Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64710, mayo 2025 (Publicado May. 15, 2025)

State of knowledge of the Glomeromycota of Costa Rica

María Mabel De Jesús-Alarcón1; https://orcid.org/0000-0002-2575-0372

Laura Yesenia Solís-Ramos2*; https://orcid.org/0000-0002-8935-5507

Antonio Andrade-Torres3*; https://orcid.org/0000-0001-9387-0483

1. Maestría en Ciencias en Ecología y Biotecnología, Instituto de Biotecnología y Ecología Aplicada, Universidad Veracruzana.

CAUVER 173 Ecología y manejo de la biodiversidad. Xalapa, Veracruz, México. mdejesusalarcon@gmail.com

2. Biotecnología de Plantas y Hongos Micorrícicos Arbusculares (Biotec-PYHMA), Escuela de Biología y Centro de

Investigación en Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa Rica, San Pedro de Montes de Oca,

San José 11501-2060, Costa Rica. laura.solisramos@ucr.ac.cr (*Correspondonce).

3. Biotecnología y Ecología de Organismos Simbióticos, CAUV-173 Ecología y Manejo de la Biodiversidad, INBIOTECA

(Instituto de Biotecnología y Ecología Aplicada), Universidad Veracruzana, Av. de las Culturas Veracruzanas No. 101,

Col. E. Zapata, Xalapa 91090, Veracruz, México. aandrade@uv.mx (*Correspondonce).

Recibido 30-VIII-2024. Corregido 30-I-2025. Aceptado 07-IV-2025.

ABSTRACT

Introduction: Arbuscular mycorrhizal fungi (AMF) play a pivotal role in plant nutrition and soil stability, also

play an important role for sustainable agriculture and the restoration of degraded areas. In Costa Rica, research

has been conducted on the diversity of AMF in select ecosystems. However, a comprehensive compilation of

these studies is currently lacking.

Objective: To compile an updated list of arbuscular mycorrhizal fungi (AMF) in Costa Rica, organised by vegeta-

tion types and life zones. This will facilitate a more comprehensive understanding of their diversity and serve as a

foundation for future research in taxonomy, distribution, ecology and biotechnological applications.

Methods: A comprehensive literature search was conducted in Google Scholar, Scopus and Web of Science,

utilising Spanish and English terms related to arbuscular mycorrhizal fungi (AMF) and Costa Rica, spanning the

period from 1971 to 2024. The selection criterion included studies that identified species based on morphological

or molecular criteria. The following information was integrated into a database.

Results: We compiled 57 studies published from 1975-2024 and after screening 549 initial records, 115 records

corresponding to species level were confirmed. 60 AMF species belonging to five orders were identified, rep-

resenting 16 % of the global diversity known as Glomeromycota. The families Glomeraceae (23 species) and

Acaulosporaceae (19 species) were the best represented. Furthermore, it was observed that studies have been

conducted in only four of the 12 life zones described for Costa Rica, and include agroecosystems and trap crops

(consortia) or pure crops (monospecific).

Conclusions: It is evident that there has been a degree of advancement in the comprehension of the ecology,

diversity and distribution of AMF. Nevertheless, further ecological and taxonomic research is required, encom-

passing both morphological and molecular analyses, to facilitate the expansion of knowledge concerning the

distribution and diversity of AMF in Costa Rica.

Keywords: Arbuscular mycorrhizal fungi; distribution; ecological diversity; taxonomy; life zones.

https://doi.org/10.15517/rev.biol.trop..v73iS2.64710

SUPPLEMENT

SECTION: MUSEUM

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

INTRODUCTION

Arbuscular mycorrhizal fungi (AMF) are a

fungal group that is widely distributed in most

terrestrial ecosystems (Schüβler et al., 2001),

which are associated with 72 % of vascular

plants (Brundrett & Tedersoo, 2018). AMF

belonging to the phylum Glomeromycota were

previously classified within the Zygomycota

(Morton & Benny, 1990), but in recent decades,

they have been recognized as an independent

and ancient phylum of the kingdom Fungi

(Schüßler et al., 2001). These fungi are obli-

gate symbionts of terrestrial plants and play

an important role in nutrient uptake, particu-

larly phosphorus absorption, improving plant

growth and the tolerance to different types of

stress (Brundrett & Tedersoo, 2018; Smith &

Read, 2010; Wang & Qiu, 2006). They also con-

tribute to soil stabilization (Rillig & Steinberg,

2002), and facilitate plant establishment and

recruitment through multiple hyphal networks

that connect plant-plant (Van der Heijden &

Horton, 2009; Van der Heijden et al., 2015).

They also play an important role in agro-

ecosystems because of their biotechnological

use as biofertilizers (Jeffries & Barea, 2012).

This highlights their importance for sustaina-

ble agriculture and the restoration of degraded

areas (Carrillo-Saucedo et al., 2022).

The phylum Glomeromycota includes

approximately 371 species distributed in three

classes, five orders, 16 families, and 48 genera

(Goto & Jobim, 2024; Wijayawardene et al.,

2022). A number of compilations of AMF

diversity have been made at multiple levels,

providing valuable information on their bio-

geographic distribution and ecological roles.

At the biogeographic region level, Stürmer and

Kemmelmeier (2021) documented 221 AMF

species in the Neotropics, representing 69% of

the global diversity described for the phylum

Glomeromycota. The study encompassed 11

biomes and 52 ecological regions, emphasizing

the ecological significance of AMF in diverse

ecosystems, including tropical rainforests and

grasslands. However, it should be noted that the

study did not include data for Costa Rica.

RESUMEN

Estado del conocimiento de los Glomeromycota de Costa Rica

Introducción: Los hongos micorrícicos arbusculares (HMA) son clave para la nutrición vegetal y la estabi-

lidad del suelo, también juegan un papel importante en la agricultura sostenible y la restauración de áreas

degradadas. En Costa Rica, su diversidad ha sido estudiada en algunos ecosistemas, pero a la fecha no hay una

compilación integral.

