Ecological Characterization of the Flora in Reserva Ecológica Arenillas, Ecuador

Abstract

Ecuador, located in the Neotropics, has 66 protected natural areas, which represent about 13.77% of its overall territory. The Reserva Ecológica Arenillas reserve (REAr), located in southwestern Ecuador, protects an area of dry forest, coastal thorn forest, and mangroves. This dry forest is part of the Pacific equatorial core and is included the Tumbes–Chocó–Magdalena, one of the 34 biodiversity hot spots of the world. It is an extremely fragile ecosystem and therefore the need for conservation is of the utmost importance. Knowledge of the flora and their ecological characteristics is still limited, which was one of the main objectives of this work. In this study, 118 plots located in different locations of the REAr were selected in order to sample the trees, shrubs, and herbaceous plants within them. This information was supplemented with data from the literature and the GBIF; life forms were included according to Raunkiaer’s classification and their growth habits. The flora of the REAr was represented by 381 species, belonging to 77 families. The two most numerous families were the Fabaceae (51 plant species) and Malvaceae (31 species). The dominant life form was the phanerophytes with 200 species (52.5%), followed by therophytes with 104 species (27.3%), and camephytes with 22 species (5.8%). Physiognomy was dominated by the herbaceous growth (44%). The biodiversity indices of two ecosystems were studied (The deciduous forest of the Jama-Zapotillo lowland and the low forest and deciduous shrubland of the Jama-Zapotillo lowland), obtaining higher values for the deciduous forest ecosystem of the Jama-Zapotillo lowland. With these indicators, a classification of each forest type was made by performing a hierarchical cluster analysis. The information provided in this paper is particularly important for focusing conservation efforts and preventing the loss of flora diversity in these forests, which are subject to great anthropogenic pressures.

Full text

Citation: Luna-Florin, A.D.; Nole-Nole,
D.A.; Rodríguez-Caballero, E.;
Molina-Pardo, J.L.; Giménez-Luque, E.
Ecological Characterization of the
Flora in Reserva Ecológica Arenillas,
Ecuador. Appl. Sci. 2022,12, 8656.
https://doi.org/10.3390/app12178656
Academic Editors: Rainer
W Bussmann and Inayat Ur Rahman
Received: 9 July 2022
Accepted: 22 August 2022
Published: 29 August 2022
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applied
sciences
Article
Ecological Characterization of the Flora in Reserva Ecológica
Arenillas, Ecuador
Alex Dumany Luna-Florin 1, Darío Alexander Nole-Nole 2, Emilio Rodríguez-Caballero 3,
JoséLuis Molina-Pardo 3and Esther Giménez-Luque 3, *
1Carrera de Ingeniería Ambiental, Grupo de Investigación INCON, Universidad Técnica de Machala,
Machala 070102, Ecuador
2Facultad de Ciencias Exactas y Naturales, Departamento de Ciencias Biológicas y Agropecuarias,
Universidad Técnica Partícular de Loja, Loja 110107, Ecuador
3Centro de Investigación de Colecciones Científicas de la Universidad de Almería, Universidad de Almería,
04120 Almeria, Spain
*Correspondence: egimenez@ual.es
Abstract:
Ecuador, located in the Neotropics, has 66 protected natural areas, which represent about
13.77% of its overall territory. The Reserva Ecológica Arenillas reserve (REAr), located in southwestern
Ecuador, protects an area of dry forest, coastal thorn forest, and mangroves. This dry forest is part of
the Pacific equatorial core and is included the Tumbes–Chocó–Magdalena, one of the 34 biodiversity
hot spots of the world. It is an extremely fragile ecosystem and therefore the need for conservation is
of the utmost importance. Knowledge of the flora and their ecological characteristics is still limited,
which was one of the main objectives of this work. In this study, 118 plots located in different loca-
tions of the REAr were selected in order to sample the trees, shrubs, and herbaceous plants within
them. This information was supplemented with data from the literature and the GBIF; life forms were
included according to Raunkiaer’s classification and their growth habits. The flora of the REAr was
represented by 381 species, belonging to 77 families. The two most numerous families were the Fabaceae
(51 plant species) and Malvaceae (31 species). The dominant life form was the phanerophytes with
200 species (52.5%), followed by therophytes with 104 species (27.3%), and camephytes with 22 species
(5.8%). Physiognomy was dominated by the herbaceous growth (44%). The biodiversity indices of
two ecosystems were studied (The deciduous forest of the Jama-Zapotillo lowland and the low forest
and deciduous shrubland of the Jama-Zapotillo lowland), obtaining higher values for the deciduous
forest ecosystem of the Jama-Zapotillo lowland. With these indicators, a classification of each forest
type was made by performing a hierarchical cluster analysis. The information provided in this paper
is particularly important for focusing conservation efforts and preventing the loss of flora diversity in
these forests, which are subject to great anthropogenic pressures.
Keywords: biodiversity; floristic composition; tropical dry forest; species diversity
1. Introduction
Ecuador is one of the 17 mega-diverse countries of the world [
1
]. It has 17,748 con-
firmed species of native flora, and it is estimated that with the continuation of studies of
Ecuadorian flora, the total number of vascular plants could reach 25,000 species [
2
]. A large
proportion of these species (1600) are included in the IUCN Red List. Furthermore, more
species are added to the Red List every year. In 2019, a total of 354 new species were
included in the list.
Dry ecosystems are one of the most valuable types of ecosystems in Ecuador. Together
with dry ecosystems of northern Peru they form the Tumbesian Region. This is one of the
areas of South America with the greatest number of endemic species, while at the same
time being one of the most threatened regions [
3
]. Among the different dry ecosystems in
the Tumbesian region in Ecuador, the equatorial dry forest is among the most fragile [
4
].
Appl. Sci. 2022,12, 8656. https://doi.org/10.3390/app12178656 https://www.mdpi.com/journal/applsci

Appl. Sci. 2022,12, 8656 2 of 17
This is a unique ecosystem of the world and is located in the south of the country, in the
regions of Esmeraldas, Manabí, Santa Elena, Guayas, El Oro, and Loja.
One of the main remnants of the equatorial dry forest in the south of Ecuador is the
Reserva Ecológica Arenillas reserve (henceforth REAr). Moreover, it is the only ecological
reserve that conserves mangroves and tropical dry forests in the southwestern region of the
country. In spite of its ecological relevance, the REAr presents conservation problems, such
as the fragmentation of its ecosystems, the expansion of agriculture and cattle ranching
inside the limits of the reserve, and several climate change-related impacts [
5
]. Another
difficulty encountered by the REAr is the redefinition of its limits. It has been a military
base and an exclusion zone since the 1970s because of its strategic location on the border
with Peru [
6
]. Originally, the former military base had an area of 22,000 ha. When the
Ecological Reserve was created in 2001, the protected area comprised only 17,082.7 ha of
the original extent and in 2012, it was reduced to 13,170.03 ha. [7].
Since the creation of the natural reserve, the REAr has been studied very little from
a floristic point of view. One of the first studies was that of Ceron et al. [
8
], in which
105 species belonging to 49 families were identified. These results were similar to those
found by Estrella and Troya [
9
] (104 species grouped into 82 genera and 48 families)
and Ochoa et al. [
10
] (79 species grouped into 69 genera and 41 families). Nevertheless,
other studies that were based on higher sampling efforts have documented highly diverse
flora [
11
], indicating that increasing the sampling effort may reveal much larger numbers.
Based on these studies and other studies in different dry forest regions, it is also known
that there are evident differences in the composition and biodiversity between the different
zones of the REAr [
12
], but the factors that delimit this special differentiation in the dry
forest have not yet been explored in great detail. Thus, it becomes difficult to study and
obtain rigorous conclusions about the current lists of scientific data on the species that exist
in this ecosystem.
Overall, the REAr, due to its history, diversity of habitats, geographic location, and so-
ciological environment, is an interesting biodiversity scenario for the study of conservation,
monitoring of plant species, and the improvement of our knowledge of the main functions
of this ecosystem that is threatened by anthropic pressure. However, not much information
exists about its floral composition, diversity, and the underlying drivers. Therefore, the
objective of this work was to analyze the floristic composition, ecological characteristics,
and diversity of the ecosystems within the REAr. By doing so, we updated the information
on the flora of the REAr, increasing its value for conservation initiatives and contributing
to a proper design of future management and conservation actions.
2. Material and Methods
2.1. Study Area
This study is conducted in the REAr, which is located to the south of the equator
between the cantons of Huaquillas and Arenillas (Figure 1), within the latitudes from
3
◦
27
0
30.94” S to 3
◦
39
0
37.49” S and from 80
◦
9
0
18.65” W to 80
◦
9
0
47.93” W. This reserve has
an area of 13,170 ha and an average elevation of 120 m above sea level (masl) [
8
]. The REAr
is composed of a matrix of dry forests, desert scrubs, and mangroves. The climate of the
study area is warm-dry, with an average temperature of 24
◦
C, and rainfall varies between
300 and 500 mm/year from the lower elevated northern area of the reserve to the higher
elevated southern part.

