Tree diversity and biogeography
of four one-hectare plots in the
lowland rainforest of the Piedras
Blancas National Park
(“Regenwald der Österreicher”),
Costa Rica
Dissertation
zur Erlangung des akademischen Grades
Doktor der Naturwissenschaften
an der
Fakultät für Naturwissenschaften und Mathematik
der Universität Wien
eingereicht von
Mag. Werner Huber
Wien, Jänner 2005
Preface
It all began with a conversation with Dr. Michael Kiehn and Prof. Dr. Anton Weber
(of the University of Vienna’s Institute of Botany) about the possibility of writing theses in
Costa Rica in the “Regenwald der Österreicher” (‘Rainforest of the Austrians’). That was on
the 14th of December 1992, and just six weeks later, Anton Weissenhofer and I travelled
out to Costa Rica. We stayed with Ronald Moya Diaz and his family in their farmhouse near
La Gamba village and abutting the wonderful Esquinas forest. Indeed, it was Ronald who
first took us into the forest, leading us with a gun! “The forest is dangerous;” he said,
“beware of all the venomous snakes”.
After a few days’ exploration in this wonderful, “dangerous” forest, we tried to find a
undisturbed research plot somewhere near the farmhouse, and we did find an excellent
piece of primary forest with huge trees, lianas and natural gaps, all just forty minutes walk
from Ronald’s finca. We decided that if we came back we would make an inventory of all
the trees in this, our inland slope plot. But would we ever get back there? We went into the
Esquinas forest every day during our two weeks with Ronald’s family; we got lost, had
close contact with poisonous snakes, were molested by insects and were, like our clothes,
constantly wet. Because of the high humidity, our clothes and shoes never dried out. Anton
and I considered this fact as we thought about our futures as tropical botanists.
Back in Vienna, all these reservations were put aside by the idea of working
towards our theses in the “Regenwald der Österreicher”, and of living in Costa Rica for six
months. Prof. Dr. Wilfried Morawetz supervised our diploma theses. Prof. Michael
Schnitzler, the father of the entire “Regenwald der Österreicher” project, bought a
corrugated iron house with loamy soar. That house would be our home for the next six
months, and was the beginning of the “Tropenstation La Gamba”.
In 1996 we began our dissertations: to learn more about the Esquinas forest and its
enormous biodiversity. For a few months we worked in the gorge plot behind the Esquinas
Rainforest Lodge. We later started the ridge plot on the Fila trail, sometimes with an
assistant; for the coastal plot we used a total of six people in three teams, so that it took
three weeks rather than the six months needed for the inland slope. The information gained
led to the production of the “Field Guide to the Flowering Plants of the Golfo Dulce
Rainforest”. With the help of Prof. Dr. Anton Weber, Prof. Dr. Roland Albert, our
supervisors and Prof. Michael Schnitzler, we had the chance to realise the concept of an
Austrian field station in the “Regenwald der Österreicher”.
The Tropenstation La Gamba now comprises four buildings with over 20 beds, and six staff
from the nearby La Gamba village work there. Since 1993, many scientists and students
have used the facility to try to understand how the forest works. It has also been used by
both students and tourists who come to enjoy the station and the rainforest.
Vienna, January 2005
Werner Huber
I would like to thank….
my partner Elisabeth Lindner – for her never ending support which helped me to realize
this work
my friend and colleague Anton Weissenhofer – for his cooperation since 1993 in
establishing the project "Tropenstation La Gamba" and in our scientific work
my supervisor and “chief” Prof. Anton Weber – for giving me the freedom to work on my
thesis, and for always supporting the project "Tropenstation La Gamba"
my supervisor Prof. Roland Albert – for always supporting the project “Tropenstation La
Gamba"
Prof. Michael Schnitzler – for initializing the La Gamba project and the “Rainforest of the
Austrians”
Ing. Javier Guevara (MINAE) – for helping us to aquire permits to work in the Piedras
Blancas National Park
my colleagues – in particular Manfred Dworak, Susanne Gockner, Heidrun
Hochwallner, Dietmar Moser, Susanne Pamperl, Susanne Sontag and Nelson
Zamora (INBio)
Gundi & Gerhard Pfeiffer - for their very fascinating talks and ideas to the area species
curves
Alice Luck & Graham Tebb – for the language revision of the thesis
Esther Greter – for arranging for us to work in the coastal plot near the Golfo Dulce Lodge
Trude & Ron McAllister (Casa Orquideas) and José Antonio Bogantes (MINAE –
Chacarita) – for allowing us to use their data for a better understanding of the
climate in the Piedras Blancas National Park
the staff at the Herbario del Museo Nacional de Costa Rica and
the Costa Rican staff the Tropenstation La Gamba (Oliver Chacon, Justo Castellon
Arauz, Eduardo Pitingo Arauz Sanchez, Agustin Zuniga)
Karin Guttenbrunner, Monika Heimel and Florian Rudolf
Contents
Table of Contents
1. INTRODUCTION.................................................................................................................. 1
1.1. Geography of the Golfo Dulce Region...........................................................................................3
1.2. Geological History of the Golfo Dulce Region ..............................................................................5
1.3. Soils in the Golfo Dulce Region......................................................................................................9
1.4. Climate of the Esquinas Rainforest .............................................................................................15
1.5. Flora of the Golfo Dulce Region ..................................................................................................20
2. MATERIALS AND METHODS .......................................................................................... 25
2.1. Study Area .....................................................................................................................................25
2.2. Description of the plots .................................................................................................................26
2.3. Field Work and Plant Identification ...........................................................................................28
2.4. Diversity Indices ............................................................................................................................29
2.5. Floral Similarity and Difference of all plots (Cluster-Analysis) ...............................................32
2.6. Distribution Pattern and Geographical Affinities......................................................................33
3. RESULTS .......................................................................................................................... 34
3.1. Floristics and Diversity .................................................................................................................34
3.2. Biogeogeographical Patterns and Affinities................................................................................72
3.3. Identification of the Tree Families of the Esquinas Forest........................................................89
4. DISCUSSION................................................................................................................... 119
4.1. Floristics and Diversity ...............................................................................................................119
4.2. Distribution Pattern and Geographical Affinities....................................................................140
5. ABSTRACT ..................................................................................................................... 152
6. CURRICULUM VITAE ..................................................................................................... 157
7. REFERENCES................................................................................................................. 158
8. LISTS ............................................................................................................................... 170
1. Introduction
1. Introduction
Tropical rainforests are complex ecosystems which cover less than 10% of the world’s land
surface, and yet contain considerably more than half the world’s living species (WILSON
1988). This is even more surprising in the case of plants, for at least two thirds of the
estimated 250,000 cormophyte species grow in the tropics (KUBITZKI 1985, WHITMORE
1990). Neotropical rainforests are considered the most species-rich forests to exist
worldwide.
GLEASON & COOK (1926) pointed out that a tropical forest might contain hundreds of
species of trees, none of which can be found on an adjoining hectare. Indeed, the most
diverse forests in terms of trees are tropical lowland rainforests (CAMPELL & al. 1986,
FABER-LANGENDOEN & GENTRY 1991, GENTRY 1988a, 1988b, 1993, GRUBB & al. 1963,
GRUBB & WHITMORE 1966, HARTSHORN 1983, LIEBERMANN & al. 1996, MADSEN & ØLLGAARD
1994, RICHARDS 1952, VALENCIA & al 1994, WATTENBERG & BRECKLE 1995). However,
there are also tropical forests which are dominated by just a few tree species (e.g.
Pentaclethra macroloba in La Selva, Costa Rica) (LIEBERMANN & LIEBERMANN 1987).
Tropical rainforests are distinguished from all other terrestrial ecosystems by a very high
level of diversity on many levels (species, habitats, life-forms, etc.).
The floristic information gathered in this study was used to evaluate biogeographical
patterns.
The diversity of tropical rainforests increases as the dry season becomes shorter (ASHTON
1989) and the amout of precipitation increases up to 4,000 mm each year (GENTRY 1988b).
The high diversity of vascular plants can be seen on a large scale (e.g. in Costa Rica there
are about 9,361 plant species) (HAMMEL & al. 2004) and also on a small scale (e.g. 233
different cormophytes grow in 100 m2 of a lowland wet forest) (WHITMORE & al. 1985). The
rainforests of the Golfo Dulce region in southeast Costa Rica belong to the most interesting
and species-rich forests in Central America. So far, 2,709 species in 935 genera of 187
families of vascular plants have been recorded in the rainforests around the Golfo Dulce
(HUBER 1996, HERRERA-MCBRYDE & al. 1997, QUESADA & al. 1997, VAUGHAN 1981, WEBER
& al. 2001, WEISSENHOFER 1996), among them about 700 tree species (QUESADA & al.
1997). Preliminary studies give an idea of the extraordinary diversity of the Esquinas forest
(HUBER 1996, WEISSENHOFER 1996). One aim of this thesis is to increase knowledge about
diversity and similarity of species composition in four different habitats in the Esquinas
forest, and another is to gain some idea as to the reasons for this high diversity.
Botanical exploration in the neotropics and in Costa Rica has increased during the past 20
years and there has been a substantial increase in the available data on neotropical forest
composition and structure. Nevertheless, when considered in the context of the size and
1
1. Introduction
biological diversity of the region, detailed knowledge of the flora of the neotropics is far
from complete and many new species of angiosperms continue to be discovered and
described each year (GROOMBRIDGE 1992, PRANCE & al. 2000). However, it has reached a
level at which we can now draw significant conclusions about diversity and
phytogeographical patterns in the region.
The angiosperm floras of Central America today show most similarity to those of South
America. In reviewing this similarity RAVEN & AXELROD (1975) suggest that, even since the
origin of angiosperms sometime during the Cretaceous, Africa and South America, and
perhaps even some of the fragments of Central America yet unconsolidated, have
constituted a site of evolutionary importance. During the Tertiary there were islands, where
today the strip of land joins the Neotropics of South America and the Hol(Ne)arctic of North
America. Sweepstakes dispersal from South America during the Palaeocene, Oligocene
and Miocene has always been more pronounced for plants than for animals.
The “land bridge”, which has existed for c. 4.7 Mio. years (CARRANZA-CASTANEDA & MILLER
2003), is party the reason for Costa Rica’s status as a “corridor” for flora and fauna to travel
from North to South America and vice versa (BURGER 1980, GOMEZ 1986, RICH & RICH
1983). During the Ice Age the climate in the tropics was cooler and dryer, but also
remaining relatively wet on different sites (COLINVAUX & al, 1996, KERR 1996).
An important contribution is being made to Neotropical phytogeographic knowledge by the
websites of the herbaria of the Missouri and New York Botanical Gardens. Since the
commencement of the work of the Instituto Nacional de Biodiversidad (INBio) in Costa Rica
(1988) and the publishing of the Manual de Plantas de Costa Rican (HAMMEL & al. 2004),
Costa Rica has become one of the best-explored regions in the Neotropics.
This study describes the diversity and geographical patterns of plants (diameter in breast
height - d.b.h. ≥ 10 cm) of four one-hectare forest inventory plots sampled in the different
habitats (gorge, ridge, inland slope and coastal slope) in the Esquinas forest in Costa Rica.
Floristic inventories are basic requirements for many ecological studies of processes at the
study sites. The floristic information gained was used to evaluate biogeographical patterns.
The destruction of tropical habitats such as forests around the Golfo Dulce (e.g. Los Mogos
and La Gamba) continues at an alarming rate, especially since we have the not yet
completed the inventory of what exists. It is therefore necessary to continue to invest
considerable resources into fieldwork, since it is especially through plot research that we
can gain information about the flora, diversity and structure of the forests.
2
1.1. Geography of the Golfo Dulce Region
1.1. Geography of the Golfo Dulce Region
Adapted version of WEISSENHOFER & HUBER (2001)
The Peninsula de Osa and the Esquinas forest are situated in the Puntarenas Province in
southern Costa Rica near southwestern Panama. The geographical coordinates lie mainly
between 8°27'-8°41'N and 83°15'-83°45'W. The main sector of the Corcovado National
Park covers 424 km² and the Esquinas forest (Piedras Blancas National Park) covers 148
km² (Fig. 1.1). The altitude ranges from sea level up to 745 m on the Osa peninsula (Cerro
Rincón and Cerro Mueller in the Fila Matajambre) and up to 579 m in the Esquinas forest
(Cerro Nicuesa). Between the two parks the Golfo Dulce Forest Reserve (592 km²) has
been established, thereby forming a natural forest corridor. The region’s extremely wet
natural systems evolved in partial isolation from the drier parts of the Pacific Coast further
north.
The whole region is still tectonically active. Up to ten tremors per day have been measured
in the region, and crustal elevations have been observed.
Within the Corcovado National Park lies the drainage of the Corcovado Basin, a broad
sediment-filled oceanic embayment between Punta Llorona and Punta Salsipuedes which
extends inland from the Pacific Ocean 2-10 km eastward. The basin's low plain covers
about 100 km² with several meandering rivers, partially rimmed by upland hills which
increase in altitude and irregular relief from an undulating plateau (below 200 m) in the
northwest part of the park (north of Llorona), to 745 m in the southeast on the peninsula's
highest peaks. The rugged uplands produced by intensive tectonic activity and weathering
(causing frequent landslides) are dominated by eroded narrow ridges and long steep
slopes with dense drainage networks (TOSI 1975; HERWITZ 1981; HARTSHORN 1983,
HERRERA-MCBRYDE & al. 1997). A virtually uninterrupted sandy beach extends for 20 km,
with cliffs and pocket beaches at the northern and southern park headlands; there is a
marine cave near the southern point of the beach.
The Piedras Blancas Park consists mainly of narrow ridges and steep slopes covered with
primary forest. The Río Esquinas, named after its conspicuous meanders (in Spanish
esquinas means corner), forms the natural border to the north and the west sides of the
park. Several quebradas (streams) and small rivers pass through the land and flow into the
Río Esquinas. Floodplains within the park along the two main rivers, the Río Esquinas and
the Río Bonito, cover abandoned farmland and secondary forest at different stages of
regrowth. Due to logging, almost no flat land with primary forest remains within the park,
except small pockets along the coast and deep inside the park. The steep and rocky
1
1.1. Geography of the Golfo Dulce Region
southern border, formed by the shoreline of the Golfo Dulce, is every so often interrupted
by sand and gravel beaches, which give way to small plains. Near the mouth of the Río
Esquinas there are extensive mangrove swamps. Some small coral reefs northwest of the
Esquinas forest also belong to the park.
Fig. 1.1. Geographical location of Costa Rica and the Golfo Dulce region with Corcovado National Park
(Parque Nacional Corcovado) and the Piedras Blancas National Park (Parque Nacional Piedras
Blancas = Esquinas forest) (from WEBER & al. 2001)
4
1.2. Geological History of the Golfo Dulce Region
1.2. Geological History of the Golfo Dulce Region
Adapted version of MALZER (2001)
1.2.1. General
Costa Rica occupies a central position on the geologically young land bridge between the
two American continents. This land bridge spans a gap of some 1600 km, from the
Motagua-Polochic fault in southern Guatemala to its anchor point on the northwest tip of
South America. The Costa Rican section covers about 200 km of this distance.
Central America is the contact area of four (possibly five) megatectonic units (crustal
plates): (1) the North American plate (2) the Caribbean plate in the north of the Central
American land bridge (3) the Cocos plate and (4) the Nazca plate in the south. (1) and (2)
are separated from the others by the deep “Middle American Trench”, along which the
Cocos plate is being subducted under the North American and Caribbean plates.
1.2.2. The Golfo Dulce Region
The geographical limits of the Golfo Dulce area are the Fila Costena in the northeast, the
Río Sierpe in the northwest, the Pacific Ocean to the west and south, and the Río Colorado
to the east. The main components include the deeply eroded and incised hills (highest
point is 579 m) of the Fila Golfito and its extension towards the Osa peninsula, the Golfo
Dulce restricted basin, and the Osa peninsula (highest elevation is 782 m).
Geologically, the Golfo Dulce region belongs to the outer belt of pre-Tertiary to Paleogene
oceanic basaltic volcanics of the Chorotega block.
The NW-trending Longitudinal Fault Zone (Ballena-Celmira Fracture Zone) separates the
Golfo Dulce area from the Fila Costena and faults of the same strike, and their NNE-SSW
oriented secondaries dominate the structure and morphology of the region.
The land portion of the Golfo Dulce area is divided into a Northern zone, where the
Cretaceous-Paleogene basement is strongly weathered but not covered by younger
sediments (except in the valleys), and a Southern zone, consisting of the larger part of the
Osa peninsula, where older rocks are covered by Pliocene and Pleistocene sediments.
1.2.3. The Basement
DI MARCO & al. (1995) divide the Golfo Dulce area into three “terranes” or tectonostratigraphic units of different origin which are separated by steep faults:
5
1.2. Geological History of the Golfo Dulce Region
(1) The Golfito Terrane occupies the land between the Ballena-Celmira fault zone in the
north, the northeast shore of the Golfo Dulce and a NNW-striking fault crossing from Playa
Gallardo on the Golfo Dulce to the Esquinas River near Piedras Blancas. The terrane is
formed by oceanic basalts at the base, followed by pelagic (deep sea) carbonates and
volcaniclastics. A volcaniclastic complex without lavaflows occupies the top position. The
entire sequence has a Late Cretaceous to Paleocene age.
(2) A mass of submarine basalts with only minor inclusions of deep sea sediment makes up
the Rincón block which borders the Golfito Terrane along the NNW-fault mentioned above
and covers the northwest part of the Esquinas forest, the hills on the north shore of the
Golfo Dulce to beyond the Río Sierpe and the inner part of the Osa Peninsula. A Late
Cretaceous to Eocene age of this mass is indicated.
(3) The Osa-Cano Accretionary Complex forms the outer, Pacific part of the Osa
Peninsula. Its lithology is fundamentally different in that it consists of typical slope
sediments, turbidites and breccias, including large blocks of basalt and ophiolite having a
wide age-range from late Cretaceous to Miocene.
The Northern zone, which includes the Esquinas rainforest, belongs to a larger part of the
Rincón block, but also contains the fault-boundary with the Golfito Terrane which forms the
eastern portion.
The lack of sediments younger than Eocene in this area indicates that the rocks may have
emerged and been exposed to subaerial erosion for millions of years. It is thus not
surprising to find rock exposures only along the coast and near the bottom of mostly narrow
quebradas (streams). Only wider valleys of the Río Esquinas, the Río Bonito and the lower
part of Río Sorpresa have a fill of Holocene and possibly Pleistocene sediments. The
remaining topography, up to the highest elevations, is covered by a weathering layer of
variable thickness, generally less than 2 m on the slopes and more on the ridges.
The following are the main lithologies and their distributions:
On the north side of the Fila Golfito, southeast and northwest of the village of La Gamba,
sediment sequences of the highest layer of the Golfito Terrane are exposed in the
Quebradas Bolsa, Chorro, Achiote (SE) and Sardinal (NW). Volcaniclastics (tuffs,
siltstones, sandstones, breccias with often very large components) and thin-bedded
(pelagic to hemipelagic) limestones, mudstones and radiolarite (coloured red, sometimes
forming jasper) are seen in the profiles. The La Gamba Field Station and the nearby
Esquinas Rainforest Lodge are both located on this unit. Light green tuff crops out on the
slope north of the station, whereas fine-grained, medium-grey basalt and boulders of basalt
breccia are found in the Quebrada La Gamba to the south (PAMPERL, pers. comm.). Further
south, massive layers of grey-green, fine-grained basalt and dykes and sills of dolerite
appear in the profiles (Quebrada Bolsa, Río Sorpresa). Large amounts of basalt, often
6
1.2. Geological History of the Golfo Dulce Region
developed as pillow lava, as well as dolerite are found exposed along the coast of the Golfo
Dulce between Playa Cacao and Punta Gallardo.
Equally, all outcrops from Punta Gallardo northwest along the coast to the Esquinas and
then west into Bahia Rincón are formed by basaltic lavas, here belonging to the Rincón
block. Volcaniclastics are conspicuously absent in this unit; minor inclusions of limestone
and radiolarite are found. Inland, Quebradas Sardinal and Machaca reach into the Rincón
block. Its uniformly more erosion-resistant rocks form the highest elevations (over 550 m)
of the Fila Golfito.
The Rincón unit appears to rim the Golfo Dulce on three sides and to occupy the
northeastern third of the Osa Peninsula. Outcrops are poor, however, to the east of the Río
Rincón. In this area Pleistocene-Holocene covers the coastal plain, underlain by sediments
of the Pliocene Osa Group which also form the low hills behind the coast.
The Southern zone, consisting of the central and southern part of the Osa Peninsula, is
composed of a ‘basement’ formed by the Osa-Cano Accretionary Complex and the
unconformably overlying clastics of the Pliocene Osa Group. The accretionary complex
(following DI MARCO & al. 1995 l.c.) represents a melange consisting of a volcaniclastic
greywacke to mudstone matrix in which various types of components from centimetre to
very large (hundreds of metres) size are embedded. Oceanic basalt and volcaniclastics,
deep-sea limestone and chert, but also shallow water carbonates are found reworked and
re-deposited in this overall deep water sediment. The combination of component types and
the age range determined for either components or matrix are characteristic of each of the
three subunits of the complex. Good exposures are found along the coast of the northwest
and southeast corners of the peninsula and in riverbeds in the interior. The generally poor
outcrop conditions and post-depositional deformation of the rocks may be responsible for
the earlier attribution of the entire ‘basement’ of the peninsula to the Nicoya Ophiolite
Complex.
1.2.4. The Golfo Dulce
The Golfo Dulce is a fault-bounded, marine inlet whose shape is controlled by the NWtrending on-land extensions of the Panama Fracture Zone and their NE-striking
secondaries. The northeast boundary fault is clearly expressed as a steep wall with
echosoundings (HEBBELN et al. 1996). The southwest boundary is partially hidden under
young sediments. The southeast entrance to the gulf is formed by a shallow (60 m) sill,
which is not covered by young sediments. From there the gulf deepens rapidly to 185 m
and to 210 m close to the northwest coast. The gulf measures approximately 55 x 12 km.
The sill at the entrance impedes water circulation. As only occasional flushings occur, the
7
1.2. Geological History of the Golfo Dulce Region
Golfo Dulce has developed into a restricted and slightly anoxic marine basin. Bottom
sediments consist mainly of turbidites, which enter the gulf via permanent channels.
Sediment thickness is unknown. The northeast coast between Río Esquinas and Golfito is
flanked by a number of small reefs.
1.2.5. The Geological History of the Golfo Dulce Region
Two phases of development can be distinguished: the older pre-Cocos Ridge history, and a
younger phase characterized by the arrival, impact and shallow subduction of the Cocos
Ridge. It is likely that in the Mid-Miocene, before the arrival of the Cocos Ridge on the Nmoving, subducting Pacific Plate, the area was separated from the central volcanic arc by
the fore arc basin (Terraba trough). The components of the present-day Golfo Dulce
Region, the island complex of the Rincón block, the Golfito Terrane and the slope-trench
sequence of the Osa Accretionary Complex were not yet in their final places. The impact
(ca. 6 Mio.) of the Cocos Ridge then resulted in the shortening — by folding and thrusting
— and uplift of the fore arc basin and the welding-together of the central island arc, the now
inverted fore arc basin, and the oceanic blocks. A bundle of NW-trending dextral strike slip
faults developed, possibly as a lateral escape response to the impact of the Cocos Ridge.
These seem to have linked up with the Panama Fracture Zone. These faults determine the
latest tectonic history of the area. Subduction of the Cocos Ridge from about the MidPliocene may have partially released the compressive force of the Cocos Ridge impact and
have led to a bulging of the crust. This bulging in turn resulted in a predominantly vertical
movement on the controlling faults. Downfaulting and drowning of the Osa Peninsula in the
mid-Pliocene and its renewed rapid uplift in the Pleistocene and the opening of the Golfo
Dulce appear to have been the most recent processes in an ongoing development.
Regarding the three morphological units of the Golfo Dulce region it follows from the above
that
- the Golfito area has not experienced significant vertical movement and has been exposed
over a geologically long period of time,
- the Golfo Dulce may still subside,
- the Osa Peninsula has been subjected to enormous up and down movements of several
hundred metres within the last 2 Mio. or so.
With regard to the floristic development of the region, one would speculate that the tropical
wet forest of the Golfito area has survived the Pleistocene cool periods and may date back
to Miocene times. The forest coverage in the greater part of the Osa Peninsula, however, is
obviously much younger, since it had to re-colonize this area after its last emergence in the
Late Pleistocene. The key to the true age of the forest of the Golfo Dulce area may
eventually be found in longer sediment cores from the bottom of the Golfo Dulce.
8
1.3. Soils of the Golfo Dulce Region
1.3. Soils in the Golfo Dulce Region
Adapted version of PAMPERL (2001)
This section presents a brief survey of the characteristics and distribution of the soil orders
and soil types found in the Golfo Dulce area. Terminology follows the classification system
of the US Soil Taxonomy (continuously updated, recent version: SOIL SURVEY STAFF 1998).
1.3.1. Main soil types
Soils of tropical rainforests are diverse (for general information see e.g. MABBERLEY 1992).
Table 1. lists the main soil types found in the humid tropics (with % representation
worldwide). According to the soil map of Costa Rica (VASQUEZ MORERA 1989), three soil
orders prevail in the Golfo Dulce region: (1.3.2.) Ultisols, (1.3.3.) Inceptisols and (1.3.4.)
Entisols. The Ultisols are the most important in the region. They are followed by the
Inceptisols, while the Entisols are of minor importance.
Old infertile loamy and clayey soils: Oxisols and Ultisols
More fertile and less weathered soils:
63
of locally less leached conditions: Alfisols and Vertisols
4
on mainly alluvial lowlands: Entisols (Fluvents) and Inceptisols (Aquepts)
12
on volcanic ash: Andosols
1
on steep slopes: Inceptisols (Tropepts) and Entisols (Lithic soils)
11
Infertile sands: Spodosols and Entisols (Psamments)
7
Infertile peats: Histosols
2
Tab. 1.1. Orders and groups of soils of the humid tropics and their global representation (in %) (from
WHITMORE 1990)
1.3.2. Pedogenesis
The most important factor inducing pedogenesis of the Golfo Dulce area is the tropical
climate. The constantly high temperatures and high precipitation cause intense, deepreaching chemical weathering of the original rock and the soil itself. Through undisturbed
pedogenesis over geologically long periods of time, the highly weathered, clayey,
characteristically yellowish-red and strongly acidic Ultisols have been formed. They would
probably cover the main area of their distribution in the Golfo Dulce region as a thick,
uniform layer if it were not for the erosive force of the high precipitation rates which have
deeply dissected the landscape and still shape the topography. With each downpour, fine
soil material is eroded from the soil surface and washed away into the numerous brooks.
9
1.3. Soils of the Golfo Dulce Region
Erosion has, over thousands of years, produced steep slopes (slope gradients up to 60%
and more), which are highly dynamic in the upper layers of the soil. Lateral soil movement
is visible in many places in the form of bent tree trunks and as the “staircase relief” of the
soil surface. In ravines and lower slopes, soils are younger and less weathered. Only on
the ridges and the upper slopes do the old clay soils still remain. Thus, even if soil maps
display only the highly weathered Ultisols, younger, more moderately weathered Inceptisols
can regularly be found in the ravines.
Pedological conditions vary across relatively short distances in the steeply dissected areas
from highly weathered, aluminium-saturated clay on the hill tops to coarser, nutrient-rich
soils with high nutrient retention capacity and optimal hydrological conditions in the small
valleys. These pedological differences undoubtedly contribute to the variety of plant
habitats and to the distribution of plant species in the National Parks.
1.3.2. Ultisols
Characteristics. These soils have a well-developed, strongly weathered clayey subsoil
that does not contain appreciable amounts of parent rock material. The clay mineral kaolinit
comprises the main fraction of the soil body and determines the small nutrient budget
through its low nutrient retention capacity. Nutrients such as calcium and magnesium
cannot subsequently be derived from primary minerals, therefore these soils are low in
nutrients and have strongly acidic subsoils. Due to decomposition of leaf litter, nutrient
supply is slightly higher in the surface horizon. Vegetation growing in Ultisols is thus forced
to adapt to nutrient deficiency. However, if the roots are able to penetrate deep enough to
reach primary minerals, the plants can absorb cations and then add nutrients to the surface
horizon through subsequent litter fall.
The iron oxides haematite, goethite and ferrihydrite give the Ultisols their characteristic
yellowish-red colour.
Distribution. The highly weathered Ultisols are the oldest and most common soils in the
Golfo Dulce area. They can be found all over the region where the parent material for
pedogenesis and the developing soil have been exposed to weathering for long periods of
time. These are the areas with occurrence of Cretaceous volcanic rock material (according
to BERGOEING 1998) and not the areas of younger (Quaternary) sediments. Ultisols are
common in the centre of the Osa Peninsula and in a broad strip on the north and northeast
sides of the Golfo Dulce. About two thirds of the Corcovado National Park on the Osa
Peninsula and the major part of the Piedras Blancas National Park are covered
predominantly by Ultisols.
10
1.3. Soils of the Golfo Dulce Region
Areas with dominant occurrence of Ultisols are often very steeply dissected, with slope
inclinations up to 60 % (“Fuertemente Ondulado”). These areas include the highest
elevations (745 m on the Osa peninsula and 597 m in the Esquinas forest).
Origin. The parent materials for soil formation are variable in the region. On the Osa
Peninsula Ultisols developed mainly from Pliocene to Pleistocene marine sediments. Along
the Golfo Dulce coast, from the Osa peninsula to the area of the Piedras Blanca N. P., the
submarine basalts of the Rincon block make up the parent rock material, whereas in the
eastern part of the Piedras Blancas N. P. the more differentiated sediment sequence of the
Golfito-Terrane overlays oceanic basalts. Pedogenesis results in Ultisols irrespective of the
lithological differences. The geological basement is described in some detail in MALZER’S
(2001) account of the geological history.
Types. Depending on the soil moisture regime and the nutrient supply, Ultisols can be
classified into two suborders in the Golfo Dulce area: Udults and Humults1. From these
suborders the following soil types are found in the area.
(a) Tropudults: Ultisols with udic soil moisture regimes (dry period less then 90 cumulative
days of the year) under tropical temperatures. This is the typical soil type in the Golfo Dulce
area. The water supply for vegetation is good during most of the year.
(b) Tropohumults: Ultisols with a high content of the nutrients carbon and nitrogen in the
upper soil horizons and with tropical temperature patterns. The soil moisture regime is also
udic in this area. This soil type only has significant coverage along the Bay of Golfito. In this
region slopes reach a gradient of greater than 60%.
Significance for plants. Aluminium saturation can reach critical concentrations in the soil
solution, resulting in a toxic effect in the thin roots (FOY 1974). The aluminium load
(depending on the Ca:Al ratio) may have a selective influence on the species composition
of the local flora. A high amount of iron oxide in the soil leads to phosphorous fixation,
which generally limits plant growth (YOUNG 1976, BORGARD 1983) and may also have an
influence on species composition (GARTLAN & al. 1986).
1.3.3. Inceptisols
Characteristics. Inceptisols are moderately developed soils with slightly weathered
subsoil. The subsoil can contain appreciable amounts of primary minerals and rock
material. Since Inceptisols still contain weatherable minerals, plant nutrient supply is
1
In the international classification system of the WRB (FAO, ISRIC und ISSS, 1998) the Ultisols of the Golfo
Dulce area are classified as Nitosols (deep, well drained clay soils with characteristic structure and ped faces)
or Acrisols (strongly acid soils with high clay content, low cation exchange capacity and low base saturation).
11
1.3. Soils of the Golfo Dulce Region
moderate to high. Nutrients are bound to clay minerals with moderate to very high nutrient
retention capacity.
Secondary iron oxides (goethite) often render Inceptisols yellowish-brown. These
characteristics set the Inceptisols apart from the more highly weathered Ultisols.
Distribution. The moderately weathered Inceptisols are the second most important soil
order in the Golfo Dulce area. These soils developed on parent material exposed to
weathering for a shorter period of time and are thus classified as younger. They occur on
recently deposited sediments along the rivers and on alluvial planes (Quaternary sediments
in the map of TOURNON & ALVARADO 1997). They are also found on terrain where erosion
has washed away the formerly overlying, more strongly weathered soil material, i.e. in
lower parts of steep slopes and in ravines.
Types. Depending on the soil moisture regime, the Inceptisols occur in two suborders:
Aquepts and Tropepts. The following types are present in the Golfo Dulce area:
(a) Tropaquepts. Inceptisols that are formed under the influence of high ground-water
levels and in tropical climatic conditions. They cover the wide plain in Corcovado National
Park near the southwest coast of the Osa Peninsula including the area of Laguna
Corcovado. This area is essentially flat and marshy, with alluvial parent material for
pedogenesis.
Aquepts are characterized by poor soil aeration, where ground-water levels regularly reach
the upper portions of the soil profile for at least part of the year. The temporarily watersaturated, redoximorphic horizons show reddish, pale and black mottling, which is a
characteristic sign of the soil moisture regime in this soil type. Nutrient supply is fairly low.
In these soils vegetation must be adapted to such hydrological conditions.
(b) Eutropepts: Nutrient-rich Inceptisols (base saturation >50%) under tropical climate.
This soil type is typical in the river basins and on the northeastern, eastern and southern
shores of the Osa Peninsula. It also covers small areas of the Piedras Blancas National
Park, where it is observed in the river basins of the Río Riyito and Río Esquinas, in the
broad valley between Fila Golfito and Fila Gamba, as well as in two places around Playa
San Josecito. These soils also develop from alluvial sediments.
If developed from non-calcaric rock, tropepts are relatively acidic soils, but may
nevertheless be nutrient-rich. Over calcaric rock the soil pH is close to neutral and an
optimal nutrient supply can create a fertile plant habitat.
Soil studies in the Bosque Esquinas have shown that Inceptisols commonly associated with
Ultisols (those occurring on the lower slopes and ravines) also belong to the eutropepts
group (PAMPERL 2001). The original material for pedogenesis in this case is not alluvial, but
in situ weathered rock from the particular geological formation.
12
1.3. Soils of the Golfo Dulce Region
1.3.4. Entisols
Characteristics. The morphology and characteristics of the soils of this soil order are very
heterogeneous, as it includes all soils that are not classified into any other order. The
common qualities of these soils are immaturity and lack of any diagnostic horizon thus
classifying them into the soil order of the Entisols. The species composition is more or less
characteristic for specific soil types where corresponding vegetation displays different
modes of adaptation to the unique hydrological conditions of the soils.
Distribution. The reasonably well developed Entisols represent the third soil order in the
Golfo Dulce area. They are typical soil formations in low-lying, plain areas such as swampy
lands, coasts and mangroves. The Entisols occur in a patchy distribution along the shore of
the Golfo Dulce and make up large parts of the Río Sierpe region.
Types. Depending on soil moisture regime and texture, Entisols appear as three different
suborders, Aquents, Psamments and Orthents, with the following soil types:
(a) Tropaquents: Entisols that are influenced by high ground-water levels in tropical
climatic conditions. They are typical soils in swampy areas, frequently poorly aerated in the
upper layers and water-saturated in the lower layers. They are also commonly known as
“gleys”. The mangrove areas on the northwest coast of the Osa Peninsula and the
estuaries of the big rivers (Río Coto Colorado, Río Esquinas, etc.) contain this soil type.
Soil-forming materials are fluviatile deposits from the river load (unconsolidated sandy-silty
material). Tropaquents also cover large parts of the landscape in the swampy lowland of
the Río Sierpe.
(b) Tropopsamments: Sandy, but not stony Entisols with tropical soil temperature
regimes. They are only documented in the soil map between Puerto Jiménez and
Puntarenitas southwards in a flat band along the beach. Unconsolidated sand is
responsible for the weakly developed structure. Because of their low water-holding
capacity, extreme hydrological conditions characterize these soils.
(c) Troporthents. Entisol with no other special diagnostic characteristics than the tropical
climatic soil conditions. These are soils in a very early stage of development. They are
limited in depth by weathered or continuous hard rock. In the soil map they are only
observed in the lower course of the Río Tigre and, outside of the Golfo Dulce region, as the
prevailing soils of the Fila Costeña.
1.3.5. Soil properties and plant diversity
It is well known that plants are adapted to hydrological soil conditions and that the species
spectrum reflects the soil properties. Additionally, plants are adapted to less conspicuous
13
1.3. Soils of the Golfo Dulce Region
chemical soil parameters such as pH, nutrient supply and specific element concentrations,
as well as to growth limiting factors such as hard rock. RICHARDS (1961) demonstrates the
correlation between vegetation and soil in the tropics. Various studies in the tropics reveal
relationships such as:
- positive correlation between tree species diversity and soil fertility (e.g. magnesium
content) (ASHTON 1982);
- plant species variation in response to phosphorus availability (GARTLAN & al. 1986);
- influence of local soil properties on species diversity in western Amazonia (KORNING & al.,
1994)
An interesting study from Indonesia was carried out by KUBOTA & al. (1998), who
investigated various soil parameters and vegetation in a transect from a ridge down to the
valley. Despite soil properties that are less than half as variable as those investigated in the
Golfo Dulce region, the study shows a recognizable relationship between tree species
diversity and soil properties.
Tree species distributions (HUBER 1996a, 1996b, WEISSENHOFER 1996, 1997) in the
Piedras Blancas National Park have been compared to the results of the soil studies. The
comparison confirms a correlation between composition of the tree flora in the main
topographical positions (ravine, slope and ridge) and pedological factors in these plots
(PAMPERL 2001). Thus differences in the number of tree species occur depending on
fertility of the surface and subsurface horizons as well as on aluminium load. The results
show the highest diversity in the ridge forest, where soils are poor in nutrients, strongly
acidic and aluminium-saturated in the subsoil, but rich in calcium and magnesium in the
epipedon. On the other hand, in those parts of the ridge where the upper horizon of soils is
also poor in Ca and Mg, only a very small number of tree species was observed. The
vegetation on slopes, where soil properties much more resemble the conditions on ridges
than those in ravines, also shares more elements of the ridge forest than of the ravine
forest. A relatively low diversity in tree species occurs in ravines where the soil is rich in
nutrients.
In order to more fully understand the relation between plant diversity and soil properties a
lot of work remains to be done. The first results of the Piedras Blancas National Park study
have laid the groundwork for further studies in this area. The rainforest of the Golfo Dulce
region is extremely interesting to scientists in various disciplines. As investigations
continue, knowledge of ecological connections will be more clearly elucidated.
14
1.4. Climate of the Esquinas Rainforest
1.4. Climate of the Esquinas Rainforest
Adapted version of WEISSENHOFER & HUBER (2001)
1.4.1. Rainfall
The Esquinas forest is one of the most humid lowland forests in Costa Rica. The Esquinas
forest and its vicinity are influenced by the rain gradient caused by the mountains of the
Fila Cruces range.
On the pacific side of Central America there are distinct rainy (May to November) and dry
(December to April) seasons, with heaviest rainfalls occurring in October and November.
On the Central American isthmus, and particularly in Costa Rica, this pattern is highly
variable due to the presence and orientation of the mountain ranges.
During the months with the highest levels of precipitation (August to November), rain falls
nearly every day. Typically, rainfall occurs in short and heavy showers in the afternoon
and during the night. However, in September and October it may rain for periods of up to
24 hours.
The dry season is not as pronounced in the area. January, February and March are the
driest months and there may be no rainfall for days at a time. However, the soil rarely has
a water deficit. The average monthly rainfall for February is 176 mm (with a consistently
high humidity level) (Tab. 1.2).
Tropenstation La
Gamba
January
February
March
1998
1999
2000
2001
2002
2003
Average
154
396.5
293.5
307
178.5
243
235
79
321
77
210
185.0
192
176
193
273
232.5
176
192.0
128
193
April
c. 408
511
374
220
264.5
227
333
May
c. 464
397
486
621.5
340.0
526
470
June
462
566
612
498
476.5
496
527
July
509
470
473
583.5
642.5
714
542
August
372
742
683
499
389.0
554
570
September
521
998
1209
709.5
945.5
456
796
October
1048
645
741
781
749.5
869
817
November
810
684
640.5
925.7
410.5
609
701
December
698
410
250
477
473.0
265
420
Total
5718
6413.5
6072
6008.2
5246.5
5279
5690
Tab. 1.2. Rainfall (in mm) during 1998 to 2003 at the Tropenstation La Gamba
15
1.4. Climate of the Esquinas Rainfores
During the drier period of the year some trees drop their leaves completely and a
considerable amount of leaf litter accumulates on the forest floor. Some of the smaller
streams on steep terrain dry up, but others persist and form small pockets of water. From
December 1997 until April 1998, “El Niño” had a great affect on the region and little rain
fell during this period. Many epiphytes died, but others recovered within a few months
(Tab. 1.2).
Storms also occur, but only rarely with the force of a tornado (COEN 1983, BOZA &
MENDOZA 1981). In May 1997 a strong storm severely affected the vicinity of La Gamba.
Many large trees, mainly on steep ridges, were felled by the storm, producing large tree
gaps.
Meteorological data have been recorded since 1998 at the Tropenstation La Gamba and
complete data sets for precipitation are available for the years 1999 to 2003 (Fig. 1.2.).
The average annual precipitation at the field station was 5.690 mm, with the highest
monthly average in September. 152 mm of precipitation were measured in just a few
hours in September 2000. The driest month was February, but even in this month we
measured from 77 (2000) to 321 mm (1999) of precipitation. The data and diagram show
clearly that there is a period with less rain, but not a real dry season (precipitation less
than 100 mm each month). A somewhat drier period of some weeks may also occur in
July and/or August, giving rise to the so-called veranillo (“little summer”). The highest
rainfall normally occurs in September, October and November. During the year it rains on
between 255 and 302 days. At the Tropenstation La Gamba just 63 to 110 days (“El Niño”
year) each year are without rain. The average number of rainy days is 275.4 per year
(Tab. 1.3). However, in the year 2000 over 60% of the yearly precipitation was measured
in just 67 days. There was no rain on 93 days and less than 10 mm of rain was recorded
on 104 days (Tab. 1.4).
Month
Days with
rain
I
14
II
III
13.7
15.2
IV
22
V
26.2
VI
VII
VIII
25.3 24.8
27.7
IX
27
X
29.5
XI
XII
27
23.3
Total
275.7
Tab. 1.3. Monthly average of rainy days during the years 1998 to 2003 at the Tropenstation La Gamba
1998
1999
2000
2001
2002
2003
Total annual precipitation in mm
c. 5718
6413.5
6072
6008.2
5246.5
5279
Max. daily precipitation in mm.
126
(09/04)
133
(25/09)
152
(03/09)
117
(31/10)
135
(13/11)
123
(29/10)
No. of days without rain
c. 110
63
93
89
86
103
Tab. 1.4. Climatic data for the Tropenstation La Gamba
16
1.4. Climate of the Esquinas Rainforest
The diagrams of rainfall on the coast of the Esquinas forest measured by Ron and Trude
McAllister (Casa Orquideas) during 1985 and 1999 and at the Ranger station in the Valle
Bonito in the centre of the Esquinas forest, show similar levels of precipitation (Fig. 1.3
and 1.4). It is probable that rainfall levels are higher on the highest peaks in the Esquinas
forest (e.g. Cerro Nicuesa, 579 m) than at the field station.
27,9° 5816
38,5
33,7
23,2
20,0
Fig. 1.2. Climatic diagram of the Tropenstation La Gamba
Casa Orquideas (sea level)
[15]
5742
Valle Bonito - Ranger station (120 m)
[1,3]
Fig. 1.3. Precipitation diagram of the Casa
Orquideas (sea level
5829
Fig. 1. 4. Precipitation diagram of the Ranger
Station in the Valle Bonito (140 m
above sea level)
17
1.4. Climate of the Esquinas Rainfores
1.4.2.Temperature and humidity
Because Costa Rica is in the equatorial zone, the average temperature of the warmest
month does not exceed the average temperature of the coolest month by more than 5°C at
a given location. Temperature measurements at Tropenstation La Gamba have been made
sporadically since December 1993 and regularly since January 1998 (or May 1997) (Tab.
1.4). The average temperature is 27.8 °C, and the minimum temperature at Tropenstation
La Gamba of 20.0 °C was reached in August. The maximum temperature measured was
39 °C, which occured in December.
1998
1999
2000
2001
2002
2003
Average max.
Temp. (°C)
c. 32.87
31.2
31.8
32.2
32.3
31.7
Average min.
Temp. (°C)
c. 24
23.2
23.2
23.4
23.8
23.1
max. Temp. (°C)
38.5
(05/04)
37
(16/09)
39
(09/12)
35.5
(01/04)
min. Temp. (°C)
20
(15/08)
21
(03/05)
22 (many
days in Jan.
and Feb.)
22 (18/01;
21/01; 8/02;
27/10)
35 (30/04;
28/08; 06/11;
07/11)
21
(03/05;
17/05; 10/10)
35
(many
days)
22
(many
days)
Tab. 1.5. Temperature data for the Tropenstation La Gamba
The coolest month was November, with an average temperature of 27.0 °C. The months
with the highest average temperatures were March and April with 28.7 °C and 28.6 °C
respectively (Tab. 1.5).
Measurements inside the forest at about 300 m altitude were made between January and
May 1994 (Weissenhofer 1996). These data suggest that inside the forest the atmospheric
temperature is lower during the daytime and slightly higher at night than it is near the
Tropenstation La Gamba. The relative humidity is also consistently high, averaging 88.3%
at the station and 97.7% inside the forest (ASHAN 1996).
The drier season usually shows a greater range of diurnal temperatures (based on monthly
averages) than the rainy season. A large range, between 10 and 13°C, is usually recorded
during January, February and March and sometimes also in April and May. The highest
diurnal variation (up to 15 °C) occurred in September and December. The lowest diurnal
variation is around 2.0 °C and occurs during long, rainy periods in October and November,
when the weather remains cloudy throughout the day and night. The seasonal variation in
the mean monthly average at the field station is about 2.0 °C, and thus the diurnal
temperature variation appears to be greater than the seasonal temperature variation (Tab.
1.6).
18
1.4. Climate of the Esquinas Rainforest
Relative humidity is consistently high. Mist forms daily at dawn and sometimes at dusk and
after heavier rainfalls. Under these conditions the air temperature is lower
Month
January
Mean monthly minimum
23.2
Mean monthly maximum
32.3
Mean monthly average
27.8
February
23.1
33.1
28.1
March
23.6
33.6
28.6
April
23.4
33.1
28.2
May
23.9
31.5
27.7
June
23.3
32.4
27.8
July
23.5
31.9
27.7
August
23.5
32.1
27.8
September
23.5
32.3
27.9
October
23.5
31.1
27.3
November
23.3
30.7
27.0
December
23.2
30.3
26.8
Tab. 1.6. Mean monthly minimum, maximum and average air temperatures (in °C) at the Tropenstation
La Gamba, 1999-2003
19
1.5. Flora of the Golfo Dulce Region
1.5. Flora of the Golfo Dulce Region
1.5.1. A Brief Historical Overview
Of all the Central American countries, Costa Rica has received the most attention from
botanical collectors. Most other Central American countries were explored by Spanish
botanists in colonial days, but Costa Rica seems almost wholly to have escaped their
attention. From the 19th century onwards, however, various European botanists have
worked in Costa Rica. The names of certain plants commemorate these scientists: e.g.
Inga oerstediana (OERSTED from Denmark, 1864), Cryosophila warscewiczii (WARSCEWICZ
from Poland, 1848), Annona pittieri (PITTIER from Switzerland, 1887). Adolfo TONDUZ after
whom Ficus tonduzii is named collected more than 18,000 specimens in Costa Rica and
DURAND & PITTIER (1891-1901) began publication of the first comprehensive account of the
Costa Rican flora in their “Primitiae Costarricenses”, bringing together about 5,000 species.
Later, Central America and Costa Rica became politically attractive to the US, and US
botanists began coming to Costa Rica. ALLEN, BURGER, COOPER, DONNELL SMITH and
STANDLEY are among those botanists who worked in Costa Rica in the 20th century. Paul
STANDLEY (1937) collected more than 15,000 plant specimens and produced an annotated
list of 5,815 higher plant species in Costa Rica. He recognized the country’s rich plant
diversity: “Of all Central American countries Costa Rica possesses by far the richest flora”.
WILLIAM BURGER collected about 70,000 specimens (GENTRY 1978) for the “Flora
Costaricensis” (1971), and asked: “Why are there so many kinds of flowering plants in
Costa Rica?” (BURGER 1980). GOMEZ-LAURITO and FOURNIER (1985) counted about 860
native genera of woody plants from 192 families in Costa Rica. Knowledge of the Costa
Rican flora has increased rapidly during the last century. In 2004 (HAMMEL & al.), INBio
(Instituto Nacional de Biodiversidad) and the Missouri Botanical Garden published the
“Flora Costaricensis” after intensive work begun in 1989. Today, 9,361 species, 2,023
genera and 255 families of higher plants are known to exist in Costa Rica.
The botanical work done around the Golfo Dulce must also be considered in a historical
overview to the flora of Costa Rica. The Austrian botanist Georg CUFODONTIS begin
scientific work in this region when he visited the Esquinas forest during the Austrian Costa
Rica Expedition in 1930. In the 1950s the US botanist Paul ALLEN, who was living in Palmar
Sur, published the “Flora of the Golfo Dulce Rainforests” (1956), which is still used today.
Then in 1997 INBio published a field guide to the trees of the Osa Peninsula (QUESADA &
al.), invastigations on diversity and ecology in the Esquinas forest was made by HUBER
20
1.5. Flora of the Golfo Dulce Region
(1996) and WEISSENHOFER (1996) and in 2001 the “Introductory field guide to the flowering
plants of the Golfo Dulce Rainforests, Costa Rica” (WEBER & al. 2001) was published.
1.5.2. Flora and Diversity
One of the most distinctive features of tropical forests worldwide and Neotropical lowland
forests in particular is the high diversity among many different taxa (KRICHER 1997). In a
temperate forest it is often possible to count the number of tree species on the fingers of
one hand. In the tropics, one hectare normally contains more than 40 species of tree, and
may contain up to 300 species (GENTRY 1988a). Compared with other neotropical forests,
central American forests show a variable but generally high species richness. For example
at La Selva Biological Station in Costa Rica one species of leguminous tree, Pentaclethra
macroloba, is disproportionately abundant compared with all the other species (HARTSHORN
& HAMMEL 1994). Up to 113 species of tree (≥ 10 cm d.b.h.) were counted in one hectare of
lowland forest on Barro Colorado Island (Panama) (FOSTER & HUBBELL 1990). Forests with
high diversity (up to 149 species per hectare) in Costa Rica are found on the Atlantic plains
in the very wet Braulio Carillo National Park (up to 8,000 mm rainfall per year) at 300 m
above sea level (LIEBERMANN & al. 1996).
Costa Rica has the greatest plant diversity of all Central American countries. This prompted
BURGER (1980) to ask “Why are there so many kinds of flowering plants in Costa Rica?” In
fact there are many reasons: the favourable climate which leads to geographic
heterogeneity also supports a great number and variety of biological interactions which are
made less predictable by small-scale disruption. These factors, together with migration,
speciation, and fusion of once separate areas, have given rise to one of the world’s richest
biotas (BURGER 1980). The Golfo Dulce region especially, which is the southern pacific
region of Costa Rica, has very high biological diversity within a relatively small
geographical area (VAUGHAN 1981).
Paul ALLEN (1956) counted 134 families embracing some 1,315 species in 662 genera in
the rainforest of the Golfo Dulce. So far now 2,369 species in 961 genera of 182 vascular
plant families (ferns, fern allies and flowering plants) have been recorded for the Golfo
Dulce area (INBio data base). It is also notable that the region harbours over 700 tree
species - the greatest diversity of tree species in all of Central America - which represents
one quarter of all tree species in Costa Rica (QUESADA & al. 1997). Studies of the
Corcovado National Park showed 98 species of tree (≥ 10 cm d.b.h.) per hectare of a virgin
forest (HOLDRIDGE & al. 1971) and an unpublished work of ZAMORA & al. shows that 149 to
21
1.5. Flora of the Golfo Dulce Region
201 species of woody plants (≥ 5 cm d.b.h.) are growing in the rainforests around the Golfo
Dulce.
In recent years, 57 new species in this area have been described. Significant new species
include Costus osaensis (Costaceae), Ruptiliocarpon caracolito (Lepidobotryaceae),
Justicia peninsularis (Acanthaceae), Pleurothyrium golfodulcensis, Aiouea obscura, Ocotea
patula (Lauraceae), Inga golfodulcensis (Fabaceae-Mimosoideae) and Stemmadenia pauli
(Apocynaceae). Since the beginning of the INBio work (1989), 59 species have been newly
recorded for the flora of Costa Rica. Ironically, most of the new discoveries were along
paths or near clearings (pers. com. ZAMORA). Out of ten species in this region which are
new to science, five species also occur in our research plots in the Esquinas forest (sp.
nov. ined. - Guatteria, Myrcia 1, 2, 3 and Pradosia). Knowledge of diversity is still growing
and tropical flora remain undercollected (PRANCE & al. 2000).
1.5.3. Biogeographical Introduction
North and South America were separated during the early evolutionary history of
angiosperms and the isthmus was formed in the Late Tertiary (c. 5 Mio. Years ago). Little is
known about the history of the geographical distribution of plants in Central America and
South America (CROAT & BUSEY 1975). Plants, like animals, migrated in both directions
during the Late Tertiary.
The flora of Central America belongs to the Neotropical Floristic Kingdom (TAKHTAJAN
1986). The neotropics are the most diverse region of the world (GENTRY 1978). About 32
families only occur in the neotropics (BORHIDI 1991), including Bromeliaceae, Cactaceae,
Caryocaraceae,
Cyclanthaceae,
Heliconiaceae,
Humiriaceae,
Marcgraviaceae,
Quiinaceae, Theophrastaceae, Vochysiaceae, among others. The neotropical flora shares
a common origin with the paleotropical flora and it may be assumed, at least for the
flowering plants, that its roots are in the Paleotropical kingdom. Many families have pantropical
distribution,
including
Annonaceae,
Bignoniaceae,
Bombacaceae,
Chrysobalanaceae, Clusiaceae, Lauraceae, Malpighiaceae, Moraceae, Myristicaceae,
Proteaceae, Sapindaceae, Sapotaceae, etc. About 450 genera are pantropical (TAKHTAJAN
1986) and 3,660 are found only in the neotropics (GENTRY 1982a).