Objetivo: Compilar una lista actualizada de hongos micorrícicos arbusculares (HMA) en Costa Rica, organizada

por tipos de vegetación y zonas de vida, que amplíe la comprensión de su diversidad y sirva de base para estudios

futuros en taxonomía, distribución, ecología y aplicaciones biotecnológicas.

Métodos: Se realizó una búsqueda bibliográfica en Google Scholar, Scopus y Web of Science, empleando térmi-

nos en español e inglés relacionados con hongos micorrícicos arbusculares (HMA) y Costa Rica, en el periodo

1971-2024. Se incluyeron únicamente estudios que identificaran especies basándose en criterios morfológicos o

moleculares, integrando la siguiente información en una base de datos.

Resultados: Compilamos 57 estudios publicados entre 1975-2024 y después de depurar 549 registros iniciales, se

confirmó que 115 registros corresponden al nivel de especie. Se identifican 60 especies de HMA pertenecientes

a cinco órdenes, lo que representa el 16 % de la diversidad global conocida como Glomeromycota. Las familias

Glomeraceae (23 especies) y Acaulosporaceae (19 especies) fueron las mejor representadas. Además, se observó

que los estudios se han llevado a cabo únicamente en cuatro de las 12 zonas de vida descritas para Costa Rica,

además de algunos estudios en agroecosistemas, cultivos trampa (consorcios) y cultivos puros (monoespecíficos).

Conclusiones: Se ha avanzado en el conocimiento de la ecología, diversidad y distribución de los HMA, sin

embargo, es necesario continuar realizando investigaciones ecológicas y taxonómicas que incluyan tanto análisis

morfológicos como moleculares, con el fin de ampliar el conocimiento sobre la distribución y diversidad de los

HMA en Costa Rica.

Palabras clave: Hongos micorrícicos arbusculares; distribución; diversidad ecológica; taxonomía; zonas de vida.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64710, mayo 2025 (Publicado May. 15, 2025)

Research has been published from Mexico

and Peru, two countries which are characterised

by high biodiversity. In Mexico, Álvarez-López-

tello et al. (2023) and Polo-Marcial et al. (2021)

identified 160 AMF species, highlighting that

the majority of studies have been conducted

in agroecosystems and xerophytic shrublands,

with Oaxaca identified as a prominent bio-

logical hotspot. In Peru, Vega-Herrera et al.

(2023) reported 93 species classified according

to natural regions (Amazon, Andes and Coast),

reflecting the ecological heterogeneity of the

country and the predominance of families such

as Glomeraceae and Acaulosporaceae. At the

ecosystem-specific level, Marinho et al. (2018)

addressed AMF diversity in Costa Rica’s tro-

pical forests. This research highlighted that

tropical rainforests, a representative ecosystem

of the country, harbour a significant proportion

of AMF species, underscoring their ecological

importance in processes such as enhancing

nutrient uptake and soil stabilisation. Notably,

Costa Rican specimens have played a pivotal

role in the taxonomic classification of novel

species, underscoring the country’s significance

in mycological research. For instance, Acaulos-

pora foveata and A. tuberculata were described

using paratypes collected in Costa Rica (Mor-

ton, 1990), while Acaulospora splendida was

identified with a holotype from the country

(Sieverding et al., 1988). Beyond these contri-

butions, however, there is no compilation that

comprehensively analyses the AMF species of

Costa Rica.

Costa Rica, with its great biodiversity and

diverse geography, offers ideal conditions for

a wide diversity of AMF. Its geography, with

mountain ranges that create numerous micro-

climates and a rich variety of ecosystems, con-

tributes to this diversity (Lobo et al., 2021).

The life zone classification system, develo-

ped by Leslie Holdridge (1979), identifies 12

life zones in Costa Rica, which reflects its

ecological diversity.

Studies on the Glomeromycota in Costa

Rica began in 1975 with an ecological survey

developed by Janos (Janos, 1975), this study was

the first of its kind in the country to address

the associations between AMF and various tree

species in tropical forests. Initially, research

focused on the biotechnological potential of

AMF and their impact on plant phenology,

and the first taxonomic records were published

years later (Blanco & Salas, 1997; Fischer et

al., 1994; Sieverding et al., 1988). Most studies

have focused on the impact of AMF on plants

rather than on their taxonomic identification or

geographic distribution (Aldrich-Wolfe, 2007;

Fischer et al., 1994; Picone, 2000; Salas & Blan-

co, 2000; Sharrock et al., 2004; Shepherd et al.,

2007; Sieverding et al., 1988). This analysis is

grounded in scientific literature available in

English and Spanish. The bibliographic search

encompasses articles from the period between

1971 and 2024.

The objective of the present study was to

compile an updated list of AMF of Costa Rica,

organized by vegetation type and life zone.

These new records broaden our knowledge of

the diversity of AMF in Costa Rica and pro-

vide a basis for making decisions about future

studies on the taxonomy, distribution, ecology

and biotechnological applications of the Glo-

meromycota of Costa Rica.

MATERIALS AND METHODS

We conducted a literature search of AMF

recorded in Costa Rica using the following

databases: Google Scholar, Scopus, and Web

of Science. The terms used in the search were:

“arbuscular”, “vesicular”, “mycorrhizal”, “Glo-

meromycota”, and each family and genus of

AMF, which were included in combination

with the words “native” and “Costa Rica” both

in Spanish and English, and we also included

articles published between 1971 and 2024 to

compile a checklist of AMF.

We included articles that employ identi-

fication criteria based on the morphological

and/or molecular characteristics of arbuscular

mycorrhizal fungi up to species level and taxon

descriptions and integrated the information

in a database with the reported species name,

associated plant species, locality where it was

reported, ecological data, molecular data, life

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

zone, soil type, and the phenotypic characte-

ristics of the glomerospores considered in the

determination of the species.