Appl. Sci. 2022,12, 8656 3 of 17
Appl. Sci. 2022, 12, 8656 3 of 18
Figure 1. Map showing the location of the REAr, the ecosystems represented, and the selected sam-
pling points.
2.2. Characterization of the Richness, Composition, and Diversity of Flora Species in the REAr
According to the ecosystem classification system of Ecuador [13], we can find four
types of ecosystems in the study area: (1) The deciduous forest of the Jama-Zapotillo low-
land, (2) the low forest and deciduous shrubland of the Jama-Zapotillo lowland, (3) the
mangrove of the Jama-Zapotillo, and (4) the low desert scrubs of the Jama-Zapotillo. In
this study, we sampled 118 plots of 20 × 20 m surface areas in two of the different dry
forest types (the low deciduous forest of the Jama-Zapotillo and low forest and deciduous
shrubland of the Jama-Zapotillo) and in the mangrove (the mangrove of the Jama-Zapo-
tillo) (Figure 1). Sampling locations were selected following a random design, but loca-
tions with difficult or impossible access were discarded. In each 20 × 20 m plot, four 5 × 5
m subplots were selected to study shrubs, and four 1 × 1 m subplots to study herbs. Life
form classes were identified according to Raunkiaer’s classification [14]. Field sampling
was carried out during the rainy season to achieve a better identification of the species,
according to the flowering and fruiting times of most of the known species. The physiog-
nomies of the plants (herbaceous, shrubs, and trees) were noted in the field during sam-
pling. Each of the plots were georeferenced with a handheld GPS (GARMIN GPSMAP 65)
with a 5 m horizontal accuracy. The list of flora of the REAr created by Molina Moreira in
2017 [11] was used as the basis for the study. This is the latest and most complete study
conducted so far in the REAr, and included 291 species from 64 families. The specimens
from this area that have already been deposited in the Reinaldo Espinoza Herbarium of
Figure 1.
Map showing the location of the REAr, the ecosystems represented, and the selected
sampling points.
2.2. Characterization of the Richness, Composition, and Diversity of Flora Species in the REAr
According to the ecosystem classification system of Ecuador [
13
], we can find four types
of ecosystems in the study area: (1) The deciduous forest of the Jama-Zapotillo lowland, (2)
the low forest and deciduous shrubland of the Jama-Zapotillo lowland, (3) the mangrove
of the Jama-Zapotillo, and (4) the low desert scrubs of the Jama-Zapotillo. In this study,
we sampled 118 plots of 20
×
20 m surface areas in two of the different dry forest types
(the low deciduous forest of the Jama-Zapotillo and low forest and deciduous shrubland of
the Jama-Zapotillo) and in the mangrove (the mangrove of the Jama-Zapotillo) (Figure 1).
Sampling locations were selected following a random design, but locations with difficult or
impossible access were discarded. In each 20
×
20 m plot, four 5
×
5 m subplots were selected
to study shrubs, and four 1
×
1 m subplots to study herbs. Life form classes were identified
according to Raunkiaer’s classification [
14
]. Field sampling was carried out during the rainy
season to achieve a better identification of the species, according to the flowering and fruiting
times of most of the known species. The physiognomies of the plants (herbaceous, shrubs,
and trees) were noted in the field during sampling. Each of the plots were georeferenced
with a handheld GPS (GARMIN GPSMAP 65) with a 5 m horizontal accuracy. The list
of flora of the REAr created by Molina Moreira in 2017 [
11
] was used as the basis for the
study. This is the latest and most complete study conducted so far in the REAr, and included
291 species from 64 families. The specimens from this area that have already been deposited in
the Reinaldo Espinoza Herbarium of the Universidad Nacional de Loja (LOJA), the Herbarium
of the Universidad Técnica Particular de Loja (HUTPL), the Herbarium of the Universidad de

Appl. Sci. 2022,12, 8656 4 of 17
Guayaquil (GUAY), and the Herbarium of the Universidad de Almería (HUAL) [
15
], were
also studied.
A database was created with information on the species found in scientific literature and
those detected during field sampling. This database was complemented with the records of
the Global Biodiversity Information Facility (GBIF) [
16
] and Tropicos [
17
], and we used the
information about the ecosystem type to locate where sampling plots or literature records
were acquired according to the ecosystem classification system of continental Ecuador.
The nomenclature of the scientific names was based on Plants of the World Online,
Kew Science (https://powo.science.kew.org/; last access on 15 August 2022), and the
families were classified according to the APG IV classification system.
2.3. Data Analysis
Based on the field sampling records, we studied the plant diversity of the deciduous
forest of the Jama-Zapotillo lowland and the low forest and deciduous shrubland of the
Jama-Zapotillo lowland ecosystems. To achieve this, we selected the results of 110 plots (the
remaining eight plots belonged to the mangrove ecosystem) and calculated species richness,
abundance, and diversity. Species richness was calculated as the number of species present
per plot. Abundance was calculated as the numbers of each individual species per plot.
Two different diversity indices were used, Shannon’s index and Simpson’s index, according
to Equations (1) and (2), respectively:
H=−
S
∑
i=1
pix log2pi(1)
D=1−∑pi2(2)
where H= Shannon’s index, D= Simpson’s diversity index, pi = Abundance of specie i(N)
relative to the total number of species (N), and S= species richness [18].
With these indicators obtained at the plot level in the two sampled ecosystems,
a hierarchical cluster analysis was performed using Sorensen’s distance measure [
19
,
20
].
The Ward method was then used to achieve the highest clustering structure coefficient. The
analysis was performed using the Agnes function of the cluster ‘factoextra’ package in the
R software, version 4.2.1. [21].
This analysis was performed at the plot level and for each of the two sampled ecosys-
tems within the REAr (the deciduous forest of the Jama-Zapotillo lowland and low forest
and deciduous shrubland of the Jama-Zapotillo lowland), separately. Differences in the
different indices between the classes obtained after hierarchical classification were tested by
one-way ANOVA and HSD post hoc analyses using the ‘dplyr ’ package of the R software
version 4.2.1 [21]. The p-value was set to be <0.05.
We used the Ward method because of its strong grouping structure for our variables
(abundance, richness, and diversity) (Table 1).
Table 1. Agglomeration factors of the four clustering methods.
Deciduous Forest of the Jama-Zapotillo Lowland
Clustering Methods Average Single Complete Ward
Agglomeration factor 0.9491697 0.8425549 0.9721348 0.9872076
Low forest and deciduous shrubland of the Jama-Zapotillo lowland
Clustering methods Average Single Complete Ward
Agglomeration factor 0.8996283 0.7202592 0.9487867 0.9791087
3. Results
In the REAr, 381 plant species belonging to 77 families were identified. Physiognomy
was dominated by the herbaceous growth form, containing 167 plant species (44%), fol-