The Neotropical kingdom is divided into five floristic regions (TAKHTAJAN 1986). Costa Rica
belongs to the Caribbean Region, which also includes the tropical lowlands and coasts of
Mexico, the southernmost tropical part of the Florida Peninsula, the Bahamas and
Bermuda Islands, the Greater and Lesser Antilles, all of Central America from Mexico to
Panama, the shores of Ecuador, Colombia and Western Venezuela, and the pacific islands
of Revillagigedo, Galapagos and Cocos. The Caribbean region is divided into three
22
1.5. Flora of the Golfo Dulce Region
provinces, the Central American province, the West Indian province and the Galapageian
province. Costa Rica is part of the Central American Province, which reaches from Mexico
to the northern parts of South America (see above) and includes the highly diverse region
of the Chocó (pacific lowlands of Colombia, Ecuador and northern Peru). Typical genera for
the Caribbean region are, for example, Beilschiedea, Clidemia, Guarea, Marcgravia,
Miconia, Ocotea, Ouratea, Podocarpus, Sapium, Schlegelia, Simarouba, Trichilia.
Costa Rica divides into four phytogeographical regions (WERCKLE 1909): (1) The lowlands
of the Caribbean from sea level up to 800 m; (2) the lowlands of the Pacific region from sea
level up to 800 m; (3) the temperate region from 800 to 1,500 m; and (4) the cold region
above 1,500 m. To these should also be added the highlands of the Páramo (HARTSHORN
1983).
1.5.4. Floristic affinities
The similarity of climate and geographical situation between Southern Central America and
the Golfo Dulce rainforest leads one to expect certain floral similarities. Indeed, the flora of
the Golfo Dulce has strong affinities to that of the forests of South America (GENTRY 1978,
1982a) and of the Chocó region of north-western South America (HARTSHORN 1983,
HARTSHORN & HAMMEL 1994, HUBER 1996, STANDLEY 1937), as well as to that of the
Amazonian and Atlantic coastal rainforests of Costa Rica (ALLEN 1956). It has less affinity
with the flora of the adjacent regions of Chiriquí (Panama) or Guanacaste (the dry north of
Costa Rica) (ALLEN 1956). GOMEZ (1982, 1986), HAMMEL & GRAYUM (1982) and HAMMEL
(1986) also describe the floral affinity of Costa Rica with South America and assign this fact
specifically to the flora of La Selva (Caribbean region of Costa Rica). The pteridophytes of
La Selva, although in general more wide-ranging than the seed plants, also show definite
evidence of southern rather than northern affinities (GRAYUM & CHURCHILL 1987). An
analysis of six representative families (Cyclanthaceae, Marantaceae, Cecropiaceae,
Clusiaceae, Lauraceae and Moraceae) of the La Selva primary forests shows that about
85% of the species are known from Panama or South America, whereas only about 45%
(widespread species) also occur in Nicaragua or farther north. Only 2% of the species only
occur farther north (HAMMEL 1986). These general trends are also true for several other
large families (HARTSHORN & HAMMEL 1994). The flora of Barro Colorado Island (Panama)
seems to consist of approximately equal numbers of both North (chiefly Central) and South
American species. Some 12% of the species are endemic to Barro Colorado and 11% are
restricted to Panama and to nearby regions of Colombia and Costa Rica. About 50% are
common to both North and South America (widespread) (CROAT & BUSEY 1975). This
“pattern” of affinity to South America seems to be a logical outcome of tectonic movement,
23
1.5. Flora of the Golfo Dulce Region
volcanism, original distribution and recent dispersal, and the direction of dispersal has been
predominantly from the nearest tropical continental source to this tropical isthmian region.
About 45% of species of the six investigated families of La Selva are endemic to Central
America, and approximately 10% are endemic to Costa Rica (HARTSHORN & HAMMEL
1986). Elements of the Guiana flora are also found in the flora of Costa Rica; they may
have arrived before the formation of the isthmus of Panama (GÓMEZ 1986). The evidence
of floral distribution indicates that most migrations between North and South America took
place in the late Tertiary period (CROAT & BUSEY 1975).
The South American tree species of the Esquinas forest are mostly abundant. Some of
them, such as Chaunochiton kappleri (Olacaceae), reach their northern limit here
(QUESADA & al 1997). Other species, some recently discovered, are also found further
north. These include Couratari guianensis (Lecythidaceae), Brosimum utile (Moraceae) in
the Canton de Aguirre and Portalon-Quepos-Puntarenas (600 m), and Caryocar
costaricense (Caryocaraceae) in the Reserva Biológica Carara (WEBER et al. 2001).
There are also distinct affinities to (1) Costa Rica's mainland flora, where shared species
are found either at higher elevations (e.g. Oreomunnea pterocarpa - Juglandaceae,
Ticodendron incognitum - Ticodendraceae) or on the Atlantic slope (e.g. Ruptiliocarpon
caracolito - Lepidobotryaceae, (2) Jamaica (Ziziphus chloroxylon - Rhamnaceae) and (3)
the floras of several countries further north, including as far as Mexico (e.g. Recchia
simplicifolia - Simaroubaceae) (HERRERA-MCBRYDE & al. 1997).
It is generally acknowledged that the area surrounding the Golfo Dulce was a natural
refuge, cut off from the mainland during glacial periods. Because of this, speciation
occurred at an accelerated rate and many new species evolved in the fragmented forests.
The results were so-called “hot-spots” of biodiversity containing high numbers of endemic
species. The MINAE (Ministerio de Natural, Ambiente y Energia de Costa Rica) counts the
region around the Golfo Dulce among the richest in endemism of Costa Rican plants
(http://www.minae.go.Costa Rica/estratigia/estudio/mapa2.html, 2000). Perhaps 2 to 3% of
the flora of the Osa peninsula will be found to be endemic, which would includes 25%
endemism of the Marantaceae (HERRERA-MCBRYDE & al. 1997).
24
2. Materials and Methods
2. Materials and Methods
2.1. Study Area
Adapted version of WEISSENHOFER (in prep.)
According to the Holdridge Life Zone System (HOLDRIDGE 1967), the main life zones in the
region are the Tropical Wet Forest, the Tropical Moist Forest and the Tropical Premontane
Wet Forest. The study sites belong to the Tropical Wet Forest life zone with over 4000 mm
precipitation per year and an average temperature of 27.8°C. This is the most diverse life
zone within Costa Rica and actually only present in the Tortuguero and Sarapiqui region on
the Atlantic coast as well as in the Golfo Dulce region on the Pacific coast (HARTSHORN
1983).
For the analysis of biodiversity and biogeographical affinities four typically primary forest
sites within the PN Piedras Blancas were selected, each of the plots covering one hectare
of primary forest. The plots are geomorphologically very different (inland and coastal
Fig. 2.1. Aerial photograph of the Esquinas forest, marked with some locations:
1: "Tropenstation La Gamba", 2: Golfo Dulce Lodge, 3: Casa Orquideas, 4: Ranger Station in the
25
2. Materials and Methods
Valle Bonito, r: ridge, g: gorge, i: inland slope, c: coastal slope.
slopes, gorges and ridges) and represent the most abundant forest types in the region. All
the plots show different vegetation types and are named as follows: Coastal slope, Inland
slope, Gorge, Ridge.
As a base for our studies we used the Tropenstation La Gamba (N 8°42'61’’, W 83°12’97’’),
a field station near the village of La Gamba, which is run by an Austrian non-profit
organization. The station can be reached all the year round and lies only 4 km SW of the
Interamerican Highway. It offers good accommodation and research facilities for doing
fieldwork.
The plots were established between the years 1993 and 2000. The ridge, gorge and inland
slope plots are situated in the vicinity of the Tropenstation La Gamba. The coastal slope
plot was selected at the southern border of the Piedras Blancas National Park near the
Playa San Josesito, about a 20 min walk from the Golfo Dulce Lodge. The gorge plot is just
behind the Esquinas Rainforest Lodge (Ozelot trail and Catarata trail) and the inland plot
slope is situated on the hilltops of the Fila trail. The ridge plot can be reached by a 40 min.
walk along the Fila trail from the Tropenstation (Fig. 2.1).
2.2. Description of the plots
Adapted of WEISSENHOFER (in prep.)
2.2.1. Coastal slope
The coastal slope was established in July 2000. It is situated near the coast on the
southern border of the Piedras Blancas National Park on the Playa San Josesito between
N 8°39’35’’, W 83°15’16’’ and N 8°40’6’’, W8°15’3’’ at an altitude of 96 m –210 m. This plot
was selected because its vegetation is typical for the Golfo Dulce coastline, where the
species Schizolobium parahyba (Fabaceae-Caes.) is particularly abundant. The plot is
situated on a SW exposed and homogenous steep slope with an average inclination of
44%.
The plot shows a typical geomorphological structure that we found only in very steep
forests. When trees snap further down the hill, the root discs leave behind a round and
relatively flat section. In this section seeds and seedlings can regenerate quicker and are
not washed out as easily as in the steep parts of the forest. They are also exposed to better
26
2. Materials and Methods
light conditions and so they develop better. In this manner, terraces are built and these flat
areas allow trees to grow very tall and thick.
The plot can be approached by boat from Golfito (1 hour) and then on foot by passing the
Golfo Dulce Lodge (20 min).
2.2.2. Gorge
The gorge plot was established in March 1997. The plot is divided into 2 subplots because
a single gorge of sufficient length was not available. Section A is located along the Sendero
Catarata from about 85 m to c. 90 m altitude at N 8°41’49’’, W 83°12’19’’. The research
strip is 200 m long and 20 m wide.
Section B is located at the Sendero Ozelot at N 8°41’52’’, W 83°12’19’’ from about 85 m to
115 m altitude. The strip is 300 m long and 20 m wide. Both subplots are situated behind
the Esquinas Rainforest Lodge along two small rivers, which have water all the year round.
They can be reached in a 15 min. walk from the Tropenstation La Gamba.
We selected this site because of its characteristic gorge vegetation and easy access to
both subplots. Due to selective cutting during the last 20 years (pers. comment Agustin
Zuniga) small parts of the forest are covered with secondary vegetation. However, this will
have minimal impact on results due to the high dynamics of the forest (see WEISSENHOFER
in prep).
In general the Ozelot section is steeper and better drained than the Catarata part. The first
has inclinations of an average of 9.6%, the latter of 2.75%. Both subplots contain several
deep gorges because of the high precipitation. The flat parts are poorly drained and found
in both sections. Underground vegetation varies conspicuously between the different parts.
Average pH is 5.1.
2.2.3. Inland slope
The inland slope was established in 1993/1994 for Weissenhofer`s (1996) and the author`s
(1996a) diploma thesis and is situated 300-336 m above sea level at about N 8°41’ and W
83°12’. This forest is situated on a well-drained SE exposed slope and can be reached by a
40 min walk from the Tropenstation in a SW direction. We found this forest site with the
help of indigenous people and people from La Gamba and it is probably the nearest
primary forest of this formation to Tropenstation La Gamba.
The well-drained plot shows a characteristic plant formation of virgin slope forests in the
region, and it has inclinations of an average of 27% with a maximum of 55% in certain
places. There are several gorges due to the high precipitation. Due to landslips the plot is
27
2. Materials and Methods
extremely steep (up to 90°) in some places. In general the upper, western part is steeper
than the lower eastern part, so some flat sites can still be found here.
The vegetation in the different parts of the plot differs conspicuously.
Data from the diploma plot are also included in this study. For detailed information about
this plot see also HUBER 1996a, WEISSENHOFER 1996.
2.2.4. Ridge
The ridge plot was established in July 1998 and covers important sections along the Fila
trail of the Piedras Blancas National Park. This plot is also subdivided into two sections.
Section A (Fila dry) is situated from 199 m – 210 m altitude at approx. N 8°41’51’’, W
83°12’19’’. The research strip is 120 m long and 20 m wide. Section B (Fila wet) is situated
between 199 and 215 m altitude at N 8°41’59’’, W 83°12’26’’. This research strip is 380 m
long and 20 m wide. Due to the dense foliage it is difficult to get an exact position with a
GPS system.
The division was made mainly to accommodate the different soil conditions in the Fila area
described by PAMPERL (2001).
The ridge plot is a typical forest along the Fila trail, about 30 min walk from the
Tropenstation La Gamba. It is not easy to find primary ridge forests in the surroundings of
the park due to felling of the valuable timber tree Peltogyne purpurea (Fabaceae-Caes.),
which only grows along drier ridges.
In general ridges are well drained and the soil is drier than in other parts of the forest.
Inclinations are on average 13.7% (maximum 80%) in the wet section and 7.8% (maximum
60%) in the drier section. Average pH is 6.2.
pH (average)
Inland slope
5.4
Gorge
5.1
Ridge
6.2
Tab. 2.1. Average pH of the forest plots (PAMPERL 2001)
2.3. Field Work and Plant Identification
Four physiognomically homogenous one-hectare plots were selected, marked out and
subdivided into 10 to 100 subplots. The subplots in the gorge and on the ridge measured
500 x 20 m and were next to paths; the subplots on the coastal slope were 250 x 40 m.
28
2. Materials and Methods
Additionally one one-hectare plot in the Esquinas Forest was investigated for the master
theses of HUBER (1996a) and WEISSENHOFER (1996).
Coastal slope
(50 subplots) - each 200 m2 (20 x 10 m)
Gorge
(10 subplots) - each 100 m2 (50 x 20 m)
Ridge
(50 subplots) - each 200 m2 (20 x 10 m)
Inland slope
(100 subplots) - each 100 m2 (10 x 10 m)
All four plots were used for comparisons of floristic diversity. All living plants (trees, lianas
and hemi-epiphytes) with a diameter at breast height of 10 cm or more (d.b.h. ≥ 10 cm)
were counted. Voucher specimens of almost all species were collected and deposited in
Costa Rica at the Herbario Nacional (CR) and in Austria at the Universität Wien (WU) and
at the Biologiezentrum Linz (LI). Binoculars (Swarovski binocular SLC 7 x50 B, SLC 10x50
B) and a telescope (Swarovski ATS 80) were used for field identification. The fieldwork was
conducted from February 1998 to August 2001. All the investigated data of the plots and
their distribution are now contained on a database (MS ACCESS, MS EXCEL).
2.4. Diversity Indices
Diversity indices are single-figure numerical measures of diversity which incorporate
species richness and equitability (evenness). Different priorities in the reconciliation of the
two factors have led to the invention of a variety of indices, which are optimized in different
ways and hence have different merits. No one index could be said to be superior for all
circumstances.
2.4.1. Simpson's Index
Simpson's index (SIMPSON 1949, PIELOU E. C. 1969, 1975) assumes that the proportion of
individuals of a given species in an area adequately represents their importance to
diversity. It is given by:
D = 1/(Sum (pi2))
where D is the diversity and pi is the overall proportion of the ith species. The index ranges
from zero to the total number of species. A value of one indicates that all the individuals in
the area belong to a single species, and when D = S, every individual belongs to a different
species.
29
2. Materials and Methods
Simpson's index gives the probability that any two individuals, drawn at random from an
infinite community, will belong to different species. It is an intrinsic diversity index weighted
towards equitability.
2.4.2. Shannon-Weaver (Wiener) Index2
The Shannon-Weaver index (SHANNON & WEAVER 1949, PIELOU E. C. 1969, 1975) is very
similar to Simpson's index except for the underlying distribution. Simpson's index assumes
that the probability of observing an individual is proportional to its frequency in the habitat,
while the Shannon-Weaver index assumes that the habitat contains an infinite number of
individuals. It is given by:
H = Sum(pi - ln(pi))
This index also ranges from zero to S. One must quote the log base used, since it alters
the value; log base 2, 10 and e (natural) are commonly employed. The Shannon-Weaver
index is one of a family of intrinsic diversity indices. The evenness index E is derived from
the Shannon-Weaver index.
2.4.3. Evenness Index
Evenness is a measure of how similar the abundances of different species are. When there
are similar proportions of all species then evenness is equal to one, but when the
abundances are dissimilar (some rare and some common species) then the value
increases. One type of evenness index is derived from the Shannon-Weaver index:
E = H/ln(S)
2.4.4. Alpha Index
The log series variable alpha is commonly used as a diversity index. It assumes that the
sample is a reasonable fit to a log-series, which is often the case, and so the alpha index
appears to be robust (FISHER & al. 1943; KEMPTON & TAYLOR 1974):
α = N (1-x)/x
The total number of species, S, is obtained by adding all the terms in the series. This
reduces to the following equation:
2
The Shannon Weaver Index
Dr. Claude Shannon recently passed away. In the late 1940s Dr. Claude Shannon and Dr. Warren Weaver developed a
general model of communication. They were working in information theory, which was initially developed to separate noise
from information. The index is often called the Shannon-Wiener index. This is in acknowledgement of the contribution that the
mathematician Norbert Wiener made to the model. Norbert Wiener is renowned for his instrumental role in the development
of cybernetics. Because the model is relatively easy to use mathematically, it is one of the indices most frequently used to
measure biodiversity. The Shannon Weaver Index (H) measures, among other things, the diversity of populations. As the
value of H increases, the level of diversity also increases.
30
2. Materials and Methods
S= α[-ln(1-x)]
where x is estimated from the iterative solution of
S/N=(1-x)/x[-ln(1-x)]
and N is the total number of individuals (MAGURRAN 1988).
2.4.5. Importance Value (IVI)
The Importance Value (CURTIS & MCINTOSH 1951) measures the importance of a species in
a plot. It is the sum of relative frequency, relative density and relative dominance.
IVI = rel. frequency + rel. density + rel. dominance
The relative frequency is an index of the general dispersal of the species, based on its
presence in the sample units.
The sum of all relative frequency values for all counted species in a plot will be 100%.
The absolute frequency (MORI & al 1983, LAMPRECHT 1986) shows the presence (1) of the
species in the subplot. The sum of all presence values in the subplots is divided by the
number of subplots. This value is multiplied by 100 to show the value as a percentage.
The relative density shows the proportion of each individuals of species in the subplot.
This number is multiplied by 100 to show the value as a percentage.
The relative dominance is the proportion of the basal area in m2 of the total basal area in
a plot. This number is multiplied by 100 to show the value as a percentage.
2.4.6. Family Importance Value (FIVI)
The FIVI, defined by MORI & al. (1983), demonstrates the importance of the plant families in
a research plot. The calculation is similar to the IVI, but instead of relative frequency,
relative diversity is used. It is therefore the sum of relative diversity, relative density and
relative dominance, multiplied by 100 to show the value as a percentage. Because the
values of three different elements are combined, an overvaluation of any one of these is
avoided.
FIVI = rel. diversity + rel. density + rel. dominance
The relative diversity is the total number of species of each family divided by the total
number of species in a plot. This number is multiplied by 100 to show the value as a
percentage.
31
2. Materials and Methods
The relative density is the total number of individuals of each family divided by the total
number of individuals in a plot. This number is multiplied by 100 to show the value as a
percentage.
The relative dominance is the total amount of basal area in m2 covered by each family,
divided by the total basal area of the plot. This number is multiplied by 100 show the value
as a percentage.
2.4.7. Area-Species Curve
Area-species curve (MACARTHUR & WILSON 1967) generally fit closely to equations of the
form
S = cAz
where S = number of species, A = area, and c and z are constants. The exponent z
determines the slope of the curve and is the critical parameter in estimating extinction
rates. All one-hectare plots consist of several subplots: coastal slope (50), gorge (10),
inland slope (100) and ridge (50).
The area-species curves are also presented by means of unit subplots (tabular calculation).
The area-species curves can also be presented in linear form:
S = c.ln (a.x+b)
where S = number of species, x = area, and c, a and b are parameters of the curve.
Parameters c and b are independent of the square measure; parameter a grows
proportional to the square measure used.
2.5. Floral Similarity and Difference of all plots (ClusterAnalysis)
This method classifies together objects judged to be similar according to distance or
similarity measures. Data can be quantitative or presence/absence. The Bray-Curtis
similarity measure (PIELOU 1984) uses group-average clustering and appears to give a
useful hierarchy of clusters, which are displayed as a dendrogram and a similarity matrix.
32
2. Materials and Methods
2.6. Distribution Pattern and Geographical Affinities
The distribution of all species was investigated to show the geographical distribution
pattern. For the distribution of the species, data from Tropicos (www.mobot.org), NYBG
(http://scisun.nybg.org) and the Flora de Costa Rica (www.inbio.ac.cr) were used, as well
as different checklists and floras of Central America and South America, and monographs
on different genera, subfamilies and families (BERG 1972, BERRY & al. 1999, BRAKO &
ZARUCCHI 1993, BURGER & TAYLOR 1993, D’ARCY 1987, HENDERSON & al. 1995, MORI & al.
1997, RIBEIRO & al. 1999, STEVENS & al. 2001, WEBER & al. 2001, ZAMORA & PENNINGTON
2000).
33
3.1. Results – Floristics and Diversity
3. Results
3.1. Floristics and Diversity
The complete data are shown in the attached and lists is given in the chapter 8.lists.
Analyses of the diversity of the plots follow.
3.1.1. Life Form Distribution
Among the 2,444 individuals (indivs.) (d.b.h. ≥ 10 cm) in the four research plots studied,
four types of life form are represented (Tab. 3.1):
(1) trees: 2,124 individuals (86.9%);
(2) palms: 262 individuals (10.7%);
(3) lianas: 44 individuals (1.8%);
(4) ground-rooted hemi-epiphytes: 14 individuals (0.6%)
Coastal slope
Gorge
Inland slope
Ridge
Total
Trees
561
440
375
748
2124
Palms
8
31
142
81
262
Lianas
18
10
5
11
44
Hemi-epiphytes
1
1
5
7
14
Total
588
482
527
847
2444
Tab. 3.1. Life form distribution in the four research plots of the Esquinas Forest (indivs. with d.b.h. ≥ 10
cm)
3.1.2. Species and Family Diversity in the four plots
In total3 2,444 indivs. and 328 spp. from 69 families and 33 orders were recorded (Tab.
3.39.).
The most abundant spp. are Welfia regia (Arecaceae) with 108 indivs., Iriartea deltoidea
(Arecaceae) with 97, Symphonia globulifera (Clusiaceae) with 79, Compsoneura sprucei
with 72, Brosimum guianense (Moraceae) with 63, Qualea paraensis (Clusiaceae) with 52,
Rinorea dasyadena (Violaceae) with 48 and Brosimum utile (Moraceae), Cecropia
obtusifolia (Cecropiaceae) and Tetrathylacium macrophyllum (Flacourtiaceae) each with 46
indiv. (Tab. 3.2.).
At the research plot on the slope near the coast 588 indivs. with d.b.h. ≥ 10 cm from 108
spp. were found, in the gorge 482 indivs. from 121 spp., on the slope near the
Tropenstation La Gamba (inland slope) 527 indivs. from 133 spp. and at the ridge 847
indivs. from 179 spp (Fig. 3.1).
3
The one unidentified species in the inland slope plot is included neither in the families nor in the orders.
34
3.1. Results – Floristics and Diversity
Species
Family
Welfia regia
Iriartea deltoidea
Symphonia globulifera
Compsoneura sprucei
Brosimum guianense
Qualea paraensis
Rinorea dasyadena
Brosimum utile
Cecropia obtusifolia
Tetrathylacium macrophyllum
Arecaceae
Arecaceae
Clusiaceae
Myristicaceae
Moraceae
Vochysiaceae
Violaceae
Moraceae
Cecropiaceae
Flacourtiaceae
Coastal
Slope
0
8
36
50
30
0
0
21
6
3
Gorge
12
16
1
0
0
0
37
3
32
41
Inland
Slope
47
71
18
8
5
2
7
15
1
2
Ridge
49
2
24
14
28
50
4
7
7
0
No. of indivs. in
all plots
108
97
79
72
63
52
48
46
46
46
Tab. 3.2. The 10 most abundant species in the 4 research plots
species
individuals
Fig. 3.1. No. of indivs. and diversity of species in the four research plots 1 ha each) in the Esquinas
forest (indivs. with d.b.h. ≥ 10 cm)
The most diverse families are Fabaceae-Mim. (23 spp., 67 indivs.), Moraceae (22 spp., 250
indivs.), Clusiaceae (18 spp., 206 indivs.), Lauraceae (14 spp., 33 indivs.), Sapotaceae (14
spp., 63 indivs.), Annonaceae (12 spp., 39 indivs.), Chrysobalanaceae (12 spp., 41 indivs.),
Rubiaceae (12 spp., 40 indivs.), Melastomataceae (10 spp., 32 indivs.) and Fabaceae-Fab.
(10 spp., 35 indivs.) (Fig. 3.2).
35
3.1. Results – Floristics and Diversity
Fig. 3.2. The 10 most diverse families in the research plots
3.1.2.1 Coastal Slope
The 1ha research plot near the coast of the Golfo Dulce, the coastal slope, comprises 588
indivs. from 108 spp. 561 are trees, 8 are palms, 18 are lianas and 1 is a hemi-epiphyte.
Simpson´s index gives the value D = 0.028 (1-D = 0.972), the Shannon-Weaver index
gives the value H’ = 4.014, the Evenness index gives the value J’(E) = 0.857 and the αIndex is 38.82. The mean number of indivs. per species (sp.) is 5.440 (Tab. 3.39).
On family level the indivs. are composed of 37 families of 22 orders. The Shannon-Weaver
Fig. 3.3. Number of indivs. per sp. – Coastal slope
36
3.1. Results – Floristics and Diversity
diversity index calculated for families gives H’ = 1.463 and the Evenness index gives E´=
0.926. The mean number of spp. per family is 2.920 (Tab. 3.39).
The spp. with the highest abundances are Compsoneura sprucei (Myristicaceae) with 50
indivs., Sorocea cufodontisii (Moraceae) with 43 and Symphonia globulifera (Clusiaceae)
with 36. Thirty-eight species are represented by only one individual (Fig. 3.3).
3.1.2.1.1 Importance Value Index (IVI)
The highest IVI values are 24.17 for Brosimum utile (21 indivs.; Moraceae), 18.82 for
Symphonia globulifera (36 indivs.; Clusiaceae), 16.04 for Compsoneura sprucei (50 indivs.;
Myristicaceae), 13.86 for Sorocea cufodontisii (43 indivs.; Moraceae), 12.19 for Brosimum
guianense (30 indivs.; Moraceae), 11.65 for Carapa guianensis (9 indivs.; Meliaceae), 9.08
for Manilkara staminodella (8 indivs.; Sapotaceae), 7.89 for Caryocar costaricense (9
indivs.; Caryocaraceae), 7.55 for Schizolobium parahyba (16 indivs.; Fabaceae-Caes.) and
7.15 for Protium aracouchini (17 indivs.; Burseraceae).
The total IVI of these 10 most important spp. (239 indivs.) is 128.40. The IVI for the
remaining 98 spp. (349 indivs.) is 171.60 (Tab. 3.3).
Species
Brosimum utile
Symphonia globulifera
Compsoneura sprucei
Sorocea cufodontisii
Brosimum guianense
Carapa guianensis
Manilkara staminodella
Caryocar costaricense
Schizolobium parahyba
Protium aracouchini
the other 98
Total of the 108
Family
Moraceae
Clusiaceae
Myristicaceae
Moraceae
Moraceae
Meliaceae
Sapotaceae
Caryocaraceae
Fabaceae-Caes.
Burseraceae
No. of indivs.
21
36
50
43
30
9
8
9
16
17
349
588
IVI
24.17
18.82
16.04
13.86
12.19
11.65
9.08
7.89
7.55
7.15
171.60
300.00
Tab. 3.3. IVI – Coastal slope
3.1.2.1.1.1. Relative Density
The highest values for relative density are 8.50 for Compsoneura sprucei (50 indivs.;
Myristicaceae), 7.31 for Sorocea cufodontisii (43 indivs.; Moraceae), 6.12 for Symphonia
globulifera (36 indivs.; Clusiaceae), 5.10 for Brosimum guianense (30 indivs.; Moraceae),
3.57 for Brosium utile (21 indivs.; Moraceae), 3.06 for Lacmellea panamensis (18 indivs.;
Apocynaceae), 2.89 for Garcinia madruno (17 indivs.; Clusiaceae), 2.89 for Guarea
pterorhachis (17 indivs.; Meliaceae), 2.89 for Protium aracouchini (17 indivs.; Burseraceae)
and 2.72 for Schizolobium parahyba (16 indivs.; Fabaceae-Caes.).
The total of these 10 spp. (265 indivs.) is 45.05. The relative density for the remaining 98
spp. (323 indivs.) is 54.95 (Tab. 3.4).
37
3.1. Results – Floristics and Diversity
Species
Compsoneura sprucei
Sorocea cufodontisii
Symphonia globulifera
Brosimum guianense
Brosimum utile
Lacmellea panamensis
Garcinia madruno
Guarea pterorhachis
Protium aracouchini
Schizolobium parahyba
the other 98
Total of the 108
Family
Myristicaceae
Moraceae
Clusiaceae
Moraceae
Moraceae
Apocynaceae
Clusiaceae
Meliaceae
Burseraceae
Fabaceae-Caes.
No. of indivs.
50
43
36
30
21
18
17
17
17
16
323
588
Rel. density [%]
8.50
7.31
6.12
5.10
3.57
3.06
2.89
2.89
2.89
2.72
54.95
100.00
Tab. 3.4. Relative density – Coastal slope
3.1.2.1.1.2. Relative Frequency
The highest values for relative frequency are 5.34 for Compsoneura sprucei (50 indivs. in
23 subplots; Myristicaceae), 4.87 for Sorocea cufodontisii (43 indivs. in 21 subplots;
Moraceae), 4.41 for Brosimum guianense (30 indivs. in 19 subplots; Moraceae), 4.41 for
Symphonia globulifera (36 indivs. in 19 subplots; Clusiaceae), 3.25 for Lacmellea
panamensis (18 indivs. in 14 subplots; Apocynaceae), 3.02 for Guarea pterorhachis (17
Species
Compsoneura sprucei
Sorocea cufodontisii
Brosimum guianense
Symphonia globulifera
Lacmellea panamensis
Guarea pterorhachis
Protium aracouchini
Brosimum utile
Heisteria concinna
Protium ravenii
the other 98
Total of the 108
Family
Myristicaceae
Moraceae
Moraceae
Clusiaceae
Apocynaceae
Meliaceae
Burseraceae
Moraceae
Olacaceae
Burseraceae
No. of indivs.
50
43
30
36
18
17
17
21
15
12
329
588
No. of subplots
23
21
19
19
14
13
13
12
12
11
Rel. frequency [%]
5.34
4.87
4.41
4.41
3.25
3.02
3.02
2.78
2.78
2.55
63.57
100.00
Tab. 3.5. Relative frequency – Coastal slope
indivs. in 13 subplots; Meliaceae), 3.02 for Protium aracouchini (17 indivs. in 13 subplots;
Burseraceae), 2.78 for Brosimum utile (21 indivs. in 12 subplots; Moraceae), 2.78 for
Heisteria concinna (15 indivs. in 12 subplots; Olacaceae) and 2.55 Protium ravenii (12
indivs. in 11 subplots; Burseraceae).
The total of these 10 spp. (259 indivs.) is 36.43. The relative frequency for the remaining 98
spp. (329 indivs.) is 63.57 (Tab. 3.5).
3.1.2.1.1.3. Relative Dominance
The basal surface area of all 588 indivs. is 43.47 m2 which represents 100% of the relative
dominance.
Table 3.6. shows the 10 spp. with the largest basal surface, which occupy more than half of
the basal surface of the 108 spp. in the coastal plot.
38
3.1. Results – Floristics and Diversity
The value of relative dominance indicates the spp. with the largest basal area. Brosimum
utile (Moraceae) has a relative dominance of 17.82 (7.75 m2), Symphonia globulifera
(Clusiaceae) 8.29 (3.60 m2), Carapa guianensis (Meliaceae) 8.26 (3.59 m2), Manilkara
staminodella (Sapotaceae) 6.33 (2.75 m2), Caryocar costaricense (Caryocaraceae) 4.74
(2.06 m2), Elaeoluma glabrescens (Sapotaceae) 3.97 (1.72 m2), Pouteria durlandii
(Sapotaceae) 2.88 (1.25 m2), Brosimum guianense (Moraceae) of 2.68 (1.17 m2),
Schizolobium parahyba (Fabaceae-Caes.) of 2.51 (1.09 m2) and Tapirira myriantha
(Anacardiaceae) of 2.30 (1.00 m2).
The total of the 10 spp. (141 indivs.) is 59.78 (25.98 m2). The remaining 98 spp. (447
indivs.) have a relative dominance of 40.22 (17.49 m2) (Tab. 3.6).
3.1.2.1.1.4. Absolute Frequency
The absolute frequency shows the abundance of species in a subplot. Those species with
a higher absolute frequency are often those with a higher relative density.
Table 3.7. shows the most frequent species per sample unit.
The highest values for the absolute frequency are 92% for Compsoneura sprucei (50
indivs.; Myristicaceae), 84% for Sorocea cufodontisii (43 indivs.; Moraceae), 76% for
Brosimum guianense (30 indivs.; Moraceae), 76% for Symphonia globulifera (36 indivs.;
Species
Brosimum utile
Symphonia globulifera
Carapa guianensis
Manilkara staminodella
Caryocar costaricense
Elaeoluma glabrescens
Pouteria durlandii
Brosimum guianense
Schizolobium parahyba
Tapirira myriantha
the other 98
Total of the 108
Family
Moraceae
Clusiaceae
Meliaceae
Sapotaceae
Caryocaraceae
Sapotaceae
Sapotaceae
Moraceae
Fabaceae-Caes.
Anacardiaceae
No. of indivs.
21
36
9
8
9
4
11
30
16
9
435
588
Basal surface area [m]2
7.75
3.60
3.59
2.75
2.06
1.72
1.25
1.17
1.09
1.00
17.49
43.47
Rel. dominance [%]
17.82
8.29
8.26
6.33
4.74
3.97
2.88
2.68
2.51
2.30
40.22
100.00
Tab. 3.6. Relative dominance – Coastal slope
Clusiaceae), 56% for Lacmellea panamensis (18 indivs.; Apocynaceae), 52% for Guarea
pterorhachis (17 indivs.; Meliaceae), 52% for Protium aracouchini (17 indivs.;
Burseraceae), 48% for Brosimum utile (21 indivs.; Moraceae), 48% for Heisteria concinna
(15 indivs.; Olacaceae) and 44% for Protium ravenii (12 indivs.; Burseraceae).
Thirty-eight spp. of the 108 spp. are represented by only one individual (indiv.) and are
therefore present in only one subplot. The absolute frequency for these spp. is 4% (1/25 x
100) (Tab.3.7).
Species
Family
No. of
39
Abundance in
Absolute
3.1. Results – Floristics and Diversity
Compsoneura sprucei
Sorocea cufodontisii
Brosimum guianense
Symphonia globulidera
Lacmellea panamensis
Guarea pterorhachis
Protium aracouchini
Brosimum utile
Heisteria concinna
Protium ravenii
indivs.
50
43
30
36
18
17
17
21
15
12
Myristicaceae
Moraceae
Moraceae
Clusiaceae
Apocynaceae
Meliaceae
Burseraceae
Moraceae
Olacaceae
Burseraceae
25 subplots
23
21
19
19
14
13
13
12
12
11
frequency [%]
92
84
76
76
56
52
52
48
48
44
Tab. 3.7. Absolute frequency – Coastal slope
3.1.2.1.2. Family Diversity
On the coastal slope 108 spp. from 39 families have been found. The most diverse families
in terms of species on the coastal slope are the Moraceae (10 spp., 117 individuals
(indivs.)), the Sapotaceae (8 spp., 34 indivs.), the Lauraceae (6 spp., 14 indivs.), the
Burseraceae (5 spp., 37 indivs.), the Clusiaceae (5 spp., 57 indivs.), the FabaceaeFaboideae ( 5 spp., 10 indivs.), the Fabaceae-Mimosoideae (5 spp., 6 indivs.), the
Meliaceae (5 spp., 42 indivs.), the Myristicaceae (5spp., 56 indivs.), the Chrysobalanaceae
Fig. 3.4. Family diversity – Coastal slope
(4 spp., 7 indivs.), the Flacourtiaceae ( 4 spp., 10 indivs.), the Olacaceae (4 spp., 24
indivs.) and the Tilliaceae (4 spp., 16 indivs.) (Fig. 3.4.). 17 families are represented with
just one species each.
40
3.1. Results – Floristics and Diversity
3.1.2.1.2.1. Family Importance Value Indices (FIVI)
The highest FIVI values for the coastal slope are 53.81 for the Moraceae (117 indivs.; 10
spp.), 27.08 for the Sapotaceae (34 indivs.; 8 spp.), 23.11 for the Clusiaceae (57 indivs.; 5
spp.), 22.37 for the Meliaceae (42 indivs.; 5 spp.), 16.92 for the Myristicaceae (56 indivs.; 5
spp.), 14.83 for the Burseraceae (37 indivs.; 5 spp.), 10.56 for the Fabaceae-Caes. (23
indivs.; 3 spp.), 10.03 for the Olacaceae (24 indivs.; 4 spp.), 9.39 for the Lauraceae (14
indivs.; 6 spp.) and 8.39 for the Tiliaceae (16 indivs.; 4 spp.).
The total of these 10 families (420 indivs.) is 196.49. The FIVI for the remaining 29 families
(168 indivs.) is 103.51 (Tab. 3.8).
Family
Moraceae
Sapotaceae
Clusiaceae
Meliaceae
Myristicaceae
Burseraceae
Fabaceae-Caes.
Olacaceae
Lauraceae
Tiliaceae
the other 29 families
Total of the 39 families
No. of indivs.
117
34
57
42
56
37
23
24
14
16
168
588
No. of spp.
10
8
5
5
5
5
3
4
6
4
53
108
FIVI
53.81
27.08
23.11
22.37
16.92
14.83
10.56
10.03
9.39
8.39
103.51
300.00
Tab. 3.8. FIVI – Coastal slope
3.1.2.1.2.2. Relative Diversity
The highest relative diversity values are 9.26 for the Moraceae (117 indivs.; 10 spp.), 7.41
Family
Moraceae
Sapotaceae
Lauraceae
Burseraceae
Clusiaceae
Fabaceae-Fab.
Fabaceae-Mim.
Meliaceae
Myristicaceae
Olacaceae
the other 29 families
Total of the 39 families
No. of indivs.
117
34
14
37
57
10
6
42
56
24
191
588
No. of spp.
10
8
6
5
5
5
5
5
5
4
50
108
Rel. diversity [%]
9.26
7.41
5.55
4.63
4.63
4.63
4.63
4.63
4.63
3.70
46.30
100.00
Tab. 3.9. Relative diversity – Coastal slope
for the Sapotaceae (34 indivs.; 8 spp.), 5.55 for the Lauraceae (14 indivs.; 6 spp.), 4.63 for
the Clusiaceae (57 indivs.; 5 spp.), 4.63 for the Meliaceae (42 indivs.; 5 spp.), 4.63 for the
Myristicaceae (56 indivs.; 5 spp.), 4.63 for the Burseraceae (37 indivs.; 5 spp.), 4.63 for the
Fabaceae-Fab. (10 indivs.; 5 spp.), 4.63 for the Fabaceae-Mim. (6 indivs.; 5 spp.) and 3.70
for the Olacaceae (24 indivs.; 4 spp.).
41
3.1. Results – Floristics and Diversity
The total of these10 families (397 indivs.) is 53.70. The relative diversity for the remaining
29 families (191 indivs.) is 46.30 (Tab. 3.9).
3.1.2.1.2.3. Relative Density
The sum of all the relative density values is 100% and relates to the 588 indivs. which
correspond to the criteria.
The highest values for relative density are 19.90 for the Moraceae (117 indivs.; 10 spp.),
9.69 for the Clusiaceae (57 indivs.; 5 spp.), 9.52 for the Myristicaceae (56 indivs.; 5 spp.),
7.14 for the Meliaceae (42 indivs.; 5 spp.), 6.29 for the Burseraceae (37 indivs.; 5 spp.),
5.78 for the Sapotaceae (34 indivs.; 8 spp.), 4.08 for the Olacaceae (24 indivs.; 4 spp.),
3.91 for the Fabaceae-Caes. (23 indivs.; 3 spp.), 3.74 for the Apocynaceae (22 indivs.; 3
spp.) and 2.72 for the Tiliaceae (16 indivs.; 4 spp.).
The total relative density of these 10 families (428 indivs.) is 72.77. The relative density for
the remaining 29 families (160 indivs.) is 27.23 (Tab. 3.10).
Family
Moraceae
Clusiaceae
Myristicaceae
Meliaceae
Burseraceae
Sapotaceae
Olacaceae
Fabaceae-Caes.
Apocynaceae
Tiliaceae
the other 29 families
Total of the 39 families
No. of indivs.
117
57
56
42
37
34
24
23
22
16
160
588
No. of spp.
10
5
5
5
5
8
4
3
3
4
56
108
Rel. density [%]
19.90
9.69
9.52
7.14
6.29
5.78
4.08
3.91
3.74
2.72
27.23
100.00
Tab. 3.10. Relative density – Coastal slope
3.1.2.1.2.4. Relative Dominance
The basal surface area of all 588 indivs. is 43.47 m2 and represents 100% of the relative
dominance. The value of relative dominance shows the families with the largest basal area.
The Moraceae have a relative dominance of 24.65 (10.72 m2), the Sapotaceae 13.88 (6.04
Family
Moraceae
Sapotaceae
Meliaceae
Clusiaceae
Caryocaraceae
Burseraceae
Fabaceae-Caes.
Myristicaceae
Chrysobalanaceae
Anacardiaceae
the other 29 families
Total of the 39 families
No. of indivs.
117
34
42
57
9
37
23
56
7
9
197
588
No. of spp.
10
8
5
5
1
5
3
5
4
1
61
108
Tab. 3.11. Relative dominance – Coastal slope
42
Basal surface in [m2]
10.72
6.04
4.60
3.82
2.06
1.70
1.68
1.20
1.04
1.00
9.61
43.47
Rel. dominance [%]
24.65
13.88
10.59
8.79
4.74
3.91
3.87
2.76
2.39
2.30
22.12
100.00
3.1. Results – Floristics and Diversity
m2), the Meliaceae 10.59 (4.60 m2), the Clusiaceae 8.79 (3.82 m2), the Caryocaraceae 4.74
(2.06 m2), the Burseraceae 3.91 (1.70 m2), the Fabaceae-Caes. 3.87 (1.68 m2), the
Myristicaceae of 2.76 (1.20 m2), the Chrysobalanaceae of 2.39 (1.04 m2) and the
Anacardiaceae of 2.30 (1.00 m2).
The total of these 10 families (391 indivs.) is 77.88 (33.86 m2). The remaining 29 families
(197 indivs.) show a relative dominance of 22.12 (9.61 m2) (Tab. 3.11).
3.1.2.2. Gorge
The research plot behind the Esquinas Rainforest Lodge La Gamba comprises 482 indivs.
of 121 spp. 440 are trees, 31 are palms and 10 are lianas. There is one ground-rooted
hemi-epiphyte with d.b.h. ≥ 10 cm (Tab. 3.1).
Fig. 3.5. Number of indivs. per sp. – Gorge
Simpson´s index gives the diversity value D = 0.027 (1-D = 0.973), the Shannon-Weaver
index gives the value H’ = 4.122, the Evenness index gives the value J’(E) = 0.859 and the
α-Index is 51.905. The mean number of indivs. per species (sp.) is 3.980.
On family level, the indivs. represent 46 families from 25 orders. The Shannon-Weaver
diversity index calculated for families gives H’ = 1.551 and the Evenness index gives E´ =
0.933. The mean number of species (spp.) per family is 2.630.
The spp. with the highest number of indivs. are Tetrathylacium macrophyllum
(Flacourtiaceae) with 41 indivs., Rinorea dasyadena (Violaceae) with 37 indivs. and
Cecropia obtusifolia (Cecropiaceae) with 32 indivs. Fifty-nine spp. are each represented by
only one indiv. (Fig. 3.5).
43
3.1. Results – Floristics and Diversity
3.1.2.2.1. Importance Value Index (IVI)
The highest IVI values are 17.48 for Dussia discolor (15 indivs.; Fabaceae-Fab.), 15.42 for
Tetrathylacium macrophyllum (41 indivs.; Flacourtiaceae), 15.16 for Mortoniodendron
anisophyllum (12 indivs.; Tiliaceae), 14.96 for Rinorea dasyadena (37 indivs.; Violaceae),
11.54 for Cecropia obtusifolia (32 indivs.; Cecropiaceae), 8.53 for Calatola costaricensis
(19 indivs.; Icacinaceae), 8.24 for Perrottetia sessiliflora (11 indivs.; Celastraceae),8.00 for
Billia colombiana (3 indivs.; Hippocastanaceae), 7.28 for Dendropanax sessiliflorus (13
indivs.; Araliaceae) and 6.91 for Carapa guianensis (6 indivs.; Meliaceae).
The total of these 10 spp. (189 indivs.) is 113.52. The IVI for the remaining 111 spp. (293
indivs.) is 186.48 (Tab. 3.12).
Species
Dussia discolor
Tetrathylacium macrophyllum
Mortoniodendron anisophyllum
Rinorea dasyadena
Cecropia obtusifolia
Calatola costaricensis
Perrottetia sessiliflora
Billia colombiana
Dendropanax sessiliflorus
Carapa guianensis
the other 111
Total of the 121
Family
Fabaceae-Fab.
Flacourtiaceae
Tiliaceae
Violaceae
Cecropiaceae
Icacinaceae
Celastraceae
Hippocastanaceae
Araliaceae
Meliaceae
No. of indivs.
15
41
12
37
32
19
11
3
13
6
293
482
IVI
17.48
15.42
15.16
14.96
11.54
8.53
8.24
8.00
7.28
6.91
186.48
300.00
Tab. 3.12. IVI – Gorge
3.1.2.2.1.1. Relative Density
The relative density shows the number of indivs. of a certain species in a research plot.
The sum of all relative density values is 100%, which represents all the 482 plants.
Table 3.13. shows the 10 spp. with the highest relative density of the gorge research plot.
The highest values for relative density are 8.51 for Tetrathylacium macrophyllum (41
indivs.; Flacourtiaceae), 7.68 for Rinorea dasyadena (37 indivs.; Violaceae), 6.64 for
Species
Tetrathylacium macrophyllum
Rinorea dasyadena
Cecropia obtusifolia
Calatola costaricensis
Iriartea deltoidea
Dussia discolor
Dendropanax sessiliflorus
Mortoniodendron anisophyllum
Welfia regia
Alsophila firma
the other 111
Total of the 121
Family
Flacourtiaceae
Violaceae
Cecropiaceae
Icacinaceae
Arecaceae
Fabaceae-Fab.
Araliaceae
Tiliaceae
Arecaceae
Cyatheaceae
Tab. 3.13. Relative density – Gorge
44
No. of indivs.
41
37
32
19
16
15
13
12
12
11
274
482
Rel. density [%]
8.51
7.68
6.64
3.94
3.32
3.11
2.70
2.49
2.49
2.28
56.84
100.00
3.1. Results – Floristics and Diversity
Cecropia obtusifolia (32 indivs.; Cecropiaceae), 3.94 for Calatola costaricensis (19 indivs.;
Icacinaceae), 3.32 for Iriartea deltoidea (16 indivs.; Arecaceae), 3.11 for Dussia discolor
(15 indivs.; Fabaceae-Fab.), 2.70 for Dendropanax sessiliflorus (13 indivs.; Araliaceae),
2.49 for Mortoniodendron anisophyllum (12 indivs.; Tiliaceae), 2.49 for Welfia regia (12
indivs.; Arecaceae) and 2.28 for Alsophila firma (11 indivs.; Cyatheaceae).
The total of these 10 spp. (208 indivs.) is 43.16. The relative density for the remaining 111
spp. (274 indivs.) is 56.84 (Tab. 3.13).
3.1.2.2.1.2. Relative Frequency
The highest values for relative frequency are 3.52 for Rinorea dasyadena (37 indivs. in 10
subplots; Violaceae), 3.52 for Tetrathylacium macrophyllum (41 indivs. in 10 subplots;
Flacourtiaceae), 2.82 for Cecropia obtusifolia (32 indivs. in 8 subplots; Cecropiaceae), 2.82
for Iriartea deltoidea (16 indivs. in 8 subplots; Arecaceae), 2.46 for Alsophila firma (11
indivs. in 7 subplots; Cyatheaceae), 2.46 for Castilla tunu (10 indivs. in 7 subplots;
Moraceae), 2.46 for Dussia discolor (15 indivs. in 7 subplots; Fabaceae-Fab.), 2.46 for
Mortoniodendron anisophyllum (12 indivs. in 7 subplots; Tiliaceae), 2.46 for Perrottetia
sessiliflora (11 indivs. in 7 subplots; Celastraceae) and 2.11 Calatola costaricensis (19
indivs. in 6 subplots; Icacinaceae).
The total of these 10 spp. (204 indivs.) is 27.09. The relative frequency for the remaining
111 spp. (278 indivs.) is 72.91 (Tab. 3.14).
Species
Family
Rinorea dasyadena
Tetrathylacium macrophyllum
Cecropia obtusifolia
Iriartea deltoidea
Alsophila firma
Castilla tunu
Dussia discolor
Mortoniodendron anisophyllum
Perrottetia sessiliflora
Calatola costaricensis
the other 111
Total of the 121
Violaceae
Flacourtiaceae
Cecropiaceae
Arecaceae
Cyatheaceae
Moraceae
Fabaceae-Fab.
Tiliaceae
Celastraceae
Icacinaceae
No. of
indivs.
37
41
32
16
11
10
15
12
11
19
278
482
No. of subplots.
Rel. frequency [%]
10
10
8
8
7
7
7
7
7
6
3.52
3.52
2.82
2.82
2.46
2.46
2.46
2.46
2.46
2.11
72.91
100.00
Tab. 3.14. Relative frequency – Gorge
3.1.2.2.1.3. Relative Dominance
The basal surface area of all 482 indivs. is 36.18 m2 and represents 100% of the relative
dominance. The value of relative dominance shows the spp. with the largest basal area.
Dussia discolor (Fabaceae-Fab.) has a relative dominance of 11.90 (4.30 m2),
Mortoniodendron
(Tiliaceae)
anisophyllum
2
10.02
(3.69
m2),
Billia
colombiana
(Hippocastanaceae) 6.67 (2.41 m ), Carapa guianensis (Meliaceae) 4.25 (1.54 m2),
45
3.1. Results – Floristics and Diversity
Species
Family
Dussia discolor
Mortoniodendron anisophyllum
Billia colombiana
Carapa guianensis
Rinorea dasyadena
Terminalia bucidoides
Perrottetia sessiliflora
Tetrathylacium macrophyllum
Calatola costaricensis
Dendropanax sessiliflorus
the other 111
Total of the 121
Fabaceae-Fab.
Tiliaceae
Hippocastanaceae
Meliaceae
Violaceae
Combretaceae
Celastraceae
Flacourtiaceae
Icacinaceae
Araliaceae
No. of
indivs.
15
12
3
6
37
4
11
41
19
13
321
482
Rel.
dominance [%]
11.90
10.02
6.67
4.25
3.76
3.71
3.49
3.39
2.48
2.47
47.86
100.00
Basal surface
2
area [m ]
4.30
3.69
2.41
1.54
1.36
1.34
1.26
1.23
0.90
0.89
17.26
36.18
Tab. 3.15. Relative dominance – Gorge
Rinorea dasyadena (Violaceae) 3.76 (1.36 m2), Terminalia bucidoides (Combretaceae)
3.71 (1.34 m2), Perrottetia sessiliflora (Celastraceae) 3.49 (1.26 m2), Tetrathylacium
macrophyllum (Flacourtiaceae) of 3.39 (1.23 m2), Calatola costaricensis (Icacinaceae) of
2.48 (0.90 m2) and Dendropanax sessiliflorus (Araliaceae) of 2.47 (0.89 m2).
The total of these 10 spp. (161 indivs.) is 52.14 (18.92 m2). The remaining 111 spp. (321
indivs.) have a relative dominance of 47.86 (17.26 m2) (Tab. 3.15).
3.1.2.2.1.4. Absolute Frequency
The highest values for absolute frequency are 100% for Tetrathylacium macrophyllum (41
indivs.; Flacourtiaceae), 100% for Rinorea dasyadena (37 indivs.; Moraceae), 80% for
Cecropia obtusifolia ( 32 indivs.; Cecropiaceae), 80% for Iriartea deltoidea (16 indivs.;
Arecaceae), 70% for Dussia discolor (15 indivs.; Fabaceae-Faboideae), 70% for
Mortoniodendron anisophyllum (12 indivs.; Tiliaceae), 70% for Perrottetia sessiliflora (11
indivs.; Celastraceae), 70% for Castilla tunu (10 indivs.; Moraceae), 70% for Alsophila firma
(11 indivs.; Cyatheaceae) and 60% for Calatola costaricensis (19 indivs.; Icacinaceae)
(Tab. 3.16).
Species
Family
Tetrathylacium macrophyllum
Rinorea dasyadena
Cecropia obtusifolia
Iriartea deltoidea
Dussia discolor
Mortoniodendron anisophyllum
Perrottetia sessiliflora
Castilla tunu
Alsophila firma
Calatola costaricensis
Flacourtiaceae
Violaceae
Cecropiaceae
Arecaceae
Fabaceae-Fab.
Tiliaceae
Celastraceae
Moraceae
Cyatheaceae
Icacinaceae
No. of
indivs.
41
37
32
16
15
12
11
10
11
19
Abundance in 10
subplots
10
10
8
8
7
7
7
7
7
6
Absolute
frequency [%]
100
100
80
80
70
70
70
70
70
60
Tab. 3.16. Absolute frequency – Gorge
A considerable part of the spp. (59 of 121) are represented by only one indiv. and are
therefore present in only one subplot. The absolute frequency for these spp. is 10% (1/10 x
100).
46
3.1. Results – Floristics and Diversity
3.1.2.2.2. Family Diversity – Gorge
121 spp. out of 46 families are found in the gorge. The most diverse families in terms of
spp. are the Moraceae (9 spp., 32 indivs.), the Euphorbiaceae (7 spp., 10 indivs.), the
Fabaceae-Mim. (7 spp., 19 indivs.), the Annonaceae (6 spp., 8 indivs.), the Clusiaceae (6
spp., 15 indivs.), the Tilliaceae (6 spp., 38 indivs.), the Fabaceae-Fab. ( 5 spp., 19 indivs.),
the Flacourtiaceae ( 5 spp., 48 indivs.) and the Rubiaceae ( 5 spp., 10 indivs.).
