Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

Composition, structure and regeneration strategy

of Campnosperma panamense (Anacardiaceae) swamp forests

in Darien, Panama

Alicia Ibáñez1*; https://orcid.org/0000-0002-3724-838X

Alexis Baúles2; https://orcid.org/0009-0009-8798-3807

María Alejandra Venegas2; https://orcid.org/0009-0003-6099-1897

Celibeth Sánchez3; https://orcid.org/0009-0005-3479-531X

Rodolfo Flores4,5; https://orcid.org/0000-0002-7911-9228

Indra Candanedo2; https://orcid.org/0009-0002-5038-0606

1. Centro de Estudios y Acción Social Panameño, Panama City, Republic of Panama; ibaneza@gmail.com

(*Correspondence)

2. Technological University of Panama, Panama City, Republic of Panama; alexis.baules@utp.ac.pa,

alejandravenegas1821@gmail.com, indra.candanedo@utp.ac.pa

3. University of Panama, Darien Campus, Villa Darién, Republic of Panama; celibethsanchez2528@gmail.com

4. University of Panama, Chiriquí Campus, Chiriquí, Republic of Panama; rflores1184@hotmail.com

5. Los Naturalistas, P.O. Box 0426-01459 David, Chiriquí, Republic of Panama.

Received 19-II-2024. Corrected 18-VIII-2024. Accepted 26-III-2025.

ABSTRACT

Introduction: Orey (Campnosperma panamense) swamp forests are found on the Caribbean coast of Central

America, from Nicaragua to Panama, and in the Pacific of Colombia to Northern Ecuador. In Panama, orey

grows in monospecific stands or is the dominant species in inundated mixed forests, mainly along the coasts of

Bocas del Toro province and Comarca Ngäbe-Buglé. The species was known to occur in Darien province, in the

Pacific, although almost no information on its distribution and forest extension in the region existed.

Objective: To describe the structure and floristics of orey forests in Darien, map their extension, and propose a

model for their regeneration strategy.

Methods: This work is part of a vegetation mapping project of the Matusagaratí complex of wetlands. It includes

the use of drones, ground truthing, vegetation sampling through temporary plots, and general plant collecting. A

supervised classification of a Landsat satellite image was performed to delimit the orey forest extension. To study

the orey forest regeneration strategy, a digitalization of forest gaps in high resolution WorldView-2 and Planet

Scope images was performed. Gap frequency and turnover time for forest stands were calculated.

Results: Several monospecific orey mature forest patches were found in remote areas of the Matusagaratí com-

plex of wetlands, for a total of 1 267 hectares. A description of the floristics and structure of orey forests in Darien

is presented. A conceptual model of orey mature forest development and gap regeneration is proposed.

Conclusions: Our knowledge of the floristic composition, structure and distribution of orey forests in the

Republic of Panama has increased. For the first time, a model about their regeneration strategy is proposed. These

forests seem to be evolving to different formations. Finally, some hypotheses are proposed about how they might

respond to changing environmental conditions.

Key words: Matusagaratí; complex of wetlands; swamp forests; orey; Darien; regeneration strategy.

https://doi.org/10.15517/rev.biol.trop..v73i1.58860

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

INTRODUCTION

Tropical wetlands provide a range of ecosys-

tem services, including ground water recharge,

conservation of biodiversity, the removal of

excess nutrients from surface waters, and the

sequestration and storage of atmospheric car-

bon (Sjögersten et al., 2011). Tropical forested

swamps have been much less studied than low-

land and mountain forests, with the exception

of mangroves, mainly because of their inacces-

sibility (Ellison, 2004; López & Kursar, 2007).

Knowledge of wetlands tree diversity and ecol-

ogy at the southernmost end of the Mesoameri-

can corridor is limited. Particularly in Panama

most of these swamp forests are continuous to

coastal areas, becoming important conservation

assets that have been largely ignored. Informa-

tion on the floristic composition and ecology of

these forests is scarce, which together with the

enormous socio-economic pressures that they

experience, jeopardize their sustainability, eco-

system services and evolutionary legacy.

The genus Campnosperma Thwaites, with-

in the Anacardiaceae, comprises 15 tree and

shrub species distributed in SE Asia (5 in

Thailand to New Guinea, 1 in Sri Lanka), 5 in

Madagascar, 1 in Seychelles, 1 in Micronesia

and 2 in the Neotropics (Plants of the World

Online [POWO], 2024). Most are restricted to

inundated areas, being the dominant species

in Southeast Asian peat swamps, mainly by

Campnosperma auriculatum (Blume) Hook. f.

and Campnosperma coriaceum (Jack.) Hallier

f. In America, 2 species, also from inundated

areas, have been described: Campnosperma

gummiferum (Benth.) Marchand, restricted to

the Amazon (from S Venezuela to N Peru) and

Campnosperma panamense Standl., that grows

in the Caribbean of Central America to Pacific

Northern South America (POWO, 2024).