Appl. Sci. 2022,12, 8656 5 of 17
lowed by 134 species of shrubs (35%) and 80 species of trees (21%). The three largest
families were the Fabaceae with 51 plant species, followed by the Malvaceae with 31 species
and the Euphorbiaceae with 26 species. The dominant life forms were phanerophytes with
200 species (52.5%), followed by therophytes with 104 species (27.3%), chamephytes with
22 species (5.8%), epiphytes with 21 species (5.5%), hemicryptophytes with 21 species
(5.5%), geophytes with 7 species (1.8%) and hydrophytes with 6 species (1.6%) (Table 2).
Table 2.
Summary of the list of vascular plants of the REAr, with their scientific name, family, growth
habit, and life form, according to Raunkiaer’s classification (G, geophytes; H, hemicryptophytes;
Hy, hydrophytes; P, phanerophytes; T, therophytes; E, epiphytes).
Family Taxon Life Form Growth Habit
Acanthaceae Dicliptera unguiculata Nees T Herb
Acanthaceae Dicliptera peruviana (Lam.) Juss. T Herb
Acanthaceae Dicliptera paposana Phil. T Herb
Acanthaceae Dyschoriste repens (Nees) Kuntze T Herb
Acanthaceae Justicia comata (L.) L. T Herb
Acanthaceae Avicennia germinans (L.) L. T Herb
Acanthaceae Ruellia floribunda Hook. T Herb
Acanthaceae Ruellia blechum L. T Herb
Acanthaceae Tetramerium nervosum Nees T Herb
Acanthaceae Elytraria imbricata (Valh) Pers. T Herb
Achatocarpaceae Achatocarpus pubescens C.H.Wright P Shrub
Aistroemeriaciae Bomarea obovata Herb. P Herb
Aizoaceae Trianthema portulacastrum L. T Herb
Aizoaceae Sesuvium portulacastrum (L.) L. T Herb
Alismataceae Echinodorus bracteatus Micheli T Herb
Amaranthaceae Alternanthera echinocephala (Hook.f.) Christoph T Herb
Amaranthaceae Alternanthera ficoidea (L.) P.Beauv. T Herb
Amaranthaceae Alternanthera paronychioides A.St.-Hil. T Herb
Amaranthaceae Alternanthera pubiflora (Benth.) Kuntze T Herb
Amaranthaceae Alternanthera porrigens (Jacq.) Kuntze T Herb
Amaranthaceae Alternanthera brasiliana (L) Kuntze T Herb
Amaranthaceae Amaranthus polygonoides Roxb T Herb
Amaranthaceae Iresine angustifolia Euphrasén T Herb
Amaranthaceae Achyranthes aspera L. T Herb
Amaranthaceae Iresine diffusa Humb. & Bonpl. ex Willd T Herb
Amaranthaceae Chamissoa altissima (Jacq.) Kunth. T Herb
Amaranthaceae Gomphrena holosericea (Mart.) Moq. T Herb
Amaranthaceae Salicornia fruticosa (L.) L. T Herb
Amaryllidaceae Leptochiton quitoensis (Herb.) Sealy G Herb
Amaryllidaceae Eucrosia bicolor Ker Gaw G Herb
Anacardiaceae Spondias mombin L. P Shrub
Anacardiaceae Mangifera indica L. P Tree
Anacardiaceae Loxopterygium huasango Spruce ex Engl P Tree
Apiaceae Conium maculatum L. T Herb
Apocynaceae Pseudomarsdenia cundurango (Rchb. f.) Schltr. T Herb
Apocynaceae Prestonia mollis Kunth. T Herb
Apocynaceae Asclepias L. T Herb
Apocynaceae Asclepias L. T Herb
Apocynaceae Asclepias curassavica L. T Herb
Apocynaceae Rauvolfia littoralis Rusby T Herb
Apocynaceae Rauvolfia tetraphylla L. T Herb
Apocynaceae Cascabela thevetia (L) Lippold P Herb
Apocynaceae Nerium oleander L. P Herb
Araceae Anthurium barclayanum Engl. H Herb
Araceae Anthurium soukupii Croat H Herb

Appl. Sci. 2022,12, 8656 6 of 17
Table 2. Cont.
Family Taxon Life Form Growth Habit
Arecaceae Saribus rotundifolius (Lam) Blume P Herb
Arecaceae Adonidia merrillii (Becc) Becc. P Herb
Arecaceae Pritchardia pacifica Seem & H.Wendl P Herb
Asparagaceae Sansevieria trifasciata Prain P Herb
Asparagaceae Yucca guatemalensis Baker G Shrub
Asteraceae Acanthospermum microcarpum BLRob. T Herb
Asteraceae Synedrella nodiflora (L) Gaertn. T Herb
Asteraceae Emilia fosbergii Nicolson T Herb
Asteraceae Eclipta prostrata (L.) L T Herb
Asteraceae Viguiera dentata (Cav.) Spreng. T Herb
Asteraceae Bidens pilosa L. T Herb
Asteraceae Sphagneticola trilobata (L) Pruski P Shrub
Asteraceae Tessaria integrifolia Ruiz & Pav. P Shrub
Asteraceae Verbesina eggersii Hieron. P Shrub
Asteraceae Verbesina lloensis Hieron T Shrub
Asteraceae Barnadesia Mutis ex L.f. T Shrub
Asteraceae Bidens bipinnata L. T Shrub
Asteraceae Dasyphyllum Kunth T Herb
Asteraceae Galinsoga Ruiz & Pav. T Herb
Asteraceae Milleria quinqueflora L. T Herb
Asteraceae Pseudogynoxys chenopodioides (Kunth) Cabrera P Shrub
Asteraceae Tagetes erecta L. T Herb
Basellaceae Anredera ramosa (Moq.) Eliasson E Herb
Bignoniaceae Lundia Puerari ex DC. T Shrub
Bignoniaceae Bignonia longiflora Cav. P Shrub
Bignoniaceae Handroanthus billbergii (Bureau & K. Schum.) S.O.Grose P Tree
Bignoniaceae Jacaranda mimosifolia D.Don P Tree
Bignoniaceae Handroanthus chrysanthus (Jacq.) S.O.Grose P Tree
Bignoniaceae Fridericia dichotoma (Jacq.) P Tree
Bignoniaceae Mansoa hymenaea (DC.) A.H:Gentry P Tree
Bignoniaceae Mansoa verrucifera (Schlecht) A.H.Gentry P Tree
Bignoniaceae Tecoma castanifolia (D. Don.) Melch P Shrub
Bixaceae Bixa orellana L. P Shrub
Bixaceae Cochlospermum vitifolium (Willd.) Spreng P Tree
Boraginaceae Cordia lutea Lam. P Shrub
Boraginaceae Heliotropium angiospermum Murray T Shrub
Boraginaceae Heliotropium curassavicum L. T Shrub
Boraginaceae Heliotropium indicum L. T Shrub
Boraginaceae Tournefortia microcalyx (Ruiz & Pav.) I.M.Johnst. C Shrub
Boraginaceae Varronia cylindristachya Ruiz & Pav. P Shrub
Boraginaceae Varronia macrocephala Desv. P Shrub
Bromeliaceae Aechmea pyramidalis Benth. H Herb
Bromeliaceae Bromelia pinguin L. E Herb
Bromeliaceae Bromelia karatas L. E Herb
Bromeliaceae Tillandsia multiflora Benth. E Herb
Bromeliaceae Tillandsia usneoides (L.) L. E Herb
Bromeliaceae Tillandsia streptocarpa Baker E Herb
Bromeliaceae Tillandsia latifolia Meyen E Herb
Bromeliaceae Tillandsia recurvata (Gaudich.) Baker E Herb
Bromeliaceae Tillandsia triglochinoides C.Presl E Herb
Bromeliaceae Tillandsia disticha Kunth. E Herb
Bromeliaceae Tillandsia barclayana Baker. E Herb
Bromeliaceae Tillandsia espinosae L. B. Sm. E Herb
Bromeliaceae Tillandsia cyanea L. B. Sm. E Herb
Bromeliaceae Tillandsia caerulea Kunth E Herb
Burseraceae Bursera graveolens (Kunth) Triana & Planch P Tree