To generate the AMF checklist, we ordered

the AMF species according to the taxono-

mic classification proposed by Błaszkowski

(2012), Oehl et al. (2008) and Oehl et al.

(2011), including additional taxa proposed by

Błaszkowski et al. (2015), Błaszkowski et al.

(2017), Błaszkowski et al. (2018), Corazón-

Guivin et al. (2019), Goto et al. (2012), Jobim et

al. (2016), Marinho et al. (2018) and Symanczik

et al. (2018). We used the life zone classification

by Leslie Holdridge (1979), which considers 12

life zones (Tropical dry forest, tropical moist

forest, tropical wet forest, premontane moist

forest, premontane wet forest, premontane

rainforest, lower montane moist forest, lower

montane wet forest, lower Montane rain forest,

montane wet forest, montane rain forest and

subalpine rain paramo).

In order to illustrate the distribution of

AMF species across the various life zones iden-

tified in Costa Rica, a Sankey diagram was

constructed utilising the RStudio’s ggsankey

package. This diagram, based on presence/

absence data, aims to graphically show the

associations between AMF species and life

zones, indicating where the species have been

recorded. Furthermore, a network diagram was

constructed using the RStudio igraph package,

with the objective of visualising the connections

between life zones and Glomeromycota species.

Rarefaction and extrapolation curves:

We performed rarefaction and extrapolation

curves based on individuals and sampling cove-

rage to estimate the state of knowledge about

the AMF community (species richness) in

different life zones in Costa Rica (Chao & Jost,

2012). Individual-based curves consider the

number of individuals collected in each sam-

ple, while sample coverage-based curves assess

the proportion of the total species community

expected to have been captured as a function

of sampling effort. In order to predict the

expected number of total species and to assess

the required sampling effort, data per life zone

were used considering their sampling cove-

rage (Chao & Jost, 2012; Chao & Lee, 1992).

Rarefaction curves were constructed both for

observed data and to extrapolate the expected

number of species beyond the original sample

size, allowing estimation of expected richness

with increased sampling effort. The correspon-

ding graphs were generated using the RStudio

iNEXT package.

RESULTS

We compiled and analyzed 57 articles that

addressed aspects related to arbuscular myco-

rrhizal fungi (AMF) in Costa Rica published

from 1975 to 2024 (Despite the fact that our

research has been conducted since 1971, the

first documented report was not published

until 1975). It is important to note that not all

of these studies focused on the identification or

geographic distribution of species; only 28 %

included taxonomic records (published from

1988 to 2024), while 38 % examined the eco-

logy (the analysis of the biological, functional

and ecological interactions between arbuscular

mycorrhizal fungi (AMF) and abiotic and biotic

factors present in these ecosystems) of AMF, 14

% investigated their physiological effect on host

plants, and 19 % mentioned the Glomeromyco-

ta of Costa Rica in terms of their potential use

and importance for ecosystems but without

exploring any aspect in detail or providing spe-

cific data (Fig. 1).

Following a screening process, we inclu-

ded a total of 549 records of AMF species in

our database. Of these records, 434 were only

determined at a genus level or designated as

related species and were thus eliminated. As

a result, we kept 115 records of AMF species

with a confirmed identification, which corres-

ponded to a total of 60 species classified into

three classes, five orders, ten families, and 17

genera of Glomeromycota in Costa Rica (Table

1 and Table 2).

The class Archaeosporomycetes is repre-

sented by the order Archaeosporales, the family

Ambisporaceae, the genus Ambispora, and

three species (Table 1 and Table 2).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64710, mayo 2025 (Publicado May. 15, 2025)

The distribution of AMF studies included

only four of the 12 identified life zones (Table

2, Fig. 2 and Fig. 3) (Holdridge, 1979): tropical

moist forest, premontane wet forest, tropical wet

forest and tropical dry forest; as well as agro-

ecosystems and trap crops (consortia) or pure

cultures (monospecific) (Fig. 2 and Fig. 3). Table

two shows detailed data of the hosts (plant spe-

cies) colonized by the 60 AMF species reported

in Costa Rica from 1988 to 2024.

As demonstrated in Table 2, the Glome-

romycota species with the highest number of

reported hosts can be organised as follows (in

decreasing order): Acaulospora scrobiculata (29

hosts), Acaulospora foveata (28 hosts), Glo-

mus clavisporum (26 hosts), Ambispora leptoti-

cha (21 hosts), Acaulospora spinosa (20 hosts),

Acaulospora denticulata (19 hots), Funnelifor-

mis monosporus (17 hosts), Acaulospora ele-

gans con (16 hosts), Glomus macrocarpum (17

hosts), Acaulospora mellea (15 hosts), Glomus

microcarpum (14 hosts), Rhizoglomus clarum

(14 hosts), Acaulospora tuberculata (13 hosts),

Paraglomus occultum (12 hosts), Acaulospo-

ra laevis (12 host), Rhizoglomus aggregatum

Fig. 1. Percentage distribution of studies on arbuscular

mycorrhizal fungi (AMF) in Costa Rica (1975–2024)

categorized by topics: taxonomic classification, ecological

aspects, ecosystems, and physiological effects on plants.

Table 1

Taxonomic classification of the arbuscular mycorrhizal fungi from Costa Rica.

Class (3) Order (5) Family (10) Genus (17) Species (60)

Archaeosporomycetes Archaeosporales Ambisporaceae Ambispora 3

Glomeromycetes Glomerales Glomeraceae Dominikia 1

Funneliformis 3

Glomus 6

Rhizoglomus 7

Sclerocystis 1

Entrophosporaceae Entrophospora 5

Diversisporales Diversisporaceae Diversispora 1

Sieverdingia 1

Acaulosporaceae Acaulospora 18

Kuklospora 1

Gigasporales Gigasporaceae Gigaspora 5

Scutellosporaceae Scutellospora 1

Racocetraceae Cetraspora 1

Racocetra 2

Dentiscutataceae Dentiscutata 3

Paraglomeromycetes Paraglomerales Paraglomeraceae Paraglomus 1

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

Table 2

Checklist of the reported arbuscular mycorrhizal fungi in Costa Rica from 1988 to 2024.