RESUMEN

Composición, estructura y estrategia de regeneración de los bosques pantanosos

de Campnosperma panamense (Anacardiaceae) en Darién, Panamá

Introducción: Los bosques pantanosos de orey (Campnosperma panamense) se encuentran en la costa Caribe

de América Central, desde Nicaragua a Panamá, y desde el Pacífico de Colombia hasta el norte de Ecuador. En

Panamá, el orey crece en formaciones monoespecíficas o es la especie dominante en bosques mixtos inundables

de la costa de la provincia de Bocas del Toro y la Comarca Ngäbe-Buglé. Se tenía conocimiento que esta especie

crece también en la provincia de Darién, en el Pacífico, aunque no había información sobre su distribución y

extensión de estas formaciones..

Objetivo: Describir la composición florística y estructura de los bosques de orey de Darién, definir su extensión

y proponer un modelo sobre su estrategia de regeneración.

Métodos: Este trabajo es parte de un proyecto de mapeo de la vegetación del complejo de humedales de

Matusagaratí, e incluyó el uso de drones, verificación de campo, muestreo de vegetación por medio de parcelas

temporales y recolectas de muestras botánicas. Con el fin de delimitar la extensión del bosque de orey, se realizó

una clasificación supervisada de una imagen de satélite Landsat. Para estudiar la estrategia de regeneración del

bosque de orey se hizo una digitalización de claros del bosque en imágenes de alta resolución WorldView-2 y

Planet Scope. Se calculó la frecuencia de claros y el periodo de renovación del bosque.

Resultados: Se encontraron varios parches de bosques maduros monoespecíficos de orey en áreas remotas del

complejo de humedales de Matusagaratí, para un total de 1 267 hectáreas. Se presenta una descripción florística

y estructural de los bosques de orey en Darién. Se propone un modelo conceptual de desarrollo y regeneración

del bosque de orey por medio de claros.

Conclusiones: Nuestro conocimiento sobre la composición florística, estructura y distribución de los bosques de

orey en la República de Panamá ha aumentado. Por primera vez se propone un modelo sobre cómo se regeneran

este tipo de bosques, los cuales parecen estar evolucionando hacia formaciones diferentes. Finalmente, se presen-

tan hipótesis sobre su posible respuesta ante condiciones ambientales cambiantes.

Palabras clave: Matusagaratí; complejo de humedales; bosques de pantano; orey; Darién; estrategia de

regeneración.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

Campnosperma panamense was described

by Stanley (1920) as “panamensis”. Since then,

the species has been named in both ways in the

literature, what has brought a certain degree of

confusion about its correct name. The generic

name Campnosperma, formed by the combi-

nation of two Greek words: “kamptein” and

“sperma”, means the plant has “curved seeds”.

The ending “ma” in “sperma” is neuter, and

so is the generic name derived from it (Stern,

1966). Once the gender of a generic name has

been established, the specific epithet to be used

must agree with it (Manara, 1991). According

to the latest International Code of Nomencla-

ture (Turland et al., 2018), it is recommended

that the adjective ending in “ensis” should be

used for an epithet derived from a geographi-

cal name (if the generic name is of masculine

gender). Since the genus Campnosperma is a

neuter generic name, its specific epithet must

be neuter (Harper, 2012). Therefore, the word

used must end in “ense” which is the neutral

way of referring to Panama in Latin.

Campnosperma panamense formations

have a limited geographic range in the Carib-

bean, from Nicaragua to Panama and the Pacif-

ic, from the Panamanian region of Darien to

Northwest Colombia and Ecuador (Aguirre &

Rangel-Ch, 2005; Carrasquilla, 2005), besides

Coco Island of Costa Rica (TROPICOS, 2024).

In Nicaragua, Ellison (2004) outlined the most

relevant vegetational characteristics of swamp

forests along the Atlantic coast, emphasizing

prominent associations dominated by Ptero-

carpus officinalis Jacq., Carapa guianensis Aubl.

and Campnosperma panamense. Urquhart

(1999) examined swamp regeneration in areas

mostly dominated by Raphia taedigera (Mart.)

Mart. and Campnosperma panamense. In Costa

Rica, flooded forests along the Caribbean coast

are found mainly in the Tortuguero floodplain

and to a lesser extent in the lower Talamanca

region near Limon (Hammel et al., 2004; Webb

& Peralta, 1998). In this area, formations domi-

nated by Campnosperma panamense, Manicaria

saccifera Gaertn. and Raphia taedigera are com-

mon (Hammel et al., 2004), although their

composition varies throughout the landscape.