Appl. Sci. 2022,12, 8656 7 of 17
Table 2. Cont.
Family Taxon Life Form Growth Habit
Cactaceae Pilosocereus lanuginosus (L.) Byles & G:D. Rowley P Shrub
Cactaceae Armatocereus cartwrightianus Backeb. P Shrub
Cactaceae Armatocereus matucanensis Backeb. P Shrub
Cactaceae Opuntia ficus-indica (L.) Mill. P Shrub
Cactaceae Opuntia stricta (Haw) Haw. P Shrub
Cactaceae Opuntia quitensis F.A.C. Weber P Shrub
Cactaceae Melocactus peruvianus Vaupel P Shrub
Cactaceae Selenicereus monacanthus (Lem) D.R.Hunt P Shrub
Cactaceae Cereus hexagonus (L.) Mill. P Shrub
Cactaceae Cephalocereus Pfeiff. P Shrub
Cactaceae Cleistocactus Pfeiff. P Shrub
Cannabaceae Celtis iguanaea (Jacq) Sarg P Shrub
Cannabaceae Trema micrantha (L.) Blume P Shrub
Capparaceae Beautempsia avicenniifolia Gaudich. ex Alleiz. P Shrub
Capparaceae Capparicordis crotonoides (Kunth) Iltis & Cornejo P Shrub
Capparaceae Colicodendron scabridum (Kunth) P Tree
Capparaceae Cynophalla heterophylla (Ruiz & Pav. ex DC.) Iltis & Cornejo P Tree
Capparaceae Cynophalla flexuosa (L.) J.Presl P Tree
Caricaceae Carica papaya L. P Shrub
Caricaceae Carica parviflora (A. DC.) Solms P Shrub
Celastraceae Tricerma octogonum (L’Hér.) Lundell P Shrub
Combretaceae Conocarpus erectus L. P Tree
Combretaceae Terminalia catappa L. P Tree
Combretaceae Laguncularia racemosa (L.) C.F.Gaertn Hy Tree
Commelinaceae Commelina diffusa Burm.f. P Herb
Commelinaceae Commelina Plum. ex L. P Herb
Commelinaceae Calli cordifolia (Sw.) Andiers. & Woodson P Herb
Convolvulaceae Camonea umbellata (L.) A.R.Simões & Staples P Herb
Convolvulaceae Cuscuta acuta Engelm. P Herb
Convolvulaceae Ipomoea batatas (L.) Lam. P Herb
Convolvulaceae Evolvulus convolvuloides (Willd. ex Schult.) Stearn T Herb
Convolvulaceae Evolvulus nummularius (L.) L. T Herb
Convolvulaceae Ipomoea asarifolia (Desr.) Roem. & Schult. T Herb
Convolvulaceae Ipomoea cholulensis Kunth P Herb
Convolvulaceae Ipomoea hederifolia L. T Herb
Convolvulaceae Ipomoea regnellii Meisn. P Herb
Convolvulaceae Ipomoea quamoclit L. P Herb
Convolvulaceae Ipomoea pes-caprae (L.) R. Br. P Herb
Convolvulaceae Ipomoea nil (L) Roth P Herb
Convolvulaceae Ipomoea carnea Jacq. P Herb
Convolvulaceae Ipomoea triloba Thunb P Herb
Convolvulaceae Jacquemontia corymbulosa Benth P Herb
Convolvulaceae Jacquemontia unilateralis (Roem. & Schult.) O’Donell P Herb
Convolvulaceae Distimake aegyptius (L.) A.R.Simões & Staples P Herb
Cucurbitaceae Momordica charantia L. P Herb
Cucurbitaceae Luffa operculata (L.) Cogn. P Herb
Cucurbitaceae Cucurbita pepo L. T Herb
Cucurbitaceae Cayaponia glandulosa (Mart.) Cogn. T Herb
Cucurbitaceae Echinopepon racemosus (Steud.) C. Jeffrey P Herb
Cyperaceae Cyperus squarrosus L. H Herb
Cyperaceae Cyperus compressus L. H Herb
Cyperaceae Cyperus ligularis L. H Herb
Cyperaceae Cyperus microbolbos C.B.Clarke H Herb
Cyperaceae Fimbristylis littoralis Gaudich. Hy Herb
Erythroxylaceae Erythroxylum glaucum O. E. Schulz P Herb
Erythroxylaceae Erythroxylum acuminatum Ruiz & Pav. P Shrub
Euphorbiaceae Jatropha curcas L. P Shrub
Euphorbiaceae Euphorbia serpens Kunth. T Shrub