Fig. 3.6. Family diversity – Gorge
The Melastomataceae (4 indivs.) and the Meliaceae (18 indivs.) consist of 4 spp. and the
Arecaceae (31 indivs.), Bombacaceae (6 indivs.), Boraginaceae (12 indivs.), Lauraceae (4
indivs.), Myristicaceae (8 indivs.) and Violaceae (44 indivs.) of 3 spp. (Fig. 3.6).
3.1.2.2.2.1. Family Importance Value Indices (FIVI)
The highest FIVI values are 27.01 for the Tiliaceae (38 indivs.; 6 spp.), 20.62 for the
Fabaceae-Fab. (19 indivs.; 5 spp.), 20.16 for the Moraceae (32 indivs.; 9 spp.), 17.89 for
the Flacourtiaceae (48 indivs.; 5 spp.), 15.92 for the Violaceae (44 indivs.; 3 spp.), 13.14
for the Meliaceae (18 indivs.; 4 spp.), 12.44 for the Fabaceae-Mim. (19 indivs.; 7 spp.),
11.44 for the Cecropiaceae (36 indivs.; 2 spp.), 11.05 for the Euphorbiaceae (10 indivs.; 7
spp.) and 10.45 for the Arecaceae (31 indivs.; 3 spp.). The total of these 10 families (295
indivs.) is 160.12. The FIVI for the remaining 36 families (187 indivs.) is 139.88 (Tab. 3.17).
47
3.1. Results – Floristics and Diversity
Family
Tiliaceae
Fabaceae-Fab.
Moraceae
Flacourtiaceae
Violaceae
Meliaceae
Fabaceae-Mim.
Cecropiaceae
Euphorbiaceae
Arecaceae
the other 36 families
Total of the 46 families
No. of indivs.
38
19
32
48
44
18
19
36
10
31
187
482
No. of spp.
6
5
9
5
3
4
7
2
7
3
70
121
FIVI
27.01
20.62
20.16
17.89
15.92
13.14
12.44
11.44
11.05
10.45
139.88
300.00
Tab. 3.17. FIVI – Gorge
3.1.2.2.2.2. Relative Diversity
The highest relative diversity values are 7.44 for the Moraceae (32 indivs.; 9 spp.), 5.78 for
the Euphorbiaceae (10 indivs.; 7 spp.), 5.78 for the Fabaceae-Mim. (19 indivs.; 7 spp.),
4.96 for the Annonaceae (8 indivs.; 6 spp.), 4.96 for the Clusiaceae (15 indivs.; 6 spp.),
4.96 for the Tiliaceae (38 indivs.; 6 spp.), 4.13 for the Fabaceae-Fab. (19 indivs.; 5 spp.),
4.13 for the Flacourtiaceae (48 indivs.; 5 spp.), 4.13 for the Rubiaceae (10 indivs.; 5 spp.)
and 3.31 for the Meliaceae (18 indivs.; 4 spp.).
The total of these 10 families (217 indivs.) is 49.58. The relative diversity for the remaining
36 families (265 indivs.) is 50.42 (Tab. 3.18).
In total 27 families occur with more than one species and 19 families with just one species.
Family
Moraceae
Euphorbiaceae
Fabaceae-Mim.
Annonaceae
Clusiaceae
Tiliaceae
Fabaceae-Fab.
Flacourtiaceae
Rubiaceae
Meliaceae
the other 36 families
Total of the 46 families
No. of indivs.
32
10
19
8
15
38
19
48
10
18
265
482
No. of spp.
9
7
7
6
6
6
5
5
5
4
61
121
Rel. diversity [%]
7.44
5.78
5.78
4.96
4.96
4.96
4.13
4.13
4.13
3.31
50.42
100.00
Tab. 3.18. Relative diversity – Gorge
3.1.2.2.2.3. Relative Density
The highest values for relative density are 9.96 for the Flacourtiaceae (48 indivs.; 5 spp.),
9.13 for the Violaceae (44 indivs.; 3 spp.), 7.88 for the Tiliaceae (38 indivs.; 6 spp.), 7.47
for the Cecropiaceae (36 indivs.; 2 spp.), 6.64 for the Moraceae (32 indivs.; 9 spp.), 6.43
for the Arecaceae (31 indivs.; 3 spp.), 3.94 for the Fabaceae-Fab. (19 indivs.; 5 spp.), 3.94
for the Fabaceae-Mim. (19 indivs.; 7 spp.), 3.94 for the Icacinaceae (19 indivs.; 1 sp.) and
3.73 for the Meliaceae (18 indivs.; 4 spp.).
48
3.1. Results – Floristics and Diversity
The total relative density of these 10 families (304 indivs.) is 63.06. The relative density for
the remaining 36 families (178 indivs.) is 36.94 (Tab. 3.19).
Family
Flacourtiaceae
Violaceae
Tiliaceae
Cecropiaceae
Moraceae
Arecaceae
Fabaceae-Fab.
Fabaceae-Mim.
Icacinaceae
Meliaceae
the other 36 families
Total of the 46 families
No. of indivs.
48
44
38
36
32
31
19
19
19
18
178
482
No. of spp.
5
3
6
2
9
3
5
7
1
4
76
121
Rel. density [%]
9.96
9.13
7.88
7.47
6.64
6.43
3.94
3.94
3.94
3.73
36.94
100.00
Tab. 3.19. Relative density – Gorge
3.1.2.2.2.4. Relative Dominance
The basal surface area of all 482 indivs. is 36.18 m2.
The value of relative dominance shows the size of basal area for each family. The Tiliaceae
show a relative dominance of 14.16 (5.12 m2), the Fabaceae-Fab. 12.54 (4.54 m2), the
Hippocastanaceae 6.67 (2.41 m2), the Meliaceae 6.10 (2.21 m2), the Moraceae 6.08 (2.20
m2), the Violaceae 4.31 (1.56 m2), the Araliaceae 3.93 (1.42 m2), the Flacourtiaceae of 3.80
(1.38 m2), the Combretaceae of 3.71 (1.34 m2) and the Celastraceae of 3.49 (1.26 m2).
The total of these 10 families (235 indivs.) is 64.79 (23.44 m2). The remaining 36 families
(247 indivs.) have a relative dominance of 35.21 (12.73 m2) (Tab. 3.20).
Family
Tiliaceae
Fabaceae-Fab.
Hippocastanaceae
Meliaceae
Moraceae
Violaceae
Araliaceae
Flacourtiaceae
Combretaceae
Celastraceae
the other 36 families
Total of the 46 families
No. of indivs.
38
19
3
18
32
44
18
48
4
11
247
482
No. of spp. Basal surface area [m2]
6
5.12
5
4.54
1
2.41
4
2.21
9
2.20
3
1.56
2
1.42
5
1.38
1
1.34
1
1.26
84
12.74
121
36.18
Rel. dominance [%]
14.16
12.54
6.67
6.10
6.08
4.31
3.93
3.80
3.71
3.49
35.21
100.00
Tab. 3.20. Relative dominance – Gorge
3.1.2.3. Ridge
The research plot on a ridge behind the Esquinas Rainforest Lodge La Gamba comprises
847 indivs. of 179 spp.. 749 are trees, 81 are palms, 11 are lianas and 7 are hemiepiphytes (Tab. 3.1).
49
3.1. Results – Floristics and Diversity
Fig. 3.7. Number of indivs. per sp. – Ridge
Simpson’s index gives the value D = 0.019 (1-D = 0.981); the Shannon-Weaver index gives
the value H’ = 4.483; the Evenness index gives the value J’(E) = 0.862 and the α-Index is
70.490. The mean number of indivs. per sp. is 4.730.
At the family level, the indivs. represent 51 families from 30 orders. The Shannon-Weaver
diversity index calculated for families gives H’ = 1.562 and the Evenness index gives E´ =
0.91. The mean number of spp. per family is 3.48 (Tab. 3.39).
The most abundant spp. are Qualea paraensis (Vochysiaceae) with 50, Welfia regia
(Arecaceae) with 49 and Vochysia ferruginea (Vochysiaceae) with 38 indivs.. 72 spp. are
represented by only one individual (Fig. 3.7).
3.1.2.3.1. Importance Value Index (IVI)
The highest IVI values are 17.26 for Vochysia ferruginea (38 indivs.), 16.96 for Qualea
paraensis (50 indivs.; both Vochysiaceae), 12.32 for Welfia regia (49 indivs.; Arecaceae),
10.17 for Vochysia megalophylla (35 indivs.; Vochysiaceae), 9.07 for Calophyllum
longifolium (18 indivs.; Clusiaceae), 8.81 for Symphonia globulifera (24 indivs.; both
Clusiaceae), 7.71 for Brosimum guianense (28 indivs.; Moraceae), 7.00 for Peltogyne
purpurea (11 indivs.; Fabaceae-Caes..), 6.18 for Aspidosperma spruceanum (15 indivs.;
Apocynaceae) and 6.08 for Croton schiedeanus (25 indivs.; Euphorbiaceae).
The total of these 10 spp. (293 indivs.) is 101.56. The IVI for the remaining 169 spp. (554
indivs.) is 198.44 (Tab. 3.21).
50
3.1. Results – Floristics and Diversity
Species
Vochysia ferruginea
Qualea paraensis
Welfia regia
Vochysia megalophylla
Calophyllum longifolium
Symphonia globulifera
Brosimum guianense
Peltogyne purpurea
Aspidosperma spruceanum
Croton schiedeanus
the other 169
Total of the 179
Family
Vochysiaceae
Vochysiaceae
Arecaceae
Vochysiaceae
Clusiaceae
Clusiaceae
Moraceae
Fabaceae-Caes.
Apocynaceae
Euphorbiaceae
No. of indivs.
38
50
49
35
18
24
28
11
15
25
554
847
IVI
17.26
16.96
12.32
10.17
9.07
8.81
7.71
7.00
6.18
6.08
198.44
300.00
Tab. 3.21. IVI – Ridge
3.1.2.3.1.1. Relative Density
The highest values for relative density are 5.90 for Qualea paraensis (50 indivs.;
Vochysiaceae), 5.78 for Welfia regia (49 indivs.; Arecaceae), 4.48 for Vochysia ferruginea
(38 indivs.; Vochysiaceae), 4.13 for Vochysia megalophylla (35 indivs.; Vochysiaceae),
3.30 for Brosimum guianense (28 indivs.; Moraceae), 2.95 for Croton schiedeanus (25
indivs.; Euphorbiaceae), 2.83 for Symphonia globulifera (24 indivs.; Clusiaceae), 2.36 for
Pausandra trianae (20 indivs.; Euphorbiaceae), 2.36 for Socratea exorrhiza (20 indivs.;
Arecaceae) and 2.24 for Marila laxiflora (19 indivs.; Clusiaceae).
The total of these 10 spp. (308 indivs.) is 36.33. The relative density for the remaining 169
spp. (539 indivs.) is 63.67 (Tab. 3.22).
Species
Qualea paraensis
Welfia regia
Vochysia ferruginea
Vochysia megalophylla
Brosimum guianense
Croton schiedeanus
Symphonia globulifera
Pausandra trianae
Socratea exorrhiza
Marila laxiflora
the other 169
Total of the 179
Family
Vochysiaceae
Arecaceae
Vochysiaceae
Vochysiaceae
Moraceae
Euphorbiaceae
Clusiaceae
Euphorbiaceae
Arecaceae
Clusiaceae
No. of indivs.
50
49
38
35
28
25
24
20
20
19
539
847
Rel. density [%]
5.90
5.78
4.48
4.13
3.30
2.95
2.83
2.36
2.36
2.24
63.67
100.00
Tab. 3.22. Relative density – Ridge
3.1.2.3.1.2. Relative Frequency
The highest values for relative frequency are 3.76 for Qualea paraensis (50 indivs. in 22
subplots; Vochysiaceae), 3.25 for Vochysia ferruginea (38 indivs. in 19 subplots;
Vochysiaceae), 3.25 for Welfia regia (49 indivs. in 19 subplots; Arecaceae), 2.56 for
Brosimum guianense (28 indivs. in 15 subplots; Moraceae), 2.22 for Croton schiedeanus
(25 indivs. in 13 subplots; Euphorbiaceae), 2.22 for Socratea exorrhiza (20 indivs. in 13
51
3.1. Results – Floristics and Diversity
subplots; Arecaceae), 2.22 for Symphonia globulifera (24 indivs. in 13 subplots;
Clusiaceae), 2.05 for Pausandra trianae (20 indivs. in 12 subplots; Euphorbiaceae), 1.71
Peltogyne purpurea (11 indivs. in 10 subplots; Fabaceae-Caes.) and 1.71 for Vochysia
megalophylla (35 indivs. in 10 subplots; Vochysiaceae).
The total of these 10 spp. (300 indivs.) is 24.95. The relative frequency for the remaining
169 spp. (547 indivs.) is 75.05 (Tab. 3.23).
Species
Qualea paraensis
Vochysia ferruginea
Welfia regia
Brosimum guianense
Croton schiedeanus
Socratea exorrhiza
Symphonia globulifera
Pausandra trianae
Peltogyne purpurea
Vochysia megalophylla
the other 169 spp.
total
Family
Vochysiaceae
Vochysiaceae
Arecaceae
Moraceae
Euphorbiaceae
Arecaceae
Clusiaceae
Euphorbiaceae
Fabaceae-Caes.
Vochysiaceae
No. of indivs.
50
38
49
28
25
20
24
20
11
35
547
847
No. of subplots
22
19
19
15
13
13
13
12
10
10
Rel. frequency [%]
3.76
3.25
3.25
2.56
2.22
2.22
2.22
2.05
1.71
1.71
75.05
100.00
Tab. 3.23. Relative frequency – Ridge
3.1.2.3.1.3. Relative Dominance
The basal surface area of all 847 indivs. is 39.81 m2. The value of relative dominance
shows the size of basal area for each spp.. Vochysia ferruginea (Vochysiaceae) has a
Species
Family
Vochysia ferruginea
Qualea paraensis
Calophyllum longifolium
Vochysia megalophylla
Brosimum utile
Peltogyne purpurea
Symphonia globulifera
Welfia regia
Aspidosperma spruceanum
Carapa guianensis
the other 169
Total of the 179
Vochysiaceae
Vochysiaceae
Clusiaceae
Vochysiaceae
Moraceae
Fabaceae-Caes.
Clusiaceae
Arecaceae
Apocynaceae
Meliaceae
No. of
indivs.
38
50
18
35
7
11
24
49
15
5
595
847
Basal surface
2
area [m ]
3.79
2.91
2.15
1.73
1.68
1.59
1.50
1.31
1.14
0.86
21.15
39.81
Rel. dominance
[%]
9.53
7.30
5.41
4.34
4.22
3.99
3.76
3.29
2.87
2.17
53.12
100.00
Tab. 3.24. Relative dominance – Ridge
relative dominance of 9.53 (3.79 m2), Qualea paraensis (Vochysiaceae) 7.30 (2.91 m2),
Calophyllum
longifolium
(Clusiaceae)
5.41
(2.15
m2),
Vochysia
megalophylla
(Vochysiaceae) 4.34 (1.73 m2), Brosimum utile (Moraceae) 4.22 (1.68 m2), Peltogyne
purpurea (Fabaceae-Caes.) 3.99 (1.59 m2), Symphonia globulifera (Clusiaceae) 3.76 (1.50
m2), Welfia regia (Araceae) of 3.29 (1.31 m2), Aspidosperma spruceanum (Apocynaceae)
2.87 (1.14 m2) and Carapa guianensis (Meliaceae) of 2.17 (0.86 m2).The total of these 10
52
3.1. Results – Floristics and Diversity
spp. (252 indivs.) is 46.88 (18.66 m2). The remaining 169 spp. (595 indivs.) have a relative
dominance of 53.12 (21.15 m2) (Tab. 3.24).
3.1.2.3.1.4. Absolute Frequency
The highest values for the absolute frequency are 88% for Qualea paraensis (50 indivs.;
Vochysiaceae), 76% for Vochysia ferruginea ( 38 indivs.; Vochysiaceae), 76% for Welfia
regia (49 indivs.; Arecaceae), 60% for Brosimum guianense (28 indivs.; Moraceae), 52%
for Croton schiedeanus (25 indivs.; Euphorbiaceae), 52% for Socratea exorrhiza (20
indivs.; Arecaceae), 52% for Symphonia globulifera (24 indivs.; Clusiaceae), 48% for
Pausandra trianae (20 indivs.; Euphorbiaceae), 40% for Marila laxiflora (19 indivs.;
Clusiaceae), 40% for Peltogyne purpurea (11 indivs.; Fabaceae-Caes.) and 40% for
Species
Family
Qualea paraensis
Vochysia ferruginea
Welfia regia
Brosimum guianense
Croton schiedeanus
Socratea exorrhiza
Symphonia globulifera
Pausandra trianae
Marila laxiflora
Peltogyne purpurea
Vochysia megalophylla
Vochysiaceae
Vochysiaceae
Arecaceae
Moraceae
Euphorbiaceae
Arecaceae
Clusiaceae
Euphorbiaceae
Clusiaceae
Fabaceae-Caes.
Vochysiaceae
No. of
indivs.
50
38
49
28
25
20
24
20
19
11
35
No. in 25
subplots
22
19
19
15
13
13
13
12
10
10
10
Absolute
frequency [%]
88
76
76
60
52
52
52
48
40
40
40
Tab. 3.25. Absolute frequency – Ridge
Vochysia megalophylla (35 indivs.; Vochysiaceae) (Tab. 3.25).
A considerable number of spp. (72 out of 179) are represented by only one indiv. and are
therefore present in only one subplot. The absolute frequency for these spp. is 4% (1/25 x
100).
3.1.2.3.2. Family Diversity
The most diverse families in terms of spp. in the gorge are the Fabaceae-Mim. (14 spp., 29
indivs.), the Moraceae (14 spp., 60 indivs.), the Clusiaceae (13 spp., 79 indivs.), the
Chrysobalanaceae (9 spp., 20 indivs.), the Burseraceae (8 spp., 31 indivs.), the Lauraceae
(6 spp., 9 indivs.), the Melastomataceae (6 spp., 14 indivs.), the Myristicaceae (6 spp., 41
indivs.), the Arecaceae (5 spp., 81 indivs.), the Meliaceae (5 spp., 22 indivs.), the
Myrtaceae (5 spp., 12 indivs.), the Rubiaceae (5 spp., 22 indivs.), the Sapotaceae (5 spp.,
7 indivs.) and the Tilliaceae (5 spp., 25 indivs.) (Fig. 3.8).
53
3.1. Results – Floristics and Diversity
Fig. 3.8. Family diversity – Ridge
3.1.2.3.2.1. Family Importance Value Indices (FIVI)
The highest FIVI values are 38.66 for the Vochysiaceae (124 indivs.; 4 spp.), 29.04 for the
Clusiaceae (79 indivs.; 13 spp.), 23.32 for the Moraceae (60 indivs.; 14 spp.), 16.65 for the
Arecaceae (81 indivs.; 5 spp.), 13.88 for the Fabaceae-Mim. (29 indivs.; 14 spp.), 12.06 for
the Myristicaceae (41 indivs.; 6 spp.), 11.25 for the Fabaceae-Caes. (24 indivs.; 4 spp.),
11.21 for the Burseraceae (31 indivs.; 8 spp.), 11.02 for the Chrysobalanaceae (20 indivs.;
Family
Vochysiaceae
Clusiaceae
Moraceae
Arecaceae
Fabaceae-Mim.
Myristicaceae
Fabaceae-Caes.
Burseraceae
Chrysobalanaceae
Euphorbiaceae
the other 41 families
Total of the 51 families
No. of indivs.
124
79
60
81
29
41
24
31
20
54
304
847
No. of spp.
FIVI
4
13
14
5
14
6
4
8
9
3
99
179
38.66
29.04
23.32
16.65
13.88
12.06
11.25
11.21
11.02
10.54
122.37
300.00
Tab. 3.26. FIVI – Ridge
9 spp.) and 10.54 for the Euphorbiaceae (54 indivs.; 3 spp.).
The total of these 10 families (543 indivs.) is 177.63. The FIVI for the remaining 41 families
(304 indivs.) is 122.37 (Tab. 3.26).
54
3.1. Results – Floristics and Diversity
3.1.2.3.2.2. Relative Diversity
The highest relative diversity values are 7.73 for the Fabaceae-Mim. (29 indivs.; 14 spp.),
7.73 for the Moraceae (60 indivs.; 14 spp.), 7.18 for the Clusiaceae (79 indivs.; 13 spp.),
4.97 for the Chrysobalanaceae (20 indivs.; 9 spp.), 4.42 for the Burseraceae (31 indivs.; 8
spp.), 4.42 for the Lauraceae (9 indivs.; 6 spp.), 3.31 for the Melastomataceae (14 indivs.;
6 spp.), 3.31 for the Myristicaceae (41 indivs.; 6 spp.), 2.76 for the Arecaceae (81 indivs.; 5
spp.), 2.76 for the Meliaceae (22 indivs.; 5 spp.), 2.76 for the Myrtaceae (12 indivs.; 5 spp.),
2.76 for the Rubiaceae (22 indivs.; 5 spp.), 2.76 for the Sapotaceae (7 indivs.; 5 spp.) and
2.76 for the Tiliaceae (25 indivs.; 5 spp.).
The total relative diversity of these 14 families (452 indivs.) is 59.63. The relative diversity
for the remaining 37 families (395 indivs.) is 40.37. In total 30 families occur with more than
one species and 21 families with just one species (Tab. 3.27)
Family
Fabaceae-Mim.
Moraceae
Clusiaceae
Chrysobalanaceae
Burseraceae
Lauraceae
Melastomataceae
Myristicaceae
Arecaceae
Meliaceae
Myrtaceae
Rubiaceae
Sapotaceae
Tiliaceae
the other 37 families
Total of the 51 families
No. of indivs.
29
60
79
20
31
9
14
41
81
22
12
22
7
25
395
847
No. of spp.
14
14
13
9
8
6
6
6
5
5
5
5
5
5
73
179
Rel. diversity [%]
7.73
7.73
7.18
4.97
4.42
3.31
3.31
3.31
2.76
2.76
2.76
2.76
2.76
2.76
40.37
100.00
Tab. 3.27. Relative diversity – Ridge
3.1.2.3.2.3. Relative Density
The highest values for relative density are 14.62 for the Vochysiaceae (124 indivs.; 4 spp.),
9.55 for the Arecaceae (81 indivs.; 5 spp.), 9.32 for the Clusiaceae (79 indivs.; 13 spp.),
Family
Vochysiaceae
Arecaceae
Clusiaceae
Moraceae
Euphorbiaceae
Myristicaceae
Burseraceae
Fabaceae-Mim.
Tiliaceae
Fabaceae-Caes.
the other 41 families
Total of the 51 families
No. of indivs.
124
81
79
60
54
41
31
29
25
24
299
847
No. of spp.
4
5
13
14
3
6
8
14
5
4
103
179
Tab. 3.28. Relative density – Ridge
55
Rel. density [%]
14.62
9.55
9.32
7.07
6.37
4.83
3.66
3.42
2.95
2.83
35.38
100.00
3.1. Results – Floristics and Diversity
7.07 for the Moraceae (60 indivs.; 14 spp.), 6.37 for the Euphorbiaceae (54 indivs.; 3 spp.),
4.83 for the Myristicaceae (41 indivs.; 6 spp.), 3.66 for the Burseraceae (31 indivs.; 8 spp.),
3.42 for the Fabaceae-Mim. (29 indivs.; 14 spp.), 2.95 for the Tiliaceae (25 indivs.; 5 spp.)
and 2.83 for the Fabaceae-Caes. (24 indivs.; 4 spp.)
The total relative density of these 10 families (548 indivs.) is 64.62. The relative density for
the remaining 41 families (299 indivs.) is 35.38 (Tab. 3.28).
3.1.2.3.2.4. Relative Dominance
The basal surface area of all 847 indivs. is 39.81 m2. The Vochysiaceae show a relative
Family
No. of indivs.
Vochysiaceae
Clusiaceae
Moraceae
Fabaceae-Caes.
Arecaceae
Meliaceae
Myristicaceae
Chrysobalanaceae
Burseraceae
Apocynaceae
the other 41 families
Total of the 51 families
No. of spp.
124
79
60
24
81
22
41
20
31
17
348
847
4
13
14
4
5
5
6
9
8
2
109
179
Basal surface
area [m2]
8.69
4.99
3.39
2.47
1.73
1.57
1.56
1.47
1.25
1.20
11.49
39.81
Rel. dominance [%]
21.83
12.54
8.51
6.21
4.34
3.95
3.91
3.69
3.13
3.02
28.87
100.00
Tab. 3.29. Relative dominance – Ridge
dominance of 21.83 (8.69 m2), Clusiaceae 12.54 (4.99 m2), Moraceae 8.51 (3.39 m2),
Fabaceae-Caes. 6.21 (2.47 m2), Arecaceae 4.34 (1.73 m2), Meliaceae 3.95 (1.57 m2),
Myristicaceae 3.91 (1.56 m2), Chrysobalanaceae of 3.69 (1.47 m2), Burseraceae of 3.13
(1.25 m2) and Apocynaceae of 3.02 (1.20 m2).
The total relative dominance of these 10 families (499 indivs.) is 71.13 (28.32 m2). The
remaining 41 families (348 indivs.) have a relative dominance of 28.87 (11.49 m2) (Tab.
3.29).
3.1.2.4. Inland Slope
The research plot on a slope behind the Tropenstation La Gamba comprises 527 indivs. of
133 spp.; 375 are trees, 142 are palms, 5 are lianas and 5 are hemi-epiphytes (Tab. 3.1).
The diversity measure according to Simpson´s index gives the value D = 0.035 (1D=0.965); the Shannon-Weaver index gives the value H’=4.119; the Evenness index gives
the value J’(E)=0.841 and the α-Index is 57.953. The mean number of indivs. per sp. is
3.930.
On a family level, the indivs. represent 50 families from 27 orders. The Shannon-Weaver
diversity index calculated for families is H’=1.556 and the Evenness index is E´= 0.916. The
56
3.1. Results – Floristics and Diversity
mean number of spp. per family is 2.680. Sixty-six spp. are represented by only one indiv.
(Tab. 3.39).
The most abundant spp. are Iriartea deltoidea (Araceae) with 71, Welfia regia (Araceae)
with 47 and Marila laxiflora (Clusiaceae) with 20 indivs.. 58 spp. are represented by only
one indiv. (Fig. 3.9).
Fig. 3.9. Number of indivs. per sp. – Inland Slope
3.1.2.4.1. Importance Value Index (IVI)
The highest IVI values are 20.90 for Iriartea deltoidea (71 indivs.; Arecaceae), 19.01 for
Brosimum utile (15 indivs.; Moraceae), 17.43 for Welfia regia (47 indivs.; Arecaceae), 15.74
for Carapa guianensis (15 indivs.; Meliaceae), 9.31 for Symphonia globulifera (18 indivs.;
Species
Iriartea deltoidea
Brosimum utile
Welfia regia
Carapa guianensis
Symphonia globulifera
Marila laxiflora
Socratea exorrhiza
Humiriastrum diguense
Brosimum lactescens
Dendropanax arboreus
the other 123
Total of the 133
Family
Arecaceae
Moraceae
Arecaceae
Meliaceae
Clusiaceae
Clusiaceae
Arecaceae
Humiriaceae
Moraceae
Araliaceae
No. of indivs.
71
15
47
15
18
20
17
7
11
12
294
527
IVI
20.90
19.01
17.43
15.74
9.31
9.16
7.49
7.18
6.86
6.08
180.84
300.00
Tab. 3.30. IVI – Inland slope
Clusiaceae), 9.16 for Marila laxiflora (20 indivs.; Clusiaceae), 7.49 for Socratea exorrhiza
(17 indivs.; Arecaceae), 7.18 for Humiriastrum diguense (7 indivs.; Humiriaceae), 6.86 for
57
3.1. Results – Floristics and Diversity
Brosimum lactescens (11 indivs.; Moraceae) and 6.08 for Dendropanax arboreus (12
indivs.; Araliaceae). The total of these 10 spp. (233 indivs.) is 119.16. The IVI for the
remaining 123 spp. (294 indivs.) is 180.84 (Tab. 3.30).
3.1.2.4.1.1. Relative Density
The highest values for relative density are 13.47 for Iriartea deltoidea (71 indivs.;
Arecaceae), 8.92 for Welfia regia (47 indivs.; Arecaceae), 3.80 for Marila laxiflora (20
indivs.; Clusiaceae), 3.42 for Symphonia globulifera (18 indivs.; Clusiaceae), 3.23 for
Socratea exorrhiza (17 indivs.; Arecaceae), 2.85 for Brosimum utile (15 indivs.;
Euphorbiaceae), 2.85 for Carapa guianensis (15 indivs.; Meliaceae), 2.47 for Mabea
occidentale (13 indivs.; Euphorbiaceae), 2.28 for Dendropanax arboureus (12 indivs.;
Araliaceae) and 2.09 for Brosimum lactescens (11 indivs.; Moraceae).
The total of these 10 spp. (239 indivs.) is 45.38. The relative density for the remaining 123
spp. (288 indivs.) is 54.62 (Tab. 3.31).
Species
Iriartea deltoidea
Welfia regia
Marila laxiflora
Symphonia globulifera
Socratea exorrhiza
Brosimum utile
Carapa guianensis
Mabea occidentale
Dendropanax arboureus
Brosimum lactescens
the 123 others spp.
Total of the 133
Family
Arecaceae
Arecaceae
Clusiaceae
Clusiaceae
Arecaceae
Moraceae
Meliaceae
Euphorbiaceae
Araliaceae
Moraceae
No. of indivs.
71
47
20
18
17
15
15
13
12
11
288
527
Rel. density [%]
13.47
8.92
3.80
3.42
3.23
2.85
2.85
2.47
2.28
2.09
54.62
100.00
Tab. 3.31. Relative density – Inland slope
3.1.2.4.1.2. Relative Frequency
The highest values for relative frequency are 5.09 for Iriartea deltoidea (71 indivs. in 20
subplots; Arecaceae), 4.82 for Welfia regia (47 indivs. in 19 subplots; Arecaceae), 3.55 for
Socratea exorrhiza (17 indivs. in 14 subplots; Arecaceae), 3.30 for Symphonia globulifera
(18 indivs. in 13 subplots; Clusiaceae), 3.05 for Marila laxiflora (20 indivs. in 12 subplots;
Clusiaceae), 2.54 for Brosimum utile (15 indivs. in 10 subplots; Moraceae), 2.54 for Carapa
guianensis (15 indivs. in 10 subplots; Meliaceae), 2.54 for Brosimum lactescens (11 indivs.
in 10 subplots; Moraceae), 2.29 for Otoba novogranatensi (10 indivs. in 9 subplots;
Myristicaceae) and 2.29 for Mabea occidentale (13 indivs. in 9 subplots; Euphorbiaceae).
The total of these 10 spp. (237 indivs.) is 32.00. The relative frequency for the remaining
123 spp. (290 indivs.) is 68.00 (Tab. 3.32).
58
3.1. Results – Floristics and Diversity
Species
Family
No. of indivs.
Iriartea deltoidea
Welfia regia
Socratea exorrhiza
Symphonia globulifera
Marila laxiflora
Brosimum lactescens
Brosimum utile
Carapa guianensis
Otoba novogranatensi
Mabea occidentale
the other 123
Total of the 133
Arecaceae
Arecaceae
Arecaceae
Clusiaceae
Clusiaceae
Moraceae
Moraceae
Meliaceae
Myristicaceae
Euphorbiaceae
71
47
17
18
20
11
15
15
10
13
290
527
No. of
subplots
20
19
14
13
12
10
10
10
9
9
Rel. frequency
[%]
5.08
4.82
3.55
3.30
3.05
2.54
2.54
2.54
2.29
2.29
68.00
100.00
Tab. 3.32. Relative frequency – Inland slope
3.1.2.4.1.3. Relative Dominance
The basal surface area of all 527 indivs. is 35.53 m2.
The value of relative dominance shows the basal area of each spp.. Brosimum utile
(Moraceae) has a relative dominance of 13.59 (4.84 m2), Carapa guianensis (Meliaceae)
10.61 (3.77 m2), Humiriastrum diguense (Humiriaceae) 4.06 (1.44 m2), Welfia regia
(Arecaceae) 3.69 (1.31 m2), Vochysia megalophylla (Vochysiaceaea) 3.45 (1.23 m2),
Bombacopsis sessilis (Bombacaceae) 3.11 (1.11 m2), Symphonia globulifera (Clusiaceae)
2.59 (0.92 m2), Byrsonima crispa (Malphigiaceae) of 2.51 (0.89 m2), Iriartea deltoidea
(Arecaceae) 2.36 (0.84 m2) and Marila laxiflora (Clusiaceae) of 2.31 (0.82 m2).
The total of the 10 spp. (206 indivs.) is 48.28 (17.17 m2). The remaining 123 spp. (321
indivs.) have a relative dominance of 51.72 (18.36 m2) (Tab. 3.33).
Species
Brosimum utile
Carapa guianensis
Humiriastrum diguense
Welfia regia
Vochysia megalophylla
Bombacopsis sessilis
Symphonia globulifera
Byrsonima crispa
Iriartea deltoidea
Marila laxiflora
the other 123
Total of the 133
Family
Moraceae
Meliaceae
Humiriaceae
Arecaceae
Vochysiaceaea
Bombacaceae
Clusiaceae
Malphigiaceae
Arecaceae
Clusiaceae
No. of indivs.
15
15
7
47
5
5
18
3
71
20
321
527
Basal surface [m2]
4.84
3.77
1.44
1.31
1.23
1.11
0.92
0.89
0.84
0.82
18.36
35.53
Rel. dominance [%]
13.59
10.61
4.06
3.69
3.45
3.11
2.59
2.51
2.36
2.31
51.72
100.00
Tab. 3.33. Relative dominance – Inland slope
3.1.2.4.1.4. Absolute Frequency
The highest values for absolute frequency are 100% for Iriartea deltoidea (71 indivs.;
Arecaceae), 95% for Welfia regia (47 indivs.; Arecaceae), 70% for Socratea exorrhiza (17
indivs.; Aracaceae), 65% for Symphonia globulifera (18 indivs.; Clusiaceae), 60% for Marila
laxiflora (20 indivs.; Clusiaceae), 50% for Brosimum utile (15 indivs.; Moraceae), 50% for
Carapa guianensis (15 indivs.; Meliaceae), 50% for Brosimum lactescens (11 indivs.;
59
3.1. Results – Floristics and Diversity
Moraceae), 45% for Mabea occidentalis (13 indivs.; Euphorbiaceae) and 45% for Otoba
novogranatensis (10 indivs. Myristicaceae).
A considerable number of spp. (66 out of 133) are represented by only one indiv. and are
therefore present in only one subplot. The absolute frequency for these spp. is 5% (1/20 x
100) (Tab. 3.34).
Species
Iriartea deltoidea
Welfia regia
Socratea exorrhiza
Symphonia globulifera
Marila laxiflora
Brosimum utile
Carapa guianensis
Brosimum lactescens
Mabea occidentale
Otoba novogranatensi
Family
Arecaceae
Arecaceae
Arecaceae
Clusiaceae
Clusiaceae
Moraceae
Meliaceae
Moraceae
Euphorbiaceae
Myristicaceae
Abundance in 20 No. of
subplots
indivs.
20
71
19
47
14
17
13
18
12
20
15
15
15
15
11
11
9
13
9
10
Absolute
frequency [%]
100
95
70
65
60
50
50
50
45
45
Tab. 3.34. Absoute frequency – Inland slope
3.1.2.4.2. Family Diversity
The most diverse families in terms of spp. on the inland slope are the Clusiaceae (12 spp.,
55 indivs.), the Fabaceae-Mim. (11 spp., 13 indivs.), the Moraceae (8 spp.ecies, 41
indivs.), the Sapotaceae (7 spp., 20 indivs.), the Arecaceae (5 spp., 142 indivs.), the
Lauraceae (5 spp., 6 indivs.), the Melastomataceae (5 spp., 14 indivs.), the Myristicaceae
(5 spp., 27 indivs.), the Annonaceae (4 spp., 4 indivs.), the Burseraceae (4 spp., 12
indivs.), Chrysobalanaceae (4 spp., 11 indivs.) and the Meliaceae (4 spp., 28 indivs.) (Fig.
3.10).
Fig. 3.10. Family diversity – Inland slope
60
3.1. Results – Floristics and Diversity
3.1.2.4.2.1. Family Importance Value Indices (FIVI)
The highest FIVI values are 37.52 for the Arecaceae (142 indivs.; 5 spp.), 30.68 for the
Moraceae (41 indivs.; 8 spp.), 25.32 for the Clusiaceae (55 indivs.; 12 spp.), 20.93 for the
Meliaceae (28 indivs.; 4 spp.), 18.20 for the Sapotaceae (20 indivs.; 7 spp.), 14.32 for the
Myristicaceae (27 indivs.; 5 spp.), 14.02 for the Fabaceae-Mim. (13 indivs.; 11 spp.), 11.32
for the Vochysiaceae (12 indivs.; 3 spp.), 7.55 for the Burseraceae (12 indivs.; 4 spp.) and
7.31 for the Chrysobalanaceae (11 indivs.; 4 spp.).
The total of these 10 families (361 indivs.) is 187.17. The FIVI for the remaining 39 families
(166 indivs.) is 112.83 (Tab. 3.35).
Family
Arecaceae
Moraceae
Clusiaceae
Meliaceae
Sapotaceae
Myristicaceae
Fabaceae-Mim.
Vochysiaceae
Burseraceae
Chrysobalanaceae
the other 39 families
Total of the 49 families
No. of indivs.
142
41
55
28
20
27
13
12
12
11
166
527
No. of spp.
FIVI
5
8
12
4
7
5
11
3
4
4
70
133
37.52
30.68
25.32
20.93
18.20
14.32
14.02
11.32
7.55
7.31
112.83
300.00
Tab. 3.35. FIVI – Inland slope
3.1.2.4.2.2. Relative Diversity
The highest relative diversity values are 8.57 for the Clusiaceae (55 indivs.; 12 spp.), 8.26
for the Fabaceae-Mimosoideae (13 indivs.; 11 spp.), 6.02 for the Moraceae (41 indivs.; 8
spp.), 5.26 for the Sapotaceae (20 indivs.; 7 spp.), 3.76 for the Lauraceae (6 indivs.; 5
spp.), 3.76 for the Arecaceae (142 indivs.; 5 spp.), 3.76 for the Melastomataceae (14
indivs.; 5 spp.), 3.76 for the Myristicaceae (27 indivs.; 5 spp.), 3.01 for the Annonaceae (4
Family
Clusiaceae
Fabaceae-Mimosoideae
Moraceae
Sapotaceae
Myristicaceae
Melastomataceae
Lauraceae
Arecaceae
Meliaceae
Chrysobalanaceae
Burseraceaeae
Annonaceae
the other 37 families
Total of the 49 families
No. of indivs.
55
13
41
20
27
14
6
142
28
11
12
4
154
527
No. of spp.
12
11
8
7
5
5
5
5
4
4
4
4
59
133
Tab. 3.36. Relative diversity – Inland slope
61
Rel. diversity [%]
9.02
8.27
6.02
5.26
3.76
3.76
3.76
3.76
3.01
3.01
3.01
3.01
44.36
100.00
3.1. Results – Floristics and Diversity
indivs.; 4 spp.), 3.01 for the Burseraceaeae (12 indivs.; 4 spp.), 3.01 for the
Chrysobalanaceae (11 indivs.; 4 spp.) and 3.01 for the Meliaceae (28 indivs.; 4 spp.). The
total of these 12 families (373 indivs.) is 55.64. The relative diversity for the remaining 37
families (154 indivs.) is 44.36. In total 27 families occur with more than one species and 22
families with just one species (Tab. 3.36).
3.1.2.4.2.3. Relative Density
The highest values for relative density are 26.94 for the Arecaceae (142 indivs.; 5 spp.),
10.44 for the Clusiaceae (55 indivs.; 12 spp.), 7.78 for the Moraceae (41 indivs.; 8 spp.),
5.31 for the Meliaceae (28 indivs.; 4 spp.), 5.12 for the Myristicaceae (27 indivs.; 5 spp.),
4.36 for the Sapotaceae (20 indivs.; 7 spp.), 3.23 for the Euphorbiaceae (17 indivs.; 3
spp.), 2.66 for the Melastomataceae (14 indivs.; 5 spp.), 2.47 for the Fabaceae-Mim. (13
indivs.; 11 spp.), 2.28 for the Burseraceae (12 indivs.; 4 spp.), 2.28 for the Vochysiaceae
(12 indivs.; 3 spp.) and 2.28 for the Araliaceae (12 indivs.; 1 spp.).
The total of these 12 families (393 indivs.) is 75.15. The relative density for the remaining
37 families (134 indivs.) is 24.85 (Tab. 3.37).
Family
Arecaceae
Clusiaceae
Moraceae
Meliaceae
Myristicaceae
Sapotaceae
Euphorbiaceae
Melastomataceae
Fabaceae-Mim.
Burseraceae
Vochysiaceae
Araliaceae
the other 38 families
Total of the 50 families
No. of indivs.
142
55
41
28
27
20
17
14
13
12
12
12
134
527
No. of spp.
5
12
8
4
5
7
3
5
11
4
3
1
65
133
Rel. density [%]
26.94
10.44
7.78
5.31
5.12
4.36
3.23
2.66
2.47
2.28
2.28
2.28
24.85
100.00
Tab. 3.37. Relative density – Inland slope
3.1.2.4.2.4. Relative Dominance
The basal surface area of all 527 indivs. is 35.53 m2. The value of the relative dominance
shows the size of basal area of each family. The Moraceae have a relative dominance of
17.19 (6.11 m2), the Meliaceae 12.76 (4.53 m2), the Arecaceae 7.00 (2.49 m2), the
Vochysiaceae 6.90 (2.45 m2), the Clusiaceae 6.31 (2.24 m2), the Myristicaceae 5.63 (2.00
m2), the Sapotaceae 5.26 (1.87 m2), the Humiriaceae of 4.07 (1.45 m2), the Fabaceae-Mim.
of 3.70 (1.31 m2) and the Bombacaceae of 3.12 (1.11 m2). The total of these 10 families
(350 indivs.) is 71.94 (25.56 m2). The remaining 39 families (177 indivs.) have a relative
dominance of 28.06 (9.97 m2) (Tab. 3.38).
62
3.1. Results – Floristics and Diversity
Family
Moraceae
Meliaceae
Arecaceae
Vochysiaceae
Clusiaceae
Myristicaceae
Sapotaceae
Humiriaceae
Fabaceae-Mim.
Bombacaceae
the other 39 families
Total of the 49 families
No. of indivs.
41
28
142
12
55
27
20
7
13
5
177
527
No. of spp.
8
4
5
3
12
5
7
1
11
1
76
133
Basal surface area [m2] Rel. dominance [%]
6.11
17.19
4.53
12.76
2.49
7.00
2.45
6.90
2.24
6.31
2.00
5.63
1.87
5.26
1.45
4.07
1.31
3.70
1.11
3.12
9.97
28.06
35.53
100.00
Tab. 3.38. Relative dominance – Inland slope
3.1.2.5. Survey of Diversity
Tab. 3.39 is a summary of all data relating to plot diversity.
No. of indivs.
No. of spp.
No. of genera
No. of families
Shannon-Weaver H´ Log (Base 2.718)
Shannon-Weaver J´ or Eveness E
Simpson´s Diversity D
Simpson´s Diversity 1/D
Simpson´s Diversity 1-D
Alpha Diversity
Mean no. of indivs. per sp.
Family Shannon-Weaver H´ (Log Base 10)
Family Shannon-Weaver J´ or Eveness E
Mean no. of spp. per family
No. of cases where only one indiv. per sp.
No. of indivs. of the most-represented sp.
Coastal slope Gorge
588
482
108
121
79
90
39
46
4.014
4.122
0.857
0.859
0.028
0.027
35.227
37.21
0.972
0.973
38.82
51.905
5.440
3.980
1.463
1.551
0.926
0.933
2.77
2.63
38
59
50
41
Inland slope
527
133
97
49
4.119
0.841
0.035
28.51
0.965
57.953
3.930
1.556
0.916
2.71
66
71
Tab. 3.39. Summary of data relating to plot diversity
80
70.5
57.9
Value
60
40
51.9
38.8
gorge
Coastal slope
20
0
Sample
Fig. 3.11. Alpha index results
63
inland
ridge
Ridge
847
179
116
51
4.476
0.862
0.019
53.35
0.981
70.490
4.730
1.562
0.910
3.51
72
50
Total
2,444
328
194
69
4.73
108
108
3.1. Results – Floristics and Diversity
Value
Ridge
Gorge
Inland slope
Coastal slope
Sample
Fig. 3.12. Shannon-Weaver indices of all plots
0.04
Value
0.03
0.0351
0.0283
Coastal slope
0.02
0.0268
inland
gorge
0.0187
ridge
0.01
0.00
Sample
Fig. 3.13. Simpson´s index results
3.1.3. Area-species curves of all research plots
The area-species curves show the relationship between the number of spp. and increasing
surface area. The species-area curves of the plots do not level out. This indicates that an
increase in the size of the research area will lead to the discovery of increasing speciesdiversity. Due to the different sizes of the subplots (Gorge: 1.000 m2 (50 x 20 m), Ridge and
Coastal slope: 200 m2 (20 x 10 m), Inland slope: 100 m2 (10 x 10 m)) the distances
between points on the curves are different (Fig. 3.14).
The form of the area-species curves depends on the arrangement of the subplots. If we
unite more subplots we will get a means curve (Fig. 3.15).
64
3.1. Results – Floristics and Diversity
With the log curve we can calculate the increasing diversity of trees on a given area (Fig.
3.16). At the beginning the curve is a root function, and it starts to be a log curve between
470 m2 (ridge) and 2,080 m2 (gorge).
Inland slope: y=36.1353 * x ^.867537 (x < 550 m2)
y = 71.3738·Ln (0.545297·x + 1.05242) (550 m2 ≤ x ≤ 10.000 m2)
Gorge: y= 28.2999* x^0.700831 (x < 2.080 m2)
y = 87.6568·Ln(0.285829·x + 1.12044) (2.080 m2 ≤ x ≤ 10.000 m2)
Coastal slope: y= 35.2614* x ^ 0.785024 (x < 650 m2)
y = 48.2739·Ln(0,824766·x + 1.14724) (650 sqm. ≤ x ≤ 10.000 m2)
Ridge: y=50.7017* x ^ 0.849483 (x < 470 m2)
y = 96.9397·Ln(0.54422·x + 1.06177) (470sqm. ≤ x ≤ 10.000 m2)
200
200
180
180
160
160
Ridge
Gorge
140
120
100
140
120
100
80
80
60
40
20
60
40
20
0
0
200
180
200
160
140
120
160
100
100
80
60
80
180
140
120
60
40
40
20
20
0
0
0
2000
4000
6000
8000
10000
0
2000
4000
6000
Cumulative area [m2]
Cumulative area [m2]
Fig. 3.14. Area-species curves (data points) of all plots
65
8000
10000
3.1. Results – Floristics and Diversity
200
180
Gorge
160
200
180
Ridge
160
140
140
120
120
100
100
80
80
60
60
40
40
20
20
0
0
0
10
20
30
40
50
60
70
80
90
0
100
10
20
30
40
50
60
70
80
90
100
160
200
180
160
140
140
120
120
200
180
100
100
80
60
80
60
40
40
20
20
0
0
0
10
20
30
40
50
60
70
80
90
0
100
2
Cummulative area [100 m ]
10
20
30
40
50
60
70
80
90
100
Cummulative area [100 m2]
Fig: 3.15. Area-species curves (means) of all plots
Gorge
200
180
Ridge
160
140
120
100
80
60
Y=50.7017* x ^ 0.849483 (0<x<0.4)
y = 96.9397·Ln(0.54422·x + 1.06177) (x>0.4)
Y= 28.2999* x^0.700831(0<x<3)
y = 87.6568·Ln(0.285829·x + 1.12044) (x>3)
40
20
0
200
180
160
140
120
100
80
60
Y=36.1353 * x ^ 0.867537 (0<x<0.5)
y = 71.3738· Ln (0.545297·x + 1.05242 (x>0.5)
Y= 35.2614* x ^ 0.785024 (0<x<0.6)
y = 48.2739·Ln(0.824766·x + 1.14724) (x>0.6 )
Cumulative area [1,000 m2]
Cumulative area [1,000 m2]
Fig: 3.16. Area-species curves (log) of all plots
66
40
20
0
3.1. Results – Floristics and Diversity
450
450
Gorge
350
Ridge
350
250
250
150
150
Y= 28.2999* x^0.700831(0<x<3)
y = 87.6568·Ln(0.285829·x + 1.12044) (x>3)
50
0
10
20
30
40
50
60
70
80
90
Y=50.7017* x ^ 0.849483 (0<x<0,4)
50
y = 96.9397·Ln(0.54422·x + 1.06177) (x>0,4)
0
100
10
20
30
40
50
60
70
80
90 100
450
450
350
350
250
250
150
150
Y= 35.2614* x ^ 0.785024 (0<x<3)
y = 48.2739·Ln(0,824766·x + 1.14724) (x>0,6)
50
0
10
20
30
40
50
60
70
80
90
Y=36.1353 * x ^ 0.867537 (0<x<0,5)
50
y = 71.3738· Ln (0.545297·x + 1.05242 (x>0,5))
0
100
10
20
30
40
50
60
70
80
90
100
Cumulative area [1,000 m2]
Cumulative area [1,000 m2]
Fig. 3.17. Asymptotic model of species-area curves of all plots for 10 ha
The results of the calculations for the plots show that we can expect following numbers of
species in ten:
Hectare
Gorge
1
2
3
4
5
6
7
8
9
10
121
169
199
222
240
255
267
279
288
297
Coastal slope
108
139
157
170
181
189
197
203
209
214
Tab. 3.40. Calculated diversity of 10 ha of all sites
67
Inland slope
134
177
204
223
239
251
262
271
279
287
Ridge
179
240
277
303
324
341
356
368
379
389
3.1. Results – Floristics and Diversity
3.1.4. Beta-Diversity: Floristic Similarities and Differences beteween all
Research Plots
16 spp. (c. 5 %) of all 328 spp. occurred in all 4 plots (Tab. 3.41), 44 spp. (c. 13 %) were
found in three, 75 spp. (c. 23 %) in two and 195 (c. 59 %) spp. in just one plot.
Species
Iriartea deltoidea
Family
Arecaceae
Protium panamense
Burseraceae
Cecropia obtusifolia
Cecropiaceae
Chrysochlamys grandifolia
Clusiaceae
Symphonia globulifera
Clusiaceae
Acacia allenii
Fabaceae-Mimosoideae
Humiriastrum diguense
Humiriaceae
Carapa guianensis
Meliaceae
Guarea grandifolia
Meliaceae
Trichilia septentrionalis
Meliaceae
Brosimum lactescens
Moraceae
Brosimum utile
Moraceae
Perebea hispidula
Moraceae
Otoba novogranatensis
Myristicaceae
Virola koschnyi
Myristicaceae
Apeiba tibourbou
Tiliaceae
Tab. 3.41. All species that occur in all four research plots
68
3.1. Results – Floristics and Diversity
3.1.4.1. Bray-Curtis Cluster Analyses
The Bray-Curtis cluster analyses show the similarities and differences between of the floral
compositions of the research plots. The plot in the gorge had the a floral composition most
at variance to the other plots. The two most similar plots (40.61%) in terms of floral
composition were the ridge and the inland slope (Tab. 3.42 and Fig. 3.18).
Step
Clusters
Distance
Similarity
Joined 1
Joined 2
1.00
3.00
59.39
40.61
3.00
4.00
2.00
2.00
73.09
26.91
1.00
3.00
3.00
1.00
79.58
20.42
1.00
2.00
Similarity Matrix
coastal slope
gorge
inland slope
ridge
coastal slope
*
13.46
26.91
26.48
gorge
*
*
20.42
18.66
inland slope
*
*
*
40.61
ridge
*
*
*
*
Tab. 3.42. Bray-Curtis cluster analyses
Gorge
Ridge
Slope - inland
Slope - coast
0
100
50
Similarity [%]
Fig. 3.18. Bray-Curtis cluster diagram
69
3.1. Results – Floristics and Diversity
3.1.4.2. Comparison of the Distribution of Species in each Plot
328 spp. were found in total. Sixteen spp. were present in all plots and 195 spp. were
represented by only one indiv. Fig. 3.19 shows the number of spp. that share different sites
(plots).
In total:
289 spp. are distributed in the gorge, on the inland slope and on the ridge;
291 spp. are distributed in the gorge, on the coastal slope and on the ridge;
267 spp. are distributed on the inland slope, on the coastal slope and in the gorge;
276 spp. are distributed on the inland slope, on the coastal slope and on the ridge.
Gorge
Inland slope
2.8
20.4
(8)
(59)
Coastal slope
Gorge
20.3
14.9
(59)
(43)
(33)
(21)
10.7
17.3
26
Inland slope
(20)
Coastal slope
15.9
25.8
(44)
(69)
9.0
16.7
(24)
(11)
(32)
Inland slope
7.5
4.1
11.0
Ridge
Gorge
(21)
(45)
(93)
ridge
(69)
15.5
32.0
(75)
25.8
(23)
(31)
(50)
7.9
(8)
7.9
11.4
7.3
2.7
2.9
(8)
13.0
(36)
(46)
16.3
(45)
6.9
(19)
28.3
19.9
(78)
(53)
Coasal slope
Ridge
Fig. 3.19. Comparison of the distribution of species in each plot in percentages with absolute numbers
in brackets
70
3.1. Results – Floristics and Diversity
3.1.4.3. Species found in only one plot
All plots contain species that only occur in that one plot. Some of them are typical species
for that habitat (plot).
Gorge: 54 spp. (44.6%); 137 indiv. (28.4%)
The most abundant spp. that occurred only in the gorge were:
Calatola costaricensis with 19 indivs., Perrottetia sessiliflora with 11 indivs., Alsophila firma
with 11 indivs., Fusispermum laxiflorum with 6 indivs., Dendropanax caucanus with 5
indivs., Simira maxonii with 5 indivs. and Myriocarpa longipes with 5 indivs.