In the coastal region of Chocó in W Colom-

bia and N Ecuador, Campnosperma panamense

dominates permanently flooded o swamp for-

ests, locally known as “sajales” which have been

reported to support few species (Aguirre &

Rangel-Ch, 2005; Alvarez-Dávila et al., 2016;

Del Valle, 1996).

In Panama, some information on the flo-

ristics and vegetation of forested swamps is

available for the San San Pond Sak (SSPS) wet-

land, a Ramsar Site of international importance,

located in the Northwest Caribbean coast of

the country (Centro Regional Ramsar para la

Capacitación e Investigación sobre Humedales

para el Hemisferio Occidental, 2010). The his-

torical reconstruction of peat accumulation

and geomorphology of the SSPS is related to

the formation of concentric vegetation rings

staggered in successional stages. This large peat

dome is covered by seven phasic communi-

ties, from open short-grassy fields dominated

by sawgrass Cladium P. B row n e in the cen-

ter, passing through dwarf-to-tall vegetation

dominated by Campnosperma, to Raphia palm

forests (Lawson et al., 2014; Phillips et al., 1997;

Sjögersten et al., 2011). Troxler et al. (2012)

have proposed that vegetational rings, as well

as the physiognomy of Campnosperma forests

in SSPS, are explained by nutrient availability,

in particular phosphorus. Studies on carbon

dynamics in SSPS, which includes CO2, CH4,

nutrients fluxes and water table hydrological

modeling, have been advanced by Hoyos-San-

tillan et al. (2015); Hoyos-Santillan et al. (2016)

and Sjögersten et al. (2020).

A maybe larger peat dome exists east of

SSPS, in the Caribbean side of the Comarca

Ngäbe-Buglé, within the Damani-Guariviara

Wetlands of International importance, also a

Ramsar Site. A few, but limited plant inven-

tories and descriptions of the vegetation are

available for this area (López et al., in prep.).

Additional surveys in the same region are part

of a study of carbon dynamics in peatlands,

specifically at the mouth of the Cricamola River

and in Guariviara (Hoyos-Santillan et al., 2014).

Until recently, the presence of Camp-

nosperma panamense in the Pacific coastal

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

wetlands of the Darien in Pacific Panama was

unknown. The species was first reported there

in 2000 (Smithsonian Tropical Research Insti-

tute (SCZ) Herbarium, Acc. No. 12999; Car-

rasquilla, 2005), although no information about

the extension of its formations existed.

Orey forest regeneration: Turnover with-

in forests is driven by stand development in

conjunction with factors influencing tree death

and replacement at various temporal and spa-

tial scales. Gap dynamics play an important

role in the tropical forest regeneration cycle.

Whitmore (1989) recognizes three phases in

gap succession in non-inundated mature tropi-

cal forests: (a) gap-phase, with an opening

of the forest canopy as a result of tree falls;

(b) building-phase consisting of young trees,

mostly shade-intolerant, growing rapidly to fill

the gap and attain the canopy; (c) mature-phase

formed by a canopy of large trees.

This process has been also described in

mangroves. The most important difference is

in the character of small forest gaps. Gaps in

terrestrial forests resulting from the fall of

large trees are normally elliptic, but those in

mangroves are circular and rarely involve falls

of large older trees. Instead, mangrove trees

usually die standing in small clusters of mixed

age cohorts, leaving small circular “scars” or

impressions in the forest, that form a mosaic

of small regeneration patches reflecting vari-

ous ages and stages of canopy recovery (Duke,

2001). Duke (2001) described six phases in the

mangrove regeneration strategy, starting with

a forest showing no gaps, cycles through two

creation phases (initiation and opening) and

three recovery phases (recruitment, filling and

closure), before returning to the original con-

dition. This author also hypothesized that the

creation and regeneration of these gaps prevent

mangrove forests from reaching senescence

stage, hence maintaining the youthful condi-

tions of the forests.

A pattern of circular gaps similar to those

in mangroves has been found to be characteris-

tic of not only mangroves but several monospe-

cific peat swamp forests dominated by species

such as Shorea albida Sym. in Brunei (Becek

et al., 2022), and Campnosperma panamense in

western Colombia (Lamb, 1959) and Caribbean

Panama (Lawson et al., 2014; López et al., in

prep.). This pattern is also characteristic of the

mature orey forests in Darien (this study).

The objective of this paper is to describe

the orey swamp forests in the Matusagaratí

complex of wetlands, Darien. Their extension

has been mapped for the region and informa-

tion on diversity, floristics and structural com-

position is presented. To properly understand

the role of gap regeneration in orey forest

turnover, spatial and temporal gap rates were

determined with the use of remote sensed

data. A model for their regeneration strategy

is also proposed.