Appl. Sci. 2022,12, 8656 8 of 17
Table 2. Cont.
Family Taxon Life Form Growth Habit
Euphorbiaceae Euphorbia hirta L. T Shrub
Euphorbiaceae Euphorbia heterophylla L. T Shrub
Euphorbiaceae Cnidoscolus aconitifolius (Mill.) IM Johnst. P Shrub
Euphorbiaceae Croton hirtus L’Hér. P Shrub
Euphorbiaceae Croton fraseri Mull. Arg. P Shrub
Euphorbiaceae Croton rivinifolius Kunth P Shrub
Euphorbiaceae Croton eggersii Pax P Shrub
Euphorbiaceae Croton aequatoris Croizat P Shrub
Euphorbiaceae Croton eggersii Pax P Shrub
Euphorbiaceae Croton jamesonii Müll. Arg. P Shrub
Euphorbiaceae Croton schiedeanus Schltdl. P Shrub
Euphorbiaceae Codiaeum variegatum (L.) Rumph. ex A.Juss P Shrub
Euphorbiaceae Acalypha villosa Jacq. P Shrub
Euphorbiaceae Acalypha subcastrata F.Aresch. T Shrub
Euphorbiaceae Acalypha diversifolia Jacq. P Shrub
Euphorbiaceae Acalypha cuspidata Jacq. P Shrub
Euphorbiaceae Acalypha setosa A. Rich. P Shrub
Euphorbiaceae Acalypha cuspidata Jacq. P Shrub
Euphorbiaceae Acalypha alopecuroides L. T Shrub
Euphorbiaceae Dalechampia scandens L. T Shrub
Euphorbiaceae Codiaeum variegatum (L.) Rumph. ex A. Juss. P Shrub
Euphorbiaceae Manihot esculenta Crantz G Tree
Fabaceae Senegalia polyphylla (DC.) Britton & Rose P Tree
Fabaceae
Vachellia macracantha (Humb. & Bonpl. ex Willd.) Seigler & Ebinger
P Tree
Fabaceae Aeschynomene tumbezensis J.F. Macbr. P Tree
Fabaceae Aeschynomene scoparia Kunth P Tree
Fabaceae Aeschynomene americana L. T Tree
Fabaceae Albizia multiflora (Kunth) Barneby y J.W.Grimes P Tree
Fabaceae Caesalpinia pulcherrima (L.) Sw. P Tree
Fabaceae Chloroleucon mangense (Jack.) Britton & Rose P Tree
Fabaceae Calliandra tumbeziana J.F. Macbr. T Shrub
Fabaceae Canavalia ensiformis (L.) DC. T Shrub
Fabaceae Canavalia rosea (SW) P Shrub
Fabaceae Canavalia brasiliensis Mart. ex Benth. P Shrub
Fabaceae Centrosema pubescens Benth. P Shrub
Fabaceae Cercidium praecox (Ruiz & Pav) Hawkins P Shrub
Fabaceae Chamaecrista nictitans (L.) Moench T Shrub
Fabaceae Crotalaria incana L. P Herb
Fabaceae Coursetia caribaea Ochroleuca (Jacq.) P Shrub
Fabaceae Delonix regia (Bojer ex Hook.) Raf. P Tree
Fabaceae Desmanthus virgatus (L.) Willd. T Herb
Fabaceae Desmodium scorpiurus (Sw.) Desv. ex DC. T Shrub
Fabaceae Desmodium procumbens (Mill.) C.L.Hitchc. T Shrub
Fabaceae Erythrina smithiana Krukoff P Tree
Fabaceae Erythrina velutina Willd. P Tree
Fabaceae Geoffroea spinosa Jacq. P Tree
Fabaceae Indigofera subulata Vahl ex Poir. P Tree
Fabaceae Leucaena trichodes (Jacq.) Benth. P Tree
Fabaceae Libidibia glabrata (Kunth) C.Cast. & G.P.Lewis P Tree
Fabaceae Machaerium Pers. P Tree
Fabaceae Machaerium millei Standl. P Tree
Fabaceae Mimosa albida Humb. & Bonpl. ex Willd P Tree
Fabaceae Mimosa acantholoba (Humb. & Bonpl. ex Willd.) Poir. P Tree
Fabaceae Mimosa pigra L. P Tree
Fabaceae Neptunia plena (L.) Benth. T Shrub
Fabaceae Parkinsonia aculeata L. P Tree
Fabaceae Pithecellobium dulce (Roxb.) Benth. P Tree
Fabaceae Piptadenia retusa (Jacq.) P.G.Ribeiro, Seigler & Ebinger P Tree

Appl. Sci. 2022,12, 8656 9 of 17
Table 2. Cont.
Family Taxon Life Form Growth Habit
Fabaceae Piscidia carthagenensis Jacq. P Tree
Fabaceae Pithecellobium excelsum (Kunth) Mart. P Tree
Fabaceae Prosopis juliflora (Sw.) DC. P Tree
Fabaceae Prosopis pallida (Willd.) (Humb. & Bonpl. ex Willd.) Kunth P Tree
Fabaceae Phaseolus vulgaris L. T Herb
Fabaceae Pseudosamanea guachapele (Kunth) Harms P Tree
Fabaceae Samanea saman (Jacq.) Merr. P Tree
Fabaceae Senna mollissima (Humb. & Bonpl. ex Willd.) H.S.Irwin & Barneby P Tree
Fabaceae Senna oxyphylla (Kunth) H.S.Irwin & Barneby P Tree
Fabaceae Galactia striata (Jacq.) Urb. P Herb
Fabaceae Vigna caracalla (L.) Verdc. T Herb
Fabaceae Schizolobium parahyba (Vell.) S.F. Blake P Tree
Fabaceae Stylosanthes scabra Vogel P Herb
Fabaceae Stylosanthes guianensis (Aubl.) Sw. P Herb
Fabaceae Inga edulis Mart. P Tree
Lamiaceae Hyptis atrorubens Poit. T Shrub
Lamiaceae Tectona grandis L. f. P Tree
Loasaceae Gronovia scandens L. P Herb
Loasaceae Mentzelia aspera L. T Herb
Loranthacea Psittacanthus divaricatus (Kunth) G.Don P Shrub
Lythraceae Adenaria floribunda Kunth P Shrub
Malpighiaceae Bunchosia plowmanii W.R. Anderson P Shrub
Malpighiaceae Malpighia glabra L. P Shrub
Malpighiaceae Malpighia emarginata DC. P Shrub
Malvaceae Abutilon pedunculare Kunth P Shrub
Malvaceae Abutilon reflexum (Lam.) Sweet P Shrub
Malvaceae Abutilon umbellatum (L.) Sweet P Shrub
Malvaceae Abutilon dianthum C. Presl P Shrub
Malvaceae Ayenia magna L. P Shrub
Malvaceae Bastardia bivalvis (Cav.) Kunth ex Griseb. P Shrub
Malvaceae Bastardia viscosa (L.) Kunth P Shrub
Malvaceae Ceiba trichistandra (A. Gray) Bakh. P Tree
Malvaceae Cienfuegosia tripartita (Kunth) Gürke T Herb
Malvaceae Corchorus orinocensis Kunth T Shrub
Malvaceae Eriotheca ruizii (K. Schum.) A. Robyns P Tree
Malvaceae Guazuma ulmifolia Lam. P Tree
Malvaceae Hibiscus escobariae Fryxell P Shrub
Malvaceae Hibiscus phoeniceus Jacq. P Shrub
Malvaceae Hibiscus rosa-sinensis L. P Shrub
Malvaceae Hibiscus pernambucensis Arruda P Shrub
Malvaceae Malachra fasciata Jacq. T Shrub
Malvaceae Malvaviscus concinnus Kunth P Shrub
Malvaceae Byttneria parviflora Benth. C Shrub
Malvaceae Byttneria glabrescens Benth. C Shrub
Malvaceae Gaya peruviana Ulbr. f. C Shrub
Malvaceae Sida acuta Burm.f. C Shrub
Malvaceae Sida ciliaris L. C Herb
Malvaceae Sida repens Dombey ex Cav. C Herb
Malvaceae Sida setosa Mart. ex Colla C Herb
Malvaceae Theobroma cacao L. P Shrub
Malvaceae Wissadula diffusa R.E.Fr. C Shrub
Malvaceae Pavonia fruticosa (Mill.) Fawc. &Rendle C Shrub
Malvaceae Triumfetta bogotensis DC. C Shrub
Malvaceae Gossypium barbadense L. P Shrub
Malvaceae Sidastrum paniculatum (L.) Fryxell T Shrub
Marantaceae Thalia geniculata L. T Shrub
Meliaceae Cedrela odorata L. P Tree
Meliaceae Azadirachta indica A. Juss. P Tree