Coastal slope: 40 spp. (37%); 167 indiv. (28.4%)
The most abundant spp. that occurred only on the coastal slope were:
Sorocea cufodontissi occur with 43 indivs., Schizolobium parahyba with 16 indivs.,
Heisteria concinna with 15 indivs., Caryocar costaricense with 9 indivs., Manilkara
staminodella with 8 indivs. and Pourouma bicolour with 8 indivs.
Ridge: 63 spp. (35.2%); 138 indiv. (16.3%)
The most abundant spp. that occurred only on the ridge were:
Isertia laevis with 11 indivs., Calyptranthes pallens with 7 indivs., Calophyllum brasiliense
with 7 indivs., Rinorea hummelii with 6 indivs., Eschweilera pittieri (cf.) with 6 indivs., Laetia
procera with 6 indivs., Abarema adenophora with 5 indivs. and Coccoloba lehmannii with 5
indivs.
Inland slope: 38 spp. (28.6%); 69 indiv. (13.1%)
The most abundant spp. that occurred only on the inland slope were:
Sloanea sp., Byrsonima crispa, Chrysophyllum cf. colombianum and Theobroma simiarum,
each with 3 indivs.
71
3.2. Biogeographical Patterns and Affinities
3.2. Biogeogeographical Patterns and Affinities
The geographical and altitudinal distribution of 312 of the 328 identified species in the four
research plots were investigated. The origins of the plant families were also investigated.
Different maps and graphs show the biogeographical connections, similarities, differences
and peculiarities of the tree flora of the investigated plots.
3.2.1. Origins of the Families found in the Research Plots
Familial origin was ascertained from CHIAPPY & al. (2000), GENTRY (1982a) and RAVEN &
AXELROD (1975). About 62% of the 69 families (plus one unidentified) found in the plots are
Amazonian-centred Gondwana families, 15% are Laurasian families, 11% are Northern
Andean-centred Gondwana families, 4% Southern Andean-centred Gondwana families, 1%
are dry-centred elements and 1% are African families. The origin of the remaining 6% are
uncertain (Fig. 3.20).
1% 1%
4%
6%
62%
11%
Dry centered
Africa
Andean-South
Unassigned family
Andean-North
Laurasia
15%
Amazonian
Fig. 3.20. Origins of families
3.2.2. Altitudinal Distribution
WERCKLÉ (1909) described four phytogeographical regions in Costa Rica: (1) Atlantic or
Caribbean region from sea level up to 800 m (A); (2) Pacific region from sea level up to 800
m (A); (3) temperate region from 800 up to 1,500 m (B); and (4) cold region above 1,500 m
(C).
The altitudes of the different research plots in the Esquinas forest are between 100 m and
350 m elevation. 76 of the species, which occur in the plots, can be found in the neotropics
between sea level and 800 m elevation, 140 occur up to 1,500 m and 96 species also
occur in higher regions (Fig. 3.21).
72
3.2. Biogeographical Patterns and Affinities
4,000 m
Cold region
No. of Species
96
1,500 m
Temperate region
140
800 m
96
Pacific and Caribbean region
Fig. 3.21. Altitudinal distribution of the species – life zones according to WERCKLÉ (1909)
HOLDRIGE (1967) stated the upper limits of altitudinal belts to be 1,000 m for tropical
lowlands, at 2,000 m for premontane, at 3,000 m for tropical lower montane and 4,000 m
for tropical montane.
A
B
C
D
lower than 1,000 m (lowland)
between 1,000 and 2,000 m (premontane)
between 2,000 and 3,000 m (lower montane)
over 3,000 m (montane)
130 spp.
146 spp.
33 spp.
3 spp.
Tab. 3.43. Altitudinal distribution of the species according to HOLDRIGE
Montane
1
3
15 11 15 15
33
2
1
1
3,000 m
Lower montane
2,000 m
Premontane
1,000 m
tot
al
39 41 70 58 130
inla
nd
Lowland
No. of Species
4,000 m
Fig. 3.22. Altitudinal distribution of the species – life zones according to Holdrigde (1967)
Of the species which occur in the plots, 130 can be found in the neotropics between sea
level and 1,000 m (lowland), and 146 occur between 1,000 and 2,000 m (premontane). 33
occur between 2,000 m and 3,000 m (lower montane) and 3 species can be found in
regions higher than 3,000 m elevation (montane) (Fig. 3.22, Tab.3.43).
73
3.2. Biogeographical Patterns and Affinities
3.2.3. Geographical Distribution
The following sections show the geographical patterns of the plants and species
investigated in each research plot.
3.2.3.1. Distribution of the Coastal Slope Species
The geographical and altitudinal distribution of 106 species from the coastal slope research
plot were investigated.
3.2.3.1.1. Altitudinal Distribution
The altitude of the coastal parts of the Esquinas forest is ranges from 60 to 150 m
elevation. Forty-one species can be found only in the lowlands (up to 1,000 m), 53 spp.
only in the premontane forests (up to 2,000 m), 11 spp. only in the lower montane forests
(up to 3,000 m) and one species only in the montane forests (higher than 3,000 m) (Fig.
3.22).
3.2.3.1.2. Regional Distribution
The recorded species have a stronger phytogeographical relationship to the south (South
America, Amazon and Chocó) than to the north (Northern Central America): 95 spp. (90%)
also occur in Panama, 77 spp. (73%) in Colombia, 64 spp. (56%) in Ecuador, 59 spp.
(56%) in Peru, 49 spp. (46%) in Brazil, 48 spp. (45%) in Bolivia, 42 spp. (40%) in
Venezuela and 29 spp. (27%) in Guayana.
106
95
59
42
48
68
77
72
49
48
35
29
44
31
do
r
C
ol
om
bi
a
Pa
na
m
C
a
os
ta
R
ic
N
a
ic
ar
ag
ua
H
on
du
ra
s
M
ex
ic
G
o
ua
te
m
al
a
Be
liz
El
Sa e
lv
ad
or
An
til
le
s
ru
Pe
11
Ec
ua
G
uy
an
as
Ve
ne
zu
el
a
Bo
liv
ia
Br
az
il
11
Fig. 3.23. Distribution of the 106 species of the coastal slope in the countries of Central and South
America (absolute figures)
The distribution to the north is much less pronounced: 72 species (68%) also occur in
Nicaragua, 48 spp. (45%) in Honduras, 44 spp. (42%) in Belize, 35 spp. (33%) in Mexico,
31 spp. (29%) in Guatemala and 11 spp. (10%) in El Salvador.
74
3.2. Biogeographical Patterns and Affinities
Eleven spp. (10%) were found on at least one of the Caribbean islands.
The dry forest of the Santa Rosa National Park shares 9 spp. (8%) and the wet lowland
forest near the Panama Canal (Panama Watershed) 70 spp. (66%).
34 spp. (32%) only occur in the Pacific region of Costa Rica. The other 70 species (66%)
also occur in the Caribbean region (Fig. 3.23).
Northern Central America
Mexico, Guatemala, Honduras, El Salvador, Nicaragua
Honduras - disjunct
Nicaragua
Costa Rica
Panama
South America
Fig. 3.24. Distribution of the coastal slope species
Forty-eight of the species in the coastal plot have a wide-ranging distribution. These
species are distributed in Central America (north of Costa Rica) and in South America.
Twenty-three species occur only in Central America. Three of them only occur between
Nicaragua and Panama, six only in Panama and Costa Rica and five only in Costa Rica.
Thirty-four species occur in South America and Central America up to Costa Rica (not in
Nicaragua or farther north). One species only occurs in Honduras and Costa Rica (disjunct
distribution) (Fig. 3.24).
3.2.3.2. Distribution of the Gorge Species
The altitudinal and geographical distribution of 119 species in the research plot located in
the gorge forest were investigated.
75
3.2. Biogeographical Patterns and Affinities
3.2.3.2.1. Altitudinal Distribution
The altitude of the gorge parts of the Esquinas forest ranges from 80 to 150 m elevation.
Thirty-nine species can only be found in the lowlands (up to 1,000 m), 63 spp. in the
premontane forests (up to 2,000 m), 15 spp. in the lower montane forests (up to 3,000 m)
and 2 spp. in the montane forests (higher than 3,000 m) (Fig. 3.22).
3.2.3.2.2. Regional Distribution
The recorded species have a stronger phytogeographical relationship to the south (South
America, Amazon and Chocó) than to the north (Northern Central America): 99 (83%)
species also occur in Panama, 75 spp. (63%) in Colombia, 70 spp. (59%) in Ecuador, 54
spp. (45%) in Peru, 30 spp. (25%) in Brazil, 37 spp. (31%) in Bolivia, 40 (34%) in
Venezuela and 23 spp. (19%) in Guayana.
Distribution to the north is much less pronounced: 75 species (63%) occur in Nicaragua, 46
spp. (39%) in Honduras, 36 spp. (30%) in Belize, 35 spp. (29%) in Mexico, 30 spp. (25%)
in Guatemala and 8 spp. (7%) in El Salvador.
Eleven spp. (9%) were found on at least one of the Caribbean islands.
The dry forest of the Santa Rosa National Park shares 12 spp. (10%) and the wet lowland
forest near the Panama Canal (Panama Watershed) 88 spp. (74%).
Fifty-five spp. (46%) only occur in the Pacific regions of Costa Rica. The other 64 species
(54%) also occur in the Caribbean region (Fig. 3.25).
119
99
70
75
75
54
40
37
46
35
30
30
36
ru
Ec
ua
do
r
C
ol
om
bi
a
Pa
na
m
C
a
os
ta
R
ic
a
N
ic
ar
ag
ua
H
on
du
ra
s
M
ex
ic
G
o
ua
te
m
al
a
Be
liz
El
e
Sa
lv
ad
or
An
til
le
s
11
Pe
Ve
ne
zu
el
a
Bo
liv
ia
Br
az
il
8
Fig. 3.25. Distribution of the 119 species of the gorge in the countries of Central and South America
(absolute figures)
Sixty-nine of the species found in the gorge have a wide-ranging distribution. These
species are distributed in Central America (north of Costa Rica) and in South America.
Twenty-nine species only occur in Central America (not in South America). Four of them
occur between Nicaragua and Panama, ten only in Panama and Costa Rica and eight only
76
3.2. Biogeographical Patterns and Affinities
in Costa Rica. Twenty-one species occur in South America and Central America up to
Costa Rica (not in Nicaragua or farther north). (Fig. 3.26).
Fig. 3.26. Distribution of the gorge species
3.2.3.3. Distribution of the Ridge Species
The altitudinal and geographical distribution of 176 species in the ridge forest research plot
were investigated.
3.2.3.3.1. Altitudinal Distribution
The altitude of the ridge part of the Esquinas forest ranges from 250 to 280 m elevation.
Seventy species can only be found in the lowlands (up to 1,000 m), 90 spp. in the
premontane forests (up to 2,000 m), 15 spp. in the lower montane forests (up to 3,000 m)
and one species in the montane forests (higher than 3,000 m) (Fig. 3.22).
3.2.3.3.2. Regional Distribution
The recorded species have a stronger phytogeographical relationship to the south (South
America, Amazon and Chocó) than to the north (Northern Central America): 148 species
(84%) occur also in Panama, 115 spp. (65%) in Colombia, 96 spp. (55%) in Ecuador, 85
spp. (48%) in Peru, 61 spp. (35%) in Brazil, 60 spp. (34%) in Bolivia, 70 (40%) in
Venezuela and 41 spp. (23%) in Guayana.
77
3.2. Biogeographical Patterns and Affinities
Distribution to the north is much less pronounced: 108 species (61%) occur in Nicaragua,
55 spp. (31%) in Honduras, 51 spp. (29%) in Belize, 44 spp. (25%) in Mexico, 36 spp.
(20%) in Guatemala and 8 spp. (5%) in El Salvador.
Eleven spp. (6%) were found on at least one of the Caribbean islands.
176
148
115
85
70
108
96
61
60
55
44
51
36
ru
Ec
ua
do
r
C
ol
om
bi
a
Pa
na
m
C
a
os
ta
R
ic
a
N
ic
ar
ag
ua
H
on
du
ra
s
M
ex
ic
o
G
ua
te
m
al
a
Be
liz
El
e
Sa
lv
ad
or
An
til
le
s
11
Pe
az
il
Br
Ve
ne
zu
el
a
Bo
liv
ia
8
Fig. 3.27. Distribution of the 176 species of the ridge in the countries of Central and South America
(absolute figures)
Fig. 3.28. Distribution of the ridge species
Nine spp. (5%) are found in the dry forest of the Santa Rosa National Park and 101 spp.
(57%) are found in the wet lowland forest near the Panama Canal (Panama Watershed).
78
3.2. Biogeographical Patterns and Affinities
Fifty-five spp. (31%) only occur in the Pacific regions of Costa Rica. The other 121 species
(69%) also occur in the Caribbean region (Fig. 3.27).
Ninety-one of the species in the ridge plot have a wide-ranging distribution. These species
are distributed in Central America (north of Costa Rica) and in South America. Forty-nine
species only occur in Central America (not in South America). Nine of them only occur
between Nicaragua and Costa Rica, 13 only in Panama and Costa Rica and 11 only in
Costa Rica. Thirty-five species occur in South America and Central America up to Costa
Rica (not in Nicaragua or farther north). One species only occurs in Venezuela and Costa
Rica (disjunct distribution) (Fig. 3.28).
3.2.3.4. Distribution of the Inland Slope Species
The altitudinal and geographical distribution of 123 species from the inland slope research
plot were investigated.
3.2.3.4.1. Altitudinal Distribution
The altitude of the inland part of the Esquinas forest ranges from to 150 m elevation. Fiftyeight species can only be found in the lowlands (up to 1,000 m), 49 spp. in the premontane
forests (up to 2,000 m), 15 spp. in the lower montane forests (up to 3,000 m) and 1 sp. in
the montane forests (higher than 3,000 m) (Fig. 3.22).
3.2.3.4.2. Regional Distribution
The recorded species have a stronger phytogeographical relationship to the south (South
America, Amazon and Chocó) than to the north (Northern Central America): 99 species
(80%) also occur in Panama, 82 spp. (67%) in Colombia, 66 spp. (54%) in Ecuador, 64
spp. (52%) in Peru, 52 spp. (42%) in Brazil, 52 spp. (42%) in Bolivia, 56 (46%) in
Venezuela and 34 spp. (28%) in Guayana.
Distribution to the north is much less pronounced: 77 species (63%) occur in Nicaragua, 46
spp. (37%) in Honduras, 36 spp. (29%) in Belize, 29 spp. (24%) in Mexico, 28 spp. (23%)
in Guatemala and 4 spp. (3%) in El Salvador.
Seven spp. (6%) were recorded on at least one of the Caribbean islands.
Three spp. (2%) occur in the dry forest of the Santa Rosa National Park and 86 spp. (70%)
occur in the wet lowland forest near the Panama Canal (Panama Watershed).
Fourty-nine spp. (40%) only occur in the Pacific regions of Costa Rica. The other 74
species (60%) also occur in the Caribbean region (Fig. 3.29). Sixty-seven of the species in
the inland plot have a wide-ranging distribution. These species are distributed in Central
America (north of Costa Rica) and in South America. Forty-one species only occur in
Central America (not in South America). Eighteen of them only occur between Nicaragua
and Panama, five only in Panama and Costa Rica and five only in Costa Rica. Twenty79
3.2. Biogeographical Patterns and Affinities
three species occur in South America and Central America up to Costa Rica (not in
Nicaragua or farther north). One species only occurs in Venezuela and Costa Rica (disjunct
distribution) (Fig. 3.30).
123
99
82
64
56
52
77
66
52
46
29
28
36
C
ol
om
bi
a
Pa
na
m
C
a
os
ta
R
ic
a
N
ic
ar
ag
ua
H
on
du
ra
s
M
ex
ic
G
o
ua
te
m
al
a
Be
liz
El
e
Sa
lv
ad
or
An
ti l
le
s
7
ua
do
r
Pe
ru
Ec
il
Br
az
ia
liv
Bo
Ve
ne
z
ue
la
4
Fig. 3.29. Distribution of the 123 species of the inland slope in the countries of Central and South
America (absolute figures)
Fig. 3.30. Distribution of the inland slope species
3.2.3.5. Distribution of the total Species
The total recorded species have a stronger phytogeographical relationship to the south
(South America, Amazon and Chocó) than to the north (Northern Central America): 251
spp. (80.4%) also occur in Panama, 197 spp. (63.1%) in Colombia, 166 spp. (53.2%) in
80
3.2. Biogeographical Patterns and Affinities
Ecuador, 146 spp. (46.8%) in Peru, 102 spp. (32.7%) in Brazil, 109 spp. (34.9%) in Bolivia,
111 (35.6%) in Venezuela and 69 spp. (22.1%) in Guayana.
Distribution to the north is much less pronounced: 186 species (59.6%) occur in Nicaragua,
312
251
197
146
111
109
186
166
109
102
69
84
75
95
24
Pe
ru
Ec
ua
do
r
C
ol
om
bi
a
Pa
na
m
C
a
os
ta
R
ic
N
a
ic
ar
ag
u
H
on a
du
ra
s
M
ex
ic
G
o
ua
te
m
al
a
Be
liz
El
Sa e
lv
ad
or
An
til
le
s
ia
az
il
Br
liv
Bo
zu
ne
Ve
G
uy
an
as
el
a
19
Fig. 3.31. Distribution of the 312 species in the countries of Central and South America (in %)
109 spp. (34.9%) in Honduras, 95 spp. (30.4%) in Belize, 84 spp. (26.9%) in Mexico, 75
spp. (24%) in Guatemala and 19 spp. (6.1%) in El Salvador. Twenty-four spp. (7.7%) were
recorded on at least one of the Caribbean islands and 7 spp. (2.2%) are also distributed in
Africa and/or Madagascar (Fig.3.31, Fig. 3.32, Fig. 3.34, Tab. 3.44).
Country
Costa Rica
Panama
Colombia
Ecuador
Peru
Brazil
Bolivia
Venezuela
Guayanas
Nicaragua
Honduras
Mexico
Guatemala
Belize
El Salvador
Antilles
Africa
Coastal slope
106
95
77
68
59
49
48
42
29
72
48
35
31
44
11
11
3
Gorge
119
99
75
70
54
30
37
40
23
75
46
35
30
36
8
11
5
Inland slope
123
99
82
66
64
52
52
56
34
77
46
29
28
36
4
7
1
Tab. 3.44. Geographical distribution of the species
81
Ridge
176
148
115
96
85
61
60
70
41
108
55
44
36
51
8
11
2
Total
312
251
197
166
146
102
109
111
69
186
109
84
75
95
19
24
7
Total [%]
100.0
80.4
63.1
53.2
46.8
32.7
34.9
35.6
22.1
59.6
34.9
26.9
24.0
30.4
6.1
7.7
2.2
3.2. Biogeographical Patterns and Affinities
South
North
Fig. 3.32. Distribution of the species in the research plots in the countries of Central and South America
(in %)
The wide-ranging species cover nearly half of all the plot species (52% in total; 45% on the
coast, 58% in the gorge, 52% on the ridge and 54% inland). About a quarter are only
distributed in Central America (28% in total; 22% on the coast, 24% in the gorge, 28% on
the ridge and 33% inland). 20% of all species (32% on the coast, 18% in the gorge, 20% on
the ridge and 19% inland) are only distributed in Costa Rica and south of Costa Rica
(Panama and South America) (Fig.3.33, Tab. 3.45).
100
Coastal slope
Gorge
80
Ridge = ident. with total
Inland slope
60
40
20
0
Wide range
Central America
Fig. 3.33. Distribution of the species in %
82
Costa Rica - South America
3.2. Biogeographical Patterns and Affinities
Coastal slope
Gorge
Ridge
Inland slope
Total
Wide range [%]
45
58
52
54
52
Central America [%]
22
24
28
33
28
Costa Rica - South America [%]
32
18
20
19
20
Tab. 3.45. Distribution of the species in %
Fig. 3.34. Distribution of the 312 species in Central and South American countries (absolute numbers)
3.2.3.5.1. Distribution between Geographical Regions
3.2.3.5.2. Wide-ranging Species
One hundred and sixty-one spp. can be found in most of the Central and South American
countries. These spp. have a wide-ranging distribution.
3.2.3.5.3. Central American Species
Eighty-six spp. are found exclusively in Central America or farther north (the US, Mexico,
Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica and Panama) and not in
83
3.2. Biogeographical Patterns and Affinities
South America. These spp. were classified into Northern, Nicaraguan-Costa RicanPanamanian, Costa Rican-Panamanian and Costa Rican endemic species.
3.2.3.5.3.1. Northern Species (Panama to Mexico)
Twenty-seven spp. are found only from Panama to the north of Central America (Mexico,
Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica and Panama).
3.2.3.5.3.2. Nicaraguan-Costa Rican-Panamanian Species
Fourteen spp. are found only in Nicaragua, Costa Rica and Panama.
3.2.3.5.3.3. Costa Rican-Panamanian Species
Twenty-four spp. are found only in Costa Rica and Panama.
3.2.3.5.3.4. Endemic and Costa Rican Species
Twenty-one spp. are only found in Costa Rica (they are endemic) and adjacent parts of
Panama. In total, 15 spp. (3.8%) are endemic to southern Costa Rica and nearby locations
in Panama (Chiriqui).
Of the endemic species, 7 (5.9%) occur in the gorge, 3 (2.8%) occur on the coastal slope, 5
(4.1%) occur on the inland slope and 6 (3.4%) occur on the ridge.
The endemic species are:
Acacia allenii, Bauhinia bahiachalensis, Caryocar costaricense (Pan. - Chiriqui), Clusia
peninsulae, Coccoloba standleyana, Duroia costaricensis, Huberodendron allenii, Licania
corniculata, Licania operculipetala, Licaria cufodontisii, Pleurothyrium golfodulcensis (Costa
Rica - San Isidro), Pradosia sp. nov., Sapium allenii, Sloanea sulcata and Ternstroemia
multiovulata.
3.2.3.5.4. Southern Species
Sixty-three spp. are found only to the south (excluding the Panamanian and Costa Rican
spp. – sections 3.2.3.5.3.3 and 3.2.3.5.3.4). Some have their northernmost limits in the
rainforest of the Río San Juan (Nicaragua).
Seventy-three spp. are found in the Amazon and 43 spp. are only found in the rainforest of
the Chocó. The floristic region of the Chocó is the Pacific part of Northern South America
and includes parts of Panama and Costa Rica.
3.2.3.5.5. Disjunction – Venezuela (Guayana) and Honduras
Three species are only found in Costa Rica/Panama and in Venezuela (Copaifera camibar,
Lacunaria panamensis and Licania glabriflora). Of these, two spp. occur exclusively in the
Guayana’s and Costa Rica. One species is found in Honduras (Hirtella papillata) and also
in Costa Rica. One hundred and sixty-one of the species in the plots have a wide-ranging
distribution. These species are found in Central America (north of Costa Rica) and in South
84
3.2. Biogeographical Patterns and Affinities
America. Eighty-six species only occur in Central America. Fourteen of them only occur
between Nicaragua and Costa Rica, 24 only in Panama and Costa Rica and 21 only in
Costa Rica. Sixty-three species occur in South America and Central America up to Costa
Rica (not in Nicaragua or farther north). One species only occurs in Venezuela and Costa
Rica and another one in Honduras and Costa Rica (disjunct distribution) (Fig. 3.35, Fig.
3.38).
Antilles
24
85
Central America
62
Nic., Pan, CR
Chocó
Disjunct
Guayana
2
43
Fig. 3.35. Geographical distribution of the species in Central and South America
85
3.2. Biogeographical Patterns and Affinities
85
South American Species
116
Fig. 3.36. Geographical distribution and relationship of the species in Central and South America
Costa Rica
Caribbean
Nicaragua, Costa Rica, Panama
213
62
22
15
Endemic
Fig. 3.37. Geographical distribution of the species in Costa Rica and Central America
86
3.2. Biogeographical Patterns and Affinities
Central America North
Mexico, Guatemala, Honduras, El Salvador, Nicaragua
Honduras - disjunct
Nicaragua
Costa Rica
Panama
Venezuela - disjunct
South America
Fig. 3.38. Distribution of the species of all plots (312 spp.)
3.2.3.5.6. Distribution in different Neotropical Sites
3.2.3.5.6.1. Pacific Region of Costa Rica Only
Ninty-nine spp. (31.3%) only occur in the Pacific regions of Costa Rica. The other 213 spp.
(68.3%) also occur in the Caribbean region (Fig. 3.37).
3.2.3.5.6.2. Amazon Species
In total 141 spp. (45%) occur in the rainforests of the Amazon. Fifty-six of these spp. (53%)
are found on the coastal slope, 62 spp. (50%) on the inland slope, 85 spp. (48%) on the
ridge and 54 spp. (45%) in the gorge (Fig. 3.35, Fig. 3.37).
53
50
48
45
coast
gorge
45
inland
ridge
total
Fig. 3.37. Distribution of the species of the different plots in the Amazon in %
87
3.2. Biogeographical Patterns and Affinities
3.2.3.5.6.3. Northern Pacific Costa Rica – Tropical Dry Forest (Santa Rosa
National Park)
In total only 20 spp. occur in the dry forests of northern Costa Rica. 9 spp. (8.5%) are found
on the coastal slope, 4 spp. (3.3%) on the inland slope, 9 spp. (5.1%) on the ridge and 12
spp. (10.1%) in the gorge (Fig. 3.38).
8.5
10.1
5.1
6.4
3.3
coast
gorge
inland
ridge
total
Fig. 3.38. Distribution of the species of the different plots in the tropical dry forest
of the Santa Rosa National Park in Northern Costa Rica in %
3.2.3.5.6.4. Panama – Watershed of the Canal (Tropical Rainforest)
One hundred and sixty-four spp. (52.6%) also occur in the rainforests of the Panama Canal
watershed.
3.2.3.5.6.5. Caribbean Island Species
Twenty-four spp. (7.7%) also occur on at least one of the Caribbean Islands.
88
3.3. Identification of the Tree Families of the Esquinas Forest
3.3. Identification of the Tree Families of the Esquinas Forest
With these keys it should be possible to identify all the trees and some big lianas in the
Esquinas Forest. Because of the large number of tropical plant families, identification can
be difficult. Three adapted and modified keys are presented here, all based on GENTRY
(1996) and WEBER & al. (2001). The first ("Survey key") is a short version of the
subsequent "Annotated Key", in which the characters and families (and/or particular
genera) are described in greater detail. The third key, “special habits and ‘spot’ characters
of some trees”, allows identification of plants with conspicuous features.
3.3.1. Survey key to the Families
A. Leaves compound, opposite
A.1.
Leaves bipinnate to biternate: Bignoniaceae
A.2.
Leaves simply pinnate: Bignoniaceae, Fabaceae, Sapindaceae
A.3.
Leaves 3-foliate and palmately compound: Bignoniaceae, Caryocaraceae,
Hippocastanaceae
B. Leaves compound, alternate
B.1.
Leaves bipinnate: Fabaceae, Sapindaceae, Araliaceae
B.2.
Leaves simply pinnate:
B.2a.
Parallel venation: Arecaceae, Cycadaceae
B.2b.
Rank odour: Meliaceae, Proteaceae, Fabaceae
B.2c.
Odour of essential oils or turpentine: Anacardiaceae, Burseraceae
B.2d.
Sweetish odour in trunk: Meliaceae
B.2e.
Punctations: Fabaceae
B.2f.
Bitter taste: Simaroubaceae
B.2g.
Spines: Fabaceae, Sapindaceae, Arecaceae, Cycadaceae
B.2h.
Latex: Sapindaceae, Anacardiaceae, Burseraceae, Fabaceae
B.2i.
Even pinnate: Fabaceae, Meliaceae, Sapindaceae, Arecaceae,
Cycadaceae
B.2j.
Miscellaneous useful features: winged rachis: various families; terminal "bud" on
rachis: Meliaceae; cylindrical pulvinuli: Fabaceae, Simaroubaceae; aborted
rachis tip: Sapindaceae; apical tendril: Fabaceae
B.2k. Nondescript: Meliaceae - Trichilia, Sabiaceae, Anacardiaceae - Tapirira,
Simaroubaceae
89
3.3. Identification of the Tree Families of the Esquinas Forest
B.3.
Leaves 3-foliolate, alternate: Sapindaceae - Allophylus, Fabaceae – Erythrina
B.4.
Leaves
palmately
compound,
alternate:
Arecaceae, Rutaceae,
Araliaceae,
Bombacaceae, Caricaceae, Cecropiaceae, Sterculiaceae, Cochlospermaceae
C. Leaves simple, opposite or whorled
C.1.
Stipules or stipule scars: Rubiaceae, Quiinaceae, Malpighiaceae, Chloranthaceae,
Vochysiaceae, Rhizophoraceae
C.2.
Latex: Apocynaceae, Clusiaceae
C.3.
Punctations: Myrtaceae, Melastomataceae, Lythraceae, Rutaceae, Clusiaceae,
Loranthaceae, Myrsinaceae, Rhizophoraceae
C.4.
3(-7)-veined leaves with parallel cross veins: Melastomataceae
C.5.
Odour of essential oils: Monimiaceae, Lauraceae, Verbenaceae
C.6.
Glands on twig at petiole base: Vochysiaceae
C.7.
Serrate margins: Hippocrateaceae, Violaceae, Elaeocarpaceae, Rhizophoraceae,
Loganiaceae,
Flacourtiaceae,
Verbenaceae,
Monimiaceae,
Brunelliaceae,
Chloranthaceae, Acanthaceae, Euphorbiaceae
C.8.
Miscellaneous:
Lythraceae,
Acanthaceae,
Loganiaceae,
Nyctaginaceae,
Verbenaceae,
Myrtaceae,
Melastomataceae,
Hippocrateaceae,
Rhamnaceae,
Caprifoliaceae, Oleaceae, Malpighiaceae, Clusiaceae, Elaeocarpaceae
D. Leaves simple, alternate
D.1.
Latex:
Sapotaceae,
Moraceae,
Euphorbiaceae,
Olacaceae,
Apocynaceae,
Myristicaceae, Anacardiaceae, Chrysobalanaceae
D.2.
Conical terminal stipule: Moraceae, Magnoliaceae, Polygonaceae
D.3.
Odour of essential oils: Magnoliaceae, Hernandiaceae, Annonaceae, Myristicaceae,
Lauraceae, Canellaceae, Anacardiaceae, Burseraceae, Fabaceae,
Araliaceae, Icacinaceae
D.4.
Palmately veined or 3-veined:
a) Malvalean Pulvinus: Tiliaceae, Sterculiaceae, Bombacaceae, Malvaceae,
Elaeocarpaceae, Bixaceae, Euphorbiaceae
b) Without swollen pulvinus: Ulmaceae, Urticaceae, Euphorbiaceae, Caricaceae,
Cochlospermaceae, Flacourtiaceae, Hernandiaceae, Araliaceae,
Rhamnaceae, Rhizophoraceae, Olacaceae, Fabaceae
D.5.
Strong
bark:
Annonaceae,
Lecythidaceae,
Thymelaeaceae,
Boraginaceae,
Fabaceae, Malvales, Urticales
D.6.
Unequal petioles: Araliaceae, Capparidaceae, Euphorbiaceae, Sterculiaceae,
Bombacaceae, Urticaceae
90
3.3. Identification of the Tree Families of the Esquinas Forest
D.7.
Petiole glands: Chrysobalanaceae, Combretaceae, Euphorbiaceae, Flacourtiaceae,
Rhamnaceae
D.8.
Serrate
margins:
Elaeocarpaceae,
Actinidiaceae,
Celastraceae,
Euphorbiaceae,
Fagaceae,
Clethraceae,
Dilleniaceae,
Flacourtiaceae,
Humiriaceae,
Icacinaceae, Lacistemataceae, Fabaceae, Myrsinaceae, Ochnaceae,
Rhamnaceae, Rosaceae, Sabiaceae, Solanaceae, Symplocaceae,
Theaceae, Theophrastaceae, Violaceae
D.9.
Thickened or flexed petiole apex: Elaeocarpaceae, Euphorbiaceae, Flacourtiaceae,
Meliaceae
D.10. Punctations: Flacourtiaceae, Rutaceae, Myrsinaceae, Theaceae
D.11. Stipules: Celastraceae, Chrysobalanaceae, Dichapetalaceae, Erythroxylaceae,
Euphorbiaceae, Flacourtiaceae, Rosaceae, Violaceae
D.12. Lepidote
or
stellate
trichomes:
Annonaceae,
Capparidaceae,
Clethraceae,
Dilleniaceae, Euphorbiaceae, Icacinaceae, Fagaceae, Malvales,
Solanaceae, Styracaceae
D.13. Leaves parallel-veined or lacking secondary veins: Podocarpaceae, Theaceae,
Arecaceae
D.14. Parallel
tertiary
venation:
Clusiaceae,
Icacinaceae,
Lacistemataceae,
Lecythidaceae, Myristicaceae, Sapotaceae, Ochnaceae, Olacaceae
D.15. Spines or spine-tipped leaves: Celastraceae, Euphorbiaceae, Flacourtiaceae,
Moraceae, Nyctaginaceae, Olacaceae, Rhamnaceae, Rosaceae,
Solanaceae, Theophrastaceae, Urticaceae
D.16. None
of
above:
Bignoniaceae
-
Crescentia,
Amphitecna,
Boraginaceae,
Capparidaceae, Celastraceae - few Maytenus, Chrysobalanaceae,
Combretaceae, Dichapetalaceae, Ebenaceae, Euphorbiaceae,
Flacourtiaceae, Humiriaceae, Icacinaceae, Fabaceae, Moraceae,
Myricaceae, Olacaceae, Onagraceae, Polygonaceae, Rhamnaceae,
Sabiaceae, Solanaceae, Violaceae
3.3.2. Annotated Key
A. Leaves compound, opposite
This is the easiest category. Only one important family (Bignoniaceae) plus a few other
small families and miscellaneous genera or species are characterised by opposite
compound leaves. If in doubt as to whether the leaf is compound or not, look for the axillary
bud. In deciduous species, thick twigs tend to indicate compound leaves. In fallen leaflets,
asymmetric leaf bases often suggest origin from a compound leaf.
91
3.3. Identification of the Tree Families of the Esquinas Forest
A.1. Leaves bipinnate, opposite
Bignoniaceae - Jacaranda
A.2. Leaves simply pinnate, opposite
Useful differentiating characters include: the rachis (angled and more or less grooved
above in Bignoniaceae, Brunelliaceae, and Juglandaceae; conspicuously jointed);
presence or absence of interpetiolar line or ridge (absent in only Rutaceae, Juglandaceae,
Sapindaceae); type of marginal serrations and pubescence of the leaflets; and presence
and type of stipules.
Bignoniaceae - Tecoma stans, leaflets petiolulate and sharply serrate.
Brunelliaceae - Brunellia hygrotermica, leaflets closely serrate, stipules between the
petioles; interpetiolar ridge with small subulate stipule-like projections.
Fabaceae - Platimiscium spp., typical legume pulvinus and pulvinuli; odour of green beans.
Juglandaceae - Leaflets asymmetric (Oreomunnea pterocarpa) and lower leaflets smaller
(Alfaroa guanacastensis), twigs and petiole hispid or the twigs with conspicuous
round white lenticels; typically rank, walnut-like odour.
Sapindaceae - Matayba opposotifolia, margins entire, petiolules short, thick-based.
A.3. Leaves 3-foliolate, opposite
Bignoniaceae (most neotropical trees of the family) - Leaflet bases rounded to cuncate,
not tapering into petiolules; pubescence usually of stellate trichomes or lepidote
scales.
Caryocaraceae - Caryocar costaricense, branchlets with a conspicuous rubiac-like
terminal stipule; (leaflets serrate or serrulate) and characterized by distinctive gland
pair at apex of petiole (unique in opposite-leaved taxa);
Hippocastanaceae - Billia colombiana, vegetatively very similar to Caryocar but lacks
petiolar glands and elongate terminal stipules; completely entire leaflets. Leaflets
turn reddish when they are about to fall.
Rutaceae - Amyris brenesii, with conspicuously punctate leaflets; acute to acuminate,
petiolulate.
Verbenaceae - Vitex cooperi, twigs somewhat square, leaf bases usually gradually
tapering into indistinct petioles is unique among opposite 3-foliolate taxa.
92
3.3. Identification of the Tree Families of the Esquinas Forest
B. Leaves compound, alternate
Several families in this group are very difficult to separate on the basis of sterile characters,
especially some Sapindaceae (Cupania, Matayba, Talisia, etc.), some Meliaceae (Trichilia)
and a few nonaromatic Anacardiaceae (Tapirira).
B.1. Leaves bipinnate or biternate, alternate
Fabaceae - Bipinnate leaves and spines on trunk, branches, or rachises are unique to
mimosoid legumes; like nonspiny bipinnate legumes these taxa are characterized
by the typical cylindrical legume pulvinus and pulvinulus and (in mimosoids) by
development of an often elaborate gland on the dorsal side of the lower petiole or
sometimes between the lower rachises.
Meliaceae - Melia azedarach, a cultivated tree now also growing wild, characterized by thin
serrate leaflets; young growth often conspicuously whitish; looks much more like
Araliaceae than Meliaceae, but lacking characteristic ligule-like base.
Sapindaceae - Dilodendron costaricense with many small serrate leaflets and the typical
aborted rachis apex of the family.
B.2. Leaves simply pinnate, alternate
B2a. Leaflets with parallel venation
Arecaceae - The only woody monocots with pinnately compound leaves, the leaf segments
unmistakable in their parallel venation.
Cycadaceae - Zamia fairchildiana, similar only to palms from which the leaflets differ in
being more coriaceous; short spine on the petiole and/or rachis, with spines shorter
and thicker-based than in pinnate-leaved spiny palms.
B2b. Rank odour
Fabaceae - Most legumes have a characteristic, more or less rank green bean-like odour.
Easy to recognize by distinctively round, swollen petiolules (whole length of
petiolule uniformly cylindric) and petiole base; Connaraceae (lianas and
occasionally treelets) lack the odour but are otherwise indistinguishable vegetatively
from Fabaceae. Some Picramnia species (Simaroubaceae) have similarly pulvinate
petioles and petiolules, but often with bitter taste and the strongly alternate leaflets
becoming much smaller toward leaf base.
Meliaceae - Cedrela odorata, somewhat garlic-like odour, always entire leaflets.
Proteaceae - Roupala montana, canned-meat odour of leaves, mostly with compound
juvenile leaves; the trunk slash has a characteristic odour resembling low-quality
canned beef; the leaflets are extremely asymmetrical, usually with one side flat
(even concave), the other margin serrate.
93
3.3. Identification of the Tree Families of the Esquinas Forest
B2c. Odour of essential oils or turpentine
Anacardiaceae and Burseraceae - These two families usually have a fairly strong
turpentine-like or mango-like odour.
Anacardiaceae - Usually more weakly turpentine-odoured or with a somewhat sweetish
mango-like odour; sometimes with a watery latex which dries black (wounded trunks
often stained with black); may be considered a derived version of Burseraceae
differentiated by the technical character of one anatropous ovule per locule or a
single ovule in ovary.
Burseraceae - Usually strongly turpentine-odoured, even in bark; almost always with milky
latex either in the twigs or as few widely scattered droplets from the bark slash, the
latex drying whitish around trunk wounds; technically separated from
Anacardiaceae by pendulous, epitropous ovules of which there are two per locule.
Rutaceae - Zanthoxylum, spines on trunk or stems (unique in simply pinnate taxa) and
typically punctate leaflets, at least along at margin below.
B2d. Sweetish odour in trunk only; excluding mango-like odours
Meliaceae - Most Meliaceae are characterized by a faint but distinctly sweetish odour from
the trunk slash (but Cedrela odorata has a very different rank, garlic-like odour).
B2e. Punctations
Look both against strong light and out of it; punctations are often restricted to the sinuses of
marginal teeth or serrulations.
Fabaceae - A few genera of legume have punctate leaves (Myroxylon balsamum,
Hymenaea courbaril, Copaifera aromatica and some Lonchocarpus, etc.), the
punctations often at least partly linear.
Rutaceae - Most species of Rutaceae are punctate, at least along leaflet margin; most
pinnate-leaved Rutaceae are species of Zanthoxylum and most of these have
spines on the trunks, branchlets or leaves.
B2f. Very bitter taste
Simaroubaceae - Most Simaroubaceae are characterized by a bitter taste when the twig is
chewed.
B2g. Spines (rare in simply pinnate taxa)
Arecaceae - Several pinnate-leaved palm genera have spiny trunks and/or leaves.
Cycadaceae - Zamia fairchildiana has short spines on the petiole and/or rachis.
Fabaceae - Machaerium, mostly lianas but also trees, usually only with paired stipular
spines and usually with red latex.
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3.3. Identification of the Tree Families of the Esquinas Forest
Rutaceae - Zanthoxylum, always trees in Neotropics; thick spines on trunk typical, often
also with spines on petioles and leaflets.
B2h. Latex (rare in compound-leaved taxa)
Burseraceae - Latex sometimes present in twigs, almost always in exceedingly
inconspicuous, scattered droplets in stem slash, these typically continuing to exude
and forming cloudy white drippings below trunk wounds.
Fabaceae - Red latex in a number of papilionate genera (Dussia, Machaeritan,
Pterocarpus, Swartzia).
B2i. Even-pinnate leaves
Fabaceae – Inga, unique in being even-pinnate and with glands between all leaflets;
Cassia often with glands between basal pair or pairs of leaflets, and several other
caesalpinioid genera, all with typical pulvinus and pulvinuli and often with typical
legume odour.
Meliaceae - Most Meliaceae except Trichilia are even-pinnate, especially Guarea with its
typical apical "bud".
Sapindaceae - Most Sapindaceae are basically even-pinnate but with alternate leaflets
and a very characteristic aborted rachis apex at base of what often appears to be a
terminal leaflet.
B2j. Miscellaneous useful characters for genera or common species with
pinnately compound alternate leaves
(1) Winged rachis : Individual species of many genera and families,
Fabaceae - Several unrelated genera have winged rachis: Inga, even-pinnate with
glands between all leaflet pairs.
Meliaceae - Guarea pterorhachis, even-pinnate with many thick leaflets and broad
coriaceous rachis wings.
Simaroubaceae - Quassia amara, characterized by mostly 5-foliolate leaves and
bitter taste.
(2) Terminal "bud" of unfolding leaflet pair at tip of rachis
Meliaceae - Guarea
(3) Uniformly cylindrical pulvinuli and pulvinus - Typical of nearly all legumes. Most
legumes have the typical legume green bean odour; Ruptiliocarpon caracolito has
unifoliolate leaves with legume-like pulvinulus.
(4) Naked rachis apex - Tree Sapindaceae (especially Cupania and Matayba) with pinnate
leaves almost always have a very characteristic aborted rachis apex extended as a
small projection at base of what appears to be a terminal leaflet.
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3.3. Identification of the Tree Families of the Esquinas Forest
B2k. Nondescript (odd-pinnately compound, alternate, no spines, odour,
punctations, latex, etc.)
Anacardiaceae - Odourless anacards (e.g. Tapirira) that also lack obvious latex are very
nondescript and especially easy to confuse with Trichilia. Often there is at least a
faint trace of a mango-like odour. The most common species dries with a
characteristic reddish tint.
Meliaceae - Trichilia, unfortunately the most common genus of Meliaceae, is atypical in the
family, having odd-pinnate leaves. The leaflets are entire, and there is usually a
sweetish odour from the trunk slash.
Simaroubaceae - Nonbitter simaroubs are often characterized by legume-like cylindrical
pulvinuli. Picramnia can be distinguished from legumes by the typical alternate
leaflets, progressively smaller toward base of the rachis.
B.3. Leaves 3-foliolate, alternate
Alternate, consistently 3-foliolate leaves are not very common although they may occur as
variants in basically pinnately compound-leaved individuals (or species or genera).
Caryocaraceae – Anthodiscus, usually blunt apex and crenate leaflet margins.
Fabaceae – Erythrina, usually with spiny trunks and branchlets; with the typical legume
cylindrical pulvinus and pulvinuli; margin always entire.
Rutaceae - Several genera have 3-foliolate leaves, at least in some species; all are
characterized by the pellucid punctations and most have a more or less citrus-like
vegetative odour.
Sapindaceae – Allophylus,
usually acute or acuminate apex and toothed (or entire)
margins; a few species have simple leaves.
B.4. Leaves palmately compound, alternate
Araliaceae - Schefflera, hemiepiphytic, characterized by the rank or medicinal odour and
the thickly triangular ligule projecting up from the more or less clasping petiole base.
Arecaceae (Palmae) - Fan palms are the only arborescent plants in this study with
palmately compound leaves with parallel-veined segments.
Bombacaceae - Most Bombacaceae have palmately compound leaves, always with a
Malvalean pulvinus at petiole apex. Several genera have spines on the trunk (at
least when young, a unique combination except for Jacaratia); whether with or
without spines, Bombacaceae are often unusually large emergents with distinctively
swollen thick trunks. One spineless genus has the leaflets continuous with the
digitately parted petiole apex (unique in Malvales).
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3.3. Identification of the Tree Families of the Esquinas Forest
Caricaceae – Jacaratia is the only palmately compound-leaved tree with milky latex in the
region; they have spiny trunks and resemble Bombacaceae except for the latex and
lack of a pulvinus.
Sterculiaceae - Herrania purpurea has palmately compound leaves, usually large,
conspicuously hairy and borne on pachycaul treelet with maroon cauliflorous
flowers and cacao-like fruit; Sterculia allenii also has Bombacaceae-like compound
leaves; both have Malvalean pulvinus and stipules.
C. Leaves simple, opposite or whorled
C.1. Stipules (or stipule scars) present
Chloranthaceae - The more or less swollen node has a stipule-like sheath; the plants are
easily distinguished by the strong Ranalean odour and the serrate leaves.
Malpighiaceae - The tree genera have intrapetiolar stipules in the axil between the petiole
and twig (looking like ligular dorsal projection from petiole base), these differing
from the few Rubiaceae with similar stipules by being fused (usually bifid in
Rubiaceae); interpetiolar lines are also usually present. The main familial vegetative
characteristic is the presence of malpighiaceous or T-shaped hairs, these almost
always visible at least on the petioles and young twigs.
Quiinaceae - The other main family with interpetiolar stipules, these always separate
(usually fused in Rubiaceae) and often rather long and subfoliaceous; differs from
Rubiaceae in the usually serrate or serrulate leaf margin (deeply incised in juveniles
of few species).
Rhizophoraceae - Rhizophora, restricted to coastal mangroves, is utterly distinctive as the
only mangrove with stilt roots; the other opposite-leaved genera are less striking,
with Cassipourea elliptica in the leaves usually obscurely and remotely denticulate
or serrate (unlike Rubiaceae) and the small, narrowly triangular, early-caducous
stipule usually sericeous.
Rubiaceae - Interpetiolar stipules are present at least 99 % of the time; if stipules are not
readily apparent check terminal bud to see if it is enclosed by caducous stipules.
These should be in a plane at right angles to the two uppermost leaves and leave
an interpetiolar line when they fall.
Vochysiaceae - Vochysia, rather thick-based stipules.
C.2. Latex
Apocynaceae (Some genera alternate-leaved and many are climbers) - Latex white and
free-flowing (red in some species with alternate leaves), lacks the typical guttifer
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3.3. Identification of the Tree Families of the Esquinas Forest
terminal bud (i.e. the petioles of terminal leaf pair not hollow-based with terminal
bud growing from within cavity, except in a few species with very profuse latex).
Clusiaceae (Guttiferae) - Very distinctive in typical terminal bud and coloured latex; latex
commonly yellow, cream, or orange, usually flowing slower than in Apocynaceae.
Terminal bud characteristically from between the hollowed-out leaf bases; typical
terminal bud not developed only when latex strongly coloured; latex white only when
the leaf bases form conspicuous chamber. The latex may not be very obvious; try
breaking a leaf and twig as well as the trunk slash; stilt roots are quite common.
C.3. Punctations
Clusiaceae - Vismia, usually by orange latex (or a moist orangeish area just inside bark
where latex should be), some have punctate leaves.
Lythraceae - Adenaria and Lafoensia, leaves thinner than in most Myrtaceae and with
more ascending secondary veins and absence of collecting vein.
Melastomataceae - Mouriri completely lacks the ascendine veins of other meastomes,
looks almost exactly like Myrtaceae and may have punctations; it differs from
Myrtaceae in the somewhat jointed nodes.
Myrtaceae - Usually further characterized by more or less parallel and close-together
secondary and tertiary venation ending in a submarginal vein. Many have smooth,
white, peeling bark; some have aromatic leaves. The only possible confusion comes
from a very few guttifers that have punctations (but also latex, unknown in
Myrtaceae) or from some Myrtaceae that are not obviously punctate (also beware
Mouriri (see above)).
Rhizophoraceae - Rhizophora, keyed out above on account of the terminal stipule, can
have leaf punctations.
Rutaceae - Ravenia rosea, leaves with punctations, characterized by sheathing guttifer-like
petiole bases in which the apical bud is protected; differs from Clusiaceae in lacking
latex and the small glandular punctations.
C.4. Leaves 3(-7)-veined, with parallel cross veins more or less
perpendicular to main veins
Melastomataceae - The very characteristic leaf venation makes this one of the easiest
families to identify. Also beware of Mouriri which lacks the typical venation and
looks almost exactly like Myrtaceae.
C.5. Odour of essential oils (only two Ranalean families are
characterized by aromatic opposite leaves)
Chloranthaceae - Hedyosmum, very characteristic swollen nodes with stipule-like sheath.
Lauraceae - Caryodaphnopsis burgeri, most notably peculiar 3-veined.
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3.3. Identification of the Tree Families of the Esquinas Forest
Monimiaceae - Siparuna, usually puberulous or with lepidote scales; Mollinedia often lacks
obvious odour but is characterized by very distinctive leaf with few very separated
marginal teeth.
Verbenaceae - usually aromatic but the odour is clearer and sweeter (often somewhat
minty) and less pungent than in the Ranales; aromatic members of these families
usually have tetragonal branchlets unlike the Ranales.
C.6. Glands on twig at base of petiole
Vochysiaceae - Very characteristic glands from the fallen stipules or stipule bases
characterize most Vochysiaceae (except Vochysia).
C.7. Serrate (serrulate) margins
The combination of opposite simple leaves and serrate margins is rare and found only in
eight woody neotropical lowland families besides the Quiinaceae, Melastomataceae and
Chloranthaceae, which are easily recognized (see above).
Brunelliaceae - Brunellia hygrotermica, the laeflets are closely serrate and with
prominulously reticulate venation below and numerous secondary veins making
obtuse angle with midvein; strong interpetiolar line.
Chloranthaceae - Hedyosmum, as noted above unmistakable in the sheathing node and
strong Ranalean odour.
Elaeocarpaceae - Sloanea is characterized by a mixture of alternate and opposite leaves,
even on the same branch, but the leaves are almost never uniformly opposite; also
very distinctive in the flexed, but non-pulvinate petiole apex and strictly pinnate
venation. The margins vary from almost entire to rather shallowly and coarsely
subdentate; species with more serrate leaves tend to be more pubescent and some
of the pubescent species have conspicuous persistent leafy stipules.
Hippocrateaceae - Mostly lianas but a few are trees and Cheiloclinium can be both a tree
and have serrate leaves; it is characterized by tertiary venation more or less parallel
and perpendicular to midvein.
Monimiaceae - Mollinedia has the teeth usually very widely separated (typically only one
or two per side) and rather sharp; Siparuna is also frequently toothed but easy to
recognize by the Ranalean odour.
Rhizophoraceae - Cassipourea, secondary veins few and brochidodromous strikingly far
from margin; margin mostly remotely serrulate; caducous triangular terminal stipule
pair leaving interpetiolar line.
Verbenaceae - Typically with more or less tetragonal stem and aromatic odour. Most
woody Verbenaceae are entire but usually serrate-leaved Callicarpa, with
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3.3. Identification of the Tree Families of the Esquinas Forest
conspicuously floccose indument on leaf undersides and twigs, is a small tree.
Interpetiolar lines lacking.
Violaceae - Rinorea usually has opposite leaves and is one of the most common
understorey-tree genera of many forests. Characterized by the nodes noticeably
jointed, the typically short petioles, and the tendency to have a small acute stipuleenclosed apical bud immediately subtended by oblique, whitish-margined,
interpetiolar ridge.
C.8.
Miscellaneous
opposite
simple-leaved
trees
(lacking
latex,
essential oils, serrate margins)
Acanthaceae - Bravaisia integerrima, tree with very weak wood, although several genera
include shrubby or small tree species. Most Acanthaceae characterized by
conspicuously jointed nodes, swollen when fresh and contracted when dried. Most
Acanthaceae have an obvious interpetiolar line. Except for the spiny-margined
species, the acanths in this region all have entire or merely serrulate, but never truly
serrate, leaves. Cystoliths (look like short black lines) often present on upper leaf
surface (also in Urticaceae).
Caprifoliaceae - Viburnum, leaves with few strongly ascending veins, puberulous at least
below, sparsely and bluntly serrulate or more or less bluntly few-toothed toward
apex.
Clusiaceae - Occasionally lacks apparent latex-like Chrysochlamys, but with the typical
hollowed-out Clusiaceae petiole bases that form a protective chamber for the
developing bud; a few Vismia species (which lack the typical hollowed petiole base)
may not always show the orange latex but there is always a hint of orange colour
under the bark where the latex should be.
Elaeocarpaceae - Sloanea is characterized by a mixture of alternate and opposite leaves,
even on same branch, almost never uniformly opposite; also very distinctive in the
flexed but non-pulvinate petiole apex and strictly pinnate venation. Margins
sometimes bluntly and irregularly toothed (see also above).
Hippocrateaceae - Two genera are sometimes trees, one (Cheiloclinium cognatum)
usually with finely crenate-serrate margins (also distinguished by conspicuously
parallel tertiary venation more or less perpendicular to midvein), the other (Salacia)
with large very thick-coriaceous entire leaves with immersed fine venation and
drying a characteristic dull olive.
Lythraceae - Usually with tetragonally angled young twigs and/or longitudinally exfoliating,
often reddish twig bark in older branchlets; interpetiolar lines or ridges in Lafoensia
(with close-together secondary veins prominulous above and below, each adjacent
pair separated by a well-developed intersecondary).