MATERIAL AND METHODS

Site description: The study was conducted

in the Matusagaratí complex of wetlands, in

Darien Province, Pacific Panama. Fieldwork

was carried out during 2022 and 2023 (Febru-

ary and March of each year). The Tuira and

Balsas rivers are the backbone of these wet-

lands, one of the largest complex in Central

America, covering around 55 750 ha (Ibáñez et

al., in prep.). The hydrology and vegetation of

this area has been recently studied (Candanedo,

2021; Carol et al., 2020, Carol et al., 2021, Carol

et al., 2022; Carol et al., 2024; Ibáñez et al., in

prep.). The wetlands include river margins and

adjacent floodplains, which contain different

vegetation types such as swamp and inundated

forests, scrublands and herbaceous formations

(Ibáñez et al., in prep.). All these communi-

ties are periodically flooded, and their struc-

ture and species composition defined by their

position in the landscape and the nature and

duration of the inundation, which can be from

estuarine waters related to daily tides, spring

tides, fluvial and/or rainfall flooding (Carol et

al., 2022; Carol et al., 2024; Centro de Estudios

y Acción Social Panameño [CEASPA], 2015;

Grauel, 2004; Ibáñez & Flores, 2020). Rainfall

averages around 2 500 mm/year, with a mean

annual temperature of 21.6 to 24 °C. There is

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

a pronounced seasonality, with a strong dry

season from January to April, and a wet sea-

son from May to December (CEASPA, 2015;

Grauel, 2004).

Orey forests in the Matusagaratí wet-

land were mapped. Fig. 1 shows the extent of

mature orey monospecific forests in the study

area. Besides mature orey monospecific for-

ests which are the objective of this work, there

are several other formations in which orey

also dominates or grows, such as stunted orey

forest and orey scrub, which are preliminary

described in Ibáñez et al., (in prep.). Fig. 2 and

Fig. 3 illustrate different views of the mature

orey forests under study.

Orey forests in Darien remain flooded

with freshwater all year round, where tip-

up pools are also common (Fig. 2B). Recent

hydrological studies show that inundation is

mainly due to rainfall, although water from the

river influences the lower ground strata. The

water table fluctuates in the dry or wet seasons

around 1 m relative to the surface (Carol et al.,

2024). A peat layer in these forests has been

recorded to be of variable depth (30-80 cm)

(Hoyos et al., in prep.).

As mentioned in the introduction Camp-

nosperma panamense (orey) forests in Darien

show a characteristic gap pattern, which has

been preliminary studied (Fig. 3).

Drone recording and ground truthing:

As part of a general vegetation survey of the

wetland region, drone flights were carried out.

A Phantom 4 PRO V 2.0 drone was used to

film and photograph the different ecosystems

and vegetation types in the wetland, as well as

the stages of gap formation and regeneration in

orey forest (Fig. 3). Ground truthing of the area

was later carried out in order to corroborate the

identity of the dominant species in the different

vegetation types (Ibáñez et al., in prep.).

Fig. 1. Orey forest extension in the Matusagaratí complex of wetlands and study areas. A. Inundated area of the Matusagaratí

complex of wetlands (yellow) and orey forests (dark red). B. Field study region in the Balsas river and location of orey

research plots (green). C. Research area for the regeneration study (WorldView-2 image).

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

Satellite image analyses: To map the main

vegetation types of the Matusagaratí complex of

wetlands, a supervised classification of a Land-

sat 5 image from 1998 was carried out with the

program ERDAS IMAGINE 2018. The results

of this mapping process will be published in a

separate paper (Ibáñez et al., in prep.). In order

to define more precisely orey mature forest

patches, a WorldView-2 image from 2017 was

used as a base for further delimiting the orey

patches found with the Landsat analysis.

Study plots: Three 0.1 ha (20 × 50 m) tem-

porary plots were established in mature orey

forest in the Balsas river, near an old logging

site known as Cacerete (Fig. 1A, 1B).

Gap mapping techniques: To study the

orey mature forest regeneration strategy, one of

the forest patches recognized in the field was

selected as study site (2.93 km2) (Fig. 1C). The

high resolution WorldView-2 image from Janu-

ary 2017 was used to initially digitize orey gaps

Fig. 2. Orey forest from the field. A. Orey (Campnosperma panamense) forest. Orey trees in the background, the palm Euterpe

oleracea in the understory, at front. B. Tip-up pool in the forest. C. Orey flowering branch. Photos ©Alicia Ibáñez.

Fig. 3. Aerial view of orey forest and gap formation stages. A. Orey forest from the air. B. Initiation. C. Opening. D.

Intermediate stage between opening and recruitment. E. Recruitment. Photos ©Alexis Baúles.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

in different states of development, with ArcGIS

Pro (Fig. 4). New gaps were mapped and the

stages of all gaps evaluated over a 6-year period,

from February 2017 to February 2023, using

Planet Scope satellite images obtained from

Planet Labs (2023) (Fig. 4). The best possible

Planet images, those less cloudy, were selected,

from the dates: 16 February 2017, 9 Decem-

ber 2017, 13 October 2018, 30 January 2019,

3 November 2020, 20 June 2021, 7 December

2022 and 2 February 2023. They were all geore-

ferenced previous to analyses.