Appl. Sci. 2022,12, 8656 10 of 17
Table 2. Cont.
Family Taxon Life Form Growth Habit
Moraceae Ficus citrifolia Mill. P Tree
Moraceae Ficus elastica Roxb. ex Hornem. P Tree
Moraceae Artocarpus altilis (Parkinson) Fosberg P Shrub
Moringaceae Moringa oleifera Lam. P Tree
Muntingiaceae Muntingia calabura L. P Herb
Musaceae Musa x paradisiaca L. T Herb
Myrtaceae Eugenia biflora (L.) DC. P Herb
Nyctaginaceae Boerhavia coccinea Mill. T Herb
Nyctaginaceae Boerhavia erecta L. T Shrub
Nyctaginaceae Pisonia floribunda Hook. f. P Shrub
Nyctaginaceae Pisonia ambigua Heimerl P Shrub
Nyctaginaceae Pisonia aculeata L. P Shrub
Nyctaginaceae Bougainvillea peruviana Bonpl P Shrub
Nyctaginaceae Bougainvillea spectabilis Willd. P Shrub
Nyctaginaceae Cryptocarpus pyriformis Kunth P Herb
Nyctaginaceae Mirabilis violacea (L.) Heimerl T Herb
Nyctaginaceae Mirabilis nyctaginea (Michx.) MacMill. T Herb
Nymphaeaceae Nymphaea ampla (Salisb.) DC. Hy Herb
Nymphaeaceae Nymphaea pulchella DC. Hy Tree
Oleaceae Priogymnanthus apertus (B.Ståhl) P.S.Green P Shrub
Onagraceae Ludwigia linifolia (Vahl) R.S.Rao T Shrub
Onagraceae Ludwigia octovalvis (Jacq.) P.H. Raven T Shrub
Opilaceae Agonandra excelsa Griseb. P Herb
Orchidaceae Oncidium hyphaematicum Rchb.f E Herb
Orchidaceae Epidendrum bracteolatum C. Presl E Herb
Orchidaceae Notylia replicata Rchb.f. E Herb
Orchidaceae Rodriguezia strobelii Garay E Herb
Orchidaceae Zelenkoa onusta (Lindl.) M.W.Chase & N.H.Williams E Herb
Orchidaceae Encyclia aspera (Lindl.) Schltr E Herb
Orchidaceae Campylocentrum micranthum (Lindl.) Rolfe E Herb
Oxalidaceae Oxalis dombeyi A.St.-Hil. G Herb
Oxalidaceae Oxalis microcarpa Benth. G Herb
Passifloraceae Passiflora biflora Lam. P Herb
Passifloraceae Passiflora eggersii Harms P Herb
Passifloraceae Passiflora suberosa L. P Herb
Phyllanthaceae Phyllanthus niruri L. T Shrub
Phytolaccaceae Achatocarpus pubescens C.H.Wright P Tree
Piperaceae Peperomia pellucida (L.) Kunth T Herb
Plumbaginaceae Plumbago scandens L. T Herb
Poaceae Cynodon dactylon (L) Per. H Herb
Poaceae Echinochloa colonum (L.) Link H Herb
Poaceae Lasiacis divaricata (L.) Hitchc H Herb
Poaceae Paspalum vaginatum Sw. H Herb
Poaceae Sporobolus pyramidatus (Lam.) Hitchc. H Herb
Poaceae Panicum trichoides Sw. H Herb
Poaceae Panicum polygonatum Schrad. H Herb
Poaceae Panicum maximum (Jacq.) H Herb
Poaceae Olyra latifolia L. H Herb
Poaceae Setaria palmifolia (J. Koenig) Stapf H Herb
Poaceae Chloris virgata Sw. H Herb
Polygalaceae Asemeia violacea (Aubl.) J.F.B.Pastore & J.R.Abbott T Herb
Polygalaceae Polygala paniculata L. T Herb
Polygonaceae Triplaris cumingiana Fisch. & C.A. Mey. P Tree
Polygonaceae Coccoloba ruiziana Lindau P Tree
Portulacaceae Portulaca oleracea L. H Herb
Primulaceae Bonellia sprucei (Mez) B.Ståhl & Källersjö P Tree
Pteridaceae Adiantum raddianum C. Presl G Herb
Pteridaceae Acrostichum aureum L. H Herb

Appl. Sci. 2022,12, 8656 11 of 17
Table 2. Cont.
Family Taxon Life Form Growth Habit
Rhamnaceae Scutia spicata (Humb. & Bonpl. ex Schult.) Weberb. C Tree
Rhamnaceae Gouania mollis Reissek C Tree
Rhamnaceae Sarcomphalus thyrsiflorus (Benth.) Hauenschild P Tree
Rhizophoraceae Rhizophora ×harrisonii Leechm. Hy Tree
Rhizophoraceae Rhizophora mangle L. Hy Tree
Rubiaceae Simira ecuadorensis (Standl.) Steyerm. P Tree
Rubiaceae Randia armata (SW.) DC. P Shrub
Rubiaceae Sphinctanthus aurantiacus (Standl.) Fagerl. P Shrub
Rubiaceae Ixora coccinea L. P Shrub
Rubiaceae Sommera purdiei Standl. P Shrub
Rubiaceae Duroia hirsuta (Poepp.) K.Schum. P Tree
Rubiaceae Spermacoce remota Lam. P Tree
Rubiaceae Morinda citrifolia L. P Tree
Rutaceae Zanthoxylum martinicense (Lam.) DC. P Tree
Rutaceae Zanthoxylum rigidum Humb. & Bonpl. ex Willd. P Tree
Rutaceae Amyris balsamifera L. P Shrub
Rutaceae Citrus ×sinensis (L.) Osbeck P Tree
Rutaceae Citrus ×limon (L.) Osbeck P Tree
Rutaceae Zanthoxylum fagara (L.) Sarg. P Tree
Sapindaceae Sapindus saponaria L. P Tree
Sapindaceae Serjania mucronulata Radlk. P Shrub
Sapindaceae Cardiospermum corindum L. P Herb
Scrophulariaceae Capraria peruviana Benth C Shrub
Solanaceae Solanum peruvianum L. T Herb
Solanaceae Browallia americana L. T Herb
Solanaceae Solanum filiforme Ruiz & Pav. T Herb
Solanaceae Solanum nigrum L. T Herb
Solanaceae Solanum pimpinellifolium L. T Herb
Solanaceae Lycianthes ecuadorensis Bitter T Herb
Solanaceae Lycianthes cyathocalyx (Van Heurck & Müll. Arg.) Bitter T Herb
Solanaceae Witheringia solanacea L’He’r C Herb
Solanaceae Lycium nodosum Jacq. P Herb
Solanaceae Capsicum annuum L. C Herb
Talinaceae Talinum paniculatum (Jacq.) Gaertn. H Shrub
Talinaceae Talinum fruticosum (L.) Juss. T Shrub
Tropaeolaceae Tropaeolum harlingii Sparre P Shrub
Urticaceae Laportea aestuans (L.) Chew T Shrub
Verbenaceae Citharexylum quitense Spreng. T Tree
Verbenaceae Citharexylum gentryi Moldenke T Tree
Verbenaceae Lantana sprucei Hayek C Shrub
Verbenaceae Lantana fucata Lindl. C Shrub
Verbenaceae Lantana camara L. C Shrub
Verbenaceae Lantana trifolia L. C Shrub
Verbenaceae Lantana verticillata verticillata (L.) Nicolson P Herb
Verbenaceae Lantana velutina M. Martens & Galeotti C Shrub
Verbenaceae Lantana horrida Kunth C Shrub
Vitaceae Cissus erosa Rich. P Herb
Zygophyllaceae Kallstroemia pubescens (G.Don) Dandy T Herb
Forest Classification
Based on the hierarchical grouping performed on the similarities of species richness,
abundance, Shannon index, and Simpson index (Supplement S1 in Supplementary Material),
we identified three dry forest types in the deciduous forest of the Jama-Zapotillo lowlands
ecosystem. Type I (black line) contains 12 sample plots, type II (red line) contains 26 sample
plots and type III (green line) contains 19 sample plots (Figure 2).

