X. foveatum (Pupulin 1900) *
 13

species were observed only once, and

they are fairly rare elements in the
orchid flora of Cahuita, even when they
are quite common elsewhere (i.e.,
Dimerandra emarginata, Polystachya
foliosa, Trigonidium (now Maxillaria)
egertonianum.
THESE ARE TWO DIFFERENT COUNT-
RIES! As Costa Rica is roughly the size
of Denmark, or West Virginia, one might
suppose that the orchid floras of the two
coasts of the country, that are about 125
miles (150 km) apart, would be grossly
the same, especially considering that
the southern Caribbean and the central
Pacific coasts, leaving out the driest
areas of northern Pacific Costa Rica, are
both wet regions. Our data show that
franco pupulin

franco pupulin

this easy supposition is basically wrong

and that the total composition of the
orchid floras along the coastal lowlands
is substantially different on the two sides
of the continental divide, underscoring
the importance of geology and tectonic
events in orchid species distribution, even
on the small geographic scale of Costa
Rica.
Although the National Parks of Cahuita
(Caribbean) and Manuel Antonio (central
Pacific) share some, less than 50%, of the
orchid genera, the coincidence at the level
of species is only about 10%. Many of the
14
genera that we recorded in our studies are
exclusive to only one of the two regions.
So, Acianthera, Brassia, Coryanthes,
Encyclia, Epistephium, Galeottia,
Macradenia, Maxillaria, Palmorchis,
Psychopsis, Oncidium (as Psygmorchis),
Rodriguezia and Stelis were only recorded
from the lowlands of the lower Talamanca
region; on the contrary, species of
Anathallis, Aspasia, Bulbophyllum,
15
[13] The elevated walkway at Cahuita Na-
tional Park. Courtesy of the MINAE.
[14] Epiphytic flora in the cativo forest at
Cahuita National Park.
[15] Christina M. Smith collecting a speci-
men of Epidendrum angustisegmentum
at Cahuita National Park.

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Maxillaria (as Camaridium, Heterotaxis
and Maxillariella), Coeliopsis, Lockhartia,
Prosthechea, Sacoila, Sobralia, Trizeuxis
and Xylobium were found along the
central Pacific coast but not the southern
Caribbean. Even in those genera that were
documented from both the coastal regions
(namely Brassavola, Campylocentrum,
Catasetum, Caularthron, Dimerandra,
Epidendrum, Gongora, Ionopsis, Leochilus,
Kefersteinia, Maxillaria (as Mormolyca
and Trigonidium), Nidema, Notylia,
Oncidium, Polystachya, Rhetinantha,
Scaphyglottis, Stanhopea, Trichocentrum
and Vanilla), they mostly differ in
composition at the species level. Actually,
the two opposite coastal floras only share
16 17
nine species in the genera Catasetum
(with Ctsm. maculatum), Caularthron
(Cau. bilamellatum), Dimerandra (Dim.
emarginata), Epidendrum (Epi. isomerum,
Epi. nocturnum, Epi. strobiliferum),
Ionopsis (Inps. satyrioides), Maxillaria
(Max. hedwigae, Max. egertoniana),
Polystachya (Pol. foliosa), Trichocentrum
(Trt. ascendens), and Vanilla (Vl.
planifolia).
Of particular interest is that the two
orchid contingents of the Caribbean
and the Pacific are often composed of
unique “sister” species, as in the cases
of Brassavola grandiflora/nodosa,
Campylocentrrum fasciola/tyrridion,
Kefersteinia costaricensis/orbicularis,
Leochilus labiatus/scriptus, Nidema
franco pupulin