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3.3. Identification of the Tree Families of the Esquinas Forest
Malpighiaceae - A few genera of shrubs and small trees lack obvious intrapetiolar stipules
(as do the lianas); Malpighia and many species of Bunchosia have neither obvious
stipules nor interpetiolar lines and are often characterized instead by a pair of
ocellar glands near base of lamina below. Like the species with stipules, they are
also vegetatively characterized by the typical T-shaped trichomes at least on
petioles and young branchlets.
Melastomataceae - Mouriri looks much more like Myrtaceae in vegetative condition than
like typical 3-7-veined Melastomataceae. It differs from Myrtaceae most notably in
the jointed nodes.
Myrtaceae - In some Myrtaceae the punctations are not very evident. They are usually
characterized (as are the punctate-leaved taxa) by the straight, often quite closetogether secondary and intersecondary veins that end almost perpendicular to a
well-developed submarginal collecting vein.
Nyctaginaceae - Rather nondescript and easily be confused with Psychotria or similar
Rubiaceae, even when in flower or fruit, except for lacking stipules. The best sterile
character is the reddish-brown pubescent terminal bud. The combination of
somewhat succulent, often variously sized and/or subopposite blackish-drying
leaves and rufescent terminal bud immediately indicates Nyctaginaceae.
Oleaceae - Chionanthus panamensis, petioles usually somewhat thickened at base (cf.
Sapindaceae petiolules), the leaf blade either pubescent or else quite narrow and
oblong; twigs lacking interpetiolar lines, often with scattered, round, raised white
lenticels. In flower unmistakable in having only 2 anthers and very narrow petals.
Verbenaceae - Usually with tendency to tetragonal branchlets and raised petiole
attachments; leaf base typically attenuate onto petiole and in many species (most
Citharexylum) with an elongate gland in the laminar attenuation on either side of
petiole apex. Leaves, at least of forest taxa, usually rather membranaceous and
somewhat aromatic.
D. Leaves simple, alternate
This "grab bag" category constitutes by far the largest and generally the most nondescript
group. In preceding groups any sterile woody plant should be identifiable to family; in this
group there will be many plants which end up as family indets, unless they are fertile and
technical characters are used.
Look for (in approximate order of importance): latex, odour of essential oils (Ranalean
odour), conical terminal stipules (usually = Moraceae), 3-veined base (frequently suggests
Malvales), punctations (and the undersurface texture which accompanies punctations in
Myrsinaceae), serrate margins (uncommon in tropical forest species), strong bark (pull a
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3.3. Identification of the Tree Families of the Esquinas Forest
leaf off a twig to see if a strip of bark comes with it; also check the twig bark itself), petiole
length and flexion, glands at tip of petiole (usually Euphorbiaceae or Flacourtiaceae),
whether petioles are thickened at base or apex or of unequal lengths, spines.
D.1. Latex
Look carefully, breaking the midveins or petioles of several leaves as well as young twigs.
Check both trunk and leaves since sometimes obvious latex is apparent only in one or the
other; note whether the trunk slash has discrete latex droplets, how these are arranged,
and what colour the latex is.
Apocynaceae - Relatively few Apocynaceae have alternate leaves. Alternate-leaved
Apocynaceae usually have white and milky free-flowing latex but this may be
yellowish in Aspidosperma spruceanum. The species of Aspidosperma are
extremely difficult to distinguish vegetatively from Sapotaceae.
Cecropiaceae - Pourouma bicolor, leaves palmately lobed, conical terminal stipule and
sap dark brown to black.
Chrysobalanaceae - Very rarely with a distinct trace of reddish latex, this not always
visible in individual trees. Look for stipules on the young twigs or their scars; lack of
Ranalean odour separates from Myristicaceae, the only source of potential
confusion.
Euphorbiaceae - White to cream milky latex typically present, often caustic and harmful to
eyes. Pausandra and some Croton have bright red latex. Note: Although latex is
considered characteristic of Euphorbiaceae, many species have no latex at all.
Serrate (or serrulatc) leaf margins, long petioles with flexed apices and often of
different lengths, and a pair of glands near petiole apex are good indicators of
Euphorbiaceae and are unique to this family among species with alternate simple
leaves and latex.
Moraceae - Latex of many of the species a unique tan shade (exactly the colour of "cafe
con leche"), but many other species with milky white latex (usually only watery in
Trophis), and in a few varying to tannish yellow (some Naucleopsis). Conical
terminal stipules and the scar from these stipules usually obvious (and definitive for
Moraceae). Leaf venation very characteristic with the brochidodromous lower
secondary veins closer together and/or joining midvein at different angle from
others.
Myristicaceae - Usually with red latex (only in trunk), this sometimes rather watery at first
but almost always soon becoming obviously red, especially when drying. Very easy
to distinguish from other families with occasional species or genera with red latex by
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3.3. Identification of the Tree Families of the Esquinas Forest
the typical myristicaceous branching, lack of petiolar glands, and presence of
Ranalean odour.
Olacaceae - Latex present only in Heisteria and Minquartia, usually present only in leaves
and petioles, white and milky (usually) to somewhat watery. Look for a slightly
longish, distinctively curved (putatively U-shaped) and somewhat apically thickened
petiole; leaves of most species of this family have a characteristic greyish or
tannish-green colour when dry. The margins are always entire.
Sapotaceae - Latex (in Neotropics) always white and milky; sometimes not very apparent
but almost always visible in either trunk slash or leaves (if not both). Leaves
typically with base of petiole enlarged (petiole more or less pop-bottle shaped) and
with numerous parallel secondary veins. Some genera lack the typical petiole but
these mostly have finely parallel tertiary and secondary veins (the extremes with
leaves similar to Clusia); margins always entire and latex of slash usually emerging
in discrete droplets. Never with conical terminal stipule or glands on petiole.
D.2. Conical terminal stipules
These are most distinctive of Moraceae. However, the stipule is not always obvious and in
Trophis can only be considered present by stretching the imagination; also note that other
families may have young leaves which are superficially somewhat similar to the Moraceae
stipules.
Magnoliaceae - Talauma has a Moraceae-like terminal stipule that falls to leave a
conspicuous nodal ring, but is aromatic and nonlactiferous.
Moraceae - The combination of milky latex and conical terminal stipule (that falls to leave a
distinct scar) is definitive for, and almost universal in, Moraceae. The exception is
Trophis where neither latex nor stipule may be discernible, where recognizable as
Moraceae only by the typical leaf venation.
Polygonaceae - Coccoloba and Triplaris have conical terminal stipules but these rupture to
form an ochrea rather than falling cleanly as in Moraceae.
Theaceae and Myrsinaceae - A number of genera of Theaceae and Myrsinaceae and
related families have young leaves rolled at branch apex and are superficially
similar to the conical terminal stipule of Moraceae.
D.3. Odour of essential oils (Ranalean odour)
Most Ranalean plants have alternate simple leaves and more or less conspicuous rank or
turpentiney odours, which may be hard to detect. As a group these are easy to recognize
by their "primitive " odour, not to be confused with the licore-like smell of a common
epiphyllous leafy liverwort. Frequently a twig split longitudinally will give off a more easily
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3.3. Identification of the Tree Families of the Esquinas Forest
detectable odour than the leaves themselves. In some Lauraceae with little or no leaf
odour, the bark slash is aromatic; in others the reverse may be true.
Families with Ranalean odours - All with completely entire margins except for a very few
somewhat lobed-leaved (but never serrate) species:
(a) Piperaceae - Swollen nodes with shoot proceeding from leaf axil. Distinctive spicate
inflorescence; odour tends to be peppery; leaf base often strikingly asymmetrical.
Usually shrubs; when trees (usually small), typically with prominent stilt roots.
(b) Magnoliaceae - Talauma gloriensis, complete rings around twig at nodes from the
distinctive caducous stipule that completely covers terminal bud (like Moraceae),
the petiole is conspicuously grooved above.
(c) Hernandiaceae - Three-veined leaves are unique in aromatic Ranalean taxa except for
a very few atypical Lauraceae. Also distinctive among Ranales are long ,
sometimes of various lengths of the the petioles. The vegetative odour is ranker
than in most Lauraceae in which 3-veined taxa also differ in shorter petioles. Differ
from similar Araliaceae in lacking conspicuously smaller short-petioled leaves and in
basal lateral vein pair curving upward rather than being straight or curving outward.
(d) Annonaceae, Myristicaceae, and Lauraceae - which are easy to tell apart when
fertile, but can be confusing when sterile.
Typically, Myristicaceae have relatively long oblong leaves with dull surfaces, short
petioles, and many close-together parallel secondary veins. Typically Lauraceae
have short elliptic leaves with glossy shiny surfaces, relatively long petioles, and
relatively glossy shiny surfaces, relatively long petioles, and relatively few, often
strongly ascending and not strictly parallel, secondary veins. Although there is little
room for confusion between Myristicaceae and Lauraceae, Annonaceae are
intermediate and overlap with both vegetatively. Both Myristicaceae and
Annonaceae (but not Lauraceae) are characterized by myristicaceous branching
with the lateral branches at right angles to the trunk and the evenly spaced leaves
2-ranked along these or along their lateral branches; in Myristicaceae, especially,
the lateral branches tend to be clustered and appear to have an almost whorled
arrangement ("myristicaceous branching"). Lauraceae never have such a phyllotaxy
and their leaves are often irregularly spaced along the branches with a definite
clustering towards the branchlet apex. Very many Lauraceae are distinguished by
the way the leaf blade gradually merges with the petiole apex, typically with at least
the hint of an involution of the margin and sometimes with a distinctly involute
auricle on each side. Lauraceae leaves typically have shinier surfaces than do the
other two families, and the pubescence, when present, is usually sericeous with
appressed simple trichomes or softly rufescent. Myristicaceae trichomes are either
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3.3. Identification of the Tree Families of the Esquinas Forest
stellate or 2-branched (T-shaped), frequently very conspicuous, and usually
rufescent (to whitish on the leaf undersurface). Annonaceae, as usual, are
intermediate but stellate (or lepidote). Trichomes are rare (mostly Duguetia) and
sericeous pubescence is common only in Xylopia. Most Annonaceae have strong
bark ("cargadero" = useful for tying cargo), a feature not found in Myristicaceae or
Lauraceae. Lauraceae twigs are typically green while those of Myristicaceae are
brownish; Annonaceae commonly have either green or brown twigs; all the greentwigged Annonaceae have strong bark but only some of the brown-twigged ones
do. The odour of Lauraceae is usually either spicy, and almost sweetish or foetid
and unpleasant; that of Annonaceae tends to be slightly rank; that of Myristicaceae
is usually more pungently turpentiney and typically not very strong.
Canellaceae – Pleodendron macranthum, highly aromatic elliptic leaves
At least five other taxa with simple alternate leaves have odours that might be confused
with the Ranalean group: Most simple-leaved species of Anacardiaceae (Mangifera and
Anacardium) have a more strongly turpentiney odour, as do the few simple-leaved species
of Burseraceae. The latter also have prominently flexed petiole apices indicating their
compound-leaved affinities. Araliaceae have aromatic leaves and some are reminiscent of
some Lauraceae; they differ prominently in their varying petiole lengths. Dendrobangia
(Icacinaceae) has a more medicinal odour than is typical of Ranalean families and is also
characterized by a grooved petiole, appressed-stellate indumentum, and black-drying
colour. Alternate-leaved weedy Asteraceae are mostly not strongly aromatic.
D.4. Leaves palmately 3(-9)-veined at base (and alternate and simple)
The majority of taxa with palmate basal veins (here referred to as "3-veined") belong to one
of two quite unrelated main groups: Malvales (Malvaceae, Tiliaceae, Bombacaceae,
Elaeocarpaceae, Sterculiaceae) and Hamamelidae (especially Ulmaceae, Urticaceae). The
Malvalean woody taxa have petioles with a distinctive swollen apical pulvinus; the
Hamamelidae and other three-veined families do not.
D.4a. Petioles with apical pulvinar thickening (or with leafy stipules) (=
Malvales)
Perhaps the main palmately veined group of plants, as an order also characterized by
strong barkfibres, by stellate (or lepidote) trichomes and the very distinctive petiole apex
which is more or less swollen and pulvinar. Only Elaeocarpaceae and most Malvaceae lack
the typical pulvinus, the former distinctive in their foliaceous stipules, the latter in their
mostly herbaceous habit. Bixa is not usually included in Malvales but has a similar, though
shorter, pulvinus. The bark slashes of tree Malvales all tend to have a mucilaginous
105
3.3. Identification of the Tree Families of the Esquinas Forest
secretion which can be felt when fresh or seen as globules after a few hours. Although
recognition based on vegetative characters is easy, separation of the individual Malvalean
families without flowers is frequently problematic. When sterile, Tiliaceae, Sterculiaceae,
and simple-leaved Bombacaceae are reliably differentiated only by first knowing the
genera. Bombacaceae are all trees and the simple-leaved ones are entire (very weakly
sublobed in Ochroma); Malvaceae are mostly herbs and subshrubs, with the woody
species in the region serrate-margined; Tiliaceae (mostly serrate) and Sterculiaceae (trees
mostly entire except Guazuma) include both large trees and small weedy shrubs.
Bixaceae - Bixa, closer to Flacourtiaceae than Malvales, has a distinct apical pulvinus
similar to that of the Malvales, but shorter. It is also characterized by scattered,
reddish, peltate scales below (but lacks the typical Malvalean stellate trichomes).
Bombacaceae - The relatively few simple-leaved genera, exclusively large trees, are best
characterized by fused filaments, a feature shared in Malvales only with Malvaceae,
which differ in being mostly herbs and shrubs, and with some Sterculiaceae. The
only definitive difference from Malvaceae is the absence of spinulose pollen,
although the stamen tube often differs from Malvaceae in being fused only at base.
Elaeocarpaceae - The three genera with 3-veined leaves are distinctive in having the order
in persistent foliaceous stipules and in lacking the typical Malvalean pulvinus.
Malvaceae - Essentially the herbaceous counterpart of Bombacaceae with which they
share the distinctive feature of fused filaments. The most definitive difference is
spinulose pollen, a feature never found in Bombacaceae. Mostly differing from
Sterculiaceae and Tiliaceae in combination of more broadly ovate leaves with
serrate or lobed margins and from most other Malvales in less developed pulvinus.
Flowers distinctive by numerous stamens with filaments fused around style into
staminal column and/or an epicalyx.
Sterculiaceae - Tree genera differ from most Tiliaceae in being entire-leaved (or palmately
lobed or compound), except Guazuma which has leaves more jaggedly serrate than
in any Tiliaceae. Shrub Sterculiaceae (i.e. most Malvalean shrubs) have serrate
leaves (see Tiliaceae above for distinguishing characters). The flowers can have
fused or distinct filaments, the former differing from Malvaceae and most
Bombacaceae in having 2-celled anthers.
Tiliaceae - Most serrate Malvalean trees are Tiliaceae (see also Sterculiaceae Guazuma);
entire-leaved Tiliaceae (except Mortoniodendron) have the lower leaf surface
canescent, a character combination not found in simple-leaved Bombacaceae and
only in a few Theobroma species in Sterculiaceae (from which entire-leaved Apeiba
species can be differentiated by longer, more slender petioles). Flowers
characterized by multiple stamens arranged in single whorl and with free filaments.
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3.3. Identification of the Tree Families of the Esquinas Forest
D.4b. Petioles lacking apical pulvinus
Araliaceae - Characterized by leaves with rank odour and of dramatically different sizes,
and with petioles of various lengths. Three-veined species of Dendropanax differ
from Hernandia in having more or less wrinkled, usually tannish-drying twig bark
and main lateral vein pair straight or curving slightly outward rather than upward.
Oreopanax usually either epiphytic or with leaves palmately lobed and
conspicuously tannish-pubescent below.
Caricaceae - Some milky latex usually present.
Cochlospermaceae – Cochlospermum vitifolium, palmately lobed with serrate margins, a
combination unique among trees of this region.
Euphorbiaceae - Conspicuously 3-veined Euphorbiaceae mostly have glands at apex of
petiole (sometimes also with latex, see above) or at base of lamina (usually in axils
of basal vein pair below), or have stellate or peltate trichomes (Croton), or are
deeply palmately lobed.
Flacourtiaceae - Lunania mexicana, have a very characteristic pair of glands at petiole
apex (euphorbs with similar glands differ in having latex or leaves larger and more
broadly ovate).
Hernandiaceae - Leaves long-petioled and entire, usually rank-smelling (but the odour not
clearly Ranalean). Very similar to Araliaceae.
Rhamnaceae - Zizyphus chloroxylon, conspicuously 3-veined leaves.
Ulmaceae - Most taxa with pinnate venation, but Trema and Celtis have 3-veined alternate
leaves, the petioles always of equal lengths; the common Trema has asperous
leaves with fine, close-together teeth; Celtis is often spiny and has leaves with
coarse rather irregular teeth, but the most common erect species has entire leaves
recognizable by the noticeably asymmetric base that characterizes most Ulmaceae.
Urticaceae - Close to Ulmaceae but leaves usually with cystoliths in upper surface and/or
with stinging hairs, in tree taxa always serrate.
D.5. Strong bark
Pull off a leaf and see if a long strip of bark comes off with it. All neotropical species with
strong bark fibres have alternate, mostly simple, leaves, and this is a very useful character
for several families, some of them (e.g. Thymelaeaceae) otherwise nondescript.
Annonaceae - Keyed out above under plants with primitive odours; if odour not apparent,
can be identified by the strong, often greenish, twig bark, entire leaf margins, and
vertical fibre lines in a very shallow bark slash.
Lecythidaceae - Differs from other strong-barked families in bark of trunk peeling off in
layers rather than as single unit. Faint but characteristic "huasca" odour. Leaves
107
3.3. Identification of the Tree Families of the Esquinas Forest
nearly always with serrate or serrulate margins and distinctive secondary (and
usually intersecondary) veins that turn up and fade out at margins.
Malvales and Urticales - The Malvalean and Urticalean families, keyed out above on
account of 3-veined leaves, are also characterized by strong bark fibres, the entire
trunk bark peeling off when pulled (as opposed to peeling in layers in
Lecythidaceae).
Thymelaeaceae - Very distinctive in the thick homogeneous bark that strips as a unit from
entire twig; the only family with thick strong non-layered homogeneous bark.
D.6. Unequal petioles
Araliaceae - Leaves with rank vegetative aroma.
Capparidaceae - Petioles unequal only when leaves terminally clustered; leaves more
oblong and/or petioles more wiry than in other taxa with unequal petioles.
Euphorbiaceae - The combination of serrate leaf margins with conspicuously differentlength petioles having flexed apices is definitive for Euphorbiaceae; nonserrate
Sagotia racemosa with unequal petioles also have the flexed petiole apex.
Sterculiaceae – Sterculia, although Sterculia petioles are conspicuously unequal, the
genus is keyed out above on account of the Malvalean pulvinus.
D.7. Petiole glands present
Chrysobalanaceae - Some Chrysobalanaceae species have a pair of lateral glands at
extreme apex of petiole or at extreme base of leaf blade below; they can usually be
recognized by small inconspicuous stipules on young twigs.
Combretaceae - Most tree Combretaceae (except most Terminalia) have a distinctive pair
of glands on upper petiole surface, also characterized by leaves clustered at tips of
ascending short-shoot branchlets or branch tips.
Euphorbiaceae - All taxa with pair of glands near petiole apex have latex and/or are
conspicuously 3-veined (see above).
Flacourtiaceae - The genera of Flacourtiaceae with glands at apex of petiole have
conspicuously 3-veined leaves (see above).
Rosaceae - Prunus subcorymbosa, leaves have distinctive large ocellate glands near base
of lamina below and a strong odour.
D.8. Leaves with serrate (or serrulate) margins
Actinidiaceae - Numerous straight parallel secondary veins; surface frequently roughpubescent; petiole base not enlarged, unlike Sabiaceae; trichomes simple, unlike
Clethra.
108
3.3. Identification of the Tree Families of the Esquinas Forest
Celastraceae - Twig usually irregularly angled from decurrent petiole base and often
zigzag and/or greenish when fresh.
Clethraceae - Clethra mexicana, distinctive in the densely tannish-stellate tomentum of the
leaf undersurface, remotely serrate or serrulate, sometimes only toward apex.
Elaeocarpaceae - Some Sloanea spp. have remotely serrate or serrulate margins; they
are recognizable by the flexed petiole apex and tendency to have both opposite and
alternate leaves.
Euphorbiaceae - Only a few arborescent euphorbs have pinnately veined leaves with
eglandular equal petioles, lack latex, and have serrate margins. These very
nondescript taxa include Alchornea, Richeria (the margin only slightly crenulate,
leaves cuneate and petiole base slightly enlarged), and Acidoton, a shrub.
Fabaceae - The vanishingly few truly simple-leaved legume genera are generally
characterized by olive-drying leaves with serrulate margins.
Fagaceae - Characterized by clustered terminal buds with scales; round white lenticels.
Inconspicuously serrate or serrulate, cuneate to short petiole.
Flacourtiaceae - Many serrate (-serrulate)-leaved pinnate-veined Flacourtiaceae are
characterized by very small pellucid punctations; stipules are always present but
usuall, early caducous and leaving inconspicuous scar. Slightly zigzag twigs are
another frequent character. Xylosma lacks punctations but is frequently spiny.
Humiriaceae - Most genera (except entire Vantanea barbourii) have crenate margins and
festooned-brochidodromous venation. Young leaves at shoot apex rolled into
narrow cone, inner bark red or dark red.
Icacinaceae - Calatola costaricensis, have groove on top of the often somewhat twisted
petiole and black-drying leaves.
Lecythidaceae - Nearly all Lecythidaceae (keyed out above on account of their strong
bark) have serrate or serrulate margins.
Myrsinaceae - A very few mostly shrubby Myrsinaceae have finely serrate leaf margins;
like other members of the family they are characterized by the typical, usually
nonpellucid punctations (see below). Serrate Myrsinaceae can be differentiated
from similarly punctate Theaceae (Ternstroemia) by the more elongate punctations
that are pellucid in bud.
Ochnaceae - Always serrate or serrulate, usually with caducous stipules leaving annular
scar; three leaf types - one with secondary veins marginally curved and becoming
almost submarginal (with several of these marginal extensions paralleling each
other at a given point), or with close, rigidly parallel, secondary veins and finely
parallel tertiary veins perpendicular to secondaries.
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3.3. Identification of the Tree Families of the Esquinas Forest
Sabiaceae - Most Meliosma species have conspicuously serrate, or at least serrulate
margins, usually with numerous, fairly straight, secondary veins. Similar to
Saurauia, but more coriaceous and the petiole base thickened and often woody;
trichomes simple unlike Clethra.
Solanaceae - Leaves of many Solanum species distinctively irregularly, broadly, and
shallowly toothed, usually also with stellate or dendroid trichomes and/or prickles.
Symplocaceae - Symplocos limoncillo, characterized by festooned-brochidodromous
venation, the leaves usually loosely and rather irregularly reticulate below with
trichoms.
Theaceae - Gordonia, Pelliciera and Ternstroemia have more or less serrate leaves
(although this can vary even within a species), at least inconspicuously near apex.
Secondary venation often immersed and non-apparent. Ternstroemia (usually only
inconspicuously serrate near apex) has well-developed petiole but is distinctively
punctate with blackish glands.
Theophrastaceae - Clavija, consisting mostly of pachycaul treelets, always has narrowly
obovate to oblanceolate leaves, typically with strongly spiny-serrate margins; when
not obviously serrate-margined the margin usually distinctively cartilaginous or the
plant reduced to a small erect subshrub.
Violaceae - Very nondescript and often impossible to differentiate from Flacourtiaceae
vegetatively.
Stipules
present,
but
usually
caducous;
leaves
usually
membranaceous; Gloeospermum leaves often dry light green with a paler central
area.
D.9. Thickened and/or flexed petiole apices
Elaeocarpaceae - Sloanea, usually recognizable by the highly unusual mixture of opposite
and alternate leaves, most species also distinctive in the large, unusually thin
buttresses; a few large-leaved species have distinctive leafy stipules.
Euphorbiaceae - Many Euphorbiaceae have flexed petiole apices but most are 3-veined
and/or serrate and/or have latex and/or petiolar glands. A few entire-margined nonlactiferous Euphorbiaceae that lack petiolar glands have thickened or flexed
petioles; taxa that do not always have conspicuously different-lengthed petioles
include Sagotia.
Flacourtiaceae - A few flacourts (Carpotroche, Mayna) have flexed petiole apices and
more or less entire margins; their petioles tend to be shorter and/or less variable in
length than the above euphorbs.
D.10. Punctations
Flacourtiaceae - Some genera (Casearia, Xylosma) are pellucid-punctate, sometimes with
almost linear punctations, but the majority of their species are serrate-margined.
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3.3. Identification of the Tree Families of the Esquinas Forest
Usually have stipules or stipule scars, unlike Myrsinaceae which also differ in
having usually dark non-pellucid punctations.
Myrsinaceae - Punctations usually non-pellucid (except in bud), usually elongate, and
often reddish or blackish; associated with distinctive pale green "matte"
undersurface; stipules completely absent unlike Flacourtiaceae. Trichomes, when
present, nearly always more or less branched, unlike other punctate taxa.
Rutaceae - Only a few (mostly non-aromatic) genera have simple leaves, these almost
always narrowly obovate to oblanceolate with cuneate bases and clustered at
branch apices or at apex of pachycaul treelet.
Theaceae - Some punctate Theaceae (Ternstroemia) have entire leaves; these differ from
Myrsinaceae in having rounder punctations, which are blackish even in juvenile
leaves and buds.
D.11. Stipules present
Chrysobalanaceae - The stipules are typically small and inconspicuous and are usually
visible only on young twigs. Many Chrysobalanaceae have very characteristic
leaves with close-together, rigidly parallel, secondary veins and a whitish
undersurface, but Hirtella and some Licania species are very nondescript. The
whole family is usually recognizable by having red inner bark with a gritty-sandy
texture.
Dichapetalaceae – Dichapetalum, has conspicuous stipules, sometimes with a very
unusual serrate or fimbriate margin.
Erythroxylaceae – Erythroxylon, when present, faint venation lines paralleling the midvein
below are very typical; stipules triangular and brownish or tannish, often
longitudinally striate.
Euphorbiaceae - Several entire-margined nondescript euphorb genera have distinctive
more or less caducous stipules (e.g. Margaritaria with reddish slightly zigzag
puberulous twigs and conspicuous stipule scar, Sagotia with terminal stipule like
Moraceae falling to leave conspicuous scar).
Flacourtiaceae - Casearia and Lacistema, Lacistema have leaves usually subentire with
faint tendency to marginal serrulation; characterized by caducous stipule leaving
conspicuous scar, with both stipule and young twigs tending to dry blackish,
contrasting with the whitish stipule scar, and a very few nondescript Casearia
species are both entire and lack punctations; stipule scars are about their only
useful character.
111
3.3. Identification of the Tree Families of the Esquinas Forest
Rosaceae - Prunus subcorymbosa, entire-leaved, have early-caducous inconspicuous
stipules; they are usually recognizable by the pair of large dark-drying glandular
ocelli near base of lamina below.
D.12. Lepidote scales and/or stellate trichomes
Annonaceae - Duguetia and some Annona species have stellate trichomes or lepidote
scales but should key out above under primitive odour.
Capparidaceae - Capparis, conspicuous tannish scales in many species including some
that have uniform petioles; a characteristic patelliform gland just above the leaf axil
on young twigs is frequently apparent.
Clethraceae - Clethra mexicana, distinctive in its densely white-stellate leaf undersurface,
but most species more or less serrate or serrulate (see above). Its margin is entire
and differs from area Styrax in having longer, laxer arms on twig trichomes, and in
lacking the scattered rufescent trichomes of the more strongly reticulate leaf
undersurface.
Euphorbiaceae - Pinnate-veined euphorbs characterized by lepidote scales and/or stellate
trichomes include Hieronyma, Pera, some Croton.
Fagaceae - Margins usually more or less serrulate; when entire the mostly stellate
trichomes are a useful indicator.
Icacinaceae – Dendrobangia, leaves characteristically membranaceous and black-drying.
Malvales - Most Malvales have stellate trichomes or scales but are keyed out above by the
pulvinar petiole apex.
Styracaceae - Characterized by densely white-stellate or lepidote leaf undersurface,
usually also rufescent with reddish-stellate hairs, especially on twigs; similarly
densely white below. Solanaceae lack the rufous-stellate twig pubescence and are
usually spiny.
D.13. Leaves parallel-veined or lacking secondary veins
Gramineae - Bamboos have parallel-veined leaves, the plants distinctive in having
segmented, often hollow stems with characteristic swollen nodes.
Podocarpaceae - Leaves very coriaceous, linear-oblong with a strong midvein, completely
lacking secondary veins or with a few faint longitudinal veins paralleling midvein.
Zingiberales
-
Cannaceae,
Costaceae,
Heliconiaceae,
Marantaceae,
Musaceae,
Zingiberaceae; like most monocots parallel-veined, often very large leaves.
112
3.3. Identification of the Tree Families of the Esquinas Forest
D.14. Parallel tertiary venation
Lecythidaceae - Strong bark (the only combination of strong bark and parallel tertiary
venation). Brochidodromous genera lacking the family's typical upcurved veins and
serrate margins have tertiary veins closely parallel and perpendicular to midvein.
Myristicaceae - Compsoneura sprucei, the tertiary veins are conspicuously finely parallel
and perpendicular to the midvein and the primitive odour is not always apparent.
Olacaceae - Most genera have finely parallel tertiary venation more or less perpendicular
to the midvein or secondary veins; usually there is a small amount of latex in
petiole.
Sapotaceae - Many Sapotaceae have conspicuously parallel tertiary venation or Clusiatype venation. They have latex (and are keyed out above), it is sometimes not very
conspicuous, especially during periods of water stress.
D.15. Spines or spine-tipped leaves
Euphorbiaceae - The family that can have virtually any characteristic has only a few spiny
members, including spiny-trunked Hura (with latex).
Flacourtiaceae - Casearia and Xylosma sometimes have branch spines, the latter
sometimes with very striking branched spines covering trunk.
Moraceae - The only spiny Moraceae in the region are Maclura and Poulsenia, both with
milky latex (see above).
Nyctaginaceae - Pisonia actually has opposite leaves but in spiny taxa they are mostly
clustered on short shoots and the disposition is not evident.
Olacaceae - Ximenia americana, has branch spines and leaves clustered at lateral branch
tips, drying olive to blackish, usually retuse at apex.
Rhamnaceae - Ziziphus chloroxylon, densely branched, spiny branches.
Solanaceae - The small thick-based spines that characterize many species of Solanum are
actually prickles and may be present on leaves as well as on twigs and branches;
spiny members of the family have stellate trichomes (and are keyed out above), in
addition usually recognizable by the rank tomato-like odour of crushed leaves.
Urticaceae - The few spiny-trunked Urticaceae have serrate or incised leaf margins.
D.16. None of the above
Anacardiaceae - Anacardium and Mangifera, aromatic leaves.
Bignoniaceae - Crescentia and Amphitecna are totally unbignoniaceous vegetatively
having simple alternate leaves. Crescentia has the narrowly obovate leaves in
characteristic fascicles alternating along thick branches; Amphitecna has elliptic to
113
3.3. Identification of the Tree Families of the Esquinas Forest
obovate coriaceous leaves, poorly demarcated from woody based petioles, drying
greyish with pale secondary veins below.
Capparidaceae - Capparis that lack different-length petioles have the leaves 2-ranked,
usually with a raised patelliform axillary gland.
Celastraceae - A few lowland Maytenus species have entire leaves, these mostly drying
olive with paler inconspicuous secondary veins.
Chrysobalanaceae - Although small stipules are present, they are usually inconspicuous
and early-caducous; even if stipules not apparent, recognizable by the grittytextured, red inner bark.
Combretaceae - Leaves usually apically clustered; except for Terminalia, usually with
petiole glands. Alternate-leaved taxa typically with leaves clustered and pagoda
branching form or bark very smooth and white.
Dichapetalaceae - Tree Dichapetalaceae usually have serrate stipules (Dichapetalum) or
uniformly terete tannish-puberulous thickish petiole, usually in part with distinctive
scars from fallen petiole-borne inflorescence.
Ebenaceae - Tropical species with trunk slash characterized by black bark ring; leaves
typically with large darkish glands on lower surface usually scattered along (but
removed from) midvein.
Euphorbiaceae - A notoriously heterogeneous and difficult to recognize family. "Left-over"
genera include Drypetes (asymmetric base and prominulous tertiary venation),
Margaritaria (caducous stipules and a characteristic twig apex).
Fabaceae - Rare simple-leaved legumes have the entire petiole cylindrically pulvinate (cf.
pulvinulus of leaflets of compound leaves). They usually have asymmetric bases,
frequently serrulate margins, and dry a distinctive light olive-green; Lecointea are
often remotely serrate. Unifoliolate legumes (e.g. some Swartzia) are more common
and easy to recognize by the apical and basal pulvinular area of the 2-parted
"petiole" with typical cylindrical pulvinus of the single leaflet forming its apical part.
Flacourtiaceae - While Flacourtiaceae have stipules, these are often not very evident; a
few Casearia species with non-obvious stipules have entire margins and lack
punctations.
Humiriaceae - Vantanea barbourii, has coriaceous, more or less obovate leaves, usually
drying a dark reddish colour; young leaves rolled at shoot apex.
Icacinaceae - Groove on top of the often twisted petiole; Calatola, usually with at least a
few serrations, lack conspicuously parallel tertiary venation, black-drying.
Moraceae - Trophis usually lacks conical stipule and the latex is watery, not milky, but the
leaf venation is typically moraceous.
114
3.3. Identification of the Tree Families of the Esquinas Forest
Olacaceae - Usually characterized by curved (sometimes almost U-shaped) green petiole,
slightly thicker toward apex; most taxa have slight latex in petiole and/or finely
parallel tertiary venation.
Onagraceae - A few Ludwigia species are subarborescent; they are restricted to swampy
areas and have exfoliating reddish bark.
Polygonaceae - The whole family very easy to recognize by presence of an ochrea, an
irregular broken sleeve of stipular tissue that covers the node above petiole base.
Sabiaceae - A few Meliosma species are essentially entire; like their more numerous
serrate-leaved congeners they are recognizable by the thickened, sometimes
subwoody petiole base (cf. Sapotaceae).
3.3.3. Special habits and "spot characters" of some trees
With this list, in combination with the descriptions of the families and genera, it should be
possible to identify a number of trees exhibiting the most conspicuous vegetative
characters.
Ant domatia on petioles or leaf base
Fabaceae - Tachigali
Melastomataceae - Tococa, a few Clidemia
Ant domatia in hollow twigs (stems)
Cecropiaceae - Cecropia
Flacourtiaceae – Tetrathylacium
Lauraceae – Ocotea
Fabaceae – sometimes Ormosia, Lonchocarpus, Macherium, Senna and Inga
Moraceae - Ficus
Piperaceae - Piper
Polygonaceae - Coccoloba, Triplaris
Ant domatia in swollen thorns
Fabaceae - Acacia
Spines - Mostly useful at level of individual species.
Trees with white milky latex
Apocynaceae – Lacmellea, Stemmadenia, Tabernaemontana, Peschiera
Caricaceae - Carica
115
3.3. Identification of the Tree Families of the Esquinas Forest
Euphorbiaceae – Hura, Sapium
Moraceae – Castillea, Ficus, Brosimum, Batocarpus, Naucleopsis, Olmedia, Perebea,
Poulsenia, Pseudolmedia, Sorocea, Trophis
Olacaceae – Heisteria, Minquartia – at least in the petiole and youngest twigs
Sapotaceae – Chrysophyllum, Manilkara, Micropholis, Pouteria
Clusiaceae - Clusia
Trees with conspicuously-coloured latex
Apocynaceae – Aspidosperma
Clusiaceae – Calophyullum, Clusia, Garcinia, Symphonia, Tovomita, Vismia
Cochlospermaceae - Cochlospermum
Euphorbiaceae - Croton, Hippomane, Pausandra, Richeria
Fabaceae – Dussia, Machaerium, Pterocarpus, Swartzia
Myristicaceae – Compsoneura, Virola, Otoba
(black sap: Pourouma, Prioria)
Trees with spines on leaves, trunks, twigs, branch-tips or at ends of twigs
or short shoots
Apocynaceae – Lacmellea
Arecaceae – Acrocima, Astrocaryum, Bactris, Cryosophila, Raphia
Bombacaceae - Ceiba
Caricaceae - Jacaratia
Euphorbiaceae - Hura
Ferns (Tree ferns) - Cyathea, Alsophila
Flacourtiaceae - Xylosma, few Casearia
Fabaceae - Acacia, Erythrina, Machaerium
Moraceae - Maclura, Poulsenia
Olacaceae - Ximenia
Rhamnaceae - Zizyphus
Rubiaceae - Chomelia, Randia
Rutaceae - Zanthoxylum
Ulmaceae - Celtis
Urticaceae – Urera
Unbranched pachycaul growth form
Theophrastaceae – Clavija
Myrsinaceae – Cybianthus
116
3.3. Identification of the Tree Families of the Esquinas Forest
Sapindaceae - Talisia
Strongly fenestrated trunks
Apocynaceae –Aspidosperma myristicifolium
Celastraceae - Perrottetia sessiliflora
Dichapetalaceae – Stephanopodium costaricense
Fabaceae – Lecointea amazonica
Olacaceae - Minquartia guianensis
Rubiaceae – Guettarda sanblasensis, Macrocnemum roseum
Sapotaceae – Pouteria torta
Stranglers or hemiepiphytic shrubs or trees
Araliaceae – Schefflera, Oreopanax aff. capitatus
Burseraceae - Bursera standleyana
Cecropiaceae - Coussapoa
Clusiaceae - Clusia
Ericaceae – Cavendishia, Satyria, Psammisia ramiflora
Marcgraviaceae – Marcgravia, Marcgraviastrum subsessilie, Sarcoptera sessiliflora,
Souroubea
Melastomataceae – Blakea, Topobea
Moraceae - Ficus
Scrophulariaceae – Schlegelia
Flowers and fruits produced directly from the trunk or branches (cauliand ramiflory)
Bignonicaeae – Crescentia cujete, Amphitecna
Caricaceae – Carica caulifora
Clusiaceae - Garcinia
Flacourtiaceae – Carpotroche platyptera
Lecythidaceae – Grias cauliflora
Fabaceae – Swartzia, Zygia
Melastomataceae – Bellucia pentamera, Henriettea
Moraceae – Castilla tunu
Myrtaceae - Plinia
Sapotaceae - Pouteria
Sterculiaceae – Theobroma, Herrania purpurea
117
3.3. Identification of the Tree Families of the Esquinas Forest
Trees with stilt roots
Acanthaceae – Bravaisia integerrima
Arecaceae – Socratea exorrhiza, Iriartea deltoidea
Bombacaceae – Pachira aquatica
Cecropiaceae – Cecropia, Pourouma bicolor
Chloranthaceae – Hedyosmum
Clusiaceae – Clusia, Symphonia globulifera, Tovomita
Euphorbiaceae – Alchorneopsis floribunda
Moraceae – Ficus
Piperaceae – Piper
Rhizophoraceae - Rhizophora
118
4.1. Discussion – Floristics and Diversity
4. Discussion
4.1. Floristics and Diversity
The rainforests around the Golfo Dulce among the most species-rich plant communities in
Central America (QUESADA & al 1997, WEBER & al. 2001). Species diversity is a very
important indicator of the level of diversity of plant and animal communities (VARESCHI
1980). Tree species inventories at defined study sites and in minimum diameter classes
provide a reliable instrument with which measure the diversity level of a study site. Habitat
diversity gradients can be used to examine biogeographic relationships. Information about
the diversity of a given site is a prerequisite for effective conservation (NAMKOONG 1995).
Species richness is used as a measure of taxonomic diversity because it is a simple,
useful, widely used and widely understood parameter (GASTON 1996).
The clearest indication from the data on tree diversity is that tree diversity is much greater
in (neo)tropical forests than in the temperate zones (LATHAM & RICKLEFS 1993). Warm, wet
climates, which are now restricted to equatorial latitudes, predate the first angiosperms and
characterized immense landmasses throughout the angiosperm radiation (TERBORGH 1973,
MORLEY 2000).
4.1.1. Reasons for the high diversity of woody plants in the tropics
The number of woody species in tropical forests tends to increase with the rate of
precipitation, forest stature, soil fertility, rate of canopy turnover and time since catastrophic
disturbance, and decrease with seasonality, latitude and diameter at breast height. High
rainfall and low seasonality in the tropics favour two key groups of natural plant enemies
(insects and fungi) that are directly responsible for promoting high rates of densitydependent plant mortality (GIVNISH 1999). Host-specific pests reduce recruitment near
reproductive adults (the Janzen-Cornell effect) and produce hyper-diverse communities
(WRIGHT 2002). Tree diversity is also higher in well-drained forests (CAMPELL & al. 1986,
DUIVENVOORDEN 1996) and declines with altitude in tropical forests in the paleo- and
neotropics (AIBA & KITMAYA 1999, GENTRY 1988a, LIEBERMANN & al. 1996, LOVETT 1996,
TERBORGH 1992). Tree diversity also differs enormously from one tropical forest to another
and even from one habitat to another (Tab. 4.1), a fact which may provide clues to the
causes of tropical tree diversity. CONDIT & al. (2002) recognized that the forest in western
Amazonia (Yasuni National Park, Ecuador and Manu Biosphere Park, Peru) is very
diverse, but that the species composition changes very little over distances of more than
1,000 kilometres. The tree species counts in any one locale are high, but each locale turns
119
4.1. Discussion – Floristics and Diversity
Site
Country
Volcán Barva
Costa Rica
Altitude
[m]
2600
San Ramon
Costa Rica
1000
San Ramon
Costa Rica
Palo Verde Wildlife Refuge
Precipitation
[mm/year]
3300
1 ha
No. of
individuals /ha
654
4000
1 ha
486
42
DI-STEFANO, J.F. & al. (1995)
1000
4000
1 ha
764
67
Di-STEFANO, J.F. & al. (1995)
Costa Rica
100
2000
4 ha
875
68
HARTSHORN (1983)
Ft. Sherman
Panama
50
3000
1 ha
647
74
PYKE & al. (2001)
Nadkarni plots, Monteverde
Costa Rica
1450
2300
1 ha
585
76
HABER (2000)
Gamboa
Panama
75
2700
1 ha
485
78
PYKE & al. (2001)
BCI, Forest dynamics plot
Panama
40
2656
1 ha
425
91
PYKE & al. (2001)
San Ramon
Costa Rica
1000
4000
1 ha
436
94
WATTENBERG & BRECKLE (1995);
Penas Blancas
Costa Rica
750
2000
1 ha
406
104
HABER (2000)
Esquinas - coastal slope
Costa Rica
150
6000
1 ha
588
108
this study
Plot size
No. of
Literature
spp./ha
29
LIEBERMANN & al. (1996)
Corcovado
Costa Rica
100
3500
1 ha
354
108
HARTSHORN (1983)
San Gerado
Costa Rica
1150
n.o.
1 ha
489
115
HABER (2000)
Barra Colorado Island
Panama
40
2656
1 ha
No information
116
GENTRY unpubl. in FOSTER & HUBBEL
(1990)
La Selva Biological Station
Costa Rica
35
4000
2 ha
845
118
HARTSHORN (1983)
Esquinas - gorge
Costa Rica
100
6000
1 ha
482
121
this study
Esquinas - inland slope
Costa Rica
300
6000
1 ha
527
133
this study
La Selva
Costa Rica
100
4000
1 ha
551
149
LIEBERMAN & al. (1996)
Esquinas – ridge
Costa Rica
300
6000
1 ha
847
179
this study
Northeast Atlanic lowland swamp
Costa Rica
20
4000
16.4 ha
7134
225
WEBB & PERALTA (1998)
Noel Kempff, Las Torres 1
Bolivia
350
1500
1 ha
n.o.
40
SEIDL (unpubl.) in SMITH & KILLEEN (1995)
Amboró, Cerro Bravo
Bolivia
2500
1000
1 ha
n.o.
43
VARGAS (1995)
Estacion Biológica del Beni, Maniqui
Bolivia
200
1800
1 ha
n.o.
49
PALACIOS & al. (unpubl.) in SMITH &
120
4.1. Discussion – Floristics and Diversity
KILLEEN (1995)
Non-ridge forest, Nudo de Cajanuma, Loja
Ecuador
2900
3000
1 ha
478
59
MADSEN & ØLLGAARD (1993)
Belém, Igapó
Brazil
30
2760
1 ha
564
60
BLACK & al. (1950)
Ridge forest, Zamora-Chinchipe
Ecuador
2700
3000
1 ha
1273
67
MADSEN & ØLLGAARD (1994)
Noel Kempff, Las Torres 3
Bolivia
250
1500
1 ha
n.o.
76
ARROYO & al. (1994)
Belém, terra firme
Brazil
30
2760
1 ha
423
87
BLACK & al. (1950)
Alto Ivon, terra firme
Bolivia
200
1550
1 ha
649
94
BOOM (1986)
Transamazonica
Brazil
100
2000
1 ha
578
101
CAMPELL & al. (1986)
Ubatuba, Atlantic forest
Brazil
900
2000
1 ha
n.o.
136
TABARELLI & MANTOVANI (1999)
Pilón Lajas, Cumbre Pilón
Bolivia
300-1200
2550
1 ha
n.o.
146
SMITH & KILLEEN (1995)
Anangu
Brazil
300
3000
1 ha
734
153
KORNING & al. (1991)
Tambopata swamp
Peru
260
2000
1 ha
532
158
GENTRY (1988a)
Tambopata, upland sandy
Peru
260
2000
1 ha
567
162
GENTRY (1988a)
Río Xingu, terra firme
Brazil
200
2250
1 ha
567
162
CAMPELL & al. (1986)
Tambopata, alluvial
Peru
260
2000
1 ha
540
165
GENTRY (1988a)
Tambopata, terra firme
Peru
260
2000
1 ha
602
181
GENTRY (1988a)
Tambopata, lateritic
Peru
260
2000
1 ha
564
185
GENTRY (1988a)
Cabeza de Mono
Peru
320
3000
1 ha
544
185
GENTRY (1988a)
Cocha Cashu
Peru
200
2028
1 ha
650
201
GENTRY (1990)
Manu Park, terra firme
Peru
400
2500
1 ha
673
210
GENTRY (1988a)
Bajo Calima, Chocó
Colombia
75
8000
1 ha
675
258
FABER-LANGENDOEN & GENTRY (1991)
Mishana
Peru
100
3700
1 ha
858
289
GENTRY (1988a)
Yanamono (Explorama)
Peru
100
2500
1 ha
606
300
GENTRY (1988a)
Amazonian, Cuyabeno
Ecuador
260
3555
1 ha
693
307
VALENCIA & al. (1994)
Tab. 4.1. Tree diversity at different sites in the Neotropics (Central and South America)
121
4.1. Discussion – Floristics and Diversity
out to be much like the others in terms of species composition. In contrast, forests on the
Isthmus of Panama (and of course in Costa Rica – e.g. the Esquinas forest) change
dramatically in tree species composition from one site to the next. Forests just 50
kilometres apart in Panama are less alike than forests 1,400 kilometres apart in western
Amazonia. As a result of such high landscape variation, parts of Panama have as many or
even more tree species than parts of Amazonia. The Esquinas forest shares this high
diversity of landscape variation, and therefore the high beta-diversity, with Panamanian
forests.
An idea proposed by MCARTHUR (1972) which retains acceptance among many
contemporary ecologists is that tropical species, because of the diversity of the
assemblages in which they occur, must have either narrower niches or greater tolerance of
ecological overlap than their counterparts living in less diverse assemblages.
Another factor contributing to the high diversity in tropical forests is the “state of dynamic
equilibrium”, that may be subdivided into three phases: the gap phase, the building phase,
and the mature phase (WHITMORE 1978). It is widely recognized that continua of irradiance
conditions and microclimates exist in forests on many spatial scales, and that conditions at
a given site can be highly dynamic (CHAZDON 1988).
The biodiversity crossroads hypothesis suggests another reason for the high diversity in
the Esquinas forest: This model proposes that some forests are especially rich in tree
species because they stand at the intersection of different phytogeographic regions
(PITMAN & al. 2002, PRANCE 1994).
The diversity of tropical trees is one of the most extraordinary mysteries of tropical forests;
it has yet to receive a definitive explanation (LEIGH JR 1999).
122
4.1. Discussion – Floristics and Diversity
4.1.2. Life Form Distribution
The most common woody life forms in the forests are trees. Trees make up about 20 to
25% of all plant species in tropical forests (GENTRY & DODSON 1987). About 97.6% of all
woody plants (≥ 10 cm d.b.h.) in the Esquinas forests are trees (1 sp. cycad, 2spp. tree
ferns, 5 spp. palms and 293 spp. dicotyledonous), but woody lianas (1.8%) and groundrooting hemi-epiphytes (0.5%) also occur. In La Selva (Costa Rica), the structure and
diversity of plants (≥ 10 cm d.b.h.) in 11 one hectare-plots from 100 m up to 2,600 m above
sea level were investigated. In total, 92.8% were trees (90.1% dicotyledonous), 1.3%
palms, 1.4% tree ferns), 5% lianas and 2.2% hemi-epiphytes (incl. stranglers) (LIEBERMANN
& al. 1996). The abundance of life forms seems to vary over a gradient of humidity.
Dicotyledonous trees were the most dominant life form in all investigated plots in the
tropics.
Site
Coastal slope - Esquinas
Gorge - Esquinas
Inland slope - Esquinas
Ridge - Esquinas
La Selva
La Selva
La Selva
La Selva
La Selva
Amazonian, terra firme, Brazil
Altitude in
Trees
m
100
100
250
250
100
500
1,000
1,250
2,000
60
95.4
89.2
70.8
88.4
74.4
94.1
77.4
68.7
86.0
93.5
Palms
1.4
6.4
26.9
9.6
22.4
2.6
10.6
0.0
0.4
3.9
Tree ferns
0.0
2.3
0.2
0.0
0.0
0.5
9.4
31.3
11.7
0.0
Lianas
3.1
2.1
1.1
1.2
3.2
2.6
0.2
0.0
0.0
2.4
Hemiepiphytes
0.2
0.0
0.9
0.8
0.0
0.2
2.5
0.0
1.9
0.15
Total
588
482
527
847
473
425
530
610
477
662
Tab. 4.2. Percentage of the abundance of life forms at different sites in the neotropics (data from La
Selva from LIEBERMANN & al. 1996; Amazonian from MILLIKEN 1998)
The high representation of palms in the Esquinas forest (262 individuals; c. 10.7%) is
typical for many pluvial and wet rainforests in the neotropics (GENTRY 1988a). Palms are
disscused further in chaptermore in chapter 4.1.3.3).
Most species of tree fern are found in clearings of wet forests and rainforests, since they
are light-gap pioneers. They are rarely found in the tropical dry and tropical moist forests
(GOMEZ 1983). Tree ferns prefer the high-rainfall forests and extend from coastal areas to
these with subandine conditions (just below frost-line elevations) and tend to be found in
gaps. The diversity of species increases with increasing rainfall and temperature, while the
density of specific stands increases with a decrease in mean temperatures and an increase
in elevation (GOMEZ 1983). An exception in the Pacific lowlands is Metaxyea rostrata (a
tree fern with compact rootstalk), where stands with many individuals are observed (but not
investigated in this study, since stems do not have d.b.h. ≥ 10 cm) on ridges in the
Esquinas forest. The condition of the terrain and the humidity of the ridge and the coastal
123
4.1. Discussion – Floristics and Diversity
Wet-La Selva
Moist-Jauneche
Dry-Santa Rosa
hemi- & epiphytes
25
6
4
climbers
12
22
17
trees
22
28
21
herbs, shrubs, treelets
40
43
57
parasites, saprophytes
1
1
1
Fig. 4.1. Percentage of species of different life forms in Costa Rican florulas (after GENTRY &
DODSON 1987)
forest do not favour the occurrence of tree ferns and no individuals were counted. Tree
ferns occur in the gorge and on the inland slope due to the high humidity and dynamic
conditions of the steep terrain (11 individuals of Alsophila firma in the gorge and one
individual of Cyathea delgadii on the inland slope). Investigations in La Selva (Costa Rica)
show that tree ferns were rare at low elevations (< 1%), becoming more numerous above
750 m. Tree fern abundance was highest at 1,250 m, where they comprised 31% of the
stems (LIEBERMANN & al. 1985).
Ten individuals (9 spp.) of hemi-epiphytes were represented among the 2,444 individuals
(d.b.h. ≥ 10 cm). Hemi-epiphytes are very abundant in premontane, lower montane, and
montane rainforest especially at about 1,800 m above sea level, and they increase in
density and diversity as conditions get wetter and appear more luxuriant (HOLDRIDGE 1967,
WILLIAMS-LINERA & LAWTON 1995). Investigations of La Selva show that hemiepiphytes
(incl. stranglers) (d.b.h. ≥ 10 cm) occur from 500 m up to 2,300 m, with a peak at 1,000 m
(2.5% of all stems; 4 spp.) (LIEBERMANN & al. 1996). The increasing number of hemiepiphytes is concomitant with decreased numbers of free-climbing lianas (Gentry 1988b). It
seems that hemi-epiphytes replace free-climbing lianas in the wettest lowland forests
(GENTRY & DODSON 1987), sometimes which also appears to occur in the inland slope and
ridge, as well as in the Chocó pluvial forests (Gentry 1988b). The genus Ficus is a very
common strangler in many paleo- and neotropical forests. In the research plots of the
Esquinas forests we counted just 4 spp. of strangling Ficus, but also 5 spp. of the
124
4.1. Discussion – Floristics and Diversity
neotropical genus Clusia (Clusiaceae). The Clusiaceae is a very diverse family in the
Chocó (Columbia) (FABER-LANGENDOEN & GENTRY 1991, GENTRY 1988b), as it is in the
Esquinas forest.
Lianas are an integral and important part of the tropical forest ecosystem. In neotropical
forests lianas represent up to 20% of the plant individuals (GENTRY 1983a). Forty-two
percent of trees (d.b.h.≥ 10 cm) at San Carlos de Río Negro, Venezuela have lianas (PUTZ
1982). In the dry forests and Amazonian rainforests lianas are more abundant than in the
moist and wet forests and in the Esquinas (GENTRY 1988a,b, 1990; GILLESPIE & al. 2000,
HARTSHORN 1983). Most of the Amazonian sites reported by GENTRY (1988a) have a much
higher representation of large lianas (d.b.h. ≥ 10 cm) than the Esquinas sites, with 14 to 24
individuals and 10 to 17 species. At Bajo Calima (Chocó, Colombia), there are only 6 to 11
individuals of lianas (d.b.h. ≥ 10 cm) per site (FABER-LANGENDOEN & GENTRY 1991). In a
Braulio Carillo National Park study (LIEBERMANN & al. 1996) the lianas are shown to be very
abundant in the lowlands (2-3%) and absent above 1,000 m. The 44 individuals in the four
research plots belong to 22 spp. In the Esquinas the lianas are most common on the very
steep and well-drained coastal slope (3.1%). In the dry forest of Santa Rosa (Costa Rica)
there are 77 individuals of woody lianas (d.b.h. ≥ 2.5 cm) in 0.1 ha (GILLESPIE & al. 2000).