Species Life Zone or adapted/

modified system Associated plant species Reference

number

Archaeosporales

Ambispora appendicula (Spain,

Sieverd., N.C. Schenck)

C. Walker

Greenhouse and premontane

wet forest

Brachiaria decumbens, Cajanus bicolor 2, 4, 16

Ambispora gerdemannii (S.L.

Rose, B.A. Daniels & Trappe)

C. Walker, Vestberg &

A. Schüssler

Tropical wet forest Carapa guianensis, Dalbergia tucurensis, Dipterix

panamensis, Mimosa púdica, Pentaclethra macroloba,

Sinarouba amara, Solanum sp., Vochysia ferruginea

Ambispora leptoticha (N.C.

Schenck & T.H. Nicolson) Walker,

Vestberg &

A. Schüssler

Premontane wet forest,

tropical dry forest and tropical

wet forest.

Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Carapa

guianensis, Casearia aculeata, Cordia alliodora,

Chomelia spinosa, Dalbergia tucurensis, Dipterix

panamensis, Enterolobium cyclocarpum, Licania

arborea, Lonchocarpus spp., Luekea spp., Lysoma

seemannii, Pentaclethra macroloba, Sinarouba amara,

Spondias mombin, Tabebuia spp., Vochysia ferruginea,

Vriesea werkleana

5, 6

Diversisporales

Acaulospora bireticulata

(F.M. Rothwell & Trappe)

Agroecosystem and tropical

wet forest

Carapa guianensis, Cedrela odorata, Dalbergia

tucurensis, Dipterix panamensis, Mimosa pudica,

Pentaclethra macroloba, Sinarouba amara, Solanum sp.,

Vochysia ferruginea

6, 15

Acaulospora colossica (P.A.

Schultz, Bever & J.B. Morton)

Tropical wet forest Inga paterno, Tabebuia ochraceae 12

Acaulospora denticulata (Sieverd.

& S. Toro)

Tropical dry forest Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Chomelia spinosa, Cordia alliodora,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Spondias mombin, Tabebuia spp.

Acaulospora elegans

(Trappe & Gerd)

Tropical dry forest Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Chomelia spinosa, Cordia alliodora,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Spondias mombin, Tabebuia spp.

Acaulospora excavata

Ingleby & C. Walker

Agroecosystem Cedrela odorata 15

Acaulospora foveata

Trappe & Janos

Agroecosystem, greenhouse,

tropical dry forest, tropical

moist forest and tropical wet

forest

Allium cepa L., Astronium graveolens, Bombacopsis

quinata, Bursea simaruba, Calycophyllum

candidissimum, Carapa guianensis, Casearia aculeata,

Cedrela odorata, Ceiba pentandra, Chomelia spinosa,

Cordia alliodora, Dalbergia tucurensis, Dipterix

panamensis, Enterolobium cyclocarpum, Hyeronima

alchorneoides, Jatropha curcas, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Mimosa pudica, Pentaclethra macroloba, Psidium

guajava L., Sinarouba amara, Solanum sp., Spondias

mombin, Tabebuia spp., Vochysia ferruginea

3, 5, 6, 10,

15, 16

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64710, mayo 2025 (Publicado May. 15, 2025)

Species Life Zone or adapted/

modified system Associated plant species Reference

number

Acaulospora laevis

Gerd. & Trappe

Agroecosystem, premontane

wet forest and tropical wet

forest

Cedrela odorata, Coffea sp., Inga paterno, Vriesea

werkleana

14, 15

Acaulospora longula (Spain & N.C.

Schenck)

Premontane wet forest Not specified 2

Acaulospora mellea Spain & N.C.

Schenck

Agroecosystem, greenhouse,

tropical moist forest and

tropical wet forest

Carapa guianensis, Cedrela odorata, Ceiba pentandra,

Cordia alliodora, Dalbergia tucurensis, Dipterix

panamensis, Hyeronima alchorneoides, Jatropha curcas,

Mimosa pudica, Pentaclethra macroloba, Sinarouba

amara, Solanum sp., Vochysia ferruginea

6, 8, 10,

15, 16

Acaulospora minuta Oehl, Tchabi,

Hount., Palenz., I.C. Sánchez &

G.A. Silva

Agroecosystem Coffea sp.

Acaulospora morrowiae Spain &

N.C. Schenck,

Tropical moist forest Cedrela odorata, Ceiba pentandra

Cordia alliodora, Hyeronima alchorneoides, Zea mays

6, 8, 10

Acaulospora rehmii Sieverd. &

S. Toro

Agroecosystem and

greenhouse

Cedrela odorata, Jatropha curcas 5, 15, 16

Acaulospora rugosa J.B. Morton Greenhouse and premontane

wet forest

Jatropha curcas, Vriesea werkleana 2, 16

Acaulospora scrobiculata Trappe Agroecosystem, greenhouse,

premontane wet forest,

tropical dry forest, tropical

moist forest and tropical wet

forest

Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Carapa

guianensis, Casearia aculeata, Cedrela odorata, Ceiba

pentandra, Chomelia spinosa, Coffea sp., Cordia

alliodora, Dalbergia tucurensis, Dipterix panamensis,

Enterolobium cyclocarpum, Hyeronima alchorneoides,

Jatropha curcas, Licania arborea, Lonchocarpus spp.,

Luekea spp., Lysoma seemannii, Mimosa pudica,

Pentaclethra macroloba, Sinarouba amara, Solanum

sp., Spondias mombin, Tabebuia spp., Termilania

amazonia, Vochysia ferruginea, Vriesea werkleana

2, 5, 6, 8,

10, 13, 14,

15, 16

Acaulospora spinosa C. Walker

& Trappe

Agroecosystem, greenhouse,

premontane wet forest,

tropical dry forest and tropical

moist forest

Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Cedrela odorata, Ceiba pentandra, Chomelia