boothii/ottonis, Notylia lankesteri/pittieri,
Rhetinantha aciantha/scorpioidea
and Stanhopea ecornuta/cirrhata,
respectively.
This difference in genera and species
composition decreases progressively
when the altitudinal gradient is raised
toward higher elevations, and above
1,640–1,968 feet (500–600 m) the
overlap increases considerably. In the
Caribbean foothills, at elevations close
to 3,280 feet (1,000 m), many other
orchid groups appear that we have not
recorded in the vegetation near the
coast, among which species of Anathallis,
Lockhartia, Maxillaria (as Camaridium
and Maxillariella), Prosthechea, Sobralia
and Xylobium, among others. On the
other side, species of Encylia, Galeottia,
Maxillaria, Psygmorchis, and several
groups of Pleurothallids, which were
not recorded around Manuel Antonio
National Park, become fairly frequent in
the region of the Cerro Nara in the central
Pacific coast, which exceeds 3,280 feet
(1,000 m) in height.
REFERENCES
Cifuentes, M., and D. Masterson, editors. 1991. Parque
18
20
[16] Even though it has not been observed
within the limits of the Park, Ionopsis
utricularioides forms large populations
on the trees and shrubs just outside its
borders.
[17] Rodrigueiza compacta, the only species
recorded in the flora of Costa Rica, is
franco pupulin
19
common on the trees in the village of
Cahuita. We have not recorded it within
the Park.
[18] Psychopsis krameriana flowering in a
cativo forest a few hundred meters from
the sea. The flower is indicated with an
arrow. Only the lynx’s eye of Diego Boga-
rín could spot it among the dense foliage
of the forest. Photo by F. Pupulin. Inset:
Psychopsis krameriana (Bogarín 10418).
The flower was photographed against a
blue background within the field vehicle
of the Lankester Botanical Garden. Photo
by F. Pupulin.
[19] Kefersteinia alata, until now known only
from the type locality in western Panama.
Photo by F. Pupulin.
[20] Coryanthes horichiana. A small portion
of the original plant was grown at Lank-
ester Botanical until flowering. Meantime,
the host tree at Puerto Viejo was cut and
the large specimen died.

854 Orchids november 2018   www.AOS.org

 Nacional Cahuita: Plan Emergente. Catie-Spn-Mire-
nem, Costa Rica. 91 pp.
Elizondo, L.H., Q. Jiménez, R.M. Alfaro, and R. Cháves.
1989. Contribución a la conservación de Costa Rica. 1.
Áreas de endemismo. 2. Vegetación Natural. Fundación
Neotrópica, San José, Costa Rica.
Estrada Chavarría, A. 2014a. Listado de Plantas Vasculares
de la Región de Baja Talamanca y los Cerros Chimú y
Muchilla, Limón, Costa Rica. p. 129–199. In: J. Sánchez
González, editor. Documentación de recursos biológicos
en la región de Baja Talamanca, Limón, Costa Rica.
Museo Nacional de Costa Rica, Departamento de His-
toria Natural, San José, Costa Rica.
Estrada Chavarría, A. 2014b. Diversidad de Plantas Vascu-
lares en la región Baja Talamanca, Limón, Costa Rica.
p. 15. In: J. Sánchez González, editors. Documentación
de recursos biológicos en la región de Baja Talamanca,
Limón, Costa Rica. Museo Nacional de Costa Rica, De-
partamento de Historia Natural, San José, Costa Rica.
Kohlmann, B. 2011. Biodiversity Conservation in Costa franco pupulin
Rica — An Animal and Plant Biodiversity Atlas. p.
203–22. In: I. Pavlinov, editor. Research in Biodiversity
— Models and Application. InTech, Rijeka, Croatia.
Kohlmann, B., D. Roderus, O. Elle, A. Solís, X. Soto,
and R. Russo. 2010. Biodiversity Conservation in 21
Costa Rica: A correspondence Analysis Between
Identified Biodiversity Hotspots (Araceae, Arecaceae,
Bromeliaceae, and Scarabaeinae) and Conservation
Priority Life Zones. Revista Mexicana de Biodiversidad
81:511–559.
Pupulin, F. 1998a. Orchid florula of Parque Nacional
Manuel Antonio, Quepos, Costa Rica. Journal of Tropi-
cal Biology 46(4):961–1031.
Pupulin, F. 1998b. Orchids of Manuel Antonio National
Park. MesoAmerican Press, San Jose, Costa Rica.
Pupulin, F. 2001. Addenda Orchidaceis Quepoanis. Lank-
esteriana 1(1):1–28.
Sánchez Vindas, P. 1983. Florula del Parque Nacional
Cahuita. EUNED, Editorial Universidad Estatal a Dis-
tancia, San José, Costa Rica.
Sánchez Vindas, P. 2001. Flórula arborescente del Parque
Nacional Cahuita. EUNED, San José, Costa Rica.
Zamora, N. 2008. Unidades Fitogeográficas para la
Clasificación de Ecosistemas Terrestres en Costa Rica.
Recursos Naturales y Ambiente 54:14–20.
Zamora, N., B.E. Hammel, and M.H. Grayum. 2004.
Vegetación. p. 91–216. In: B.E. Hammel, M.H. Grayum,
C. Herrera, and N. Zamora, editors. Manual de Plantas
de Costa Rica, Vol I. Monogr. Syst. Bot. Missouri Bot.
Gard. 97.
Acknowledgments
We thank the Ministry of the
Environment (MINAE) and its National
System of Protected Areas (SINAC) for
granting unrestricted access to Cahuita
National Park during our study. The officers
at the Park were greatly collaborative,
and among them we want to mention at
least Mirna Cortés and Mario Cerdas for
their special help. Gratitude is extended
to the Vice-Presidency of Research,
University of Costa Rica, that financed
our research from 2012 to 2016 through
the project 814-B2-159, La flora de
orquídeas del Parque Nacional Cahuita,
Limón. Thanks are due to our present
and former colleagues, Noelia Belfort,
Melissa Díaz, Melania Fernández, Adam
Karremans, Darha Solano, and Christina
22
Smith, for their friendly company during
the collections at Cahuita, and for their [21] Researchers scanning floral detail of Cahuita orchids in the hotel room.
invaluable help in documenting and [22] Lankester Digital Composite Plate of Maxillaria uncata, based on Pupulin 8650, Cahuita
naming the orchid flora of the Park. National Park.
— Franco Pupulin is a Senior Research