The different gap regeneration stages were

defined as follow (adapted from Fig. 2 in Duke,

2001): A: closed canopy, B: the gap is initiating

and leaves turn brown (not seen in the images),

C: trees died but still standing, D: dead stumps,

fallen branches on gap floor, seedlings starting,

E: regeneration of forest inside the gap, F: trees

in gaps approaching maximal canopy height

(Fig. 3, Fig. 4, Fig. 5, Table 1). The number of

days each gap was in any of the four stages (C,

D, E, F) was counted and the mean calculated

for each stage (Table 1).

Vegetation census: To characterize orey

mature forest (forest structure and species diver-

sity), vegetation inventories were conducted in

the plots; all stems ≥ 5 cm in diameter at breast

height (DBH) were mapped, measured and

marked. Voucher samples of all species were

collected, and their identities verified at the

Herbarium of the University of Panama (PMA),

following TROPICOS (2024) nomenclature.

To describe the forest structure, the most

common descriptors reported in the literature

Fig. 4. Orey forest with gaps at various stages of recovery. A. Different stages of gap formation and recovery as seen in the

WorldView-2 satellite image. B. Digitalized gaps at different stages of recovery in the study area. Composite images of bands

4, 3 and 2 (RGB) were used in order to obtain the greatest spectral difference of gaps.

Table 1

Gap phases, characteristics and estimated age in orey forest in Darien.

Gap Phase Gap Characteristics Estimated Age

A. Closed canopy No gap presence -

B. Initiation Leaves brown, still in the trees unknown

C. Opening Bare branches and twigs, dead trees standing 1.5 years

D. Recruitment Dead stumps, fallen branches on gap floor, seedlings starting 2.3 years

E. Growth Saplings and small trees growing in gap 4.2 years

F. Closure Trees in gaps approaching site maximal canopy height 3.6 years

TOTAL 11.6 years

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

were used. The basal area of the tree species

found in the plots was calculated from the

DBH data. The relative abundance (RA, %)

was defined as the number of individuals of a

species in proportion to the total number of

individuals in the entire sample. Similarly, the

relative frequency (RF, %) was calculated as the

number of quadrants in which a species is pres-

ent; the relative dominance (RD, %) as the basal

area of a species in proportion to the total basal

area, and the Importance Value Index (IVI) as

the sum of RA + RF + RD, following Dallmeier

et al. (1992) (Table 2). To further assess forest

structure, diametric distribution graphs were

obtained (Fig. 6).

Sampling effort was evaluated based on the

species accumulation curve, using the number

of estimated species versus the number of indi-

viduals (Fig. 7). The curve was calculated using

a random order of the individuals with the pro-

gram iNEXT (Chao et al., 2016). The expected

maximum diversity was calculated with the

non-parametric estimator Chao-1, which uses

the numbers of singletons and doubletons to

estimate the number of undetected species,

as undetected species information is mostly

concentrated on those low frequency counts

(Chao, 1984). The program SpadeR was used

(Chao et al., 2015).

Planet Scope imagery limitations: The

major benefit of Planet Scope imagery is the

availability of at least one image per year

through the 6-year period of study, such that

Fig. 5. Six phases in orey forest gap creation and recovery cycle. Adapted from Duke (2001).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

we were able to approximately date gaps and

determine gap rate and turnover time. How-

ever, there are several disadvantages of using

that imagery, including relatively low resolu-

tion (compared to WorldView-2) and visibility

interference (e.g. edge shadow, cloud shadow).

Gap frequency and turnover time: Gap

frequency and turnover time for the study

region over a 6 year period were calculated.

Gap frequency (Rg, % yr-1) was defined as the

percentage of gap area (Ag) per sampled region

(As), i.e. Rg = 100 x Ag/As x 1 / 6 yr for all gap

area observed during the 6-year study period.

Fig. 4B shows the distribution of gaps through-

out the study area in the period studied.

The average rate of forest turnover due

to gaps (Tg) is the inverse of gap frequency.

This metric reflects the time required for gaps

to impact all the orey forest, solely based on

their rate of occurrence. Table 1 shows an esti-

mated age for the different gap stages according

to our data.

Table 2

List of tree species (DBH ≥ 5 cm) and their abundance indexes.