spinosa, Cordia alliodora, Enterolobium cyclocarpum,

Hyeronima alchorneoides, Jatropha curcas, Licania

arborea, Lonchocarpus spp., Luekea spp., Lysoma

seemannii, Spondias mombin, Tabebuia spp., Vriesea

werkleana

5, 6, 7, 8,

15, 16

Acaulospora spinosissima Oehl,

Palenz., Sánchez-Castro, Tchabi,

Hount. & G. A. Silva

Agroecosystem Cedrela odorata 15

Acaulospora splendida Sieverd.,

Chaverri & I. Rojas

Greenhouse Quercus costaricensis 1

Acaulospora tuberculata Janos

& Trappe

Agroecosystem, greenhouse

and tropical wet forest,

Carapa guianensis, Cedrela odorata, Dalbergia

tucurensis, Dipterix panamensis, Jatropha curcas,

Mimosa pudica, Pentaclethra macroloba, Sinarouba

amara, Solanum sp., Vochysia ferruginea

6, 15, 16

Kuklospora colombiana (Spain &

N.C. Schenck) Oehl & Sieverd

Agroecosystem Cedrela odorata 15

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

Species Life Zone or adapted/

modified system Associated plant species Reference

number

Sieverdingia tortuosa (N.C.

Schenck & G.S. Sm.) Błaszk.,

Niezgoda & B.T. Goto.

Tropical moist forest Cedrela odorata, Ceiba pentandra, Cordia alliodora,

Hyeronima alchorneoides

8, 10

Diversispora versiformis (P. Karst.)

Oehl, G.A. Silva & Sieverd

Greenhouse and premontane

wet forest

Tithonia diversifolia, Vriesea werkleana 11

Entrophosporales

Entrophospora claroidea (N.C.

Schenck & G.S. Sm.) Błaszk.,

Niezgoda, B.T. Goto & Magurno

Greenhouse, ropical moist

forest and premontane wet

forest

Brachiaria decumbens, Cajanus bicolor, Ceiba

pentandra, Vriesea werkleana

Entrophospora colombiana Spain

& N.C. Schenck

Premontane wet forest,

tropical wet forest

Carapa guianensis, Dalbergia tucurensis, Dipterix

panamensis, Mimosa pudica, Pentaclethra macroloba,

Sinarouba amara, Solanum sp., Vochysia ferruginea,

Vriesea werkleana

2, 6

Entrophospora etunicata

(W.N. Becker & Gerd.) Błaszk.,

Niezgoda, B.T. Goto & Magurno

Agroecosystem, premontane

wet forest and tropical moist

forest

Cedrela odorata, Coffea sp., Cordia alliodora,

Hyeronima alchorneoides, Vriesea werkleana, Zea mays

9, 14

Entrophospora infrequens (I.R.

Hall) R.N. Ames & R.W. Schneid.

Greenhouse Brachiaria decumbens, Cajanus bicolor 4

Entrophospora lutea (L.J. Kennedy,

J.C. Stutz & J.B. Morton) Błaszk.,

Niezgoda, B.T. Goto & Magurno

Tropical moist forest Termilania amazonia 13

Glomerales

Funneliformis geosporum (T.H.

Nicolson & Gerd.) C. Walker & A.

Schüssler

Agroecosystem, premontane

wet forest and tropical wet

forest

Carapa guianensis, Cedrela odorata, Dalbergia

tucurensis, Dipterix panamensis, Mimosa pudica,

Pentaclethra macroloba, Sinarouba amara, Solanum sp.,

Vochysia ferruginea, Vriesea werkleana

2, 15

Funneliformis monosporus (Gerd.

& Trappe) Oehl, G.A. Silva &

Sieverd

Tropical dry forest Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Chomelia spinosa, Cordia alliodora,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Spondias mombin, Tabebuia spp.

Funneliformis mosseae (T.H.

Nicolson & Gerd.) C. Walker

& A. Schüssler

Agroecosystem, premontane

wet forest

Cedrela odorata, Vriesea werkleana 2, 15

Glomus brohultii (Sieverd. &

Herrera)

Premontane wet forest Not specified 2

Glomus clavisporum (Trappe)

R.T. Almeida & N.C. Schenck

Agroecosystem, tropical dry

forest, tropical moist forest

and tropical wet forest

Allium cepa L., Astronium graveolens, Bombacopsis

quinata, Bursea simaruba, Calycophyllum

candidissimum, Carapa guianensis, Casearia

aculeata, Cedrela odorata, Chomelia spinosa, Cordia

alliodora, Dalbergia tucurensis, Dipterix panamensis,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Pentaclethra macroloba, Psidium guajava L., Sinarouba

amara, Spondias mombin, Tabebuia spp., Vochysia

ferruginea

3, 5, 6, 15

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Species Life Zone or adapted/

modified system Associated plant species Reference

number

Glomus microcarpum

(Tul. & C. Tul.)

Tropical dry forest Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Chomelia spinosa, Cordia alliodora,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Spondias mombin, Tabebuia spp.

Glomus macrocarpum

(Tul. & C. Tul.)

Tropical dry forest and ropical

moist forest

Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Chomelia spinosa, Cordia alliodora,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Spondias mombin, Tabebuia spp., Termilania amazonia

5,13

Glomus rubiforme (Gerd. &

Trappe) R.T. Almeida & N.C.

Schenck

Tropical wet forest Carapa guianensis, Dalbergia tucurensis, Dipterix

panamensis, Pentaclethra macroloba, Sinarouba amara,

Vochysia ferruginea

Glomus taiwanense (C.G. Wu &

Z.C. Chen) R.T. Almeida & N.C.

Schenck ex Y.J. Yao

Greenhouse Jatropha curcas 16

Dominikia indica (Błaszk., Wubet

& Harikumar) Błaszk., G.A. Silva

& Oehl

Agroecosystem Coffea sp. 14

Rhizoglomus aggregatum (N.C.