www.AOS.org   november 2018 Orchids 855

 Orchids of the Cahuita National Park
(A) Acianthera ellipsophylla; (B) Brassia caudata; (C) Brassavola gran-
diflora; (D) Campylocentrum fasciola; (E–F) Catasetum maculatum,
male and female flowers; (G) Cohniella ascendens; (H) Coryanthes
horichiana; (I) Dimerandra emarginata. Photographs by F. Pupulin,
except (H) by D. Bogarín.
(A) Gongora species (Bogarín 0000); (B) Gongora species (Bogarín
0000); (C) Ionopsis satyrioides; (D) Ionopsis utricularioides; (E) Kefer-
steinia alata; (F) Leochilus labiatus; (G) Macradenia brassavolae; (H)
Maxillaria uncata; (I) Mormolyca hedwigae. Photographs by F. Pupulin
(G not from Cahuita).
856 Orchids november 2018   www.AOS.org
(A) Encyclia alata; (B) Epidendrum angustisegmentum; (C) Epi-
dendrum eburneum; (D) Epidendrum flexuosum; (E) Epidendrum
fulfordianum; (F) Epidendrum isomerum; (G) Epidendrum nocturnum;
(H) Epidendrum strobiliferum; (I) Galeottia grandiflora. Photographs by
F. Pupulin.
(A) Nidema boothii; (B) Notylia lankesteri; (C) Polystachya foliosa; (D)
Polystachya masayensis; (E) Psychopsis krameriana; (F) Psygmor-
chis pusilla; (G) Rhetinantha aciantha; (H) Rodriguezia compacta; (I)
Scaphyglottis minutiflora. Photographs by F. Pupulin.
Professor at the University of Costa
Rica, where he works as the Head of
Research at the Lankester Botanical
Garden. He is a Research Associate with
the Harvard University Herbaria and The
Marie Selby Botanical Gardens. Franco
has mostly worked on the floristics of
Costa Rica and monographed several
groups of Neotropical orchids, mainly in
the Oncidiinae, Pluerothallidinae, and
Zygopetalinae. His current projects focus
on Flora Costaricensis and the systematics
of Neotropical orchids, as well as on
biogeography and the evolution of floral
traits in the Orchidaceae. Diego Bogarín is
a researcher at Lankester Botanical Garden
and a Research Associate at the herbarium
UCH of the Universidad Autónoma de
Chiriquí, Panama. Since 2015, he has been
a Ph.D. candidate at Leiden University,
The Netherlands. Diego is interested
in the biology, ecology, evolution and
systematics of the Orchidaceae. Currently,
his main research projects are focused
on bioinformatics, genomics, taxonomy
and in situ conservation through the 23
development of orchid floristic inventories
in national parks.

25

[23] Orchids of the Cahuita National Park;

(A) Stanhopea ecornuta; (B) Stelis spe-
cies (Pupulin 8640); (C) Stelis species
(Pupulin 8640); (D) Trigonidium egertoni-
anum; (E) Vanilla cf. mexicana; (F) Vanilla
planifolia. Photographs by F. Pupulin.
[24] A large plant of Vanilla cf. mexicana
fruiting within the Park.
[25] Vanilla planifolia has been recorded
flowering both in the dense forest inside
the coast, as well as along the beach, ex-
posed to the full sun on the white sands.
The species shows broad phenotypic
variation in Cahuita National Park.
24

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