Because of their very slow diameter increase, the presence of thick lianas is considered an
excellent indicator of undisturbed forest (BUDOWSKI 1965).
4.1.2.1. Coastal slope
The coastal slope seems to have drier soil conditions than all the other Esquinas research
plots. As in other dry or moist forests the abundance of lianas (18 individuals -3.06%; 9
spp.) is higher than in wet forests, while the abundance of hemiepiphytes (1 individual) and
tree ferns (0 individuals) is lower. Just 16 trees bear hemi-epiphytes (2.7%) (HUBER in
prep.). The high abundance of thick lianas indicates a low dynamism.
4.1.2.2. Gorge
Due to the steep terrain, the subsequently high dynamic conditions and the high humidity,
the gorge shares features typical of montane forests (more individuals of tree-ferns and
hemi-epiphytes and fewer individuals of palms and lianas than in lowland wet forests),
which is reflected by the presence of 11 individuals of tree ferns (Alsophila firma), 10
individuals of lianas and just 31 individuals of palms. In the hectare-plots of La Selva
(Costa Rica) up to 750 m, between zero (0) to two tree ferns were found, whereas at 1,000
m about 50 individuals (9.4%) and at 1,250 m about 190 individuals (31% of the stems)
were counted (LIEBERMANN & al. 1985, 1996). Investigations by HUBER (in prep.) in the
Esquinas plots show that in the very wet and shady forest in the gorge 13% of the trees
125
4.1. Discussion – Floristics and Diversity
(d.b.h.≥ 10 cm) bear hemi-epiphytes but none of these had ground-reaching roots with
d.b.h.≥ 10 cm. In La Selva there were also no hemi-epiphytes up to 500 m but about 13
individuals at 1,000 m (LIEBERMANN & LIEBERMANN 1985, LIEBERMANN & al 1996). The
reason for these results is probably the very high dynamism.
4.1.2.3. Inland slope
As in the gorge, the steep terrain and the dynamic and climatic conditions of the inland
slope are favourable to the growth of tree ferns. One individual tree fern (Cyathea delgadii)
was counted, and just 6 individuals (1.13 %) of lianas. Due to the high succession rate in
the steep plot there were many gaps (WEISSENHOFER 1996) and fewer lianas than in the
other plots in the Esquinas. The five hemi-epiphytes (1%) on the inland slope represent
another characteristic of tropical premontane rainforests.
4.1.2.4. Ridge
The ridge shows features typical of tropical wet lowland forests. The high abundance of
palms (c. 10%), the abundance of lianas (c. 1.2%) and the absence of tree ferns are all
characteristics of tropical wet forest in the lowlands. The abundance of hemi-epiphytes (7
individuals, 0.8%) and the abundance of lianas reflect stabile conditions, such as occur in
many lowland forests on flat terrain or slender slopes.
4.1.2. Species and Family Diversity of the Research Plots
Diversity, as measured by species richness, is very high in the Esquinas forest. The four
investigated plots in the Esquinas belong to different habitats (slope – inland and coast,
ridge and gorge). The ridge, with 179 spp. and 847 individuals, is the most diverse plot in
the Esquinas. Plot diversity in African forests shows that stem diversity is greater on ridge
tops than valley sites (LOVETT 1996) and denser forests are more diverse than sparser
forests (ARTES & al. 2003).
The investigated plots in the Esquinas forest belong to the most species-rich forest of
Central America and the plot on the ridge is the most diverse investigated plot in Central
America (Tab. 4.1). In the four research plots a total of 328 spp. of plants (d.b.h. ≥ 10 cm)
were found i.e. about half of all suspected tree species of the Golfo Dulce region.
Unpublished investigations of ZAMORA & AGUILAR. in the forests around the Golfo Dulce
(Tab. 4.3) reflect the high tree diversity of this region.
In Central America the most species-rich forests are in the wet Pacific and Caribbean
lowlands of Costa Rica. The world’s most species-rich forests are in South America. The
126
4.1. Discussion – Floristics and Diversity
most species-rich forest plots are on the foothills of the Amazonian Andes (Peru, Ecuador
and Colombia) and in the Chocó (Colombia), with up to 307 spp. per ha (Tab. 4.1).
Site
Bahia de Chal
Agujas
Sirena
Esquinas
No. of individuals
1161
1098
881
871
No. of species
201
189
149
188
Tab. 4.3. Diversity of plant species in one-hectare plots (d.b.h. ≥ 5 cm) in the Golfo Dulce rainforests
(ZAMORA & AGUILAR unpubl.)
4.1.3.1. Species Diversity
4.1.3.1.1. Importance of species and Importance Value (IVI)
Due to the combination of three indices the IVI (importance value index) gives a very good
idea about the importance of certain tree species. The most important species on the
coastal slope are the Moraceae Brosimum utile (IV 24.17) and the Clusiaceae Symphonia
globulifera (IV 18.82), and on the inland slope the palm Iriartea deltoidea (IV 20.90) and the
Moraceae Brosimum utile (IV 19.01) are the most important. Iriartea deltoidea is also a very
important tree in the Amazon region of Ecuador (IVI 26.87; KORNING & al. 1991).
In the gorge the Fabaceae Dussia discolor (IV 17.48) and the Flacourtiaceae
Tetrathylacium macrophyllum (IV 15.42) and on the ridge two species of Vochysiaceae,
Vochysia ferruginea (IV 17.26) and Qualea paraensis (IV 16.96) are the most important
trees. As we can see, no single tree species dominates one plot. Only Brosimum utile is an
important tree in two plots (coastal and inland slope). Brosimum utile is a common tree in
the wet region of the Pacific lowlands of Costa Rica and is found mainly on well-drained
slopes and ridges (HARTSHORN 1983).
4.1.2.1.1. 1. Relative Density
As in many other highly diverse lowland forests, which are moist or wet, no single species
is dominant. However the palms Iriartea deltoidea and Welfia regia are very common trees
(see chapter 4.1.3.3) and the Clusiaceae Symphonia globulifera (79 individuals) and the
Myristicaceae Compsoneura sprucei (72 individuals) occur in all investigated plots. The
densities of the investigated plants in the plots are shown in chapter 3. The ten most
abundant spp. constitute about 27% of all investigated individuals (3% of all spp.) in all
plots. The ten most abundant spp. on the coastal (9.3% of all spp.) and inland (7.5% of all
spp.) slopes account for 45% of the individuals; in the gorge this is 43% (8.3% of all spp.)
and on the ridge 36% (5.6% of all spp.). These results are comparable with other
investigations in species-rich forests (FABER-LANGENDOEN & GENTRY 1991, LIEBERMANN &
al. 1996, MADSEN & ØLLGAARD 1994, MILLIKEN 1998).
127
4.1. Discussion – Floristics and Diversity
In the species-rich Amazonian rainforest of Anangua, the ten most abundant species per
hectare constitute 50% of the individuals and 6.5% of all species (total 153 spp.) (KORNING
& al. 1991). In the Anangua plot two species, Quararibea ochrocalyx and Iriartea deltoidea,
constitute 39.9% of all individuals. In the lowland wet forest of La Selva (Costa Rica) the
Fabaceae - Caesalpinioideae Pentaclethra macroloba dominates with 14.4% to 17.3% of
all stems in all types of virgin forest. In the dry forest of Palo Verde the Fagaceae Quercus
oleoides dominates with 34% and in the mountain rainforests of Costa Rica (Cerro de la
Muerte) the Quercus costaricensis dominates with about 41% (HARTSHORN1983).
Dominance of one single species is found in the Esquinas forest only in special habitats
such as mangroves (Rhizophora mangle) or in secondary growth (Cecropia obtusifolia or
Trichospermum grewiifolium).
More obvious is the great abundance of rare species. Many such rare species are found in
tropical lowland rainforests (ASHTON 1989) as well as in the Esquinas forest. 136 spp.
(54%) out of 252 spp. at Bajo Calima (Chocó, Colombia) were only recorded in a single
occurance (FABER-LANGENDOEN & GENTRY 1991). In the transect of the northern slope of
the Vulcan Barva, Costa Rica (LIEBERMANN & al. 1996) and in the premontane forest of the
Cordillera Tilaran, Costa Rica, 36.2% and 36% respectively (WATTENBERG & BRECKLE
1995) of all counted spp. were represented by only one individual. In the gorge nearly half
(c. 49%) of all spp. were recorded from a single occurence and on the coastal slope this
figure was 35%. The rare species play a major role in the diversity in tropical forests. The
high percentage of tree species which are represented by only one individual per hectare
supports the previous observation concerning highly dispersed tree species. It indicates a
high heterogeneity level of forest stand (LAMPRECHT 1986) and supports the assumption
that the number of species recorded in the Esquinas would notably increase if the size of
the study plots increased (see chapter 4.1.3.).
4.1.2.1.1.2. Relative and Absolute Frequency
The frequency values of the most abundant tree species indicate an almost even
distribution throughout the plots. Just three spp. occur in all subplots and two in more than
90% of all subplots on each site (see chapter 3).
The inland slope might easily be taken for a palm garden. The three species with the
highest relative frequency are the palms Iriartea deltoidea – 5.09% (100% abs. freq.),
Welfia regia – 4.83% (95% abs. freq.) and Socratea exorrhiza – 3.56% (70% abs. freq.). In
the premontane rainforest of the Cordillera Tilaran, Costa Rica, only one species (Iriartea
deltoidea) occurs in more than 90% (5.68% rel. freq.) of all subplots (WATTENBERG &
BRECKLE 1995).
128
4.1. Discussion – Floristics and Diversity
In the gorge, Tetrathylacium macrophyllum (3.52% rel. freq.) and Rinorea dasyadena
(3.52% rel. freq.) occur in all subplots. Tetrathylacium macrophyllum occurs more
frequently in clumps near the stream.
No one species occurs in all subplots on the ridge and the coastal slope. On the ridge the
palm Welfia regia also shows a tendency to occur in clumps.
In general, clumping is not evident within our samples, because such clumping would result
in species being ranked differently according to their density and frequency in the sample,
and this is mostly not the case. The frequency results are similar in other highly diverse
forests in the neotropics which indicate that just one or very few spp. occur in nearly all
subplots (HARTSHORN & HAMMEL 1994, KORNING & al. 1991, MADSEN & ØLLGAARD 1994,
MILLIKEN 1998, SMITH & KILLEEN 1995).
4.1.2.1.1.3. Relative Dominance
The basal area of the coastal plot (43.47 m2) is the largest due the abundance of thick and
tall trees (WEISSENHOFER in prep.). The other plots have similar basal areas (35.53 to 39.81
m2).
On the coastal and inland slopes the most dominant species is Brosimum utile. On the
coastal slope especially, there are more than twice as many individuals (21) of Brosimum
utile than of the second most dominant species. On the coastal slope about 6 species
account for 50% of the basal area, but in the gorge (9 spp.) and on the ridge and inland
slope (11 spp. each) this number is higher. In the gorge, inland slope and ridge no one
species reaches the dominance of Brosimum utile on the coastal slope. A ranking of the
species according to their basal area or relative dominance shows a pronounced
discrepancy with the ranking by density. A high ranking in basal area may be achieved
either by many average-sized or small individuals or by a few very large individuals. In the
gorge Billia colombiana, with just 3 individuals, has a greater basal area and relative
dominance than the 37 individuals of Rinorea dasyadena. Thick trees on the ridge and
coastal and inland slopes are often found to be Brosimum utile.
In the Esquinas rainforests the plots are on slopes and ridges, and so the basal area is
much higher (35.53 to 43.47 m2 per hectare) than that of the La Selva (Costa Rica) flat
lowland plots (23.49 to 27.06 m2 per hectare) (HARTSHORN & HAMMEL 1994, LIEBERMANN &
al.1996), and the Amazonian lowland forest plots (22.2 to 34.1 m2 per hectare) (KORNING &
al. 1991, MILLIKEN 1998). In dry forests of Central America (17.7 to 23.3 m2 per hectare;
d.b.h ≥ 2.5 cm) and in secondary forests in La Selva (c. 10.7 m2 per hectare), the total
basal area is also smaller than in primary rainforests of Central America. In the montane
forests the basal area increases. In upper montane rainforests of an Ecuadorian ridge
forest (2,700 m) the basal area is 39.5 m2 per hectare (MADSEN & ØLLGAARD 1994). In
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4.1. Discussion – Floristics and Diversity
Costa Rica HEANEY & PROCTOR (1990) reported the highest basal area of 51.2 m2 per
hectare near the summit of Volcán Barva (2,600 m).
4.1.3.1.2. Species Diversity Indices
The number of species present in vegetation (its richness) and the way in which individuals
are distributed among species (its evenness) are often combined into a single index to
represent both these characteristics (its diversity). There are many different indices to
demonstrate and compare the richness, evenness and diversity of different sites.
The Simpson diversity index (D) gives very similar results within the Esquinas plots (1-D =
between 0.965 and 0.981). Other tropical montane rainforests with lower species richness
(75 and 90 spp.) in Ecuador have a Simpson’s index of 0.97 and 0.96 (MADSEN &
ØLLGAARD 1994). Forests in the Amazon region of Ecuador with a very high richness of 153
species have 0.92 (KORNING & al. 1991). STOCKER & al. (1985) concluded that although
commonly applied, Simpson’s index is inappropriate to measure diversity in species-rich
tropical rainforests because it only represents evenness, and contains little information
about richness.
The Shannon-Weaver diversity index (H’) contains information about both richness and
evenness and appears to be the most appropriate diversity measure for use with highdiversity forests (STOCKER & al. 1985). With this index it is possible to compare the diversity
of forests with the different sizes of research plots.
The H’ index shows the same trend as species richness. The plot on the ridge with 179
species (847 individuals) has the highest H’ with 4.476. The gorge with 121 species (482
individuals) gives an H’ index of 4.122. This is slightly higher than the inland slope index of
4.119 (133 species, 527 individuals). The different of the species abundance in the gorge is
less than in all other plots (Fig. 3.5) (Evenness E 0.859) and it therefore has a higher H’
index (Fig. 3.12). The coastal slope with 108 species (588 individuals ) shows the lowest
value of H’ at 4.014 because species richness here is less than in the other plots. The
Evenness index for the inland slope is 0.841 less than in the other plots. This is therefore
due to the heterogeneity of abundance of the species (Fig. 3.9).
In the Caribbean lowlands of Costa Rica the Shannon-Weaver diversity index ranges from
H’ = 2.556 (2,600 m above sea level – 29 spp. and 645 individuals) to H’ = 4.508 (300 m
above sea level – 551 individuals and 149 spp.) and the Evenness (E) ranges from 0.759
to 0.901 (LIEBERMANN & al. 1996). The highly diverse montane rainforests in Ecuador range
from H’ = 4.2 to 4.26. Investigations in the watershed of the Panama Canal (lowland forest)
in 40 plots (each 1 ha) show about 800 spp. (d.b.h. ≥ 10 cm) in total which represent a
regional flora with exceptional β-diversity. The Shannon-Weaver diversity index (H’) ranges
from 2.65 to 4.06 (PYKE & al. 2001). Investigations in the tropical lowland rainforests of
130
4.1. Discussion – Floristics and Diversity
North Queensland (Australia) show H’ = 2.4 to 3.6 (STOCKER & al. 1985). This comparison
with other tropical rainforests emphasizes the high diversity of trees in the Esquinas forest.
The range found in Fisher’s α in the Esquinas was from 38.82 (coastal slope) to 70.49
(ridge) which demonstrates the species richness of the plots. Fisher’s α index ranges in
these plots from 16.15 to 54.76 (PYKE & al. 2001). The high species richness in plots (1 ha)
of Western Amazonia and on the terra firme forests of Central Amazonia is visible in the
average Fisher’s α index, which ranges from 99.7 to 126.3. The diversity of tree species in
Eastern Amazonia (average α = 20.8 to 56.0) and in the forests on the Guiana Shield (α =
7.5 to 51.9) is mostly lower than that in the Esquinas plots. The plot with the highest
species richness (322 spp. per hectare) and a Fisher’s α index of 221.18 is located in
Western Amazonia (STEEGE & al. 2000 and 2003). As we can see, the forests with the
highest diversity in tree species in the neotropics, and probably in the world are located in
Western Amazonia (Ecuador).
Gentry (1990) assumed Central American forests to be much poorer in tree species than
Amazonian ones. As we can see, this is true for the Western Amazonia forests but not for
the Central and Eastern Amazonian forests (Tab. 4.1).
The results of this study and comparisons with other regions show that the Central America
forest with the highest diversity in tree species is probably the Esquinas forest.
4.1.3.2. Reasons for the high tree species diversity in the Esquinas forest
Mature (old growth) primary forests have the greatest species richness (PYKE & al. 2001).
In this study only plots in primary forests were investigated.
Denser forests are more diverse than sparser forests (STEEGE & al. 2003). The ridge with
847 individuals belongs to the densest forests.
The Esquinas forest has strong spatial structure at the landscape scale and has therefore
many microhabitats and niches. Highly dynamic forests on a geographical gradient
contribute to the maintenance of high diversity, which is what happens in the Esquinas
forest.
The forest is said to be in a “state of dynamic equilibrium” that may be subdivided into three
phases: the gap phase, the building phase, and the mature phase (WHITMORE 1978). Gap
formation, and successive development in gaps is very important for high diversity,
especially in highly dynamic forests (WHITMORE 1975, 1978, GRUBB 1977, DENSLOW 1980,
1987, HUBBELL & FOSTER 1986, HUBBELL & al. 1990). The Esquinas forest is very dynamic
and therefore all phases between gap and climax were found in all plots.
High annual precipitation levels without a dry season support a high tree diversity.
Investigations in the watershed of the Panama Canal show that the highest diversity of
trees is found in the forests with the highest precipitation levels during the dry season
131
4.1. Discussion – Floristics and Diversity
(PYKE & al. 2001). Strong seasonality with a dry season is a strong predictor of tree density
and of maximum tree α-diversity Amazonia (STEEGE & al. 2003). Precipitation in the
Esquinas is high (c. 6,000 mm) and there is no significant dry season.
High edaphic heterogeneity and diversity of soil conditions support high β- plant diversity
(CONDIT & al. 2002). PAMPERL (2001) observed a high level of soil heterogeneity in the
research plots.
The biodiversity crossroads hypothesis proposes that some forests are especially rich in
tree species because they stand at the intersection of different phytogeographic regions
(PITMAN & al. 2002, PRANCE 1994).
Distribution patterns from the Esquinas forests show a strong affinity to the Chocó
phytogeographical region (coastal Colombia and adjacent Ecuador). The Chocó has one of
the world’s highest diversities of plants (GENTRY 1982b, 1986).
4.1.3.3. Family Diversity
Family diversity follows the same pattern as species diversity. Fifty-one families were
counted in the most species-rich plot on the ridge. While on the species-rich ridge the
mean is 3.51 species per family, in the gorge, where 121 species out of 46 families were
found, the mean is 2.63 species per family. These results are due to the many species out
of Fab.-Mimosaceae (12 spp.), Moraceae (12 spp.) and Clusiaceae (13 spp.) that were
found on the ridge. In the gorge (121 spp.), however, the most diverse family, the
Moraceae, was represented by just 9 species.
Tab. 4.4. shows the total values for the Esquinas sites and compares them with other
selected localities in the Neotropics. The Esquinas plots fall within the range of neotropical
lowland forests. The plot on the ridge with 51 families and 116 genera shows a very high
diversity of families compared with the other sites. The research plot at 300 m above sea
level at La Selva (55 families) shows the highest diversity in Central America (LIEBERMANN
& al. 1996). All plots in the South of Costa Rica with plants d.b.h. ≥ 5 cm have more
families (ZAMORA unpubl.) than the Esquinas plots plants with d.b.h. ≥ 10 cm. Obviously,
there are many more plant families that do not reach d.b.h. ≥ 10 cm in the Esquinas. The
most diverse forest in terms of families is also in the lowlands.
In total there are more plant families in Central America than in Amazonian forests. In La
Selva there are 121 families and in the Ducke forests (Brazil, near Manaus) there are 104.
The Manaus area flora is richer in trees than that of La Selva (Costa Rica) or Barro
Colorado Island (Panama) (GENTRY 1990).
In total, 69 families occur in all four plots in the Esquinas. In the 14 plots on the northern
slope of the Vulcan Barva from 30 m at the base to 2,600 m above sea level near the
132
4.1. Discussion – Floristics and Diversity
summit, 91 families were counted. This result reflects the high diversity of the forests in
Costa Rica.
Site
Country
Belém (Igapó)
Brazil
No. of No. of
No. of No. of
Source
trees families genera species
10 cm 564
28
51
60
BLACK & al. 1950
Río Colorado
Bolivia
10 cm
588
31
64
78
SMITH & KILLEEN 1995
Belém (terra firme)
Brazil
10 cm
423
31
65
87
BLACK & al. 1950
Alto Ivon (terra firme)
Bolivia
10 cm
649
28
61
94
Boom 1986
Transamazonica
Brazil
10 cm
578
30
77
101
CAMPELL & al. 1986
La Selva – 100 m
Costa Rica
10 cm
475
45
115
LIEBERMANN & al. 1996
Cumbre Pilón
Bolivia
10 cm
647
37
146
SMITH & KILLEEN 1995
La Selva – 300 m
Costa Rica
10 cm
551
55
149
LIEBERMANN & al. 1996
Río Xingú
Brazil
10 cm
567
33
162
CAMPELL & al. 1986
Esquinas Forest
Costa Rica
10 cm
527
39-51
Central Amazonian
Brazil
10 cm
645
34
95
201
MILLIKEN 1998
Anangua
Ecuador
10 cm
728
53
141
228
BASLEV & al. 1987
Mishana
Peru
10 cm
842
50
275
GENTRY 1988b
Yanamono
Peru
10 cm
580
58
283
GENTRY 1988a
Amazonian
Ecuador
10 cm
693
46
138
307
VALENCIA & al. 1994
Sirena, Corcovado
Costa Rica
5 cm
881
58
117
149
unpubl. ZAMORA &
AGUILAR
Agujas, Osa
Costa Rica
5 cm
1098
65
139
189
unpubl. ZAMORA &
AGUILAR
Bahia de Chal, Golfo Dulce Costa Rica
5 cm
1161
55
141
201
unpubl. ZAMORA &
AGUILAR
Esquinas
5 cm
871
62
144
188
unpubl. ZAMORA &
AGUILAR
Costa Rica
d.b.h.
92
83
79-116 108-179 this study
Tab. 4.4. Comparison of one-hectare forest inventories established in humid lowland tropical forests in
Central and South America
Central American and Amazonian tree communities also show differences in their floristic
makeup, although the variations in diversity are more striking than the differences in familial
representation. Leguminosae are the most diverse family in all lowland neotropical forest
plots. In Central Amazonian and Central American sites the Burseraceae, Sapotaceae,
Lauraceae and Chrysobalanaceae, are along with Leguminosae and Moraceae, the most
diverse families. In Central America, the Chocó and in the Esquinas forest the Clusiaceae
was among the most diverse families in all plots. The Lecythidaceae, Vochysiaceae and
Myrtaceae in Amazonia are very important and diverse families (PRANCE 1990); in Central
America and the Esquinas just a few species from these families occur in the plots.
Angiosperms dominate tropical forests (PRANCE 1977), especially the tree flora. Two
species of tree ferns grow in the Esquinas plots, as well as one gymnosperm (Podocarpus
guatemalensis); monocotyledons are represented by palms.
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4.1. Discussion – Floristics and Diversity
Leguminosae
The most common family in all Central American and tropical South American lowland
forests are the legumes (WEBB & PERALTA 1998, STEEGE & al. 2000, PRANCE 1990). In the
Amazon the abundance of Leguminosae is twice as high as those of the two next-most
abundant families (Arecaceae and Lecythidaceae) (STEEGE & al. 2000). Data from Reserva
Ducke (near Manaus, Central Amazon) listed the legumes (43 spp.) after the Sapotaceae
(48 spp.) (PRANCE 1990). The Leguminosae are always the most diverse family (up to 49
species) in the Brazilian Atlantic forest plots (TABARELLI & MANTOVANI 1999).
In mountain rainforests the Leguminosae are less diverse. Thirty different families were
counted in two plots (1 ha each) in the Andean region of Southern Ecuador (at 2,700 m and
2,900 m above sea level), although the Leguminosae were not among them (MADSEN &
ØLLGAARD 1994). The same phenomenon it was observed in Central America, on the
northern slope of the Vulcan Barva. The Leguminosae were absent in the plots over 1,750
m above sea level, but they were one of the most abundant families on the lowland plots
(LIEBERMANN & al. 1996).
Leguminosae also dominate other Central American lowland forests. In the swamp forest in
northeastern Costa Rica the Leguminosae are the most diverse family (36 spp. out of a
total of 225 spp.) (WEBB & PERALTA 1998). In the tropical dry forests in Central America the
Fabaceae are the dominant tree and shrub family at most sites (GILLESPIE & al. 2000). By
comparison, the somewhat drier and more seasonal moist forests of BCI (Barro Colorado
Island) and Central Amazonia contains more species in Leguminosae, which tend to be
more xerophilous (HAMMEL & GRAYUM 1982).
Clusiaceae
In the Amazon and Atlantic lowland forests of Brazil and in the premontane forests of
Bolivia the Clusiaceae are either almost absent, or are less diverse and a less important
family (MILLIKEN 1998, SMITH & KILLEEN 1995, TABARELLI & MANTOVANI 1999). On the other
hand, in the mountain rainforests of southern Ecuador the Clusiaceae are an important tree
family (MADSEN & ØLLGAARD 1994). On many Central and South American mountains the
dominant trees near the timber line belong to Clusia (pers. observation). On an altitudinal
gradient in the Andes, Lauraceae replace Leguminosae as the most species-rich family at
intermediate elevations. Guttiferae (Clusiaceae) also contribute to the diversity of middle
elevation forests. The wet lowland forest of the Chocó has features and taxa, such as the
Clusiaceae, more characteristic of upland forests (GENTRY 1988a). This is another indicator
of the biogeographical relationship between the Esquinas forests and the Chocó.
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4.1. Discussion – Floristics and Diversity
Arecaceae
Palms (Arecaceae d.b.h ≥ 10 cm) are among the most abundant species in many other
neotropical rainforests (BOOM 1986, BASLEV & al. 1987, CAMPELL & al. 1986, GENTRY
1990). The great abundance of palms (262 individuals – 27%) reflects the vegetation of
lowland tropical wet forest. In the Esquinas, the most abundant species are the palms
Welfia regia (108 individuals) and Iriartea deltoidea (97 individuals). In Costa Rica
HARTSHORN (1988) reported high numbers of tree palms, especially in La Selva. Tree
palms make up 26 percent of the individuals (≥ 10 cm d.b.h.) in La Selva (LIEBERMANN &
LIEBERMANN 1985). Here a great abundance of palms was found in the lowlands (25.5% of
all stems), but they were very rare (< 1%) above 1,000 m (LIEBERMANN & al 1996).
Investigations on the Vulcan Barva show that the abundance of palms decreases with
altitude. Palms are very abundant in the lowlands and prefer wet conditions. The palm
Welfia regia is the most common tree in La Selva (LIEBERMANN & al. 1996) and, along with
Iriartea deltoidea, it is also dominant in the lowland rainforest of Los Katios, Colombia
(ZULUAGA-RAMIREZ 1987). It requires a habitat with relatively high rainfall and is thus totally
absent from dry forests (VANDERMEER 1972 & 19834). In the Esquinas, 27% of all
individuals on the wet inland slope are palms, but on the drier coastal slope just 1.3% are
palms. Investigations in Amazonia show a high density of understorey palms in moist
Amazonian rainforests. They also show that, in the western Amazon and in floodplain
forest in the eastern Amazon the palms are the family with the greatest number of
individuals (KHAN & DE CASTRO 1988). However, they are less important in the Guiana
Shield, in central and eastern Amazonia (TER STEEGE & al. 2000) and in tropical dry forests
in Central America (GILLESPIE & al. 2000).
In the lowland rainforest in Beni (Amazonian region of Bolivia) the palm Iriartea deltoidea
also dominates (SMITH & KILLEEN 1995). Along with Socratea exorrhiza (40 individuals),
Iriartea deltoidea and Welfia regia trees seem to be especially abundant in those forests in
which small light gaps are frequently formed, that is, where the turnover rate of light gaps is
high (VANDERMEER 1983). In the Esquinas forest the Welfia trees are very abundant in the
light, dynamic forest types on the ridge (unpubl. PRINZ & al. 2002) and on the inland slope,
but almost absent on the coastal slope and rare in the gorge. Iriartea deltoidea is very
abundant on the inland slope (71 individuals) and also common in the gorge (unpubl. PRINZ
& al. 2002), but rare on the ridge and absent on the coastal slope.
Especially common are stilt-rooted palms (Iriartea deltoidea, Welfia regia, Socratea
exorrhiza). These palms are much more shade-intolerant (pioneers) and must grow quickly
on an inverted cone to take advantage of canopy openings (HARTSHORN 1983).
4
Due to insufficient investigations, VANDERMEER noted that Welfia georgii ( = W. regia) is rare in the southern
Pacific lowlands.
135
4.1. Discussion – Floristics and Diversity
4.1.3.2.1. Family Importance Indices (FIVI)
The important families differ between the research plots and, interestingly, no one family
dominates all four plots. The Moraceae, however, which is the most important family on the
coastal slope, is also important in the other plots. The Clusiaceae are also important on the
ridge and on the coastal and inland slopes. The Vochysiaceae is the most important family
on the ridge, but not one of the ten leading families in the gorge or on the coastal slope. In
the gorge the Tiliaceae is the most important family and on the inland slope the Arecaceae
is the leading family. In general we can see that the ten most important families hold nearly
two thirds of the total FIVI in all plots, and if we count the Leguminosae (Mimosaceae,
Caesalpiniaceae and Fabaceae) in the gorge as a single family, they would be the most
important, as they are in many Amazonian forests (KORNING & al. 1990, MILLIKEN 1998).
4.1.2.1.1. Relative Diversity
The Moraceae is the most diverse family in the gorge (7.44%), coastal slope (9.26%) and
ridge (7.73%). Fourteen spp. occur on the ridge. Naturally, if we were to count the
Leguminosae (Mimosaceae, Caesalpiniaceae and Fabaceae) as single family, it would be
the most diverse family in each plot. The Esquinas plots follow the same trends as those in
many
other
neotropical
lowland
forests.
Sapotaceae,
Lauraceae,
Myristicaceae,
Burseraceae and Meliaceae are also among the 10 families with the highest relative
diversity, except in the gorge. In the very dynamic gorge, Euphorbiaceae (5.78%), Tiliaceae
(4.96%) and Annonaceae count, along with Clusiaceae, Moraceae and Leguminosae, as
very diverse families. The families Rubiaceae and Flacourtiaceae also occur in the gorge
with 5 spp. (4.13%) each. Chrysobalanaceae are very diverse on the ridge (5 spp.; 4.97%)
and on the inland slope (4 spp.; 2.86%). The high diversity of Clusiaceae gives the
Esquinas its characteristic makeup. The Clusiaceae are among the four most diverse
families in all plots. On the inland slope the considerably diverse Clusiaceae is represented
by 12 tree and hemi-epiphytic species (rel. diversity 8.57%).
4.1.2.1.2. Relative Density
The inland plot resembles a palm garden: 142 individuals are palms; about every fourth
tree in the research plot. The palms are not very diverse (5 spp.) and are not very thick
individuals (7% of the rel. dominance), but they have high relative density (26.94%). The
palms are also very abundant on the ridge (9.55 %) and in the gorge (6.64%). On the
coastal slope the palms are amost absent. Just 8 individuals of Iriartea deltoidea were
found on the coastal slope. Moraceae are the most common family on the coastal slope
(19.9%) and they are also common in the other plots. Few species (4 spp.), but many
individuals (124), make up the high density of the Vochysiaceae on the ridge (14.62%).
Vochysiaceae are a common family in Central Amazonia (GENTRY 1990) but not in Central
136
4.1. Discussion – Floristics and Diversity
American forests (LIEBERMANN & al. 1996). In the lowlands of La Selva the palms are also
the most abundant family, and from 300 up to 1,500 m above sea level the Rubiaceae are
among the most abundant (LIEBERMANN & al. 1996). Rubiaceae, however, is not an
abundant tree family in the Esquinas plots. Clusiaceae and Myristicaceae are among the
most abundant families on the ridge and on the coastal and inland slopes, but not in the
gorge. The special makeup of the gorge is also seen in the relative density. The most
common families are the Flacourtiaceae (48 individuals; 9.96%) and the Violaceae
(individuals; 9.13%). Tiliaceae (44 individuals; 7.88%) and Cecropiaceae (38 individuals;
7.47%) are often common in gaps, secondary growth and in neotropical forests.
4.1.2.1.3. Relative Dominance
The Moraceae have the highest relative dominance on the coastal (10.72 m2; 24.65%) and
inland slopes (6.11 m2; 17.19%) and they are among the dominant families on the ridge
(3.39 m2; 8.51%) and in the gorge (2.20 m2; 6.08%). The 124 individuals of Vochysiaceae
have a high relative dominance of 21.83% (8.69 m2) on the ridge. The Clusiaceae are
represented by many individuals on the ridge (4.99 m2; 12.54%), and on the coastal (3.82
m2; 8.79%) and inland slopes (2.24 m2; 6.31%) and are therefore fairly dominant. In the
gorge the Tiliaceae (5.12 m2; 14.16%) is the most dominant family. Just three individuals of
Billia colombiana (Hippocastanaceae) cover 2.41 m2 and provide 6.67% of the relative
dominance in the gorge. Though the palms are very common on the inland slope (relative
dominance 7.00%) (2.49 m2) they are less so on the ridge ( 4.34%, 1.73 m2). Moraceae
and Meliaceae are often large trees, and therefore have a high relative dominance in
tropical lowlands.
4.1.3.2.2. Family Diversity Indices and mean number of species per
family
The Shannon-Weaver (H’) and Eveness (E’) indices are not useful for the comparison of
family diversity due to the low differences of the values.
The mean number of species per family in the Esquinas plots is 2.63 in the gorge and up to
3.51 on the ridge. This correlates more or less with the results of the LIEBERMANN & al.
(1996) investigation in La Selva.
4.1.4. Area-species curves of all research plots
The area-species relationships show that habitat diversity increases as area sampled
increases. The tree species recorded in the research plots reflect only a part of the overall
tree diversity of the Esquinas rainforest, as demonstrated by the area-species curve (Fig.
3.14, Fig. 3.15), which continues to climb steadily beyond the 1 ha of sampling. The high
percentage of tree species represented by only one individual per hectare supports the
137
4.1. Discussion – Floristics and Diversity
previous observation that these species are widely dispersed. It indicates a high
heterogeneity level of the forest stand (LAMPRECHT 1986) and supports the assumption that
the number of species would increase notably if the study plots were extended. For forests
with lower species diversity like Río Colorado, Bolivia (SMITH & KILLEEN 1995), which has
78 spp. per hectare, a plot size of 1 ha is sufficient to characterize the structure and
composition of the locality.
The 1 ha plot sizes used in the present study were too small to provide an adequate
sample of local species richness in high diversity areas (LIEBERMANN & al. 1996). Areaspecies curves are a practical guide for the minimum size of research plots (Madsen &
Øllgaard 1994).
Local species richness can be estimated by extrapolating area-species curves. These
relationships are important in ecological study because they provide insight into community
structure, and the mathematical expressions of the model are used for predicting species
richness on a larger scale, and extinction rates caused by habitat destruction (SAGAR & al.
2003).
Tab. 3.40 and the model in Fig. 3.16, Fig. 3.17 show a calculation of increase in species
richness caused by enlarging the research plots to 10 ha (100.000 m2). Even between 9
and 10 ha the increase in species richness is obvious.
The current plot size can give a representative picture of species, family and floristic
composition, although one hectare is not large enough to allow enumeration of most
species in a lowland tropical rainforest.
4.1.5. Floristic Similarity and Differences of all Research Plots, and β
Diversity
The β or differentiation diversity in the Esquinas is very high. The 328 species are not
homogeneous or regularly distributed in the four Esquinas plots. The four plots are of
different habitat types and habitat type influences species distribution (CONDIT & al. 2002).
The floristic similarity of the plots is shown in the dendrogram of the Bray-Curtis cluster
analysis (Fig. 3.18) and in Tabxx. The inland slope and the ridge share 82 species and
show a similarity of 40.6%. This floristic similarity is due to similar habitat type: the upper
100 m of the inland slope is on a ridge, and some parts of the ridge plot include slope
forests.
The gorge and the coastal slope share 31 species. These two habitat types show the
greatest floristic difference within the plots with 79.6%. The gorge forest is the most
dissimilar habitat type and shares just 55.4% of its species with the other plots, while the
inland slope shares 71.4% of its species. The inland slope also comprises the ridge forest
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4.1. Discussion – Floristics and Diversity
and gorge forest habitat types. The results from the Esquinas sites are comparable with
research carried out in Panamanian forests. Tree species composition in forests on the
Isthmus of Panama changes dramatically from one site to the next. Forests just 50
kilometers apart in Panama are less alike than forests 1,400 kilometers apart in the
western Amazon (CONDIT & al. 2002). The Esquinas forest and Panamanian forests have
the same high diversity of landscape and therefore the same high β diversity.
Species found in only one plot
Despite the geographical similarity of the four 1 ha plots in the present study, 194 spp.
were recorded only in one plot. In a study in Caribbean Costa Rica involving 14 plots (23.4
ha) from lowland to 2,600 m above sea level, LIEBERMANN & al. (1996) found that 203 of
the 561 spp. were recorded in a single 1 ha plot. The floristic composition of the Esquinas
lowland plots seems to be more varied than that of the widely distributed plots in the
Caribbean region of Costa Rica. Some of these species especially are common to the plots
and are useful as indicator species. Calatola costaricensis (19 individuals) was only found
near streams in the gorge, Sorocea cufodontissi (43 individuals), Schizolobium parahyba
(16 individuals) and Heisteria concinna (15 individuals) were only found on the coastal
slope, and Isertia laevis (11 individuals) was only found on the ridge. No species with a
similarly narrow distribution was found on the inland slope.
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4.2. Discussion – Distribution Pattern and Geographical Affinities
4.2. Distribution Pattern and Geographical Affinities
The origin and the recent distribution of the Esquinas tree flora demonstrate substantial
affinities with the Central and South American (Amazonian and Chocó) floras.
4.2.1. Origin of the Families found in the Research Plots
Since the end of the Cretaceous there has been the opportunity for relatively direct floristic
interchange between South America and tropical North America via island hopping along
the proto-Antilles (GENTRY 1982a). The uplift of the Andes has led to an evolutionary
explosion in a number of Gondwanan families. The formation of Central America occurred
about 4.7 Mio. years ago (CARRANZA-CASTANEDA 2003), and this permitted lowland tropical
Gondwana elements to move northwards on the newly formed land bridge. The southerly
migration of Laurasian taxa in lowland forests was less significant.
In Costa Rica and Panama the Laurasian-derived taxa are the dominant elements of
montane forests and the South American-derived taxa dominate the lowlands. Eighty-four
percent of the total Panamanian flora is of Gondwanaland origin (Gentry 1985). In the
tropical rainforest of northern Central America in Veracruz (Mexico) just 37 out of 116
families (32%) belong to Amazonian-centred taxa, but 42 (36%) belong to Laurasian taxa
(CHIAPPY & al. 2000). The Amazonian-centred taxa show little speciation or endemism in
Panama (GENTRY 1985) and Central America. The majority of the genera considered have
more species in South America than in Central America. The larger land mass of South
America and the greater pool of variability there could be reasons for this (CROAT & BUSEY
1975).
Trees (especially canopy trees) and lianas in the Neotropics are mostly Amazoniancentred taxa, while the extra-Amazonian ones are chiefly shrubs, epiphytes and palmettos.
Tree taxa in Panama and Costa Rica are also dominated by Amazonian-centred
Gondwana families. Taxa of the Gondwanaland families dominate (77%) the tree flora of
the Esquinas plots. Sixty-two percent of the Gondwana-derived families in the Esquinas
plots have Neotropical centres of diversity in Amazonia, a fact which can be seen from their
distribution within the Esquinas plots. Amazonian-centred plant families show little
speciation or endemism in Central America. These taxa generally decrease in diversity
from east to west across the Isthmus of Panama and north-westwards through Central
America (Gentry 1985). Examples are Caryocaraceae, which has 25 neotropical species
with two reaching Costa Rica and one occurring in the Esquinas plots (Caryocar
costaricense), and Humiriaceae which has 50 species, five reaching the Chocó and four
Costa Rica (one occurs in the Esquinas forest). Vochysiaceae and Myristicaceae show
similar distribution patterns, which suggests that these families are recent arrivals in
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4.2. Discussion – Distribution Pattern and Geographical Affinities
Central America. They probably arrived subsequent to the formation of an isthmian land
connection about 4.7 million years ago, a conclusion supported by palynological evidence
(GRAHAM 1987, GENTRY 1983).
The Andean north-centred taxa are the second major Neotropical group. They are
predominantly understorey shrubs or treelets (Rubiaceae, Melastomataceae, Myrsinaceae
and others), palmettos (Heliconiaceae, Marantaceae and others) and epiphytes
(Orchidaceae, Gesneriaceae, Ericaceae and others), but also trees and hemi-epiphytes
like Guttiferae (Clusiaceae) and Araliaceae. In the Esquinas plots, 12 spp. of Rubiaceae,
10 spp. of Melastomataceae and 18 spp. of Clusiaceae were counted. These families are
not as well represented in Amazonian forests, as in central and northern South America
(Chocó). The Andean north-centred taxa are very well represented in Costa Rica and
Panama, where 22% of their species occur (GENTRY 1982d). These taxa are concentrated
in Panama as well as in the Costa Rican lowland and middle elevation wet forests (GENTRY
1985). In the tropical rainforest of Veracruz (Mexico) 31 (27%) out of 116 families belong to
Andean north-centred taxa (CHIAPPY & al. 2000). However many of the genera of the
Andean montane flora have not extended their ranges north across lowland Panama
(HABER 2000).
These groups of plants are characteristic of the specific floristic composition of the Chocó
region (GENTRY 1982a, FORERO & GENTRY 1989). The Clusiaceae family is a typical
element of the Andean north-centred region and of the Chocó, and is also the second
richest family in the Esquinas plots with 18 spp. This circumstance demonstrates the
biogeographical affinities of the Golfo Dulce rainforests to the Chocó region, one of the
world´s most diverse regions in terms of plants (GENTRY 1982a, GENTRY 1986, FORERO &
GENTRY 1989), birds (TERBORGH & WINTER 1982), and butterflies (BROWN 1982).
Fifteen percent of the tree flora of the Esquinas plots are of Laurasian origin: Theaceae,
Magnoliaceae and Ulmaceae, which are typical elements in the montane forests of Costa
Rica, are also found here. The Boraginaceae (Cordia spp.), also a Laurasian family, is
better represented in the lowland than the upland of Central America, and occurs in the
Esquinas plots with 3 species of the genus Cordia. In the lowland rainforest of Barro
Colorado Island (BCI) only 4.5% of the total native flora (out of 1,210 seed plant species),
belongs to putatively Laurasian families.
The reason for this relatively high occurrence of Laurasian elements in the lowland
rainforest of the Esquinas plots could be the very high precipitation (up to 6,000 mm per
year) and the topology of the Esquinas, which is similar to that of montane regions.
Because of these abiotic factors typical elements of montane rainforests like tree ferns,
many epiphytes and hemiepiphytes also occur in the Esquinas (Huber in prep.). The
southern Andes-centred taxa (Myrtaceae, Podocarpaceae, Proteaceae) and dry area141
4.2. Discussion – Distribution Pattern and Geographical Affinities
centred taxa (Capparidaceae) are poorly represented in Central America as a whole and
therefore in the Esquinas forest (4% of all families).
4.2.2. Altitudinal Distribution
Two species in the Esquinas plots were also found in tropical montane rainforests above
3,000 m in South America (HOLDRIGE 1967). Only about 30% occur exclusively in the
lowland up to 800 m, 41% up to 1,000 m. In total, about 70% (above 800 m) or 59% (above
1000 m) of the investigated species are “euryözisch” species which also occur in montane
forests. MACARTHUR (1972) argued that tropical species, because of the diversity of the
assemblages in which they occur, must have either narrower niches or greater tolerance of
ecological overlap, than their counterparts living in less diverse assemblages. LIEBERMANN
& al. (1996) found that on a large-scale altitudinal gradient in Costa Rica, on the Caribbean
slope of the Cordillera Central from sea level to 2,600 m, tree species occurring in very
high-diversity assemblages in the lowland did not differ in their altitudinal breadth from tree
species growing in stands of lower diversity higher on the gradient; at all altitudes some
species with narrow distributions and some with wide ranges were found. No one species
was distributed over the entire range of altitudes. Just one species (Ardisia palmana –
Myrsinaceae) occurred over a range of about 2,000 m (from 45 m to 2,000 m) and Billia
colombiana (Hippocastanaceae) had a range from 300 to 2,000 m. No other lowland
species had a range over 1,500 m above sea level. About 45% had an altitudinal range of
about 200 m. A wide altitudinal range of many highland taxa and a mixture of lowland and
upland taxa was reported on tepuis of Venezuelan Guayana (HUBER 1988). Because of the
altitudinal occurrence of the species in the Esquinas plots, MACARTHUR’s (1972) idea in this
case cannot be confirmed. The altitudinal occurrence of the species in the steep and wet
gorge is especially conspicuous. Just 33% of the species are “stenözisch” and occur only in
lowlands. On the coastal and inland slopes and on the ridge the occurrence of exclusively
lowland species is higher (39% to 47%). Three species are also found above 3,000 m and
two of them are found in the gorge. The high precipitation and steep slopes (as in montane
forests) favour the occurrence of many montane elements (species) and therefore there is
a higher level of taxa of Laurasian origin in the Esquinas. The same phenomenon is
obvious in the occurrence of birds in the Esquinas. Many middle-elevation birds occur in
the Golfo Dulce forests, but not on the Atlantic slope of Costa Rica (STILES 1983). The
limiting factor of many tropical plant species is frost. In Costa Rica the critical temperature
line above which killing frosts occur lies between 1,500 m and 1,800 m above sea level in
the tropical premontane altitudinal belt (HARTSHORN 1983). Fifty-nine species of those
investigated in the Esquinas plots are also found above 1,800 m. Thus it appears that
ecological influences like precipitation, non-occurrence of a dry season and geomorphic
142
4.2. Discussion – Distribution Pattern and Geographical Affinities
structure of the forests are very important for the altitudinal occurrence of plant species.
The ecological plasticity against low temperature is very high in many tropical plant
species.
4.2.3. Regional Distribution
This study contains a survey of the phytogeographical relationship between the
investigated plants in the Esquinas plots. The diagram (Fig. 3.27) shows the strong affinity
of the species of the Esquinas plots to South America. The Esquinas species are more
common in Colombia than in Nicaragua, and they are more common in Bolivia (which is
further away) than in Honduras. The following sections will discuss this pattern of
distribution in detail.
4.2.3.1. Wide-ranging Species
STANDLEY (1937) assumed that in the Costa Rican “terra caliente” (lowlands) the majority of
the species especially the trees, would have a relatively wide range. Some of the trees, he
predicted, would even extend southwards to the Amazon basin. In the Esquinas plots an
average of 52% of the species were wide-ranging. This is comparable with studies of the
occurrence of certain plant families in La Selva, of which 45% are widespread (HAMMEL
1986), and of the trees on BCI, 47% of which are widespread (CROAT 1978). Forty-nine
percent of the 576 plant species of the Río Palenque, Ecuador (DODSON & GENTRY 1978)
and 60% of the total Ecuadorian flora (RENNER & al. 1990) are also widespread in the
Neotropics. However, of the 1,227 spp. of the Guayana Shield flora just 26.8% are wideranging (KELLOFF & FUNK 2004). The Guayana Shield has a distinctive flora, possibly the
result of development under isolation (HUBER 1995). Twenty-nine percent of the total flora
of Nicaragua are also wide-ranging species (STEVENS & al. 2001). The wide-ranging
species are the most common plant species (especially trees) in the Central American flora
of tropical lowland rainforests. Since the existence of the American land bridge (late
Tertiary), plants and animals have been able to migrate from south to north and vice versa.
GENTRY (1978, 1982b) has pointed out that many species are common to the wet lowlands
of both Central and South America. When GENTRY (1990) did his work focussing on the
differences between Central American and Amazonian floras, he was impressed by their
similarities.
4.2.3.2. South America and Amazon
The Amazon harbours a very significant portion of the world’s diversity (STEEGE & al. 2003)
and many trees occurring in Central America originated in the Amazonian forests, the main
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4.2. Discussion – Distribution Pattern and Geographical Affinities
centre of tree diversity in the Neotropics. This explains the very strong affinity of the
species of the Esquinas plots to South America (an average of about 20% of the
investigated plants do not occur north of Costa Rica). This is similar to a subset of the flora
of La Selva (HAMMEL 1986) and the trees of BCI (CROAT 1978), each of which was shown
to contain 20% from South America. In their introductions to the flora of Costa Rica,
STANDLEY (1937) and (GOMEZ 1986) pointed out the relationship of the Costa Rican flora to
that of South America. About 45% of the species also occur in the Amazonian region.
This pattern of affinity with South America seems the logical outcome of any scenario of
Costa Rican phytogeographical history, including tectonic movement, volcanism, original
range, recent climate and dispersal from the nearest tropical continental source to the
tropical isthmian region of Central America.
4.2.3.3. Central America
It is interesting to note the floristic relationship between the Esquinas plots and
neighbouring areas. The number of Esquinas species that occur exclusively from Panama
to the north (the Central American elements) is 27.5% of all species in the Esquinas plots.
Investigated in La Selva from the Caribbean lowlands in Costa Rica are restricted to Costa
Rica and/or to areas north of Costa Rica (HAMMEL 1986). CROAT (1978) found 21% of
Central American (Panama to Mexico) tree species on BCI. Significantly more tree species
than other life forms (epiphytes, herbs etc.) show this pattern of distribution. The total flora
of Nicaragua shows more affinity to the north than to the south (26% to 19.6%), except in
the rainforest in the Caribbean lowlands (16.3% to 28.8%). The plants of the dry regions of
pacific lowlands and of the northern montane forests show a stronger affinity to the north.
The savannahs containing Caribbean pine (Pinus caribbea) in the north of Nicaragua and
south of Honduras make the distribution of South American rainforest elements difficult.
Nicaragua is the southernmost limit for many Laurasian elements, because of the tropical
lowlands of the Río San Juán (STEVENS & al. 2001). Just 32% of the species (109 spp.) of
the Esquinas plots, including the wide-ranging species, also occur in Honduras. Of those,
just 27 species do not occur in South America. These comparisons again show the close
relationship of the Esquinas plot species to South America.
4.2.3.4. Endemic Species
The wide endemics (distribution from Nicaragua to Panama) make up to 19% (incl. 3.8%
endemics to the south of Costa Rica) of the total Esquinas plot species.
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4.2. Discussion – Distribution Pattern and Geographical Affinities
Fig. 4.2. Map of potential vegetation in northern South and Central America, with focus on Costa Rica
(adapted by Pamperl)
About 20% of these are the families noted by HAMMEL (including herbs, hemi-epiphytes and
trees) in his 1986 study of La Selva. On BCI about 16% of all species and 20% of trees
(CROAT 1978) are wide endemics (incl. endemics restricted to Panama and these
extending to Costa Rica and/or northern Colombia), and 4.6% of Nicaraguan flora are wide
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4.2. Discussion – Distribution Pattern and Geographical Affinities
endemics (STEVENS & al. 2001). The rate of endemism in Costa Rica is in general higher in
epiphytes (e.g. Orchidaceae), herbs (e.g. Marantaceae) and shrubs (e.g. Rubiaceae), but
lower in trees (BURGER 1980, HERRERA-MCBRYDE & al. 1997). These are mostly Andean
north-centred taxa, which show a very pronounced endemism in Central America. The
northward invasion of lowland Gondwana taxa of canopy trees and lianas into Central
America leads to their ecological dominance in lowland tropical forests throughout the
region, despite little significant speciation in Central America (GENTRY 1982). BASLEV &
RENNER (1989) found that in Ecuador the occurrence of endemics at mid- and high
elevation was much greater than in the lowlands. He also encountered a low-density rate of
endemism in tree species in the Ecuadorian lowlands of the Amazonian and Coastal
regions (PITMAN & al. 2002, JØRGENSEN & LEÓN-YÁNEZ 1999). In Ecuador, the dry forest
area contains more endemic plant species (20% of the local flora from Capeira) (DODSON &
GENTRY 1991) than the adjacent moist forest at Jauneche (15%) (DODSON & al. 1985). Very
high rates of endemism were found in northern South America, in Venezuelan Guayana
and on islands. About 40% of the 9,411 species in Venezuelan Guayana are endemic to
the Guayana Shield (BERRY & al. 1995). In isolated regions (e.g. islands – the Caribbean
islands have 58.3% species endemism) (SANTIAGO-VALENTIN & OLMSTEAD 2004) or
Pleistocene refugia (HAFFER 1979) the rate of endemism is higher (MAYR & O’HARA 1986).
On the Pacific slope, the tropical wet forest only extends from the Parrita area south to the
western part of Chiriquí, Panama. Drier forests and savannahs isolate this area from the
main body of tropical wet forest on the Caribbean slope to the north and south, and the
Cordillera Talamanca to the east (Fig. 4.2). This isolation probably dates back to the
Pleistocene refugia and is reflected in the distribution of animal groups like butterflies, birds
and amphibians as well as plants. The very high rate of endemism in plants (e.g. 23% of
Marantaceae species, 25% of Orchidaceae species, 21% of the genus Inga, 18.9% of
Lauraceae) (HERRERA-MCBRYDE & al. 1997, HAMMEL & al. 2004, ZAMORA & PENNINGTON
2000) and in butterflies (Ithomiinae and Heliconiini) hints at Pleistocene refugia and
supports the arguments of STILES (1983) and HAFFER (in HEPPNER 1991).