Family Species NRA RF BA RD IVI

Anacardiaceae Campnosperma panamense 171 38.26 13.64 9.35 72.66 124.55

Arecaceae Euterpe oleracea 234 52.35 13.64 2.76 21.47 87.46

Arecaceae Elaeis oleifera (Kunth) Cortés 4 0.89 9.09 0.35 2.75 12.73

Aquifoliaceae Ilex guianensis (Aubl.) Kuntze 10 2.24 9.09 0.18 1.38 12.71

Euphorbiaceae Alchornea grandis Benth. 7 1.57 9.09 0.03 0.20 10.86

Moraceae Ficus cf. popenoei Standl. 3 0.67 9.09 0.04 0.35 10.11

Apocynaceae Lacmellea panamensis (Woodson) Markgr. 2 0.45 9.09 0.02 0.14 9.68

Rhizophoraceae Cassipourea elliptica (Sw.) Poir. 10 2.24 4.55 0.06 0.45 7.24

Araliaceae Dendropanax arboreus (L.) Decne. & Planch. 2 0.45 4.55 0.03 0.23 5.23

Malvaceae Pseudobombax septenatum (Jacq.) Dugand 1 0.22 4.55 0.03 0.21 4.98

Euphorbiaceae Alchornea sp. 1 1 0.22 4.55 0.01 0.09 4.86

Chrysobalanaceae Parinari chocoensis Prance 1 0.22 4.55 0.00 0.03 4.80

Melastomataceae Henriettea succosa (Aubl.) DC. 1 0.22 4.55 0.00 0.03 4.80

TOTAL 447 100 100 12.87 100 300

N: number of individuals, RA: relative abundance, RF: relative frequency, BA: basal area, RD: relative dominance, IVI:

importance value index.

Fig. 6. Diametric distributions of orey forest in Darien. A. Only Campnosperma panamense included. B. All tree species

included.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

RESULTS

Several patches of dense orey forest were

identified in the study region, totaling an area

of 1 267 hectares. Two of these patches were

visited and studied by means of plots (Fig. 1,

Fig. 2, Fig. 3).

General findings: A total of 447 indi-

viduals (815 stems), including trees and palms

greater than 5 cm DBH, were recorded in 0.3

ha. We found 13 species, in 12 genera and 11

families. Two species, Campnosperma pana-

mense and the palm Euterpe oleracea Mart.,

made up to 90.60 % of the total individuals

sampled and show the highest IVI, 124.55 and

87.46 respectively. They are both the dominant

species in this kind of wetland forest (Table

2). While Campnosperma panamense forms

the canopy, which reaches c. 20 m, the multi-

stemed palm Euterpe oleracea is the dominant

species in the understory and reaches heights

of 2-12 m. Common associates are the shrubs:

Tococa guianensis Aubl., Ardisia sp. 1, Ardi-

sia sp. 2, Psychotria hoffmannseggiana (Roem.

& Schult.) Müll. Arg., Psychotria poeppigiana

Müll. Arg., Palicourea triphylla DC.; herbaceous

plants: Monotagma plurispicatum (Körn.) K.

Schum., Spathiphyllum phryniifolium Schott;

lianas and epiphytes: Schradera sp., Thora-

cocarpus bissectus (Vell.) Harling, Anthurium

clavigerum Poepp., Monstera pinnatipartita

Schott, Philodendron fragrantissimum (Hook.)

G. Don and numerous fern species.

Vernacular names and uses of orey:

Campnosperma panamense receives different

names in its area of occurrence: the name

recorded by our team in Darien was “sajo”

(only known in the locality of Camogantí),

same as in Western Colombia (Del Valle, 1996).

The formation is called “sajal” in Darien and

Colombia. In other regions of Panama it is

called orey, orí (Bocas del Toro) (Phillips et al.,

1997), degetda tain (Comarca Ngäbe-Buglé)

(Flores et al., 2021) and nusmas (Guna Yala)

(TROPICOS, 2024).

No uses were recorded for this species in

Darien. Carrasquilla (2005) reports that its

wood is used for paper pulp. In the Comarca

Ngäbe-Buglé it is used as building material to

construct houses (beams, posts and walls) and

canoes. Its bark is cooked and used in tradi-

tional medicine (Flores et al., 2021). Bioactive

flavonoids with antimalarial and anti Leishma-

nia activity have been isolated from its leaves

(Weniger et al., 2004).

Diametric distribution and stand struc-

ture: The analysis of diameter distributions for

different tree populations allows us to evaluate

the ecological and conservation conditions of

the forest. The diameter distribution for Camp-

nosperma panamense (Fig. 6A) showed highest

proportion (36 %) of individuals in the middle

Fig. 7. Species-individuals curve.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

classes (20-30 cm DBH), while 85 % had diam-

eters from 10-40 DBH. Only 15 individuals

(9 %) belonged to the lowest class (5-10 cm).

A distribution that included all tree species

showed similar results, although with higher

individuals in the lowest class (Fig. 6B).

Sampling effort: The number of tree spe-

cies increased as new individuals were sampled

(Fig. 7). However, most increment appears

up to the 447 trees sampled, after which the

extrapolation curve shows few species addi-

tions, and seems to almost stabilize. This indi-

cates that our inventory captured much of the

orey forest tree diversity.