Appl. Sci. 2022,12, 8656 12 of 17
Appl. Sci. 2022, 12, 8656 13 of 18
Figure 2. A dendrogram (Using the ward.D algorithm and Euclidean distance) of the grouping of
the plots sampled in the deciduous forest of the Jama-Zapotillo lowland ecosystem. The grouping
of the plots into the three forest types, according to the calculated diversity indices, is illustrated.
The mean values of the diversity indices showed significant differences between forest
types I and III. Type I presented the highest species richness (6.16) and abundance (18.58).
The type I forest was also the ecosystem with the highest values of the two different diver-
sity indices, but no significant differences were found between this ecosystem and ecosys-
tem type III. Forest type III had the lowest values of both diversity indices. The highest val-
ues of richness, abundance, and diversity that were found in forest type I (Figure 3) fits with
the specific features that define a conserved forest, with some relevant species such as
Handroanthus chrysanthus, Eriotheca ruizii, Bursera graveolens, and Ceiba trischistandra. The
maximum height that was determined in this forest was 27 m, which was represented by
the presence of the species Ceiba trischistandra.
Figure 3. Comparison of the mean values of each diversity index in the three forest types determined
in the deciduous forest of the Jama-Zapotillo lowland ecosystem.
Figure 2.
A dendrogram (Using the ward.D algorithm and Euclidean distance) of the grouping of the
plots sampled in the deciduous forest of the Jama-Zapotillo lowland ecosystem. The grouping of the
plots into the three forest types, according to the calculated diversity indices, is illustrated.
The mean values of the diversity indices showed significant differences between forest
types I and III. Type I presented the highest species richness (6.16) and abundance (18.58).
The type I forest was also the ecosystem with the highest values of the two different diversity
indices, but no significant differences were found between this ecosystem and ecosystem
type III. Forest type III had the lowest values of both diversity indices. The highest values
of richness, abundance, and diversity that were found in forest type I (Figure 3) fits with
the specific features that define a conserved forest, with some relevant species such as
Handroanthus chrysanthus,Eriotheca ruizii,Bursera graveolens, and Ceiba trischistandra. The
maximum height that was determined in this forest was 27 m, which was represented by
the presence of the species Ceiba trischistandra.
Appl. Sci. 2022, 12, 8656 13 of 18
Figure 2. A dendrogram (Using the ward.D algorithm and Euclidean distance) of the grouping of
the plots sampled in the deciduous forest of the Jama-Zapotillo lowland ecosystem. The grouping
of the plots into the three forest types, according to the calculated diversity indices, is illustrated.
The mean values of the diversity indices showed significant differences between forest
types I and III. Type I presented the highest species richness (6.16) and abundance (18.58).
The type I forest was also the ecosystem with the highest values of the two different diver-
sity indices, but no significant differences were found between this ecosystem and ecosys-
tem type III. Forest type III had the lowest values of both diversity indices. The highest val-
ues of richness, abundance, and diversity that were found in forest type I (Figure 3) fits with
the specific features that define a conserved forest, with some relevant species such as
Handroanthus chrysanthus, Eriotheca ruizii, Bursera graveolens, and Ceiba trischistandra. The
maximum height that was determined in this forest was 27 m, which was represented by
the presence of the species Ceiba trischistandra.
Figure 3. Comparison of the mean values of each diversity index in the three forest types determined
in the deciduous forest of the Jama-Zapotillo lowland ecosystem.
Figure 3.
Comparison of the mean values of each diversity index in the three forest types determined
in the deciduous forest of the Jama-Zapotillo lowland ecosystem.

Appl. Sci. 2022,12, 8656 13 of 17
As similarly observed in the deciduous forest of the Jama-Zapotillo lowland, the
hierarchical grouping of the low forest and deciduous shrubland of the Jama-Zapotillo
lowland discriminated three forest types based on the similarity of the indicators: species
richness, abundance, Shannon index, and Simpson index. Type I (black line) contains
27 plots, type II (red line) contains 5 plots and type III (green line) 21 plots. (Figure 4).
Appl. Sci. 2022, 12, 8656 14 of 18
As similarly observed in the deciduous forest of the Jama-Zapotillo lowland, the hi-
erarchical grouping of the low forest and deciduous shrubland of the Jama-Zapotillo low-
land discriminated three forest types based on the similarity of the indicators: species rich-
ness, abundance, Shannon index, and Simpson index. Type I (black line) contains 27 plots,
type II (red line) contains 5 plots and type III (green line) 21 plots. (Figure 4).
Figure 4. A dendrogram (Using the ward.D algorithm and Euclidean distance) of the grouping of
the plots sampled in the low forest and deciduous shrubland of the Jama-Zapotillo lowland ecosys-
tem of the REAr. The grouping of the plots into three forest types, according to the calculated diver-
sity indices, is illustrated.
Significant differences in species richness (p-value = 1.27 × 10−6), abundance (p-value
< 2 × 10−16), and Shannon index (p-value = 0.00712) were identified among the three forest
types identified for this ecosystem. Forest type III had the highest species richness (7.00),
abundance (20.20), alpha diversity with Simpson’s index (0.72), and Shannon index (1.63),
which represent characteristics that define a well preserved forest (Figure 5). In type III,
the following species stood out: Cochlospermum vitifolium, Handroanthus chrysanthus, Caes-
alpinia glabrata, and Geoffroea spinosa. The maximum height that was determined in this
forest was 13 m.
Figure 5. Comparison of the mean values in the three forest types of the low forest and deciduous
shrubland of the Jama-Zapotillo lowland ecosystem.
Figure 4.
A dendrogram (Using the ward.D algorithm and Euclidean distance) of the grouping of the
plots sampled in the low forest and deciduous shrubland of the Jama-Zapotillo lowland ecosystem
of the REAr. The grouping of the plots into three forest types, according to the calculated diversity
indices, is illustrated.
Significant differences in species richness (p-value = 1.27
×
10
−6
), abundance
(p-value < 2
×
10
−16
), and Shannon index (p-value = 0.00712) were identified among the
three forest types identified for this ecosystem. Forest type III had the highest species rich-
ness (7.00), abundance (20.20), alpha diversity with Simpson’s index (0.72), and Shannon
index (1.63), which represent characteristics that define a well preserved forest (Figure 5). In
type III, the following species stood out: Cochlospermum vitifolium,Handroanthus chrysanthus,
Caesalpinia glabrata, and Geoffroea spinosa. The maximum height that was determined in this
forest was 13 m.
Appl. Sci. 2022, 12, 8656 14 of 18
As similarly observed in the deciduous forest of the Jama-Zapotillo lowland, the hi-
erarchical grouping of the low forest and deciduous shrubland of the Jama-Zapotillo low-
land discriminated three forest types based on the similarity of the indicators: species rich-
ness, abundance, Shannon index, and Simpson index. Type I (black line) contains 27 plots,
type II (red line) contains 5 plots and type III (green line) 21 plots. (Figure 4).
Figure 4. A dendrogram (Using the ward.D algorithm and Euclidean distance) of the grouping of
the plots sampled in the low forest and deciduous shrubland of the Jama-Zapotillo lowland ecosys-
tem of the REAr. The grouping of the plots into three forest types, according to the calculated diver-
sity indices, is illustrated.
Significant differences in species richness (p-value = 1.27 × 10−6), abundance (p-value
< 2 × 10−16), and Shannon index (p-value = 0.00712) were identified among the three forest
types identified for this ecosystem. Forest type III had the highest species richness (7.00),
abundance (20.20), alpha diversity with Simpson’s index (0.72), and Shannon index (1.63),
which represent characteristics that define a well preserved forest (Figure 5). In type III,
the following species stood out: Cochlospermum vitifolium, Handroanthus chrysanthus, Caes-
alpinia glabrata, and Geoffroea spinosa. The maximum height that was determined in this
forest was 13 m.
Figure 5. Comparison of the mean values in the three forest types of the low forest and deciduous
shrubland of the Jama-Zapotillo lowland ecosystem.
Figure 5.
Comparison of the mean values in the three forest types of the low forest and deciduous
shrubland of the Jama-Zapotillo lowland ecosystem.