4.2.3.5. Disjunction
4.2.3.5.1. Guayana Shield - Venezuela and Guyana
Three species occur exclusively in Costa Rica and Panama or in Costa Rica and
Venezuela (disjunct distribution), yet there are no obvious affinities of the Esquinas plants
to Guayana. Reasons for this distribution pattern could be:
there are elements of the Guayanan flora that may have arrived before the formation of the
Panama Isthmus
the pattern is an artefact caused by inadequate sampling.
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4.2. Discussion – Distribution Pattern and Geographical Affinities
About 35% of the total species in the Esquinas plots were also found in Venezuela, and
about the same amount were found in Honduras.
KELLOFF & FUNK (2004) were surprised that as much as 25% of the flora of the Kaieteur
Falls (Venezuela) might have an affinity to that of Central America.
4.2.3.5.2. Honduras
Only one species showed a disjunct distribution pattern: Hirtella papillata was collected
both in Costa Rica (Golfo Dulce region) and Honduras. However the data from Honduras is
likely to be erroneous, in which case this species is endemic to the Golfo Dulce forests.
4.2.4. Pacific Region of Costa Rica Only
The pacific and Caribbean lowland rainforests of Costa Rica are about 100 km apart. The
backbone of Costa Rica, the high mountains of the Cordillera Talamanca (Cerro Chirippó
3,830 m), divides the two lowland regions. The Caribbean lowlands extending from
southern Mexico to Panama and thence to northern Colombia are covered with moist to
wet tropical forests. In recent history there has been no direct connection between these
two ever-wet lowlands in Central America.
About 31% of all investigated plants of the Esquinas plots do not occur on the Caribbean
slopes of Costa Rica. The differences between the pacific wet lowlands and the main body
of tropical wet forest in Central America on the Caribbean slope are also reflected in the
avifauna (many sedentary tropical birds common on the Atlantic slope do not occur in the
Golfo Dulce region) (STILES 1983), in the amphibian fauna (e.g. Dendrobatidae) (SCOTT &
al. 1983) and in the amphibian and reptilian fauna (SAVAGE 2002) e.g. Basiliscus spp. (VAN
DEVENDER 1983). The lowland wet forests around the Golfo Dulce are different to their
counterpart on the Atlantic slope of Costa Rica due to the formation of the Central
American land bridge, the recent and paleo-climatic conditions and their subsequent
settlement by migration of plants and animals.
4.2.5. Panama – Watershed of the Canal
Only half of the plants that are found in the Esquinas plots also occur some 400 km away in
the highly diverse moist lowland and montane forest of the Panama Canal watershed. The
different climate of the watershed (short dry season and less precipitation), along with its
location on the Atlantic side of Panama, explains why the affinity is not greater between the
two areas.
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4.2. Discussion – Distribution Pattern and Geographical Affinities
4.2.6. The Caribbean Islands
The percentage (7.7%) of plants species which the Esquinas plots share with the
Caribbean Islands is much lower than the percentage they share with the Caribbean slope
of Central America. About 38% of all plant species occurring on BCI are also found in the
West Indies (CROAT 1978). These species reached the West Indies (or migrated from
there) by long-distance dispersal (HARTSHORN & HAMMEL 1994).
4.2.7. Chocó
The Chocó region of pacific Colombia is one of the rainiest regions (10,000 mm or more
annual rainfall) in the world (FORERO & GENTRY 1989). The moist forest flora of adjacent
north-western Ecuador and western Panama (the Darien region) is almost identical in
species composition. The rainforests around the Golfo Dulce and in Darien (Panama) are
the only wet forests still to exist on the Pacific side of Central America. HARTSHORN (1983)
recognized the strong floristic affinities of the forests of the Corcovado to the Colombian
Chocó. In fact about 14% (43 spp.) of the Esquinas plots species occur exclusively in the
Chocó. Caryocar, Chaunochiton, Couratari, Huberodendron, Parkia and Peltogyne all
reach their northern limits in southern Costa Rica. Unfortunately, floristic knowledge about
the Chocó is far from complete, because political troubles and terrorism inhibit scientific
work. GENTRY (1986) suspects that there are more yet-to-be-discovered plant species in
the Chocó than anywhere else in the world. Many interesting features (Tab. 4.5) are typical
of both the Chocó and the Golfo Dulce forests:
Chocó (GENTRY 1986)
The Moraceae are the most important family
canopy trees (DODSON & GENTRY 1978)
High densities of both small (d.b.h 2.5 to 10 cm)
and medium-sized trees (d.b.h. ≥ 10 cm)
Replacement of free-climbing lianas (the normal
Neotropical forest components) by hemi-epiphytic
climbers
High floristic diversity
Unusual prevalence of families like Guttiferae,
Arecaceae, Myrtaceae, Melastomataceae and
Moraceae (on good soil)
Esquinas plots
Moraceae are the second most important family (22
spp.)
Many treelets of Arecaceae, Rubiaceae and
Melastomataceae
Up to 26% of all trees in the gorge forest carry hemiepiphytic plants (HUBER in prep.)
Up to 179 spp. of plants (d.b.h. ≥ 10 cm) per hectare
Guttiferae (Clusiaceae) are the third most important
family (18 spp.). Arecaceae, for example, are
represented by 45 spp. in the whole Golfo Dulce
forests
Zoochory, especially mammal-dispersed genera
Many species with unusually large fruits
Some of the largest and most coriaceous leaves in
the world
Tab. 4.5. Typical features of the Chocó and the Golfo Dulce forests
Typical for the Chocó forests (GENTRY 1982b, 1986, 1988b), but also for the Esquinas, are
many features that would seem more characteristic of cloud forests than lowland forests.
The exceedingly high diversity (and density) of woody hemiepiphytes and palms gives the
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4.2. Discussion – Distribution Pattern and Geographical Affinities
Chocó forest much of its characteristic aspect (GENTRY 1986), and the Esquinas forests are
very similar in this respect. For example, Hedyosmum (Chloranthaceae), Panopsis
(Proteaceae),
Meliosma
(Sabiaceae),
Talauma
(Magnoliaceae)
and
Podocarpus
(Podocarpaceae) are all taxa which occur in the Chocó lowlands and highlands as well as
in the Esquinas plots. Several Andean species of palms occur in the Chocó at lowland
elevations. HENDERSON & al. (1995) proposed that this was due to high rainfall, and that
rainfall might be more important than elevation in influencing palm distributions.
Bignoniaceae, which, as well as containing trees, is usually the main Neotropical liana
family, is poorly represented in the Chocó pluvial forests compared to other Neotropical
lowland sites. In the Esquinas one tree species and one unidentified liana species of
Bignoniaceae were found.
The vegetation of the “Bosque muy húmedo tropical” (ESPINAL & MONTENEGRO 1963) in the
Chocó region, Colombia (precipitation of more than 4,000 mm) consists of species that also
occur in flat parts of the Esquinas forest, and others related to these. The characteristic
species are Ceiba pentandra, Ochroma lagopus, Pachira aquatica, Apeiba tibourbou, Piper
anducum, Inga spp., Vitex gigantea and Rollinia microsepala. The main areas of this type
of forest in the Golfo Dulce region have been destroyed. Again, this composition of a forest
in the Chocó region shows the very close biogeographical relationship with the Esquinas
forest.
4.2.8. Dry forests
The seasonal moist and dry forest (NP Santa Rosa) located on the Pacific slope in northern
Costa Rica shows a completely different floristic composition to that of the Esquinas plots.
Just 20 spp. occur in both locations which are about 400 km apart. The montane forests of
Monteverde (Costa Rica) show the same floristic composition as the NP Santa Rosa. Only
11 species out of the 992 that make up the entire Guanacaste flora reach the Monteverde
region above 1,200 m (HABER 2000). This distribution pattern is similar to that of Ecuador:
Many of the trees in Río Palenque (wet lowland forest) do not occur in the drier part of
Pacific Ecuador, just a few kilometres away. DODSON & GENTRY (1978) suggest that most
tropical plant species have geographically ample, but ecologically restricted, distributions.
4.2.9. Differences in phytogeographical relationships within the
Esquinas plots
The differences in the phytogeographical relationships within the Esquinas plots are not
obvious, although there are differences in terms of species distribution. Widespread and
exclusively Central American species are less common in the coastal slope plot than in
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4.2. Discussion – Distribution Pattern and Geographical Affinities
the other plots, as this plot shows more affinity to South America. While the species in the
gorge (more premontane and montane affinity) are more wide-ranging, they have less
South American affinity. The gorge and the coastal slope plots have least similarity in terms
of floristic composition of all research plots in the Esquinas. This floristic pattern also
reflects the differences in the phytogeographical relationships of these two plots. The
habitat type influences species distribution. For example, tree species common in dry areas
reappear on rapidly draining soils in wet areas (CONDIT & al. 2002). About 10% of the
plants in the steep gorge and 8.5% in the steep coastal forest occur in the dry forest of
Guanacaste, Costa Rica, more than in the other Esquinas research plots. Due to the
similarity of the gorge and montane forests in terms of precipitation levels and
oreographical conditions, many more montane species occur in the gorge than in the other
plots.
In the inland slope plot there are clearly more Costa Rican and Costa Rican/Panamanian
species than in any other. The ridge plot (179 different species) shows the highest diversity
of the research plots but the species do not show significant biogeographical differences
compared with the other plots.
4.2.10. Diversity and biogeographical aspects
Some forests are extraordinarily rich in tree species because they stand at the intersection
of different phytogeographic regions. This is the so called “biodiversity crossroads
hypothesis” (PITMAN & al. 2002, PRANCE 1994). The Golfo Dulce region is located in the
Caribbean region and the Central American province (TAKHTAJAN 1986). The biodiversity
crossroads hypothesis does not explain the high diversity of trees in the Esquinas plots,
because they are located within a single, well-defined phytogeographic region on the
margin of Central America. Neighbouring regions, however, such as those with a significant
dry season, and those with montane (and therefore floristically different) forests, do
influence their diversity. Many tree species in the Esquinas plots occur in montane regions
(e.g. Cordillera Talamanca). Trees originally from the dry regions to the north and south are
also found in the Esquinas.
4.2.11. Final comments on the distribution patterns of plants
The extent of tropical floristic knowledge was largely unknown in 1992 (GENTRY). Some
species are far from being locally endemic - they have merely been inadequately collected.
NELSON & al. (1990) have shown that the maps of concentrations of endemism on which
the conservation programme for plants of the Brazilian Amazonian region was based, are
exactly coincident with maps of collection density. In the meantime different floras have
150
4.2. Discussion – Distribution Pattern and Geographical Affinities
been completed (e.g. Nicaragua, Costa Rica, Venezuela, Guayana) and now there is better
information about distribution patterns of Neotropical plants and investigations into
biogeographical patterns are more meaningful.
151
5. Abstract
5. Abstract
The rainforests around the Golfo Dulce, Costa Rica are among the most species-rich plant
communities in Central America (ALLEN 1950, QUESADA et al. 1997).The present study
deals with the diversity and biogeographical patterns of woody plants (d.b.h. ≥ 10 cm) in
four ecologically different one-hectare plots (inland slope, coastal slope, gorge and ridge) in
the Esquinas forest (Piedras Blancas National Park) of the southern Pacific region of Costa
Rica. An effort is made to reach a better understanding of the composition, diversity,
variability within the plots and the biogeographical affinities of these forests.
The forest inventory data were collected between 1993 and 2001 from plots in pristine
forest parts, located between 85 and 336 m above sea level, about 8° 41'N / 83° 13'W,
near the “Tropenstation La Gamba” and the “Golfo Dulce Lodge”. The average annual
precipitation of 5,690 mm (1998 – 2003), the annual mean temperature of nearly 28°C and
the absence of a pronounced dry season have resulted in the establishment of a “perhumid
tropical lowland wet forest” (Life Zones system of HOLDRIDGE & al. 1971).
In the four one-hectare plots all woody plants (trees, palms, lianas and ground-rooted hemiepiphytes) with d.b.h. ≥ 10 cm were identified, counted, and the diameters and height of the
trunks were measured. Altogether 2,444 trunks were measured, representing 328 species
and 69 families. On the coastal slope 588 individuals (108 spp.) were recorded, in the
gorge 482 individuals (121 spp.), on the inland slope 527 individuals (133 spp.) and on the
ridge 847 individuals (179 spp.). Figures for the four life forms were trees are: (2,124
specimens), palms (262), lianas (44) and ground-rooted hemi-epiphytes (14). Palms were
found to be very abundant on the inland slope (c. 27%) and on the ridge (c. 10%). Lianas
accounted for 3% (18 specimens) of all individuals recorded on the coastal slope, and
ground-rooted hemi-epiphytes accounted for 1% on the inland slope (5 individuals).
As to species distribution, sixteen species occurred in all four plots and 195 species were
counted in only one plot each. The palms Welfia regia (108 indiv.) and Iriartea deltoidea (97
indiv.) proved to be the most abundant species. The high representation of palms in the
Esquinas forest (262 individuals; c. 10.7%) is typical of many pluvial and wet rainforests in
the neotropics.
In all four research plots, the Fabaceae-Mimosoideae (23 species), the Moraceae (22 spp.)
and the Clusiaceae (18 spp.) proved the most diverse families. This is in agreement with
the fact that in all Central Amazonian and Central American lowland forests Leguminosae
and Moraceae are the most diverse families. The high diversity of Clusiaceae is
characteristic of Central America and the Chocó in NW South America. The congruence in
152
5. Abstract
the most diverse families indicates the close biogeographical relationship between the
Cental American forests (including the Esquinas forests) and the Chocó.
The coastal slope comprised 38 spp. represented by just one individual. On the other
extreme, one species (Compsoneura sprucei, Myristicaceae) was represented 50
individuals. Brosimum utile (Moraceae) had the highest Importance Value Index (IVI) of
24.17. Compsoneura sprucei showed the highest relative density (8.5%) and relative
frequency (5.34%) and Brosimum utile with many large trees had by far the highest relative
dominance (17.82%). The deciduous canopy tree Schizolobium parahyba (Fabaceae) was
the most conspicuous species on the coastal slope (16 indiv.). The Alpha index H (alpha) =
38.82, the Shannon-Weaver H’ = 4.014 and Simpson’s diversity D = 0.028. The families
with the highest Family Importance Indices (FIVI) were the Moraceae (53.81; 117
individuals; 10 spp.), Sapotaceae (27.08; 8 spp.; 34 individuals) and Clusiaceae (23.11; 5
spp.; 57 individuals).
In the gorge, 59 spp. were represented by only one individual, while Tetrathylacium
macrophyllum (Flacourtiaceae) was represented by 41 individuals. This understorey tree
had the highest relative density (8.51%), the relative frequency (3.52%) and absolute
frequency (100%), but the 15 individuals of the big canopy tree Dussia discolour
(Fabaceae-Fab.) had the highest relative dominance (11.90%) and, therefore, the highest
IVI (17.48). The Alpha index was 51.905, the Shannon-Weaver H’ = 4.122 and the
Simpson’s diversity D = 0.027. The families with the highest FIVI were the Tiliaceae (27.01;
38 individuals, 6 spp.), Fabaceae-Faboideae (20.62; 19 individuals; 5 spp.) and Moraceae
(20.16; 32 individuals; 9 spp.).
The plot on the ridge proved to be the most diverse one, comprising 179 spp., 72 species
(about 40% of all spp.) of which were represented by only one individual. Qualea paraensis
(Vochysiaceae) was represented by 50 individuals and the palm Welfia regia by 49. Qualea
paraensis had the highest relative density (5.90%), relative frequency (3.76) and absolute
frequency (88%), and the 38 individuals of Vochysia ferruginea (Vochysiaceae) had the
highest relative dominance (9.53%). The two species of Vochysiaceae, Vochysia
ferruginea and Qualea paraensis, had therefore the highest IVI (17.26 and 16.96). The
Alpha index H (alpha) = 70.490, the Shannon-Weaver H’= 4.483 and Simpson’s diversity D
= 0.019. The families with the highest FIVI were the Vochysiaceae (38.66; 124 individuals,
4 spp.), Clusiaceae (29.04; 79 individuals; 13 spp.) and Moraceae (23.32; 60 individuals;
14 spp.). Fabaceae-Mimosoideae were represented by 14 spp. and 29 individuals.
In the plot of the inland slope 142 individuals (about 27%) of the recorded trees were
palms. Eighty-five species occurred in the plot with only one individual, while the palm
Iriartea deltoidea was represented with 71 individuals. Iriartea deltoidea was also the
leading species in terms of IVI (20.90), relative density (13.47%), relative frequency
153
5. Abstract
(5.08%) and absolute frequency (100%). With just 15 individuals, Brosimum utile
(Moraceae) had the highest relative dominance (13.59%). The Alpha index was 57.953, the
Shannon-Weaver H’= 4.119 and the Simpson’s diversity D = 0.035. At family level, the
Clusiaceae were represented by 12 spp. (55 individuals) and the Fabaceae-Mimosoideae
by 11 spp. (13 individuals). The families with the highest FIVI were the Arecaceae (37.52;
142 individuals, 5 spp.), Moraceae (30.68; 41 individuals; 12 spp.) and Clusiaceae (25.32;
55 individuals; 12 spp.).
In total, 16 spp. occurred in all four research plots, while 195 spp. (59.45%) were
represented only by one individual. The Bray-Curtis cluster analysis yielded a floristic
similarity of 40.61% for the inland slope and the ridge. The coastal slope and the gorge had
the least floral similarity (13.46%). The gorge forest proved the most dissimilar habitat type,
sharing just 55.4% of its species with the other plots, while the inland slope shared 71.4%
of its species with other plots. The Esquinas forest has a high landscape diversity and,
therefore, a high β-diversity.
No single tree species dominated one plot. Only Brosimum utile, a common tree in the wet
region in the Pacific lowlands of Costa Rica and found mainly on well-drained slopes and
ridges, proved an important tree in two plots (coastal and inland slope).
The 10 most abundant species constituted about 27% of all investigated individuals (3% of
all spp.) in all plots. The floristic composition of the Esquinas plots seems to be more varied
than that of comparable plots in the Caribbean region of Costa Rica. Some particular
species are useful as indicator species: Calatola costaricensis (19 individuals) was only
found near streams in the gorge, Sorocea cufodontisii (43 individuals), Schizolobium
parahyba (16 individuals) and Heisteria concinna (15 individuals) were only found on the
coastal slope, and Isertia laevis (11 individuals) was only found on the ridge plot.
The high percentage of tree species represented by only one individual per hectare
indicates a high heterogeneity level of the forest. This supports the assumption that the
number of species in the Esquinas would notably increase with increase of the size of the
study plots. Newly developed area-species curves generated for all plots demonstrate
graphically the increase in species per area with increasing study site size and allow the
prediction of the species number that can be expected in a certain habitat. Calculations for
10 ha plots yield the following numbers: ridge plot: 389 spp., gorge: 297 spp., coastal
slope: 214 spp., and inland slope: 287 spp. The current plot size (1 ha) can give a rough
picture of species number, family representation and floristic composition, but it is not large
enough to give an accurate estimation of the species number present in a tropical lowland
rainforest.
Diversity, in terms of species richness, is very high in the Esquinas forest. The investigated
plots support the view that the Esquinas forest belongs to the most species-rich forests of
154
5. Abstract
Central America. The plot on the ridge is, indeed, the most diverse plot ever investigated in
Central America. The comparison of the various diversity indices (Simpson, ShannonWeaver, α-Index etc.) demonstrates this very precisely in the form of numerical graphs.
The most important reasons for the high tree species diversity in the Esquinas forest
include:
•
The forest is strongly spatially structured at the landscape level and provides, therefore,
many microhabitats and niches.
•
The forest is very dynamic. All phases of forest regeneration, from gap phases to climax
forest are present in all four plots.
•
High annual precipitation levels without a dry season support a high diversity of trees.
•
High edaphic heterogeneity and diversity of soil conditions support a high β- diversity.
•
Distribution patterns of taxa from the Esquinas forest show a strong affinity to the Chocó
region (coastal Colombia and adjacent Ecuador), which ranks among
the world’s
richest regions in terms of plant diversity.
The biogeographical patterns and affinities of 312 species recorded in the four research
plots were analysed. 62% percent of the families have their origins in Amazonia, which is
the main centre of tree diversity in the Neotropics. 15% are of Laurasian and 11% of NorthAndean origin. The origin and the recent distribution of the Esquinas tree flora demonstrate
substantial affinities with the Central and South American (Amazonian and Chocó) floras.
The high precipitation and steep slopes (as in montane forests) favour the occurrence of
many montane elements (species) and therefore a high proportion of the taxa in the
Esquinas of Laurasian origin. Just 130 spp. occur only in the lowlands (below 1,000 m),
146 spp. also occur in premontane forests (above 1,000 m), 34 spp. also in lower montane
forests (above 2,000 m) and 2 species. were also found in montane forests (above 3,000
m).
Of the 312 species, more have ranges extending to the south (Panama: 251, Colombia:
197 and Ecuador: 166) than to the north (Nicaragua: 186; Honduras; 109). About 51%
(161) are wide-ranging species and occur in both South and Central America. Many
species are common to the wet lowlands of both Central and South America. 86 species
are found exclusively in Central America, and 15 of these are endemic to the south of
Costa Rica. 63 species are found only to the South. 99 species (31.3%) occur only in the
Pacific regions of Costa Rica and 213 (68.3%) occur also in the Caribbean region. The
lowland wet forests around the Golfo Dulce are different to their counterparts on the
Atlantic slope of Costa Rica due to the formation of the Central America land bridge, the
recent and paleo-climatic conditions and their subsequent settlement by migration of plants
and animals.
155
5. Abstract
In total, 141 species (45%) also occur in the rainforests of the Amazon, but just 20 species
occur in the dry forests of northern Costa Rica. Nicaragua is the southernmost limit for
many Laurasian elements and some species exclusively occur in the Chocó and reach
their northern limits in southern Costa Rica. The floristic composition of the forests in the
Esquinas forest show a very close biogeographical relationship with the Chocó region. The
differences in the phytogeographical relationships within the Esquinas plots are not
obvious, although there are differences in terms of species distribution between the plots.
156
5. Abstract
6. Curriculum vitae
Name:
Werner Huber
Mother:
Margarete Huber
Father:
Engelbert Schörghuber
Date of birth:
29th December 1961
Place of birth:
Linz/Upper Austria
Nationality:
Austria
EDUCATION
1967 – 1977 Primary and Secondary School
1988
Abitur (roughly equivalent to GCSEs)
1988 – 1996
University of Vienna, Master in Natural Science
157
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169
8. Lists
8. Lists
No.
Family
Species
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
Anacardiaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Annonaceae
Apocynaceae
Apocynaceae
Apocynaceae
Araliaceae
Araliaceae
Araliaceae
20
Arecaceae
21
Arecaceae
22
Arecaceae
23
24
25
26
27
28
29
Arecaceae
Arecaceae
Bignoniaceae
Bignoniaceae
Bombacaceae
Bombacaceae
Bombacaceae
Tapirira myriantha
Annona amazonica
Annona pittieri
Cymbopetalum costaricense
Duguetia confusa
Guatteria amplifolia
Guatteria chiriquiensis
Guatteria recurvisepala (aff.)
Guatteria sp.nov.
Rollinia pittieri
Unonopsis pittieri
Unonopsis theobromifolia
Xylopia sericophylla
Aspidosperma spruceanum
Lacmellea panamensis
Odontadenia cf. cognata
Dendropanax arboreus (cf.)
Dendropanax caucanus
Dendropanax sessiliflorus
Euterpe precatoria var.
longevaginata
Iriartea deltoidea
Oenocarpus mapora ssp.
mapora
Socratea exorrhiza
Welfia regia
sp. 1 (liana - unident.)
Tabebuia chrysantha
Bombacopsis sessilis
Ceiba pentandra
Huberodendron allenii
Coastal slope
Gorge
9
0
0
5
3
0
0
0
0
4
0
0
0
3
18
1
7
0
1
1
0
1
0
0
0
1
3
1
0
1
1
0
0
0
0
0
5
13
Inland
slope
0
1
0
0
0
1
1
1
0
0
0
0
0
2
2
0
12
0
0
0
0
8
No. of species per Species in how many
family
plots
1
3
12
1
12
1
12
1
12
2
12
2
12
2
12
3
12
1
12
1
12
1
12
1
12
1
3
1
3
3
3
1
3
3
3
1
3
3
Ridge
Total
8
0
0
0
1
12
0
1
0
0
0
0
1
15
2
0
3
0
9
18
1
1
5
4
13
2
5
1
4
1
1
1
20
22
1
22
5
23
6
1
7
5
2
16
71
2
97
5
4
0
0
1
9
10
5
2
0
0
1
0
0
0
0
3
12
0
2
0
1
1
17
47
0
0
5
0
0
20
49
0
0
6
0
0
40
108
1
2
11
1
1
5
5
2
2
4
4
4
3
3
1
1
2
1
1
170
8. Lists
No.
Family
Species
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
Bombacaceae
Boraginaceae
Boraginaceae
Boraginaceae
Burseraceae
Burseraceae
Burseraceae
Burseraceae
Burseraceae
Burseraceae
Burseraceae
Burseraceae
Capparidaceae
Caricaceae
Caryocaraceae
Cecropiaceae
Cecropiaceae
Cecropiaceae
Celastraceae
Celastraceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae
Clusiaceae
Clusiaceae
Ochroma pyramidale
Cordia collococca
Cordia cymosa
Cordia megalantha
Protium aracouchini
Protium cf. schippii
Protium costaricense
Protium glabrum
Protium panamense (cf.)
Protium ravenii
Tetragastris panamensis
Trattinickia aspera
Capparis pittieri
Jacaratia dolichaula
Caryocar costaricense
Cecropia obtusifolia
Cecropia peltata
Pourouma bicolor
Crossopetalum eucymosum
Perrottetia sessiliflora
Hirtella americana
Hirtella lemsii
Hirtella papillata
Hirtella triandra
Hirtella trichotoma
Licania arborea
Licania corniculata
Licania glabriflora
Licania hypoleuca
Licania operculipetala
Licania sparsipilis
Maranthes panamensis
Calophyllum brasiliense
Calophyllum longifolium
Coastal slope
Gorge
0
0
0
0
17
0
4
0
1
12
3
0
0
0
9
6
0
8
0
0
3
0
1
2
0
1
0
0
0
0
0
0
0
2
4
4
7
1
0
0
0
4
1
0
0
0
0
2
0
32
4
0
0
11
0
0
0
0
0
0
0
0
0
0
0
0
0
0
171
Inland
slope
0
0
0
0
1
0
0
0
7
2
0
2
0
0
0
1
0
0
1
0
0
0
0
1
0
0
0
0
0
1
2
4
0
0
Ridge
Total
0
0
1
0
2
1
10
1
5
5
6
1
1
0
0
7
2
0
0
0
3
1
0
0
1
1
1
1
2
0
9
1
7
18
4
4
8
1
20
1
14
5
14
19
9
3
1
2
9
46
6
8
1
11
6
1
1
3
1
2
1
1
2
1
11
5
7
20
No. of species per Species in how many
family
plots
4
1
3
1
3
2
3
1
8
3
8
1
8
2
8
2
8
4
8
3
8
2
8
2
1
1
1
1
1
1
3
4
3
2
3
1
2
1
2
1
13
2
13
1
13
1
13
2
13
1
13
2
13
1
13
1
13
1
13
1
13
2
13
2
18
1
18
2
8. Lists
No.
Family
Species
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Clusiaceae
Combretaceae
Combretaceae
Cyatheaceae
Cyatheaceae
Dichapetalaceae
Dilleniaceae
Dilleniaceae
Ebenaceae
Elaeocarpaceae
Elaeocarpaceae
Elaeocarpaceae
Elaeocarpaceae
Elaeocarpaceae
Elaeocarpaceae
Elaeocarpaceae
Elaeocarpaceae
Euphorbiaceae
Euphorbiaceae
Chrysochlamys allenii
Chrysochlamys grandifolia
Clusia cylindrica
Clusia peninsulae
Clusia sp.1
Clusia sp.2
Clusia valerii
Garcinia madruno
Marila laxiflora
Marila pluricostata
Symphonia globulifera
Tovomita longifolia
Tovomita stylosa
Tovomita weddelliana
Vismia baccifera
Vismia macrophylla
Terminalia amazonia
Terminalia bucidoides
Alsophila firma
Cyathea delgadii
Stephanopodium costaricense
Doliocarpus hispidus
Doliocarpus multiflorus (cf.)
Diospyros panamense
Sloanea ampla
Sloanea brachytepala
Sloanea faginea
Sloanea guianensis
Sloanea medusula
Sloanea sp.
Sloanea sulcata
Sloanea zuliaensis
Alchornea costaricensis
Croton schiedeanus (cf.)
Coastal slope
Gorge
0
1
0
0
0
0
1
17
0
0
36
0
0
0
0
0
0
0
0
0
0
0
6
0
0
0
5
0
1
0
0
1
0
0
0
6
0
0
0
0
0
0
0
2
1
2
3
0
1
0
0
4
11
0
0
1
1
0
0
0
0
0
3
0
0
0
1
1
172
Inland
slope
5
1
1
1
1
1
1
3
20
0
18
2
0
1
0
0
0
0
0
1
1
4
0
0
0
0
0
0
0
3
0
0
0
0
Ridge
Total
1
2
0
0
0
0
1
1
19
2
24
1
0
1
1
1
1
0
0
0
0
2
0
2
1
4
0
2
0
0
1
0
0
25
6
10
1
1
1
1
3
21
39
4
79
5
3
2
2
1
1
4
11
1
1
7
7
2
1
4
5
2
4
3
1
1
1
26
No. of species per Species in how many
family
plots
18
2
18
4
18
1
18
1
18
1
18
1
18
3
18
3
18
2
18
2
18
4
18
3
18
1
18
2
18
2
18
1
2
1
2
1
2
1
2
1
1
1
2
3
2
2
1
1
8
1
8
1
8
1
8
1
8
2
8
1
8
1
8
1
9
1
9
2
8. Lists
No.
Family
Species
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
Euphorbiaceae
Euphorbiaceae
Euphorbiaceae
Euphorbiaceae
Euphorbiaceae
Euphorbiaceae
Euphorbiaceae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fab.-Caesalpinioideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Faboideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Hyeronima alchorneoides
Mabea occidentale
Pausandra trianae
Richeria obovata
Sapium allenii
Sapium aurifolium
Sapium pachystachys
Bauhinia bahiachalensis
Bauhinia glabra
Bauhinia guianensis (cf.)
Bauhinia manca
Copaifera camibar
Dialium guianense
Macrolobium hartshornii
Peltogyne purpurea
Schizolobium parahyba
Andira inermis
Dussia discolor
Dussia macroprophyllata
Lonchocarpus pentaphyllus
Machaerium floribundum
Machaerium kegelii
Machaerium seemannii
Ormosia panamensis
Pterocarous hayesii
Pterocarpus officinalis (cf.)
Abarema adenophora
Abarema macradenia
Acacia allenii
Inga acrocephala
Inga acuminata
Inga alba
Inga densiflora
Inga goldmannii
Coastal slope
Gorge
0
0
0
0
0
0
0
0
1
0
0
0
6
0
0
16
2
0
5
0
0
1
1
0
0
1
0
0
1
0
1
0
0
0
1
0
1
0
1
2
3
1
0
3
0
0
0
0
0
0
0
15
0
1
1
0
0
0
1
1
0
0
4
0
0
0
4
4
173
Inland
slope
0
13
3
1
0
0
0
0
0
0
0
2
0
2
2
0
0
2
0
0
0
0
0
0
0
0
0
0
1
1
0
1
0
0
Ridge
Total
0
0
20
9
0
0
0
0
0
0
2
2
0
9
11
0
0
0
1
1
0
0
0
2
0
0
5
2
4
0
1
1
2
1
1
13
24
10
1
2
3
1
1
3
2
4
6
11
13
16
2
17
6
2
1
1
1
2
1
2
5
2
10
1
2
2
6
5
No. of species per Species in how many
family
plots
9
1
9
1
9
3
9
2
9
1
9
1
9
1
9
1
9
1
9
1
9
1
9
2
9
1
9
2
9
2
9
1
11
1
11
2
11
2
11
2
11
1
11
1
11
1
11
1
11
1
11
2
23
1
23
1
23
4
23
1
23
2
23
2
23
2
23
2
8. Lists
No.
Family
Species
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Fabaceae-Mimosoideae
Flacourtiaceae
Flacourtiaceae
Flacourtiaceae
Flacourtiaceae
Flacourtiaceae
Flacourtiaceae
Flacourtiaceae
Flacourtiaceae
Flacourtiaceae
Hippocastanaceae
Hippocrateaceae
Hippocrateaceae
Humiriaceae
Icacinaceae
Icacinaceae
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Inga jinicuil
Inga marginata
Inga oerstediana
Inga pezizifera
Inga polita
Inga punctata
Inga sapindoides
Inga sp.1
Inga sp.2
Inga sp.5
Inga thibaudiana
Inga umbellifera
Inga venusta
Parkia pendula
Pithecellobium macradenium
Casearia arborea
Casearia tacanensis
Hasseltia floribunda
Lacistema aggregatum
Laetia procera
Lozania pittieri
Lunania mexicana
Pleuranthodendron lindenii
Tetrathylacium macrophyllum
Billia colombiana
Cheiloclinium cognatum
sp. (liana, unident.)
Humiriastrum diguense
Calatola costaricensis
Discophora guianensis
Aiouea costaricensis
Beilschmiedia alloiophylla
Beilschmiedia pendula (cf.)
Caryodaphnopsis burgeri
Coastal slope
Gorge
0
0
2
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
2
4
3
0
0
0
1
0
0
0
2
0
1
2
0
0
3
0
0
0
0
0
0
1
0
1
0
0
1
1
4
0
0
1
0
0
41
3
0
0
1
19
0
0
0
0
0
174
Inland
slope
0
0
0
0
0
0
1
1
1
1
1
1
0
2
2
0
0
0
1
0
0
0
2
2
0
0
1
7
0
1
1
0
2
1
Ridge
Total
0
2
0
2
2
0
1
0
0
0
3
1
2
0
0
3
0
0
1
6
0
0
1
0
0
1
0
9
0
0
0
0
0
0
2
2
2
5
2
1
2
1
1
1
5
3
3
2
2
5
1
4
2
6
1
2
7
46
3
1
1
18
19
1
1
2
2
2
No. of species per Species in how many
family
plots
23
1
23
1
23
1
23
2
23
1
23
1
23
2
23
1
23
1
23
1
23
3
23
3
23
2
23
1
23
1
9
3
9
1
9
1
9
2
9
1
9
1
9
1
9
3
9
3
1
1
2
1
2
1
1
4
2
1
2
1
16
1
16
1
16
1
16
2
8. Lists
No.
Family
Species
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Lauraceae
Lecythidaceae
Lecythidaceae
Lecythidaceae
Lecythidaceae
Lecythidaceae
Lecythidaceae
Lepidobotryaceae
Loganiaceae
Magnoliaceae
Malpighiaceae
Malpighiaceae
Malvaceae
Malvaceae
Marcgraviaceae
Marcgraviaceae
Melastomataceae
Melastomataceae
Melastomataceae
Melastomataceae
Melastomataceae
Melastomataceae
Melastomataceae
Melastomataceae
Melastomataceae
Cinnamomum neurophyllum
Licaria cufodontisii
Nectandra umbrosa
Ocotea insularis
Ocotea leucoxylon
Ocotea mollifolia
Ocotea pullifolia
Ocotea rubriflora
Pleurothyrium golfodulcensis
Pleurothyrium trianae
Couratari guianensis
Eschweilera integrifolia
Eschweilera longirachis
Eschweilera pittieri (cf.)
Grias cauliflora
Lecythis mesophylla
Ruptiliocarpon caracolito
Strychnos panurensis
Talauma gloriensis
Byrsonima crispa
Tetrapterys cf. seemannii
Hampea appendiculata
Malvaviscus arboreus
Marcgravia schippii
Souroubea sympetala
Conostegia lasiopoda
Henriettea odorata
Henriettea sp.
Henriettea succosa
Henriettea tuberculosa
Miconia doniana
Miconia sp.
Miconia trinervia
Mouriri gleasoniana
Coastal slope
Gorge
0
1
1
0
7
0
0
2
0
0
0
0
0
0
0
0
0
5
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
2
0
0
1
0
8
0
1
0
0
0
1
1
1
0
0
1
1
0
0
0
1
0
1
0
175
Inland
slope
0
0
0
0
0
0
0
0
1
1
1
0
0
0
2
0
3
0
2
3
0
0
0
0
0
0
0
7
0
4
0
1
1
1
Ridge
Total
1
1
4
1
0
0
1
0
1
0
0
1
0
6
1
3
4
0
9
0
0
0
0
1
1
0
8
0
1
2
0
0
1
1
1
2
5
1
8
1
1
2
2
3
1
1
1
6
11
3
8
5
11
3
1
1
1
1
1
1
9
7
1
6
1
1
3
2
No. of species per Species in how many
family
plots
16
1
16
2
16
2
16
1
16
2
16
1
16
1
16
1
16
2
16
2
6
1
6
1
6
1
6
1
6
3
6
1
1
3
1
1
1
2
2
1
2
1
2
1
2
1
2
1
2
1
10
1
10
2
10
1
10
1
10
2
10
1
10
1
10
3
10
2
8. Lists
No.
Family
Species
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
Melastomataceae
Meliaceae
Meliaceae
Meliaceae
Meliaceae
Meliaceae
Meliaceae
Meliaceae
Meliaceae
Menispermaceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Moraceae
Myristicaceae
Myristicaceae
Topobea maurofernandeziana
Carapa guianensis
Guarea grandifolia
Guarea kunthiana
Guarea pterorhachis
Trichilia cf.martiana
Trichilia hirta
Trichilia septentrionalis
Trichilia tuberculata
Anomospermum reticulatum
Brosimum alicastrum
Brosimum costaricanum
Brosimum guianense
Brosimum lactescens
Brosimum utile
Castilla tunu
Clarisia biflora
Ficus brevibracteata
Ficus bullenei
Ficus colubrinae
Ficus morazaniana
Ficus nymphaeifolia
Ficus tonduzii
Maclura tinctoria
Maquira costaricana
Naucleopsis ulei
Olmedia aspera
Perebea hispidula
Pseudolmedia spuria
Sorocea affinis
Sorocea cufodontisii
Sorocea pubivena
Compsoneura sprucei
Otoba novogranatensis
Coastal slope
Gorge
0
9
12
2
17
0
0
2
0
0
0
2
30
10
21
0
0
0
0
0
0
0
0
1
0
1
0
6
2
0
43
1
50
1
0
6
7
0
0
0
1
4
0
0
3
1
0
2
3
10
1
1
0
0
0
0
10
0
0
0
0
1
0
0
0
0
0
2
176
Inland
slope
0
15
8
0
0
2
0
3
0
0
3
0
5
11
15
1
0
0
0
0
0
0
1
0
0
0
0
4
0
1
0
0
8
10
Ridge
Total
1
5
8
0
2
0
0
5
2
2
2
2
28
4
7
0
0
0
2
1
2
2
0
0
2
1
3
2
0
2
0
0
14
3
1
35
35
2
19
2
1
14
2
2
8
5
63
27
46
11
1
1
2
1
2
2
11
1
2
2
3
13
2
3
43
1
72
16
No. of species per Species in how many
family
plots
10
1
8
4
8
4
8
1
8
2
8
1
8
1
8
4
8
1
1
1
22
3
22
3
22
3
22
4
22
4
22
2
22
1
22
1
22
1
22
1
22
1
22
1
22
2
22
1
22
1
22
2
22
1
22
4
22
1
22
2
22
1
22
1
6
3
6
4
8. Lists
No.
Family
Species
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
Myristicaceae
Myristicaceae
Myristicaceae
Myristicaceae
Myrsinaceae
Myrsinaceae
Myrsinaceae
Myrtaceae
Myrtaceae
Myrtaceae
Myrtaceae
Myrtaceae
Myrtaceae
Myrtaceae
Myrtaceae
Ochnaceae
Ochnaceae
Olacaceae
Olacaceae
Olacaceae
Olacaceae
Olacaceae
Oleaceae
Podocarpaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Proteaceae
Quiinaceae
Quiinaceae
Rhizophoraceae
Rubiaceae
Rubiaceae
Virola guatemalensis
Virola koschnyi
Virola sebifera
Virola surinamensis
Ardisia compressa
Ardisia dodgei
Parathesis aeruginosa
Calyptranthes chytraculia (aff.)
Calyptranthes pallens
Eugenia glandulosopunctata
Eugenia sp.
Myrcia sp. 1 (nov.-ined.)
Myrcia sp. 2 (nov.-ined.)
Myrcia sp. 3 (nov.-ined.)
Myrciaria floribunda
Ouratea lucens
Ouratea valerii
cf. Heisteria scandens
Chaunochiton kappleri
Heisteria acuminata
Heisteria concinna
Minquartia guianensis
Chionanthus panamensis
Podocarpus guatemalensis
Coccoloba belizensis
Coccoloba lehmannii
Coccoloba obovata
Coccoloba standleyana
Panopsis suaveolens
Lacunaria panamensis
Quiina macrophylla
Cassipourea elliptica
Borojoa panamensis
Chimarrhis latifolia
Coastal slope
Gorge
0
1
1
3
0
0
4
0
0
1
0
0
0
0
0
1
0
0
4
2
15
3
2
0
0
0
0
0
0
0
4
2
0
0
2
4
0
0
2
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
3
1
0
0
0
0
0
0
177
Inland
slope
4
3
2
0
3
0
4
2
0
0
1
0
0
0
1
0
1
0
1
0
0
3
0
1
0
0
0
0
0
1
3
2
0
1
Ridge
Total
10
3
8
3
1
4
6
2
7
0
0
1
1
0
1
0
0
1
0
0
0
0
0
0
1
5
0
1
2
0
1
5
1
0
16
11
11
6
6
4
14
4
7
1
1
1
1
1
2
1
1
1
5
2
15
6
2
1
1
5
3
2
2
1
8
9
1
1
No. of species per Species in how many
family
plots
6
3
6
4
6
3
6
2
3
3
3
1
3
3
8
2
8
1
8
1
8
1
8
1
8
1
8
1
8
2
2
1
2
1
5
1
5
2
5
1
5
1
5
3
1
1
1
1
5
1
5
1
5
1
5
1
1
1
2
1
2
3
1
3
12
1
12
1
8. Lists
No.
Family
Species
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Sabiaceae
Sabiaceae
Sapindaceae
Sapindaceae
Sapindaceae
Sapindaceae
Sapindaceae
Sapindaceae
Sapindaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Sapotaceae
Scrophulariaceae
Chione sylvicola
Coussarea hondensis
Duroia costaricensis
Faramea stenura
Gonzalagunia panamensis
Guettarda crispiflora
Guettarda sanblasensis
Isertia laevis
Rondeletia bertieroides
Simira maxonii
Meliosma donnellsmithii
Meliosma grandiflora
Cupania livida
Matayba ingifolia
Matayba oppositifolia
Paullinia costata
sp. 2 (liana, unident.)
Talisia nervosa
Vouarana guianensis
Chrysophyllum aff. parvulum
Chrysophyllum argenteum
Chrysophyllum brenesii
Elaeoluma glabrescens
Manilkara staminodella
Micropholis melinoniana
Pouteria durlandii (cf.)
Pouteria foveolata
Pouteria glomerata
Pouteria laevigata
Pouteria reticulata
Pouteria torta
Pradosia sp. nov.
Sarcaulus brasiliensis
Schlegelia parviflora (cf.)
Coastal slope
Gorge
0
0
0
0
1
0
3
0
0
0
0
0
0
0
0
0
1
0
0
3
1
1
4
8
0
11
4
0
0
0
0
0
2
1
0
1
1
2
0
0
0
0
1
5
0
2
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
1
178
Inland
slope
3
0
0
0
0
0
0
0
0
0
0
7
0
1
0
0
0
2
1
0
0
0
7
0
4
2
1
0
2
2
2
0
0
0
Ridge
Total
3
0
4
0
0
3
0
11
0
0
1
4
1
1
1
0
0
0
0
0
0
0
2
0
1
1
0
1
0
0
2
0
0
1
6
1
5
2
1
3
3
11
1
5
1
13
2
2
1
1
1
2
1
3
1
1
13
8
6
14
5
1
2
2
4
1
2
3
No. of species per Species in how many
family
plots
12
2
12
1
12
2
12
1
12
1
12
1
12
1
12
1
12
1
12
1
2
1
2
3
7
2
7
2
7
1
7
1
7
1
7
1
7
1
14
1
14
1
14
1
14
3
14
1
14
3
14
3
14
2
14
1
14
1
14
1
14
2
14
1
14
1
1
3
8. Lists
No.
Family
Species
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
Simaroubaceae
Sterculiaceae
Sterculiaceae
Theaceae
Theophrastaceae
Tiliaceae
Tiliaceae
Tiliaceae
Tiliaceae
Tiliaceae
Tiliaceae
Tiliaceae
Tiliaceae
Ulmaceae
Ulmaceae
unidentified
Urticaceae
Violaceae
Violaceae
Violaceae
Violaceae
Violaceae
Vochysiaceae
Vochysiaceae
Vochysiaceae
Vochysiaceae
Zamiaceae
total
Simarouba amara
Sterculia recordiana
Theobroma simiarum
Ternstroemia multiovulata
Clavija costaricana
Apeiba membranacea
Apeiba tibourbou
Goethalsia meiantha
Heliocarpus appendiculatus
Mortoniodendron anisophyllum
Mortoniodendron guatemalense
Trichospermum galeottii
Trichospermum grewiifolium
Ampelocera macrocarpa
Celtis schippii
unidentified
Myriocarpa longipes
Amphirrhox longifolia
Fusispermum laxiflorum
Gloeospermum diversipetalum
Rinorea dasyadena
Rinorea hummelii
Qualea paraensis
Vochysia allenii
Vochysia ferruginea
Vochysia megalophylla
Zamia fairchildiana
Coastal slope
Gorge
0
2
0
0
0
2
5
0
0
1
0
0
8
5
1
0
0
0
0
3
0
0
0
0
2
0
0
588
0
1
0
0
1
3
8
5
2
12
0
0
8
0
0
0
5
0
6
1
37
0
0
0
0
0
0
482
List 1.: Abundance of species and families in the research plots
179
Inland
slope
2
2
3
1
0
1
1
0
0
0
0
0
0
3
0
1
0
0
0
1
7
0
2
0
5
5
0
527
Ridge
Total
2
5
0
0
0
0
2
1
0
0
2
1
19
1
0
0
0
3
0
0
4
6
50
1
38
35
1
847
4
10
3
1
1
6
16
6
2
13
2
1
35
9
1
1
5
3
6
5
48
6
52
1
45
40
1
2444
No. of species per Species in how many
family
plots
1
2
2
4
2
1
1
1
1
1
8
3
8
4
8
2
8
1
8
2
8
1
8
1
8
3
2
3
2
1
1
1
1
1
5
1
5
1
5
3
5
3
5
1
4
2
4
1
4
3
4
2
1
1
8. Lists
Family
Anacardiaceae
Annonaceae
Apocynaceae
Araliaceae
Arecaceae
Bignoniaceae
Bombacaceae
Boraginaceae
Burseraceae
Capparidaceae
Caricaceae
Caryocaraceae
Cecropiaceae
Celastraceae
Chrysobalanaceae
Clusiaceae
Combretaceae
Cyatheaceae
Dichapetalaceae
Dilleniaceae
Ebenaceae
Elaeocarpaceae
Euphorbiaceae
Fab.-Caesalpiniaceae
Fab.-Fabaceae
Fab.-Mimosaceae
Flacourtiaceae
Hippocastanaceae
Hippocrateaceae
Humiriaceae
Icacinaceae
Lauraceae
Lecythidaceae
Lepidobotryaceae
Loganiaceae
Magnoliaceae
Malphigiaceae
Coastal slope
1
3
3
2
1
1
0
0
5
0
0
1
2
0
4
5
0
0
0
1
0
3
0
3
5
5
4
0
0
1
0
6
0
0
1
0
0
Gorge
1
6
0
2
3
1
3
3
2
0
1
0
2
1
0
6
1
1
0
2
0
1
7
2
5
7
5
1
0
1
1
3
2
1
0
0
1
Inland slope
0
4
2
1
5
0
1
0
4
0
0
0
1
1
4
12
0
1
1
1
0
1
3
3
1
11
3
0
1
1
1
5
2
1
0
1
1
180
Ridge
1
4
2
2
5
0
1
0
8
1
0
0
2
0
9
13
1
0
0
1
1
4
3
4
3
14
4
0
1
1
0
8
4
1
0
1
0
No. of species per family
1
12
3
3
5
2
4
3
8
1
1
1
3
2
12
18
2
2
1
2
1
8
9
9
10
23
9
1
2
1
2
15
6
1
1
1
2
Origin
g-amaz.
g-amaz.
g-amaz.
g-n-and.
g-amaz.
g-amaz.
g-amaz.
l.
g-amaz.
dry
g-n-and.
g-amaz.
l.
l.
l.
g-n-and.
g-amaz.
l.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
l.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
africa
g-amaz.
l.
g-amaz.
8. Lists
Family
Malvaceae
Marcgraviaceae
Melastomataceae
Meliaceae
Menispermaceae
Moraceae
Myristicaceae
Myrsinaceae
Myrtaceae
Ochnaceae
Olacaceae
Oleaceae
Podocarpaceae
Polygonaceae
Proteaceae
Quiinaceae
Rhizophoraceae
Rubiaceae
Sabiaceae
Sapindaceae
Sapotaceae
Scrophulariaceae
Simaroubaceae
Sterculiaceae
Theaceae
Theophrastaceae
Tiliaceae
Ulmaceae
unidentified sp.
Urticaceae
Violaceae
Vochysiaceae
Zamiaceae
Coastal slope
0
0
0
5
0
10
5
1
1
1
4
1
0
0
0
1
1
2
0
1
8
1
0
1
0
0
4
2
0
0
1
1
0
Gorge
2
0
4
4
0
9
3
1
1
0
0
0
0
2
0
0
0
5
1
2
2
1
0
1
0
1
6
0
0
1
3
0
0
Inland slope
0
0
5
4
0
8
5
2
3
1
2
0
1
0
0
2
1
2
1
3
8
0
1
2
1
0
2
1
1
0
2
3
0
Ridge
0
2
6
5
1
14
6
3
5
0
1
0
0
2
1
1
1
5
2
3
5
1
1
1
0
0
5
1
0
0
3
4
1
No. of species per family
2
2
10
8
1
22
6
3
8
2
5
1
1
4
1
2
1
12
2
7
14
1
1
2
1
1
8
2
1
1
5
4
1
Origin
unassigned
g-n-and.
g-n-and.
g-amaz.
g-amaz.
g-amaz.
g-amaz.
g-n-and.
g-s-and.
g-amaz.
g-amaz.
l.
g-s-and.
g-amaz.
g-s-and.
g-amaz.
g-amaz.
g-n-and.
l.
g-amaz.
g-amaz.
l.
g-amaz.
g-amaz.
l.
l.
g-amaz.
l.
unassigned
g-n-and.
g-amaz.
g-amaz.
unassigned
List 2.: Origin and abundance of families in the research plots
dry = Dry area-centred; g-amaz. = Gondwana-amazon-centred; g-n-and. = Gondwanda-centred Northern Andes; g-s-and. = Gondwanda-centred Southern
Andes: l. = Lauras
181
8. Lists
Class (Subclass)
Suborder
Order
Family
Arecidae
Magnoliidae
Filicales
Cycadales
Coniferales
Arecales
Magnoliales
Cyatheaceae
Zamiaceae
Podocarpaceae
Arecaceae
Annonaceae
Magnoliaceae
Myristicaceae
Menispermaceae
Sabiaceae
Lauraceae
Polygonaceae
Cecropiaceae
Moraceae
Ulmaceae
Urticaceae
Dilleniaceae
Caryocaraceae
Clusiaceae
Marcgraviaceae
Ochnaceae
Quiinaceae
Theaceae
Bombacaceae
Elaeocarpaceae
Malvaceae
Sterculiaceae
Tiliaceae
Lecythidaceae
Caricaceae
Flacourtiaceae
Violaceae
Capparidaceae
Ebenaceae
Sapotaceae
Myrsinaceae
Theophrastaceae
Chrysobalanaceae
Pterophytae
Gymnospermae
Angiospermae-Liliopsida
Angiospermae-Liliopsida
Ranunculales
Caryophyllidae
Hamameliidae
Dilleniidae
Laurales
Polygonales
Urticales
Dilleniales
Theales
Malvales
Lecythidales
Violales
Capparales
Ebenales
Primulales
Rosidae
Rosales
182
No. of sp.
No. of indiv.
2
1
1
5
12
1
6
1
2
15
5
3
22
1
1
2
1
18
2
2
2
1
4
8
2
2
8
6
1
9
5
1
1
15
3
1
12
12
1
1
262
144
11
132
2
14
33
11
60
250
10
5
14
9
206
2
2
9
1
17
21
2
13
81
23
2
74
68
1
2
66
24
1
35
8. Lists
Class (Subclass)
Suborder
Order
Family
Apiales
Sapindales
Araliaceae
Anacardiaceae
Burseraceae
Hippocastanaceae
Meliaceae
Sapindaceae
Simaroubaceae
Caesalpiniaceae
Fabaceae
Mimosaceae
Proteaceae
Lepidobotryaceae
Malpighiaceae
Vochysiaceae
Humiriaceae
Celastraceae
Dichapetalaceae
Hippocrateaceae
Icacinaceae
Euphorbiaceae
Combretaceae
Melastomataceae
Myrtaceae
Rhizophoraceae
Olacaceae
Apocynaceae
Loganiaceae
Bignoniaceae
Oleaceae
Boraginaceae
Rubiaceae
Fabales
Proteales
Polygalales
Linales
Celastrales
Euphorbiales
Myrtales
Asteridae
Rhizophorales
Santalales
Gentianales
Scrophulariales
Lamiales
Rubiales
unidentified Liliopsida
List 3. Taxonomic relationship of all counted families (after CRONQUIST 1988)
183
No. of sp.
No. of indiv.