On the other hand, the total number of

species in all 3 plots was 13. This is 76 % of the

expected diversity that according to Chao-1

index is estimated to be around 17 species. This

would also indicate that the tree inventory is

quite complete, and it adequately represents the

orey forest tree diversity. Chao-1 is a diversity

index sensitive to the number of rare species

found in the sample (species that only have one

individual “singletons” or two individuals in

the entire sample “doubletons”) (Colwell, 2009).

We found a total of 4 species in our sampling

that were registered only once, and two spe-

cies of which only two individuals were found,

which together, singletons and doubletons, rep-

resent about 46 % of species in the sample.

Gap frequency and forest turnover: A

total of 472 gaps were digitized from the sat-

ellite images for the study area in the period

under study, with a mean area of 1 046 m2.

Gap recovery in orey forests is characterized in

six phases (closed canopy, initiation, opening,

recruitment, growth and closure), and it takes

about 11.6 years for complete gap recovery

(Table 1). Approximately 16.83 % (0.49 km2)

of the orey forest regeneration study area (2.93

km2) experienced gaps over the 6-year study

period. Gap frequency was 2.78 % yr-1, (0.081

km2 or 8.1 ha/yr-1), which corresponds to a for-

est turnover time of 35 years.

DISCUSSION

Forest structure and diversity: This is the

first study of the orey swamp forests of Darien,

Panama, where 1 267 hectares of mature forma-

tions were mapped. Campnosperma panamense

forms large monospecific stands (IVI of 124.55

and 72 % of the basal area) on permanently

inundated ground with peat of around 30-80

cm deep. These formations are structurally

similar to monospecific forests in SSPS, Bocas

del Toro province, Caribbean coast of Panama,

where C. panamense showed 70 % of the basal

area (Phillips et al., 1997; Sjögersten et al.,

2011). They are also similar to the “sajo” forests

in coastal Western Colombia, with a 74-92 % of

Campnosperma in their forests (Alvarez-Dávila

et al., 2016; Del Valle, 1996). In Darien, middle

canopy and understory is dominated by the

colonial palm Euterpe oleracea, known here as

“murrapo”, which shows pneumatophores of

up to 1 m height, indicating an adaptation to

permanently waterlogged conditions. This spe-

cies does not grow in Caribbean orey forests,

where a non-colonial species of the same genus,

Euterpe precatoria Mart., is common. Euterpe

oleracea is also abundant in Pacific Colombian

sites, where it is called “naidí” (Alvarez-Dávila

et al., 2016; Del Valle, 1996).

Tree diversity in orey forests of Darien was

low compared to Panamanian lowland forests

on mineral soils, which typically contain around

100 sp/ha (e.g. Pyke et al., 2001). Low diversity

tree communities are typical of Central and

South American wetlands (Ellison, 2004; López

& Kursar, 2007; Webb & Peralta, 1998). Species

richness (13 sp ≥ 5 cm DBH in 0.3 ha, 10 sp ≥

10 cm DBH / 0.3 ha, 4-7 sp ≥ 10 cm DBH / 0.1

ha) was similar to SSPS orey forest (7 sp ≥ 10

cm DBH / 0.1 ha) (Sjögersten et al., 2011) and

Colombian “sajo” forests (9 sp ≥ 10 cm DBH /

0.5 ha) (Alvarez-Dávila et al., 2016).

In this sense, the low species richness in

the tropical swamp forests is generally due to

the inability of solid ground species to face the

stress conditions that result from the perma-

nent flooding and the acid peat soils typical of

Campnosperma forests (López & Kursar, 2003;

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

Sjögerstenet al., 2011). These patterns of low

diversity have been described by Richards

(1952) and Whitmore (1975) in South East

Asian peatland forests dominated by Campno-

sperma brevipetiolatum Volkens (New Guinea),

C. coriaceum (Malaysia) and Shorea albida in

Borneo (Whitmore, 1975).

The analysis of diameter distributions,

with a higher number of trees within the inter-

mediate size classes, indicates a lack of enough

young trees for replacement of the older ones.

This lack of regeneration could be explained by

human activities (logging) or else a natural age-

ing trend (Bermadzki et al., 1998). In Darien,

orey forests have never been logged or show any

other kind of human impact, so we can infer

they are naturally evolving to a mixed type or to

a more open formation. Inundated mixed for-

ests and more open formations, such as stunted

or scrub orey are common in the surroundings

of the main mature orey forest patches. The

unimodal diametric structure, with mostly con-

temporary individuals and the lack of enough

young trees has also been reported for Colom-

bian orey forests (Lamb, 1959), although there

it has been explained by their secondary nature

due to logging activities (Del Valle, 2000).