Schenck & G.S. Sm.) Sieverd.,

G.A. Silva & Oehl

Tropical dry forest Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Chomelia spinosa, Cordia alliodora,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Spondias mombin, Tabebuia spp.

Rhizoglomus clarum (T.H.

Nicolson & N.C. Schenck)

Sieverd., G.A. Silva & Oehl

Agroecosystem, greenhouse,

premontane wet forest,

tropical moist forest and

tropical wet forest

Carapa guianensis, Cedrela odorata, Dalbergia

tucurensis, Dipterix panamensis, Hyeronima

alchorneoides, Inga paterno, Jatropha curcas, Mimosa

pudica, Pentaclethra macroloba, Sinarouba amara,

Solanum sp., Vochysia ferruginea, Vriesea werkleana

6, 10, 12,

15, 16

Rhizoglomus fasciculatum (Thaxt.)

Sieverd., G.A. Silva & Oehl

Agroecosystem, premontane

wet forest

Coffea sp., Vriesea werkleana 14

Rhizoglomus intraradices (N.C.

Schenck & G.S. Sm.) Sieverd.,

G.A. Silva & Oehl

Greenhouse, premontane wet

forest, tropical moist forest

and tropical wet forest

Tabebuia ochraceae, Termilania amazonia, Tithonia

diversifolia, Vriesea werkleana, Zea mays

9, 11, 12, 13

Rhizoglomus microaggregatum

(Koske, Gemma & P.D. Olexia)

Sieverd., G.A. Silva & Oehl

Greenhouse Brachiaria decumbens, Cajanus bicolor 4

Rhizoglomus proliferum (Dalpé

& Declerck) Sieverd., G.A. Silva

& Oehl

Tropical wet forest Vriesea werkleana 12

Rhizoglomus vesiculiferum

(Thaxt.) Błaszk., Kozłowska,

Niezgoda, B.T.Goto & Dalpé

Premontane wet forest Vriesea werkleana 12

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

Species Life Zone or adapted/

modified system Associated plant species Reference

number

Sclerocystis coremioides Berk. &

Broome

Agroecosystem, greenhouse,

premontane wet forest,

tropical moist forest and

tropical wet forest

Allium cepa L., Carapa guianensis

Cedrela odorata, Dalbergia tucurensis, Dipterix

panamensis, Jatropha curcas, Mimosa pudica,

Pentaclethra macroloba, Psidium guajava L.,

Sinarouba amara, Solanum sp., Vochysia ferruginea,

Vriesea werkleana

3, 6, 15, 16

Gigasporales

Dentiscutata heterogama (T.H.

Nicolson & Gerd.) Sieverd., F.A.

de Souza & Oehl

Greenhouse Brachiaria decumbens, Cajanus bicolor 4

Dentiscutata scutata (C. Walker

& Dieder.) Sieverd., F.A. de Souza

& Oehl

Tropical moist forest Cordia alliodora, Hyeronima alchorneoides 10

Dentiscutata nigra (J.F. Readhead)

Sieverd., F.A. de Souza & Oehl

Tropical dry forest Astronium graveolens, Bombacopsis quinata, Bursea

simaruba, Calycophyllum candidissimum, Casearia

aculeata, Chomelia spinosa, Cordia alliodora,

Enterolobium cyclocarpum, Licania arborea,

Lonchocarpus spp., Luekea spp., Lysoma seemannii,

Spondias mombin, Tabebuia spp.

Gigaspora albida N.C. Schenck &

G.S. Sm

Greenhouse and remontane

wet forest

Brachiaria decumbens, Cajanus bicolor,

Vriesea werkleana

4, 6

Gigaspora gigantea (T.H.

Nicholson & Gerd.) Gerd. &

Trappe

Premontane wet forest and

tropical wet forest

Carapa guianensis, Dalbergia tucurensis, Dipterix

panamensis, Pentaclethra macroloba, Sinarouba amara,

Vochysia ferruginea, Vriesea werkleana

Gigaspora margarita W.N. Becker

& I.R. Hall

Agroecosystem and

greenhouse

Coffea sp., Jatropha curcas 14, 16

Gigaspora ramisporophora Spain,

Sieverd. & N.C. Schenck

Greenhouse Brachiaria decumbens, Cajanus bicolor 4

Gigaspora rosea T.H. Nicolson &

N.C. Schenck

Tropical moist forest Zea mays 9

Cetraspora pellucida (T.H.

Nicolson & N.C. Schenck) Oehl,

F.A. de Souza & Sieverd

Greenhouse, premontane wet

forest, tropical moist forest

and tropical wet forest

Allium porrum, Cajanus cajans, Capsicum annum,

Carapa guianensis, Cedrela odorata, Cordia alliodora,

Cucumis sativus, Dalbergia tucurensis, Dipterix

panamensis, Hyeronima alchorneoides, Pentaclethra

macroloba, Sinarouba amara, Vochysia ferruginea,

Vriesea werkleana, Zea mays

6, 7, 10

Racocetra castanea (C. Walker)

Oehl, F.A. de Souza & Sieverd

Premontane wet forest and

tropical moist forest

Cedrela odorata, Hyeronima alchorneoides,

Vriesea werkleana

Racocetra coralloidea (Trappe,

Gerd. & I. Ho) Oehl, F.A. de

Souza & Sieverd

Tropical wet forest Carapa guianensis, Dalbergia tucurensis, Dipterix

panamensis, Pentaclethra macroloba, Sinarouba amara,

Vochysia ferruginea

Scutellospora calospora (T.H.