Appl. Sci. 2022,12, 8656 14 of 17
4. Discussion
The detailed floristic analysis of our study reveals that, despite only focusing our
sampling on two of the four ecosystems, [
13
], the richness and diversity of the species in
this ecological reserve is greater than what was described in the last catalog of vascular flora
detailing the complete sets of ecosystems included within the REAr [
11
]. The physiognomy
of the species found in the study area was dominated by the herbaceous growth form
with 167 species (44%). This is particularly interesting, as the tree species component in
the tropical dry forests of the equatorial Pacific region is known, but knowledge of the
shrubs and herbs is very limited. Nevertheless, seasonal dry forests have a very wide
range of climatic tolerance [
22
], the predominance of herbaceous species over tree species
may be due to the climatic conditions [
23
–
25
]. In the present study, all flora species were
inventoried during the rainy season, corresponding with the onset of herbaceous flora.
Our findings are similar to those obtained in studies analyzing the floristic features
of the dry forests of the central coastal Andes mountains that cover the Tumbesian region
(58 forest families) [
24
] and other studies on Ecuador’s dry forests that report data on woody
plant species [
26
,
27
]. Among all of the families, the Fabaceae was the most abundant, fol-
lowed by the Malvaceae and Euphorbiaceae, and the dominant species in the reserve was the
Handroanthus chrysanthus. These results are in accordance with previous studies reporting the
Fabaceae as the best represented group in neotropical dry forests [
28
] and with specific studies
in this region. For example, Molina [
11
] documented that the Fabaceae and Euphorbiaceae were
the two predominant families in the REAr, and Luna et al. [
12
] cited the Fabaceae and Malvaceae
as the most representative families. In 2006, Cerón et al. [
8
] reported that the Malvaceae and
Poaceae were the most numerous families, with8 species each, while the Fabaceae had 7 species.
Species assemblages in tropical dry forests appear to be primarily controlled by al-
titude and water availability [
28
], and the life forms reflect the bioclimate of the area.
Raunkiaer [
14
] designed three main phytoclimates based on the landform spectrum, includ-
ing phanerophyte for the tropics, therophyte for xeric environments, and hemicryptophyte
for the cool temperate region. The present study identified seven different classes of life
forms in the study area. As is the case in most dry forest remnants, our study reveals that
the dominant life forms in the REAr were phanerophytes (52.5%) and terophytes (86%).
Many botanists have collected and studied the flora of Ecuador since the early 18th
century [
29
–
32
]; the findings of our study highlight the importance of conserving the forests
of the REAr and the species composition of the region. The analysis of species dominance
and diversity within the REAr shows that 35% of the tree species identified are exclusive
to the large floristic group of the Central Andes of the neotropical dry forest coast [
24
].
In the seasonal dry forests of Ecuador and Peru, 313 woody species can be found [
33
].
The presence of the Andes is one of the main causes of the isolation of the trans-Andean
Pacific coastal region, which is characterized by the high levels of floristic endemism in the
seasonal dry forests of Ecuador [33,34].
Many species are shared between dry forest formations and between the provinces
of Ecuador [
26
]. In this study, we have observed that the species richness and diversity of
REAr ecosystems depend on the dominant ecosystem type. Herbaceous species abundance
and Shannon type diversity were higher in the deciduous forest of the Jama-Zapotillo
lowland. The dominant families by number of species, density, abundance, and domi-
nance of individuals are: the Fabaceae, Mimosaceae, Moraceae, and Bombacaceae [
35
,
36
].
The species composition analysis of this dry forest was dominated by species such as
Handroanthus chrysanthus and revealed a general pattern of variation in community diver-
sity and species composition. Several authors [
26
,
34
,
37
] have reported similar composition
patterns of this species in the dry forests of the same region. The dendrograms divide the
ecosystems studied into three types that differ in richness, abundance, and diversity. This
seems to be related to the anthropic pressures exerted, with the least rich and diverse areas
being the most degraded near the border, roads, or agricultural environments.
The information provided in this study is especially relevant due to the fragility of
the tropical dry forest, which is the main ecosystem within the REAr and is the ecosystem

Appl. Sci. 2022,12, 8656 15 of 17
where our specific data collection has been performed. Indeed, the tropical dry forest is
one of the most threatened biomes in the world [
26
,
38
] and has experienced one of the
most extensive rates of habitat loss during the last few decades [
38
]. Be that as it may, it is
a much less-studied ecosystem than other tropical ecosystems, such as rainforests [
6
]. The
small extension and high fragmentation of the Ecuadorian dry forests makes them more
sensitive than those which are located in other countries [
39
]; this situation is especially
relevant in the REAr. Despite being classified as a natural reserve, the highest protection
level in Ecuador, it is a vulnerable area due to the constant pressure of extracting resources
and expanding the agricultural and ranching areas within the REAr limits. In addition,
clandestine crossings by outsiders and the deforestation of trees to create new access routes
occur in this border area. These activities exert a strong negative impact on the vegetation.
Further studies are needed on the biodiversity of the REAr and the relationships within
the ecosystems and external pressures. Furthermore, transdisciplinary initiatives that take
into account the ecological and human dimensions are a priority for safeguarding the
integrity of the REAr [
40
,
41
] along with similar regions. Strategies can be implemented
in this manner to reduce the negative impacts on these regions and provide alternative
livelihoods for local communities. This would allow for the sustainable use and conserva-
tion of the REAr’s invaluable biodiversity for future generations, as well as its associated
ecosystem services [42–44].
5. Conclusions
The present study reveals that the REAr has a greater floristic diversity than has previ-
ously been described. These new data are beneficial for the protection and conservation of
this region.
The predominant life forms are phanerophytes and terophytes, which are conditioned
by the presence of dry tropical climate in deciduous forests.
Within the dry forest zone, there is a great variety of forest types with different levels
of richness, possibly because of differences in the level anthropogenic pressure between
zones (i.e., greater impacts and more degraded areas with less richness are on the edges of
the reserve, the areas closest to access roads or agricultural zones).
Supplementary Materials:
The following supporting information can be downloaded at: https://
www.mdpi.com/article/10.3390/app12178656/s1, Supplement S1: Ecosystem type and biodiversity
and richness values of all field plots.
Author Contributions:
Conceptualization: A.D.L.-F., D.A.N.-N., E.R.-C., J.L.M.-P. and E.G.-L.; valida-
tion, formal analysis and investigation: D.A.N.-N. and A.D.L.-F.; resources: E.G.-L.; writing: A.D.L.-F.;
visualization: A.D.L.-F., E.G.-L., E.R.-C., J.L.M.-P. and D.A.N.-N.; supervision: E.G.-L. All authors
have read and agreed to the published version of the manuscript.
Funding: This research received no external funding.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: All data are available upon request.
Acknowledgments:
We express our gratitude to the park rangers and the person in charge of the
REAr, who are the direct actors in the control and care of this protected area attached to the National
System of Protected Areas, part of the Ministry of Environment, Water and Ecological Transition of
Ecuador; through the approved research permit No. 005-2018-IC-FLO/FAU-DPAEO-MAE, it was
possible to conduct this research.
Conflicts of Interest: The authors declare no conflict of interest.

Appl. Sci. 2022,12, 8656 16 of 17
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