3
1
8
1
8
7
1
9
10
23
1
1
2
4
1
2
1
2
2
9
2
10
8
1
5
3
1
2
1
3
12
1
50
18
85
3
110
10
4
57
35
67
2
8
4
138
18
12
1
2
20
81
5
32
18
9
29
43
5
3
2
13
40
1
8. Lists
Index
total No. of spp.
total No. of families
total No. of orders
Mean spp. per family
No. of indiv.
Shannon H´ (Log Base 2,718)
Shannon J´ or Eveness (E)
Simpsons Diversity (D)
Simpsons Diversity (1/D)
Simpsons Diversity (1-D)
Alpha-Index
Mean indiv. per sp.
Family Shannon H´(Log Base 10)
Family Shannon J´ or Eveness (E)
Mean spp. per family
No. of only one indiv. per sp.
No. of indiv. of the most represented spp.
Coastal slope
Gorge
Inland slope
Ridge
Total
108
37
22
2.84
588
4.014
0.857
0.028
35.714
0.972
35.227
5.44
1.463
0.926
2.92
38
50
121
46
25
2.63
482
4.122
0.859
0.027
37.037
0.973
51.905
3.98
1.551
0.933
2.63
59
41
134
50
27
2.68
527
4.119
0.841
0.035
28.571
0.965
57.953
3.93
1.556
0.916
2.68
66
71
179
51
30
3.44
847
4.483
0.862
0.019
52.631
0.981
70.490
4.73
1.562
0.910
3.48
72
50
329
70
33
4.70
2444
List 4. Indices and number of diversity
184
4.7
107
108
8. Lists
Species
Family
Abarema
Fab. - Mim
adenophora
A. macradenia
Acacia allenii
Aiouea costaricensis Lauraceae
Alchornea
Euphorbiacae
costaricensis
Cyatheaceae
Alsophila firma
Ampelocera
Ulmaceae
macrocarpa
Amphirrhox longifolia Violaceae
Fab. – Fab.
Andira inermis
Annona amazonica Annonaceae
A. pittieri
Anomospermum
Menispermacaee
reticulatum
Apeiba
membranacea (syn. Tiliaceae
A. aspera)
A. tibourbou
Ardisia compressa Myrsinaceae
A. dodgei
Aspidosperma
Apocynaceae
spruceanum
Bauhinia
Fab. – Caes.
bahiachalensis
B. glabra
B. guianensis
B. manca
Beilschmiedia
Lauraceae
alloiophylla
B. pendula
Hippocastanaceae
Billia columbiana
M
B
G
H
El
N P C
x
x
Ec
Pe
Bo
Br
V
Gy
A
Cc
E
Am
Sr
Hm Ca
0
0
0
5
x
750
200
2400
0
1
0
0
4
0
0
1
1
2
4
0
x
x
900
0
1
0
0
x
x
2100
0
11
0
0
x
800
5
0
3
1
400
1400
1000
1000
0
2
0
0
0
0
0
1
0
0
1
0
3
0
0
0
1050
0
0
0
2
x
900
2
3
1
0
x
1000
2300
700
5
0
0
8
2
0
1
3
0
2
1
4
1400
3
0
2
15
150
0
1
0
0
1000
1300
330
1
0
0
0
3
0
0
0
0
0
0
2
2200
2
0
0
0
3000
2450
0
0
0
3
2
0
0
0
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
x
x
x
x
x
x
x
x
x
x
x
x
185
x
x
x
x
x
x
x
slope
Ridge
1100
x
x
x
Inland
slope
x
x
Gorge
[m]
x
x
Alt. max Coastal
8. Lists
Species
Family
Bombacopsis
Bombacaceae
sessilis
Boroja panamensis Rubiaceae
Brosimum alicastrum Moraceae
B. costaricanum
B. gianense
B. lactenscens
B. utile
Malpighiaceae
Byrsonima crispa
Calatola
Icacinaceae
costaricensis
Calophyllum
Clusiaceae
brasiliense
C. longifolium
Calyptranthes
Myrtaceae
chytraculia
C. pallens
Capparidaceae
Capparis pittieri
Carapa guianensis Meliaceae
Caryocar
Caryocaraceae
costaricense
Caryodaphnopsis
Lauraceae
burgeri
Flacourtiaceae
Casearia arborea
C. tacanensis
Cassipourea elliptica Rhizophoraceae
Moraceae
Castilla tunu
Cecropia obtusifolia Cecropiaceae
C. peltata
Bombacaceae
Ceiba pentandra
Ulmaceae
Celtis schippii
Chaunochiton
Olacaceae
kappleri
Hippocrateaceae
Cheiloclinium
M
B
G
H
El
N P C
Ec
Pe
Bo
Br
V
Gy
A
Cc
E
Am
Sr
Hm Ca
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
0
0
5
6
x
x
x
x
x
1100
1200
500
1200
1200
1500
1000
0
0
2
30
10
21
0
0
3
1
0
2
3
0
0
3
0
5
11
15
3
1
2
2
28
4
7
0
x
2400
0
19
0
0
x
1800
0
0
0
7
x
950
2
0
0
18
x
x
1200
0
0
2
2
x
x
x
2000
1100
2400
0
0
9
0
0
6
0
0
15
7
1
5
600
9
0
0
0
x
600
1
0
1
0
x
x
x
x
x
x
1450
1980
1800
500
1600
1200
1800
740
1
0
2
0
6
0
0
1
1
1
0
10
32
4
1
0
0
0
2
1
1
0
0
0
3
0
5
0
7
2
0
0
450
4
0
1
0
1790
0
0
0
1
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
186
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
slope
Ridge
700
x
x
x
Inland
slope
x
x
Gorge
[m]
x
x
Alt. max Coastal
8. Lists
Species
Family
cognatum
Chimarrhis latifolia Rubiaceae
Chionanthus
Oleaceae
panamensis
Rubiaceae
Chione sylvicola
Chrysochlamys
Clusiaceae
allenii
Ch. grandifolia
Chrysophyllum
Sapotaceae
argentum
Ch. brenesii
Ch. colombianum
Ch. parvulum
Cinnamomum
Lauraceae
neurophyllum
Moraceae
Clarissa biflora
Clavija costaricana Theophrastaceae
Clusiaceae
Clusia cylindrica
C. peninsulae
C. valerii
Coccoloba belizensis Polygonaceae
C. lehmannii
C. obovata
C. standleyana
Compsoneura
Myristicaceae
sprucei
Conostegia
Melastomataceae
lasiopoda
Copaifera camibar Fab. – Caes.
Boraginaceae
Cordia collococca
C. cymosa
C. megalantha
Couratari guianensis Lecythidaceae
M
B
G
H
El
N P C
Ec
Pe
Bo
Br
V
Gy
A
Cc
E
Am
Sr
Hm Ca
x
x
0
0
1
0
x
3000
2
0
0
0
x
1700
0
0
3
3
x
2150
0
0
5
1
x
600
1
6
1
2
1200
3
0
0
0
1500
950
150
1
1
0
0
0
0
0
0
0
0
0
1
1500
0
1
0
0
2100
1200
1000
1000
1100
700
620
1090
250
0
0
0
1
0
0
0
0
50
1
0
0
0
0
0
3
1
0
0
1
1
1
0
0
0
0
8
0
0
0
1
1
5
0
1
14
1200
0
1
0
0
800
0
0
2
2
300
860
1600
900
700
0
0
0
0
0
4
7
1
0
1
0
0
0
1
0
0
1
0
0
0
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
187
x
x
x
x
x
x
x
x
x
x
x
slope
Ridge
850
x
x
Inland
slope
x
x
x
Gorge
[m]
x
x
Alt. max Coastal
8. Lists
Species
Family
Coussarea
hondensis
Crossopetalum
eucymosum
Croton schiedeanus
Cupania livida
Cyathea delgadii
Cymbopetalum
costaricense
Dendropanax
arboreus
D. caucanus
D. sessiliflorus
Dialium guianense
Diospyros
panamense
Discophora
guianensis
Doliocarpus hispidus
D. multiflorus
Duguetia confusa
Duroia costaricensis
Dussia discolor
D. macroprophyllata
Elaeoluma
glabrescens
Eschweilera
integrifolia
E. longirachis
E. pittieri
Eugenia
glandulosopunctata
Euterpe precatoria
Faramea stenura
Rubiaceae
M
B
G
H
El
N P C
x
x
Celastraceae
x
x
x
x
x
x
Euphorbiaceae
Sapindaceae
Cyatheaceae
x
x
x
x
x
x
x
x
x
Ec
Pe
Bo
Br
V
Gy
A
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Fab. – Caes.
x
Am
Sr
Hm Ca
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
0
0
1
0
x
1600
0
1
0
25
1500
2400
2800
0
0
5
1
0
0
0
1
0
1
0
0
1200
7
0
12
3
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
Dilleniaceae
x
x
Annonaceae
Rubiaceae
Fab. – Fab.
x
x
x
x
x
x
2300
0
5
0
0
x
x
x
2440
3200
1000
1
6
0
13
0
0
0
0
0
9
0
2
1200
0
0
1
0
x
1500
0
1
4
2
x
1200
1200
1000
600
900
1500
6
3
0
0
5
4
1
0
1
15
0
0
0
0
0
2
0
7
0
1
4
0
1
2
700
0
0
0
1
1300
0
1
0
0
700
1400
0
1
0
0
0
0
6
0
1300
0
0
6
1
1500
1000
0
0
2
1
0
0
0
0
x
x
x
x
x
x
x
Sapotaceae
x
x
Lecythidaceae
x
x
x
x
x
x
x
Myrtaceae
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
188
x
x
x
x
x
slope
Ridge
1500
Icacinaceae
x
Inland
x
x
x
Gorge
slope
x
x
Alt. max Coastal
[m]
Ebenaceae
Arecaceae
Rubiaceae
E
x
x
x
Annonaceae
Araliaceae
Cc
x
x
8. Lists
Species
Family
Ficus brevibracteata
F. bullenei
F. colubrinae
F. morazaniana
F. nymphaefolia
F. tonduzii
Fusispermum
laxiflorum
Garcinia madruno
Gloeospermum
diversipetalum
Goethalsia meiantha
Gonzalagunia
panamensis
Grias cauliflora
Guarea grandifolia
G. kunthiana
G. pterorhachis
Guatteria amplifolia
G. chiriquensis
G. recurvisepala
Guettarda crispiflora
G. sanblasensis
Hampea
appendiculata
Hasseltia floribunda
Heisteria acuminata
H. concinna
H. scandens
Heliocarpus
appendiculatus
Henriettea odorata
H. succosa
Moraceae
M
B
G
H
El
N P C
x
x
x
x
x
x
x
x
x
x x
x
x x
x x
Violaceae
x
Violaceae
x
Lecythidaceae
Meliaceae
x
x
x
Annonaceae
x
x
x
A
x
x
Cc
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
Melastomataceae
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
0
0
0
0
1
0
2
1
2
2
0
0
800
17
0
3
1
1500
3
1
1
0
1100
0
5
0
1
2000
1
0
0
0
1500
0
8
2
1
480
2300
3000
1100
800
700
1300
2900
1000
12
2
17
0
0
0
0
3
0
7
0
0
0
1
3
0
0
1
8
0
0
1
1
1
0
0
0
8
0
2
12
0
1
3
0
0
x
1700
0
4
0
0
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
2000
2400
800
1000
2
15
0
0
0
0
0
2
0
0
0
0
0
0
1
0
x
x
x
1600
0
1
0
8
x
x
x
189
x
x
x
x
x
1200
1000
0
0
0
0
7
0
0
1
x
x
x
x
x
x
x
x
x
0
0
0
0
10
6
x
x
x
slope
Ridge
0
0
0
0
0
0
x
x
x
x
x
Inland
2000
500
1700
1200
1670
2500
x
x
x
x
x
Gorge
slope
x
x
x
x
x
Alt. max Coastal
[m]
x
x
x
x
x x
x x
Hm Ca
x
x
Flacourtiaceae
Olacaceae
Sr
x
x
x
x
x
Am
x
x
x
E
x
x
x
x
Gy
x
Malvaceae
x
V
x
Rubiaceae
x
Br
x
Tiliaceae
x
Bo
x
x
Rubiaceae
Pe
x
x
Clusiaceae
Tiliaceae
x
Ec
8. Lists
Species
H. tuberculosa
Hirtella americana
H. lemsii
H. papillata
H. triandra
Hirtella trichotoma
Huberodendron
allenii
Humiriastrum
diguense
Hyeronima
alchorneoides
Inga acrocephala
I. acuminata
I. alba
I. densiflora
I. goldmanii
I. jinicuil
I. marginata
I. oerstediana
I. pezizifera
I. polita
I. punctata
I. sapindoides
I. thibaudiana
I. umbellifera
I. venusta
Iriartea deltoidea
Isertia laevis
Jacaratia dolichaula
Lacistema
aggregatum
Lacmellea
Family
M
B
G
H
Chrysobalanaceae x
x
x
x
x
x
x
x
x
x
El
N P C
Ec
Pe
x
x x
x
x
x
x
x
x
x
x
x
x
Bo
x
Br
V
x
x
x
x
Gy
A
Cc
x
x
x
x
x
Am
x
x
x
Bombacaceae
Sr
Hm Ca
x
x
x
x
Euphorbiaceae
Fab. – Mim.
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Arecaceae
Rubiaceae
Caricaceae
x
x
Flacourtiaceae
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
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
190
x
x
x
x
x
x
0
0
0
0
0
0
4
0
0
0
1
0
2
3
1
0
0
1
250
0
1
0
0
x
850
1
1
7
9
x
900
0
1
0
0
x
x
x
2100
1200
1100
2300
500
1900
2400
2500
2000
850
2000
1600
1300
1500
1500
2000
2000
1200
0
1
0
0
0
0
0
2
0
0
1
0
0
1
0
8
0
0
0
0
0
4
4
2
0
0
3
0
0
0
1
0
1
16
0
2
1
0
1
0
0
0
0
0
0
0
0
1
1
1
0
71
0
0
0
1
1
2
1
0
2
0
2
2
0
1
3
1
2
2
11
0
x
3000
0
0
1
1
x
700
18
0
2
2
x
x
x
x
x
x
x
x
slope
Ridge
0
3
0
1
2
0
x
x
Inland
1000
1000
700
300
1300
300
x
x
x
x
x
x
Gorge
slope
x
x
Alt. max Coastal
[m]
x
Humiriaceae
Apocynaceae
E
8. Lists
Species
panamensis
Lacunaria
panamensis
Laetia procera
Lecythis mesophylla
Licania arborea
L. corniculata
L. glabriflora
L. hypoleuca
L. operculipetala
L. sparsipilis
Licaria cufodontisii
Lonchocarpus
pentaphyllus
Lozania pittieri
Lunania mexicana
Mabea occidentalis
Machaerium
floribundum
M. kegelii
M. seemannii
Maclura tinctoria
Macrolobium
hartshornii
Malvaviscus
arboreus
Manilkara
staminodella
Maquira costaricana
Maranthes
panamensis
Marcgravia schippii
Marila laxiflora
M. pluricostata
Family
M
B
G
H
El
Quiinaceae
N P C
x
Flacourtiaceae
Lecythidaceae
Chrysobalanaceae x
x
x
x
Pe
Bo
Br
V
x
x
x
Ec
x
x
x x
Gy
A
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Cc
E
Am
Sr
Hm Ca
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
0
0
1
0
900
1300
800
300
300
1100
1000
900
400
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
2
0
6
3
1
1
1
2
0
9
1
500
0
1
0
1
1000
1700
1000
0
2
0
1
0
0
0
0
13
0
0
0
1700
0
1
0
0
700
1600
1300
1
1
1
0
0
0
0
0
0
0
0
0
1000
0
0
2
9
2000
0
1
0
0
1800
8
0
0
0
1000
0
0
0
2
x
700
0
0
8
1
x
x
2000
1300
1000
0
0
0
0
0
2
0
20
0
1
19
2
x
x
x
x
x
x
x
x
Lauraceae
Fab. – Fab.
x
x
x
Euphorbiaceae
x
x
x
x
Fab. – Fab.
x
x
x
x
Moraceae
x
x
x
x
x
x
x
x
x x
x x
x x
x
x
x
x
x
x
Flacourtiaceae
x
x
x
x
Sapotaceae
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
Chrysobalanaceae
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
Moraceae
Marcgraviaceae
Clusiaceae
x
x
Fab. – Caes.
Malvaceae
x
x
x
x
x
x
x
x
x
x
x
x
191
x
x
x
x
x
x
x
slope
Ridge
500
x
x
Inland
slope
x
x
Gorge
[m]
x
x
Alt. max Coastal
8. Lists
Species
Family
Matayba ingaefolia
M. oppositifolia
Meliosma
donnellsmithii
M. grandiflora
Miconia donaeana
M. trinervia
Micropholis
melinoniana
Minquartia
guianensis
Mortoniodendron
anisophyllum
M. guatemalense
Mouriri gleasoniana
Myrciaria floribunda
Myriocarpa longipes
Naucleopsis ulei
Nectandra umbrosa
Ochroma pyramidale
Ocotea insularis
O. leucoxylon
O. mollifolia
O. pullifolia
O. rubriflora
Odontadenia
puncticulosa
Oenocarpus mapora
Olmedia aspera
Ormosia
panamensis
Otoba
novogranatensis
Ouratea lucens
Sapindaceae
M
B
G
H
x
x
x
x
El
Melastomataceae
x
x
x
x
x
x
Tiliaceae
Br
V
Gy
A
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
Myristicaceae
x
x
x
x
x
E
Am
Sr
Hm Ca
x
x
x
x
x
x
x
Cc
x
Arecaceae
Moraceae
Ochnaceae
Bo
x
x
x
Apocynaceae
Fab. – Fab.
Pe
x
x
x x
x x
x
x
Olacaceae
Melastomataceae
Myrtaceae
Urticaceae
Moraceae
Lauraceae
Bombacaceae
Lauraceae
Ec
x
Sabiaceae
Sapotaceae
N P C
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
800
0
0
0
1
0
0
0
2
1
1
7
0
1
4
0
1
0
1
4
1
x
x
x
x
1200
3
0
3
0
x
1500
1
12
0
0
1600
1000
1000
1400
400
1500
1800
2300
2500
1700
1600
700
0
0
0
0
1
1
0
0
7
0
0
2
0
0
0
5
0
0
4
0
1
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
2
1
1
0
1
4
0
1
0
0
1
0
1000
1
0
0
0
1000
1300
0
0
0
0
1
0
9
3
1000
0
0
0
2
x
2000
1
2
10
3
x
1500
1
0
0
0
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
192
1
1
1600
x
x
1
0
x
x
x
0
0
x
x
x
0
0
x
x
x
200
1500
x
x
x
slope
x
x
x
slope
Ridge
[m]
1800
1800
1600
x
x
Inland
x
x
x
x
Gorge
x
x
x
x
x
Alt. max Coastal
x
8. Lists
Species
O. valerii
Panopsis
suaveolens
Parathesis
aeruginosa
Parkia pendula
Paullinia costata
Pausandra trianae
Peltogyne purpurea
Perebea hispidula
Perrottetia
sessiliflora
Pithecellobium
macradenium
Pleuranthodendron
lindenii
Pleurothyrium
golfodulcensis
P. trianae
Podocarpus
guatemalensis
Pourouma bicolor
Pouteria durlandii
P. foveolata
P. glomerata
P. laevigata
P. reticulata
P. torta
Pradosia sp. nov.
Protium aracouchini
P. costaricense
P. glabrum
P. panamense
P. ravenii
Family
M
B
G
H
x
El
N P C
x
Proteaceae
Fab. – Mim.
Sapindaceae
Euphorbiaceae
Fab. – Caes.
Moraceae
x
Bo
Br
V
Gy
x
Cc
E
Am
x
x
Sr
Hm Ca
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Fab. – Mim.
0
1
0
2700
0
0
0
2
1300
4
0
4
6
925
3100
1100
550
400
0
0
0
0
6
0
1
1
0
1
2
0
3
2
4
0
0
20
11
2
2000
0
11
0
0
200
0
0
2
0
1500
4
0
2
1
600
0
0
1
1
1400
0
2
1
0
x
2300
0
0
1
0
x
x
x
x
x
x
x
x
x
1750
800
1000
1200
300
2100
1900
400
900
1600
1200
1100
700
8
11
4
0
0
0
0
0
17
4
0
1
12
0
0
0
0
0
0
0
1
0
0
4
1
0
0
2
1
0
2
2
2
0
1
0
0
7
2
0
1
0
1
0
0
2
0
2
10
1
5
5
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Lauraceae
x
x
x
x
x
x
x
x
Podocarpaceae
x
x
x
x
x
x
x
x
Cecropiaceae
Sapotaceae
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
Burseraceae
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
193
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
slope
Ridge
0
x
x
Inland
400
x
x
Gorge
slope
x
x
Alt. max Coastal
[m]
x
x
x x
x x
x
Celastraceae
Flacourtiaceae
A
x
x
x
Pe
x
x
Myrsinaceae
Ec
8. Lists
Species
Family
P. schipii
Pseudolmedia spuria Moraceae
Pterocarpus hayesii
Fab. – Fab.
(syn. P. rohrii)
P. officinalis
Vochysiaceae
Qualea paraensis
Quiinaceae
Quiina schippii
Euphorbiaceae
Richeria obovata
Rinorea dasyadena Violaceae
R. hummelii
Annonaceae
Rollinia pittieri
Rondeletia
Rubiaceae
bertieroides
Ruptiliocarpon
Lepidobotryaceae
caracolito
Euphorbiaceae
Sapium allenii
S. laurifolium
S. pachystachys
Sarcaulus
Sapotaceae
brasiliensis
Schizolobium
Fab. – Caes.
parahyba
Schlegelia parviflora Scrophulariaceae
Simaroubaceae
Simarouba amara
Rubiaceae
Simira maxonii
Elaeocarpaceae
Sloanea ampla
S. brachyteptale
S. faginea
S. guianensis
S. medusala
S. slcata
S. zuliaensis
Socratea exorrhiza Arecaceae
Moraceae
Sorocea affinis
M
B
G
H
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
El
x
x
x
x
N P C
Ec
Pe
Bo
Br
x
x
x
x
x
V
Gy
x
x
x
x
x
x
x
x
E
Am
Sr
Hm Ca
x
x
x
x
x
x
Cc
x
x
x
A
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
194
x
x
x
x
x
x
x
x
x
Alt. max Coastal
Gorge
Inland
slope
Ridge
[m]
slope
1100
1200
0
2
0
0
0
0
1
0
1500
0
1
0
0
350
850
700
1000
1200
750
1330
1
0
4
0
0
0
4
1
0
0
0
37
0
0
0
2
3
1
7
0
0
0
50
1
9
4
6
0
1250
0
1
0
0
800
0
1
3
4
400
1700
1900
0
0
0
1
2
3
0
0
0
0
0
0
1200
2
0
0
0
900
16
0
0
0
1900
1800
300
2300
850
1500
1100
780
400
1500
1300
1100
1
0
0
0
0
5
0
1
0
1
0
0
1
0
5
0
0
0
0
3
0
0
3
0
0
2
0
0
0
0
0
0
0
0
17
1
1
2
0
1
4
0
2
0
1
0
20
2
8. Lists
Species
Family
S. cufodontisii
S. pubivena
Souroubea
Marcgraviaceae
sympetala
Stephanopodium
Dichapetalaceae
costaricense
Sterculia recordiana Sterculiaceae
Strychnos
Loganiaceae
panurensis
Symphonia
Clusiaceae
globulifera
Tabebuia chrysantha Bignoniaceae
Talauma gloriensis Magnoliaceae
Sapindaceae
Talisia nervosa
Anacardiaceae
Tapirira myriantha
Terminalia amazonia Combretaceae
T. bucidioides
Ternstroemia
Theaceae
multiovulata
Tetragastris
Burseraceae
panamensis
Tetrapterys
cf.
Malpighiaceae
seemannii
Tetrathylacium
Flacourtiaceae
macrophyllum
Theobroma
Sterculiaceae
simiarum
Topobea
Melastomataceae
maurofernandeziana
Tovomita longifolia Clusiaceae
T. stylosa
T. weddelliana
Trattinnickia aspera Burseraceae
Meliaceae
Trichilia hirta
M
B
x
G
x
H
El
x
N P C
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Pe
x
x
x
x
x
x
x
x
x
x x
x x
x x
x x
x x
x
x
x
Br
x
x
V
Gy
A
Cc
E
Am
Sr
Hm Ca
x
x
x
x
x
x
0
0
0
0
0
0
x
1700
0
0
0
1
x
700
0
0
1
0
1100
2
1
2
5
1100
5
0
0
0
2200
36
1
18
24
1800
1000
940
1500
2000
1600
0
0
0
9
0
0
2
0
0
1
0
4
0
2
2
0
0
0
0
9
0
8
1
0
1250
0
0
1
0
1000
3
0
0
6
280
0
1
0
0
x
1800
3
41
2
0
x
1000
0
0
3
0
1500
0
0
0
1
1300
2000
2000
400
3100
0
0
0
0
0
2
3
0
0
1
2
0
1
2
0
1
0
1
1
0
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
195
x
x
x
x
x
x
x
x
x
x
slope
Ridge
43
1
x
x
x
Inland
800
900
x
x
x
Gorge
slope
x
x
x
Alt. max Coastal
[m]
x
x
x
Bo
x
x
x
x
Ec
8. Lists
Species
Family
T. martiana
T. septentrionalis
T. tuberculata
Trichospermum
Tiliaceae
galeottii
T. grewiifolium
Annonaceae
Unonopsis pittieri
U. theobromifolia
Virola guatemalensis Myristicaceae
V. koschnyi
V. sebifera
V. surinamensis
Clusiaceae
Vismia baccifera
V.macrophylla
Vochysiaceae
Vochysia allenii
V. ferruginea
V. megalophylla
Vouarana guianensis Sapindaceae
Arecaceae
Welfia regia
Xylopia sericophylla Annonaceae
Zamia fairchildiana Zamiaceae
M
B
G
H
El
N P C
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Ec
Pe
Bo
Br
V
Gy
A
Cc
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
Am
Sr
x
x
x
Alt. max Coastal
[m]
slope
Gorge
Inland
slope
Ridge
x
x
x
2080
2200
280
0
2
0
0
4
0
2
3
0
0
5
2
x
x
1700
0
0
0
1
1100
1000
600
2000
1000
1750
1100
1800
800
1000
1040
1000
8
0
0
0
1
1
3
0
0
0
2
0
0
0
0
0
8
1
1
2
4
0
0
1
0
0
0
0
0
12
0
0
0
0
0
4
3
2
0
0
0
0
5
5
1
47
0
0
19
0
0
10
3
8
3
1
1
1
38
35
0
49
1
1
x
x
x
x
x
x
x
x
x
x
x
x
Hm Ca
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
E
1200
700
1680
List 5. Geographic distribution of the species
M = Mexico, B = Belize, G = Guatemala, H = Honduras, El= El Salvador, N = Nicaragua, P = Panama, C = Colombia, Ec = Ecuador, Pe = Peru, Bo = Bolivia, Br = Brazil, V =
Venezuela, Gy= Guayana, A = Antillean, C = Caribbean/Costa Rica, e = endemic, Am = Amazon, SR = P.N. Santa Rosa, HM = Honduras-Mosquito, Ca = Watershed of the
Panama-Canal
196
8. Lists
Herbarlist of the Species
All plants have been collected by Huber and Weissenhofer (H&W). Of the most of the species is one specimen in CR, WU and LI. Some of the
species were determinate in the field without specimen.
Lacmellea panamensis (Woodson) Markg. (H&W 49, H&W 64, H&W 590)
Anacardiaceae
Odontadenia cf. cognata (Stadelm.) Woodson (H&W 1820)
Tapirira myriantha Triana & Planch. (H&W 1367, H&W 1884)
Araliaceae
Annonaceae
Dendropanax arboreus (L.) Decne. & Planch. (H&W 45, H&W 218, H&W 373, H&W
1156, H&W 1594, H&W 1817)
Annona amazonica R. E. Fries (H&W 220)
Dendropanax caucanus (Harms.) Harms. (H&W 1066)
Annona pittieri Donn. Sm. (H&W 1075)
Dendropanax sessiliflorus (Standl. & A. C. Sm.) A. C. Sm. (H&W 3016, H&W 3042)
Cymbopetalum costaricense (Donn. Sm.) Saff. (H&W 589, H&W 698, H&W 1804,
H&W 1837, H&W 1842)
Arecaceae
Duguetia confusa Maas (H&W 1519, H&W 1827, H&W 1913)
Guatteria aff. recurvisepala R.E. Fr. (H&W 309)
Euterpe macrospadix Oerst. (H&W 308)
Guatteria amplifoloa Triana & Planch. (H&W 2103, H&W 2211)
Iriartea deltoidea Ruiz & Pav. (H&W 212)
Guatteria chiriquensis R.E. Fr. (H&W 37)
Oenocarpus mapora H. Karst (H&W 18)
Guatteria sp. nov. (H&W 879, H&W 1196)
Socratea exorrhiza (Mart.) H. Wendl. (H&W 275)
Rollinia pittieri Saff. (H&W 1802, H&W 1887)
Welfia regia Mast. (det. in the field)
Unonopsis pittieri Saffor (H&W 365, H&W 1057)
Bignonicaeae
Unonopsis theobromifolia N. Zamora & L. Poveda (H&W 44, H&W 1219)
Xylopia sericophylla Standl. & L. O. Williams (det. in the field)
Tabebuia chrysantha (Jacq.) G. Nicholson (H&W 805, H&W 1779)
sp. (unidentified)
Apocynaceae
Aspidosperma spruceanum Benth. Ex Muell. Arg. (H&W 1614)
197
8. Lists
Bombacaceae
Capparidaceae
Bombacopsis sessilis (Benth.) Pittier (H&W 473)
Capparis pittieri Standl. (H&W 1421, H&W 2965)
Ceiba pentandra (L.) Gaertn. (H&W 379)
Caricaceae
Huberodendron allenii Standl. & L. O. Williams (det. in the field)
Ochroma lagopus Swartz (H&W 36)
Jacratia dolichaula (Donn. Sm.) Woodson (H&W 762, H&W 2080)
Boraginaceae
Caryocaraceae
cf. Cordia sp. (H&W 1613)
Caryocar costaricense Donn. Sm. (446, 1801)
Cordia collococca L. (H&W 1186, H&W 1202)
Cordia cymosa (Donn. Smith) Standl. (H&W 1177)
Cecropiaceae
Cordia cf. megalantha S. F. Blake (H&W 1072, H&W 1119)
Cecropia obtusifolia Bertoloni (H&W 891,
Burseraceae
H&W 892, H&W 950)
Cecropia peltata L. (H&W 890)
Protium aracouchinii (Aublet) Marchand (H&W 144, H&W 258, H&W 1537, H&W 1540,
Pourouma bicolor C. Martius (H&W 1799)
H&W 1825, H&W 1889, H&W 1904)
Protium cf. schippii Lundell (H&W 1630)
Celastraceae
Protium costaricense (Rose) Engl. (H&W 57, H&W 132, H&W 257, H&W 1818, H&W
1844, H&W 1859)
Crossopetalum eucymosum (Loes. & Pitt.) Lundell (H&W 221)
Protium glabrum (Rose) Engl. (H&W 1593)
Perottetia sessiliflora Lundell (H&W 1078, H&W 1099)
Protium panamense (Rose) I.M. Johnst. (H&W 31, H&W 252, H&W 334, H&W 359,
H&W 554)
Chrysobalanacae
Protium ravenii D. Porter (H&W 508, H&W 1548, H&W 1809, H&W 1822)
Tetragastris panamensis (Engl.) Kuntze (H&W 626, H&W 1401, H&W 1429, H&W
Hirtella americana L. (H&W 1949, H&W 1959)
1561, H&W 1950)
Licania operculipetala Standl. & L.O. Williams (H&W 527)
Trattinnickia aspera (Standl.) Swart (H&W 543)
Licania sp. nov. (H&W 1178)
Licania sparsipilis S.f. Blake (H&W 358, H&W 528)
198
8. Lists
Maranthes panamensis (Standley) Prance & F. White (H&W 292, H&W 434)
Vismia macrophylla Kunth (H&W 167, H&W 1150, H&W 2108)
Hirtella americana L. (H&W 1650)
Combretaceae
Hirtella lemsii L.O. Williams & Prance (H&W 1517, H&W 1592)
Hirtella trichotoma Prance (H&W 1547)
Licania arborea Seem. (det. in the field)
Terminalia amazonica (J. F. Gmel.) Exell (H&W 1394)
Licania corniculata Prance (H&W 1553)
Terminalia bucidoides Standl. & L.O. Williams (H&W 1623)
Licania hypoleuca Benth. (H&W 1427,H&W 1509, H&W 1649, H&W 1395)
Cyatheaceae
Licania sparsipilis Blake (H&W 1651, H&W 1654)
Maranthes panamensis (Standl.) Prance & F. White (H&W 1648)
Alsophila firma Baker (D. S. Conant (det. D. Lautsch)
Cyathea delgadii Sternb. (H&W 420, H&W 1193)
Clusiaceae
Dichapetalaceae
Calophyllum brasiliense Cambess. (H&W 1551)
Calophyllum longifolium Willd. (H&W 127)
Stephanopodium costaricense Prance (H&W 147, H&W 731)
Chrysoclamys grandifolia (L. O. Williams) Hammel comb. ined. (H&W 1139, H&W
2290, H&W 2443, H&W 2500, H&W 2528,)
Dilleniaceae
Clusia cylindrica Hammel (H&W 131, H&W 522, H&W 702)
Clusia peninsulae Hammel -ined. (H&W 477, H&W 535)
Doliocarpus hispidus Standley & L.O. Williams (H&W 438)
Clusia sp.1 (det. in the field)
Doliocarpus multiflorus Standl. (H&W 192, H&W 441, H&W 465)
Clusia sp.2 (det. in the field)
Ebenaceae
Clusia valerii Standl. (H&W 529, H&W 657, H&W 729)
Garcinia madruno (Kunth) Hammel (H&W 1863, H&W 1874)
Diospyros panamense S. Knapp (H&W 1574)
Marila laxiflora Rusby (H&W 32, H&W 73, H&W 91, H&W 187)
Marila pluricostata Standl. & L.O. Williams (H&W 1312)
Elaeocarpaceae
Symphonia globulifera L. f. (H&W 27, H&W 361, H&W 1351)
Tovomita longifolia (Rich.) Hochr. (H&W 110, H&W 400, H&W 1060)
Sloanea faginea Standl. (H&W 1800)
Tovomita stylosa Hemsley (H&W 56, H&W 518)
Sloanea ampla Johnst. (H&W 689)
Tovomita weddeliana Planch. & Triana (H&W 77)
Sloanea brachytepala Ducke (H&W 641, H&W 1557)
Vismia baccifera Triana & Planch. (H&W 2520, H&W 3073)
199
8. Lists
Sloanea guianensis (Aubl.) Benth. (H&W 656, H&W 1424, H&W 1661)
Peltogyne purpurea Pittier (H&W 203, H&W 295)
Sloanea medusala K. Schum. & Pittier (H&W 1784)
Schizolobium parahyba (Vell.) S. F. Blake (det. in the field)
Sloanea sp. (H&W 198, H&W 364)
Fab./Fabaceae
Sloanea sulcata ined. Smith (H&W 1535)
Sloanea zuliaensis Pittier (det. Nelson Zamora)
Andira inerims (W. Wright) Kunth (H&W 3082, H&W 3435)
Dussia discolor (Benth.) Amsh. (H&W 483)
Euphorbiaceae
Dussia macropropyhllata (Donn. Sm.) Harms (H&W 1545, H&W 1860, H&W 2616)
Alchornea costaricensis Pax & K. Hoffm. (H&W 1118)
Lonchocarpus pentaphyllus (Poir.) DC. (H&W 1669)
Croton schiedeanus Schltdl. (H&W 2109, H&W 2133, H&W 2229, H&W 2386, H&W
Machaerium floribundum Benth. (H&W 1281)
3026, H&W 3045)
Machaerium kegelii Meissner (H&W 1242)
Hyeronima alchorneoides Allemao (H&W 467)
Macherium seemannii Benth. (H&W 1846)
Mabea occidentalis Benth. (H&W 89, H&W 262, H&W 452)
Ormosia panamensis Benth. (H&W 1554, H&W 1658, H&W 3010)
Pausandra trianae (Müll. Arg.) Baill. (H&W 42, H&W 401, H&W 618)
Pterocarpus hayesii Hemsl. (H&W 1682)
Richieria obovata (Mull. Arg.) Pax & K. Hoffm. (H&W 478, H&W 1133, H&W 1399,
Pterocarpus cf. officinalis Jacq. (H&W 1187)
H&W 1604)
Fab./Mimosaceae
Sapium allenii Huft (H&W 1126)
Sapium laurifolium (Rich.) Griseb. (H&W 788, H&W 1223)
Abarema adenophora (Ducke) Barneby & Grimes (H&W 1435)
Sapium pachystachys Schum. & Pittier (H&W H&W 861)
Abarema macradenia (Pittier) L. Rico (H&W 1245)
Acacia allenii D. H. Janzen (H&W 580, H&W 1415)
Fab./Caesalpiniaceae
Inga acrocephala Steudel (H&W 229)
Bauhinia bahiachalensis ined. Zamora (H&W 1520)
Inga acuminata Benth. (H&W 1380, H&W 1830)
Bauhinia glabra Jacq. (det. Nelson Zamora)
Inga alba Willd. (H&W 65, H&W 1384)
Bauhina guianensis Aubl. (H&W 2998, H&W 3005)
Inga densiflora Benth. (H&W 1200, H&W 1370, H&W 1627)
Bauhinia manca Standley (H&W 345)
Inga goldmannii Pittier (H&W 1098, H&W 1094)
Copaifera camibar Poveda, Zamora, Sánchez (H&W 572)
Inga jinicuil Schltdl. & Cham. ex G. Don. (H&W 1081, H&W 1129)
Dialium guianense (Aubl.) Sandwith (H&W 1892, H&W 1899, H&W 2517)
Inga marginata Willd. (H&W 1544, H&W 1632)
Macrolobium hartshornii R. S. Cowan (H&W 510, H&W 530)
Inga oerstediana Benth. ex Seem. (H&W 1833)
200
8. Lists
Inga pezizifera Benth. (H&W 1425, H&W 1560)
Hippocrateaceae
Inga polita Killip (H&W 1501)
Inga punctata Willd. (H&W 1897)
Cheiloclinium cognatum A.C. Smith (H&W 1641)
Inga sapindoides Willd. (H&W 1417)
sp. cf. (liana – unidentified)
Inga sp. (H&W 51)
Humiriaceae
Inga sp. 1 (H&W 453)
Inga sp. 2 (H&W 265)
Humiriastrum diguense Cuatrec. (H&W 490, H&W 509, H&W 532)
Inga thibaudiana DC. (H&W 552, H&W 1396, H&W 1438)
Inga umbellifera (Vahl) Steud. (H&W 156, H&W 1662, H&W 1886)
Icacinaceae
Inga venusta Standl. (H&W 1398)
Parkia pendula (Willd.) Benth. ex Walp. (H&W 213, H&W 1637)
Calatola costaricensis Standl. (H&W 3078)
Pithecolobium macradenium Pittier (H&W 142, H&W 184)
Discophora guianensis Miers. (H&W 491, H&W 1457, H&W 1788)
Flacourtiaceae
Lauraceae
Casearia arborea (Rich.) Urb (H&W 1149)
Aiouea costaricensis (Mez) Kosterm. (H&W 663)
Casearia tacanensis Lundell (H&W 1056)
Beilschmiedea pendula (Sw.) Hemsl. (H&W 280)
Hasseltia floribunda Kunth (H&W 2429)
Beilschmiedia alloiophylla (Rusby) Kosterm. (H&W 1895)
Lacistema aggregatum (Bergius) Rusby (H&W 1645, H&W 1946)
Caryodaphnopsis burgeri Zamora & Poveda (H&W 474, H&W 588)
Laetia procera (Poepp.) Eichler (H&W 1449, H&W 1498, H&W 1591)
Cinnamomum neurophyllum (Mez & Pittier) Koesterm. (H&W 670, H&W 1559, H&W
Lozania pittierii (S.F. Blake) L.B. Sm. (H&W 1077, H&W 1586)
1597)
Lunania mexicana Brandegee (H&W 1876)
Licaria cufodontisii Koesterm. (H&W 1906, H&W 1925, H&W 1957)
Pleuranthodendron lindenii (Turcz.) Sleumer (H&W 1369, H&W 1832)
Nectranda umbrosa (Kunth) Mez (H&W 1442, H&W 1442, H&W 1550, H&W 1646,
Tetrathylacium macrophyllum Poeppig (H&W 8, H&W 278, H&W 347)
H&W 1664, H&W 1948)
Ocotea insularis (Meisn.) Mez (H&W 1440, H&W 2110, H&W 2329)
Hippocastanaceae
Ocotea leucoxylon (Sw.) Laness. (H&W 1855, H&W 1879, H&W 1883, H&W 3043)
Ocotea mollifolia Mez & Pittier (H&W 1091)
Billia colombiana Planch. & Lindl. (H&W 740)
Ocotea pullifolia van der Werft (H&W 1659, H&W 3009)
201
8. Lists
Ocotea rubriflora Mez (H&W 1894)
Malvaceae
Pleurothyrium golfodulciensis W. C. Burger & N. Zamora (H&W 230, H&W 1602)
Hampea appendiculata (Donn. Sm.) Standl. (H&W 1229)
Pleurothyrium trianae (Mez) Rohwer (H&W 160)
Malvaviscus arboreus Cav. (H&W 2177)
Lecythidaceae
Marcgraviaceae
Couratari guianensis Aublet (H&W 585)
Eschweilera integrifolia (Ruiz & Pav. ex Miers) R. Knut (H&W 1666)
Marcgravia schippii Standl. (H&W 665)
Eschweilera longirachis S. A. Mori (H&W 3004)
Souroubea sympetala Gilg. (H&W H23, H&W H52)
Eschweilera pittieri Kunth (H&W1374, H&W 1397, H&W 1422, H&W 1606)
Melastomataceae
Grias cauliflora L. (H&W 313)
Lecythis mesophylla Mori (H&W 1534)
Conostegia lasiopoda Benth. (H&W 1157, H&W 1189)
Lepidobotryaceae
Henriettea odorata (Markgr.) Almeda (H&W 1542, H&W 1673)
Ruptiliocarpon caracolito Hammel & N. Zamora (H&W 32, H&W 173, H&W 1130)
Henriettea succosa (Aubl.) DC. (H&W 1653)
Henriettea sp. (H&W 293, H&W 460)
Henriettea tuberculosa (Donn. Sm.) L.O. Williams (H&W 207)
Loganiaceae
Miconia doniana Gleason (H&W 1218)
Miconia sp. (H&W 107, H&W 463)
Strychnos panurensis Sprague & Sandwith (H&W 1839)
Miconia trinervia (Sw.) D. Don. (H&W 300)
Mouriri gleasoniana Stan. ex Stan. & Steyerm. (H&W 342, H&W 1541)
Magnoliaceae
Topobea maurofernandeziana (Triana) Cogn. (H&W 769, H&W 2709)
Talauma gloriensis Pittier (H&W 363, H&W 454, H&W 495, H&W 1366)
Meliaceae
Malphigiaceae
Carapa guanensis Aublet (H&W 120)
Guarea grandifolia A.DC. (H&W 228, H&W 427, H&W 1566)
Byrsonima crispa A. Juss. (H&W 1677)
Guarea kunthiana A. Juss. (H&W 1862, H&W 1880)
Tetrapteris cf. seemannii Triana & Planch. (H&W 1690)
Guarea pterorhachis Harms (H&W 1624)
202
8. Lists
Trichilia hirta L. (H&W 1096, H&W 3006)
Perebea hispidula Standl. (H&W 1448, H&W 1504, H&W 1527, H&W 1635, H&W
Trichilia cf. martina (identified in the field)
1823, H&W 1829, H&W 1852)
Trichilia septentrionalis C. DC. (H&W 66, H&W 617, H&W 1507)
Pseudolmedia spuria (Sw.) Griseb. (H&W 1512, H&W 1896)
Trichilia tuberculata (Triana & Planchon) C. DC. (H&W 1500)
Sorocea affinis Hemsl. (H&W 244, H&W 2415, H&W 2744)
Sorocea cufodontisii W.C. Burger (H&W 1813, H&W 1816, H&W 1851, H&W 1857,
Menispermaceae
H&W 1902, H&W 1915)
Sorocea pubivena Hemsl. (H&W 72, H&W 1908)
Anomospermum reticulatum (Mart.) Eichler (H&W 1476)
Myristicaceae
Moraceae
Compsoneura sprucei (A.DC.) Warb. (H&W 1, H&W 78, H&W 1798, H&W 3033)
Brosimum alicastrum Sw. (H&W 461, H&W 2040)
Otoba novogratensis Moldenke (H&W 86, H&W H&W 336, H&W 440)
Brosimum costaricanum Liebm. (H&W 1634, H&W 1877, H&W 1905)
Virola guatemalsensis (Hemsl.) Warb. (H&W 175, H&W 494, H&W 1558, H&W 3013)
Brosimum guianense (Aubl.) Huber (H&W 1419, H&W 1803, H&W 1555, H&W 3446)
Virola koschnyi Warb. (H&W 180)
Brosimum lactescens (S. Moore) C.C. Berg (H&W 1626, H&W 1814, H&W 1821)
Virola sebifera Aublet (H&W 165, H&W 2277)
Brosimum utile (Kunth) Oken (H&W 694)
Virola surinamensis (Rol. ex Rottb.) Warb. (H&W 1552, H&W 3012)
Castilla tunu Hemsley (H&W 333, H&W 382)
Clarisa biflora Ruiz & Pavon (H&W 1074)
Myrsinaceae
Ficus brevibracteata Burger (H&W 1518)
Ficus bullenei I.M. Johnst. (H&W 875, H&W H33)
Ardisia compressa Kunth (H&W 1403)
Ficus colubrinae Standl. (H&W 874)
Ardisia dodgei Standl. (H&W 622, H&W 2052)
Ficus morazaniana Burger (H&W 3018)
Parathesis aeruginosa Standl. (H&W 26, H&W 1238)
Ficus nymphaeifolia Miller (H&W 687, H&W 1477)
Myrtaceae
Ficus tonduzii Standl. (H&W 371, H&W 1999)
Maclura tinctoria (L.) G. Don. (H&W 1951)
Maquira costaricana (Standl.) C.C. Berg (H&W 1441, H&W 1838)
Calyptranthes chytraculia (L.) Sw. (H&W 1631, H&W 3017)
Naucleopsis ulei (Warb.) Ducke (H&W 1898)
Calyptranthes pallens Griesb. (H&W 1638)
Olmedia aspera Ruiz & Pav. (H&W 1375)
Eugenia glandulosopunctata P.E. Sánchez & Poveda ined. (H&W 1893)
Eugenia sp. (H&W 227, H&W 343, H&W 531)
203
8. Lists
Myrcia sp. nov. 1(H&W 1663)
Coccoloba lehmannii Lindau (H&W 1556)
Myrcia sp. nov. 2 (H&W 3008
Proteaceae
Myrcia sp. nov. 3 (H&W 3015)
Myrciaria floribunda (Willd.) O. Berg (H&W 148, H&W 362, H&W 1436, H&W 1486)
Panopsis mucrunata (Klotzsch & H. Karst.) Pittier (H&W 597, H&W 1655)
Ochnaceae
Quiinaceae
Ouratea lucens (Kunth) Engl. (H&W 48, H&W 357, H&W 1114)
Lacunaria panamensis (Standl.) Standl. (H&W 492)
Ouratea valerii Standley (H&W 331)
Quiina macrophylla Tul. – syn. Q. schippii Standl. (H&W 226, H&W 320, H&W 1888)
Olacaceae
Rhizophoraceae
Chaunochiton kapplerii (Sagot ex Engl.) Ducke (H&W 586, H&W 896, H&W 1819)
Cassipourea elliptica (Sw.) Poiret (H&W 1261, H&W 1362, H&W 1437)
Heisteria acuminata (Humb. & Bonpl.) Engl. (H&W 1848, H&W 1900)
Heisteria concinna Standl. (H&W 1861, H&W 1875)
Rubiaceae
Heisteria scandens Ducke (H&W 264, H&W 550)
Minquartia guianensis Aubl. (H&W 297, H&W 330)
Borojoa panamensis Dwyer (H&W 1376, H&W 2053)
Chionanthus panamensis (Standl.) Stearn (H&W 1890)
Chimarrhis latifolia Standl. (H&W 515, H&W 881, H&W 1093, H&W 1185)
Chione sylvicola (Standl.) W.C. Burger (H&W 1656, H&W 2472, H&W 2570, H&W
Podocarpaceae
3080)
Coussarea hondensis (Standl.) C. M. Taylor & W.C. Burger (H&W 1140, H&W 1976,
Podocarpus guatemalensis Standley (H&W 223)
H&W 2653)
Duroia costaricensis Standl. (H&W 1603, H&W 2049, H&W 2135)
Polygonaceae
Faramea stenura Standl. (H&W 1184, H&W 1221)
Coccoloba belizensis Standley (identified in the field)
Gonzalagunia panamensis (Cav.) K. Schum. (H&W 1808)
Coccoloba lehmannii Lindau (H&W 1657)
Guettarda crispiflora Vahl (H&W 2163)
Coccoloba obovata Kunth (H&W 3007)
Guettarda sanblasiensis Dwyer (H&W 1878)
Coccoloba standleyana P.H. Allen (H&W 835, H&W 1216, H&W 1231, H&W 1381)
Isertia laevis (Triana) B. M. Boom (H&W 15, H&W 2091)
204
8. Lists
Rondeletia bertieroides Standl. (H&W 883)
Pouteria durlandii (Standl.) Baehni (H&W 241, H&W 520, H&W 1806, H&W 1850,
Simira maxonii (Standl.) Steyerm. (H&W W53, H&W 1127)
H&W 1858, H&W 1881, H&W 3038)
Pouteria foveolata T.D. Penn. (H&W 551, H&W 1909)
Sabiaceae
Pouteria glomerata (Miq.) Radlk. (H&W 1400, H&W 3014)
Pouteria laevigata (Mart.) Radlk. (H&W 323)
Meliosma donnell-smithii Urban H&W 1543
Pouteria reticulata (Engl.) Eyma (H&W 267)
Meliosma grandiflora C. Morton ex A. Gentry (H&W 161, H&W 296, H&W 322, H&W
Pouteria torta (Mart.) Radlk. (H&W 276, H&W 1402)
332, H&W 338, H&W 1420)
Pradosia sp. nov. (H&W 1205)
Sarcaulus brasiliensis (A. DC.) Eyma (H&W 1815, H&W 1845)
Sapindaceae
Scrophulariaceae
Cupania livida (Radlk.) Croat (H&W 880, H&W 1368)
Matayba ingifolia Standl. (H&W 106, H&W 219, H&W 1502)
Schlegelia parvifolia (Oerst.) Monarch.( H&W 900)
Matayba oppositifolia (A.Rich.) Britton (H&W 596)
Paullinia costata Schltdl. Cham. (H&W 2999)
Simaroubaceae
sp. (liana – unidentified)
Simarouba amara Aublet (H&W 87)
Talisia nervosa Radlk. (H&W 266)
Vourana guianensis Aublet (H&W 370)
Sterculiaceae
Sapotaceae
Sterculia recordiana Standl. - ined. (H&W 506, H&W 1625, H&W 1828)
Chrysophyllum aff. parvulum Pittier (H&W 1734, H&W 1853, H&W 1854, H&W 3034)
Theobroma simiarum Donn. Sm. (H&W 312, H&W 261)
Chrysophyllum argenteum Jacq. (H&W 1885)
Theaceae
Chrysophyllum brenesii Cronquist (H&W 1901)
Chrysophyllum cf. colombianum Aubrév.) T.D. Penn. (H&W 277, H&W 310)
Ternstroemia multiovulata Q. Jimenez, Gomez & Zamora (H&W 451)
Elaeoluma glabrescens (Mart. & Eichl.) Aubrev. (H&W 210, H&W 339, H&W 456,
H&W 555, H&W 584, H&W 1439, H&W 1773, H&W 1805, H&W 1891)
Theophrastaceae
Manilkara staminodella Gilly (H&W 1701, H&W 1882)
Micropholis melinoniana Pierre (H&W 260, H&W 455, H&W 462)
Clavija costaricana Pittier (H&W 1572)
205
8. Lists
Apeiba membranaceae Spruce ex Benth. (H&W 1796)
Tiliaceae
Apeiba tibourbou Aubl. (H&W 2304)
Gloeospermum diversipetalum Standl. & L. O. Williams (H&W 1807, H&W 1840,
Goethalsia meiantha (Donn. Sm.) Burret (H&W 1329, H&W 1443)
H&W 1910)
Heliocarpus appendiculatus Turcz. (H&W 804)
Rinorea dasyadena A. Robyns (H&W 111, H&W 112, H&W 299, H&W 426, H&W
Mortoniodendron anisophyllum (Standl.) Standl. & Steyerm. (H&W 602, H&W 1907)
488, H&W 1418, H&W 1499)
Mortoniodendron guatemalense Standl. & Steyerm. (H&W 1536)
Rinorea hummelii Sprague (H&W 1377, H&W 1608, H&W 1639
Trichospermum galeottii (Turcz.) Kosterm. (H&W 1562)
Trichospermum grewiifolium (A. Rich.) Koesterm. (H&W 825, H&W 2088)
Vochysiaceae
Ulmaceae
Qualea paraensis Ducke (H&W 157, H&W 306, H&W 482)
Ampelocera macrocarpa Forero & A.H. Gentry (H&W 519, H&W 1940)
Vochysia allenii Stand. & L. O. Williams (det. in the field)
Celtis schippii Standl. (H&W 1824)
Vochysia ferruginea Mart. (H&W 546, H&W 1423)
Vochysia megalophylla Stafleu (H&W 202, H&W 1428)
Urticaceae
Zamiaceae
Myriocarpa longipes Liebm. (H&W 33, H&W 2283)
Zamia fairchildiana L. D. Gomez (H&W 565, H&W 2642)
Violaceae
Amphirrhox longifolia (A.St.-Hill.) Spreng. (H&W 1360, H&W 1378)
Fusispermum laxiflorum Hekking (H&W 1073)
List 6. Herbarlist of the species
206
Lists
207