Nicolson & Gerd.) C. Walker &

F.E. Sanders

Greenhouse Brachiaria decumbens, Cajanus bicolor 4

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Species Life Zone or adapted/

modified system Associated plant species Reference

number

Paraglomerales

Paraglomus occultum (C. Walker)

J.B. Morton & D. Redecker

Agroecosystem and tropical

moist forest, premontane wet

forest and tropical wet forest

Carapa guianensis, Cedrela odorata, Cordia alliodora,

Dalbergia tucurensis, Dipterix panamensis, Hyeronima

alchorneoides, Mimosa pudica Pentaclethra macroloba,

Sinarouba amara, Solanum sp., Vochysia ferruginea,

Vriesea werkleana

6, 10, 15

Citations arranged in ascending chronological order; Sieverding et al. (1988)1, Vargas (1990)2, Fischer et al. (1994)3, Blanco

& Salas (1997)4, Johnson & Wedin (1997)5, Picone (2000)6, Salas & Blanco (2000)7, Lovelock et al. (2003)8, Lovelock et al.

(2004)9, Lovelock et al. (2005)10, Sharrock et al. (2004)11, Shepherd et al. (2007)12, Aldrich-Wolfe (2007)13, Aldrich-Wolfe et

al. (2020)14, Polo-Marcial et al. (2023)15, Solís-Ramos et al. (2023)16.

(12 hosts), Entrophospora colombiana (12

hosts), Gigaspora gigantea (11 hosts), Acau-

lospora minuta (11 hosts), Glomus rubiforme

(11 hosts), Cetraspora pellucida (15 hosts),

Funneliformis mosseae (10 hosts), Acaulospora

excavata (10 hosts), Rhizoglomus intraradices

(10 hosts), Sieverdingia tortuosa (9 hosts), Scle-

rocystis coremioides (14 hosts), Acaulospora

rehmii (9 hosts), Acaulospora splendida con 8

and Ambispora gerdemannii (9 hosts) (Table 2).

In terms of their distribution by life zones,

of the 60 species identified in Costa Rica (Table

1 and Table 2; Fig. 2 and Fig. 3), 24 species (40

%) were recorded in the premontane wet forest,

with three species unique to this life zone; 22

species (36 %) were reported in agroecosys-

tems, with five unique species; 23 species (38

%) were reported in greenhouses (trap crops),

with three unique species; 19 species (31 %)

were reported in the tropical moist forest, with

six unique species; 21 species (35 %) in the tro-

pical wet forest, with seven unique species and

12 species (20 %) in the tropical dry forest, with

five unique species (Fig. 3).

With respect to family composition in

the different life zones, the families Acau-

losporaceae and Glomeraceae were the most

predominant, exhibiting a wide distribution

and diversity in these ecosystems (Table 1).

The families Ambisporaceae, Entrophospho-

raceae, Racocetraceae, Dentiscutataceae, and

Paraglomeraceae showed lower richness and

abundance, and were even absent in 8 life zones

(Table 3), which may indicate specific environ-

mental conditions that limit their presence or a

lack of sufficient studies in those areas.

Table 3

Diversity and Abundance of the reported Glomeromycota in Different Life Zones and Agroecosystems of Costa Rica from

1988 to 2024.

Life zone Families Genera Species Total Abundance

Tropical dry forest 3 4 5 98

Tropical moist forest 8 11 19 55

Tropical wet forest 7 12 21 130

Premontane moist forest 0 0 0 0

Premontane wet forest 11 14 24 38

Premontane rain forest 0 0 0 0

Lower montane moist forest 0 0 0 0

Lower montane wet forest 0 0 0 0

Lower Montane rain forest 0 0 0 0

Montane wet forest 0 0 0 0

Montane rain forest 0 0 0 0

Subalpine rain paramo 0 0 0 0

Agroecosystem 6 10 22 27

Greenhouse 8 10 23 39

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

Fig. 2. Sankey diagram depicting the families of arbuscular mycorrhizal fungi identified in different life zones and production systems. The width of the bands indicates the relative

proportion of each AMF genus associated with the life zones. The species of each genus are detailed on the right.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64710, mayo 2025 (Publicado May. 15, 2025)

Fig. 3. Diagram of connection networks between life zones and reported Glomeromycota species in Costa Rica from 1988 to 2024. Life zones include Agroecosystem, Tropical Wet

Forest, Tropical Dry Forest, Premontane Wet Forest, Tropical Moist Forest and Greenhouse.

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

The analysis of the rarefaction and extra-

polation curves showed differences in the spe-

cies diversity captured according to the number

of individuals sampled in the different life

zones (Fig. 4). In the agroecosystems and pre-

montane wet forest, it is clear that diversity

showed a gradual increase without reaching an

asymptote. This indicates that more sampling

effort is required to capture the full diversity

present. The broad confidence bands indicate

uncertainty in the extrapolations (Fig. 4).

In contrast, in the tropical moist forest,

tropical wet forest, and tropical dry forest, the

curves reach a plateau, it is clear that most spe-

cies were recorded with the current sampling

(Fig. 4). The thinner confidence bands in these

cases indicate more precise estimates (Fig. 4). It

is important to mention that this analysis did

not consider trap or pure crops (greenhouses),

since they were grouped according to the life

zones where the samples were extracted from

the soil for the experiments. This analysis

Fig. 4. Individual-based rarefaction and extrapolation curves of arbuscular mycorrhizal fungi found in four different life

zones and in agroecosystems in Costa Rica from 1988-2024. a) Agroecosystem, b) Tropical moist forest, c) Premontane wet

forest, d) Tropical wet forest, e) Tropical dry forest. The dashed line represents the extrapolation curve, indicating the expected

number of species that could be found by increasing the sampling effort in the different life zones of our reference samples.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64710, mayo 2025 (Publicado May. 15, 2025)

demonstrates that AMF species diversity varies

significantly between life zones and modified

systems. In certain regions, such as agroecosys-

tems and pre-montane moist forests, the full

diversity has not yet been documented due to

inadequate sampling efforts. Conversely, in

other regions, such as tropical dry forests, the

sampling appears to be adequate to reflect the

majority of species present. Furthermore, the

confidence bands indicate divergent levels of

precision in the estimates, contingent on the

quality and quantity of the available data.