Forest regeneration: A conceptual model

of orey forest development and gap regen-

eration has been proposed. Preliminary evi-

dence shown in this study indicates that orey

mature forest regenerates in a similar way to

that reported in mangroves by Duke (2001),

in which gaps are formed by trees which die

standing in small clusters. Gap recovery in orey

forests can also be characterized in six phases

(closed canopy, initiation, opening, recruit-

ment, growth and closure).

The importance of gap creation on forest

turnover can be explored further using the rela-

tionship between area of gaps formed in each

time and the rate of gap recovery. Our results

indicate that orey forests in Darien seem to be

very dynamic with a forest turnover time of 35

years. Around 17 % of the area was in gap mode

during the 6 years of the studied period, with a

gap frequency rate of 8.1 ha/year and only takes

12 years for gaps to recover. As a comparison,

for a mangrove forest dominated by Rhizophora

L. trees in Panama, it took them around fifteen

years to achieve early gap closure (Duke, 2001).

Phillips et al. (1997), after evaluating air

photos of orey forests in SSPS from 1954, 1981

and 1992, saw striking differences in the distri-

bution of the distinctive Campnosperma canopy

within the region, and proposed that stands

develop rapidly and may be quite short-lived.

It seems that orey forests in Darien are

constantly re-newing themselves, with a high

frequency of gap formation. Assuming this

is correct, turnover in these forests might

occur entirely via gap formation rather than

via trees getting older. This is why such for-

ests may never reach the senescence phase of

stand development.

Factors influencing gap creation: There

is no accepted common cause for gap creation

in mangroves, although different theories have

been proposed, such as windstorms, lightning

strikes, frost damage, hail damage, plant patho-

gens, wood-boring insects, etc. However, the

most common gaps in mangroves are small

gaps comprising around 10-20 trees and reput-

edly caused by lightning (Duke, 2001).

In orey forests of Darien, we can hypoth-

esize that they are not caused by lightning. Pre-

liminary evidence showed an initiation stage

in which trees in the gap had dried leaves.

Although there was no sign of disease, it seems

reasonable to think about some kind of plant

pathogen. For these forests, still remains the

question of what triggers gap creation.

Conservation: Orey forests in Darien are

unique ecosystems that need recognition and

strict conservation. The orey mature forma-

tions described here are included in two pro-

tected areas, although they are not managed.

Given their remote location and inaccessibility,

at present there are no direct threats, except

those related with climate change.

Much of the importance of these forests

rests in their ecosystemic roles. First of all, the

Matusagaratí wetland has a hydrologic function

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e58860, enero-diciembre 2025 (Publicado Abr. 28, 2025)

as a buffer for coastal ecosystems, regulating

hydrological variations due to rainfall. This

function of filtering and buffering is vital for

the well-being of marine ecosystems in the

region. Second is the value of the wetland as a

reservoir of fresh water during the rainy season,

as in the dry season, the maintenance of the

caudal of the rivers is due to the subterranean

flux of water from the wetland. Also, we can

highlight the value of these forests as a reservoir

of organic carbon, as prospections have shown

the presence of peat in these forests (Hoyos et

al., in prep.).

Estuarine and continental wetlands, such

as the ones in Matusagaratí are very vulnerable

to climate change. Hydrological studies indicate

that these forests depend on rain water that

maintains the water table, so severe drought

events seem to be the main threat. Rising sea

levels is an important second issue, as orey

forests may experience severe disturbances

because of salinity. All these changes may sup-

pose the collapse of stands or even of large

areas. Such vegetation changes may already

be taking place, but they are very difficult to

distinguish as few studies are focusing on this

issue. A long-term monitoring program should

be a priority to record these phenomena and

also to study the carbon accumulation process

in the peats, in order to estimate the value of the

wetland as a carbon reservoir.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

This research has been possible thanks

to the funding provided by the Panamani-

an National Secretariat for Science, Technol-

ogy and Innovation (SENACYT), Project

PFID-FID-2021-114. We thank the Ministry

of Environment for collecting permits (SE/

AP-21-18, ARB-012-2022), the Technological

University of Panama (UTP) for the scientific

and logistical support and to CEMCIT-AIP for

funds management. Thanks to Jorge Hoyos,

Hermel López, Manuel Arcia, Kasey Clark

and Daniel Holness for their help in different

stages of the research. To the personnel of the

Herbaria at the University of Panama and the

Smithsonian Tropical Research Institute (PMA,

SCZ), and the specialists that helped with

plant identifications: Orlando Ortiz, Marco

Cedeño, Cristian López and Charlotte Tay-

lor. To our field assistants, Osiris Rodríguez,

Aurelio Flaco, Jorge Tomí, Ismael Flaco, Hayro

Cunampio, Eduardo Garabato, Alberto Arroyo

and Antonio Martínez.

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