DISCUSSION

Since the compilation made by Blanco and

Salas in 1997, where 19 species were recorded,

the present checklist shows an increase of

31 % in the recorded taxa. This illustrates Costa

Rica’s biological potential in terms of biodi-

versity, since the species identified represent

16 % of the Glomeromycota species worldwi-

de (Goto & Jobim, 2024). Of the 15 Glome-

romycota families, ten are represented in the

present checklist, with Glomeraceae being the

most abundant, followed by Acaulosporaceae,

which is consistent with the records from other,

better studied megadiverse countries such as

Mexico and Brazil (Jobim & Goto, 2016; Polo-

Marcial et al., 2023; Stürmer & Kemmelmeier,

2021). The dominant genera is Acaulospora,

which has a high representativeness in Neo-

tropical ecosystems (Stürmer & Kemmelmeier,

2021), we found that in the four life zones and

vegetation types studied of Costa Rica, the

AMF community is dominated by Acaulospora

spp. (Johnson & Wedin 1997; Lovelock & Ewel

2005; Picone, 2000). Some reported studies on

AMF have focused on determining the physio-

logical response of different hosts with AMF

cultivated strains (Blanco & Salas, 1997; Her-

nández & Salas, 2009; Salas & Blanco, 2000).

Studies identifying and evaluating native AMF

communities are scarce, for example Polo-

Marcial et al. (2023) and Solís-Ramos et al.

(2023), studied the effect of AMF communities

in Cedrela odorata and Jatropha curcas respecti-

vely. However, a high morphological richness of

AMF has been found in tropical forests and dry

grasslands (Johnson & Wedin, 1997; Lovelock

& Ewel, 2005; Picone, 2000).

Among the life zones evaluated, the pre-

montane moist forest is the best studied, since

24 of the 60 species recorded in Costa Rica are

found in this life zone. This number of species

highlights the unique vegetation types of the

region and contrasts with the previous limited

taxonomic inventories in these areas (Aldrich-

Wolfe, 2007; Aldrich-Wolfe et al., 2020; Fischer

et al., 1994; Johnson & Wedin, 1997; Lovelock

et al., 2003, Lovelock et al., 2004; Lovelock et al.,

2005; Picone, 2000; Polo-Marcial et al., 2023;

Salas & Blanco, 2000; Sharrock et al., 2004;

Shepherd et al., 2007; Sieverding et al., 1988;

Solís-Ramos et al., 2023).

The evaluation of different areas provides

relevant information for understanding the

diversity of AMF in Costa Rica. Among its

life zones, the premontane moist forest, lower

montane rain forest, premontane rain forest,

lower montane moist forest, and montane wet

forest represent underexplored regions, there is

no record of any Glomeromycota species. These

underexplored or under-inventoried areas may

host new AMF species whose value for the local

flora remains unknown.

The rarefaction curves show the limited

knowledge of species richness in the few (four)

life zones where studies have been conduc-

ted. Given that Costa Rica is recognized by

its biodiversity (Acuña, 2002), studies focu-

sed on bioprospecting for AMF diversity in

understudied life zones will help to broaden

the knowledge of this group, which has sig-

nificant potential for applications in horticul-

ture, production of ornamentals, sustainable

agriculture and forest management, and the

restoration of degraded areas. The knowledge

generated will support the development of

public policies for the successful conservation

of biodiversity and the sustainable production

in Costa Rica. For example, some species of the

genera Acaulospora, Glomus, and one Ambis-

pora are dominant in terms of the number of

hosts and vegetation types on which they have

been reported. This information makes them

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64710, mayo 2025 (Publicado May. 15, 2025)

the obvious candidates for consideration in

mycorrhizal inoculum production. However,

further ecological and taxonomic research is

required, encompassing biotechnological tech-

niques, morphological and molecular analyses,

to validate the applications of the diversity of

AMF in Costa Rica

CONCLUSIONS

We compiled data on 60 AMF species

found in Costa Rica from 1988 to 2024, which

represents 16 % of the Glomeromycota known

in the world. Twenty percent of the species

reported in the studies were classified only at

the genus level and the reports include only 4 of

the 12 life zones of the country, which suggests

a high possibility of identifying new species.

If we consider the diversity of edaphoclimatic

conditions, the variability in the terrain, and

the different types of natural vegetation that

prevail in Costa Rica, which remain to be fur-

ther explored, we can expect the record of AMF

species to expand as further research is con-

ducted. It is fundamental to perform studies on

the interactions with different hosts, as well as

to make taxonomic inventories of AMF in the

different life zones and biogeographic districts,

combining morphological analyses and mole-

cular characterizations, since this information

allows a robust confirmation of taxonomic

identity, the establishment of phylogenetic rela-

tionships, the determination of AMF species

distribution, and complete characterizations

that aid in species recognition and tracking in

situ, in order to gain a more comprehensive

view of the richness and ecology of the Glo-

meromycota of Costa Rica. The data presented

here significantly contribute to the knowledge

of the diversity and distribution of this group in

the natural life zones of Costa Rica and to the

understanding of the diversity and distribution

of AMF worldwide, and also provide a basis for

making decisions about future studies on the

taxonomy, distribution, ecology, and biotech-

nological applications of the Glomeromycota

of Costa Rica.

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

followed all pertinent ethical and legal proce-

dures and requirements. All financial sources

are fully and clearly stated in the acknowled-

gments section. A signed document has been

filed in the journal archives.

ACKNOWLEDGEMENTS

MMJA thanks the Consejo Nacional de

Ciencia y Tecnología de México (CONAH-

CyT) for the MSc scholarship (1288567) at

INBIOTECA, UV, and LYSR thanks to the

Vice-rectory of Research, at the University of

Costa Rica, for funding our project through the

research projects C4764 and C4058. We would

like to express our gratitude to the two anony-

mous reviewers who provided comments that

have contributed to the enhancement of this

manuscript.

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