ArticlePDF Available

Abstract and Figures

Neotropical dry forests are among the most diverse and threatened ecosystems worldwide. The extent and knowledge of Neotropical dry forests are quite heterogeneous with forests located in the Ecuadorian province especially diverse, threatened and poorly studied. In this work, we review patterns and conservation status of biodiversity, ecosystem processes and human perception of tropical dry forest of the Ecuadorian province. We found that patterns of biodiversity, endemism and conservation status are generally poorly studied. Overall, these forests provide habitat for at least 900 species including trees, birds, mammals, amphibians and reptiles. On average 18% of these species (range 6–25%) are endemic to the region and 25% (3–50%) are recognized as threatened. Little is known about groups such as invertebrates, fish, fungi, or herbaceous plants, and as well as about processes generating and maintaining critical ecosystem functions. Available literature points out the importance of positive ecological interactions such us plant–frugivore and plant–plant facilitation interactions in maintaining the regeneration dynamics of these forests. Faced by the formative state of knowledge about basic biodiversity patterns and ecological functions, the implementation of ecosystem risk assessment under the IUCN criteria for the Red List of Ecosystems may offer constructive means to organize, integrate and advance existing knowledge and conservation priorities for dry forests of the Ecuadorian province. With examples of existing conflicts between people and protected areas, we emphasize the importance of consultation and involvement of local communities in the development of conservation measures including new protected areas. Lastly, we reflect on some encouraging examples where ecosystem goods and services provided by these forests may be used in a sustainable manner, contributing to local communities’ income and preserving biodiversity. In this regard, we highlight how the interaction between research and innovation together with local management may lead to sustainable development and, thus, encourage these sectors to work together for the conservation of dry forests of the Ecuadorian province.
Content may be subject to copyright.
Full Terms & Conditions of access and use can be found at
http://www.tandfonline.com/action/journalInformation?journalCode=tneo20
Download by: [Inst Medit Estudios Avanzados] Date: 15 March 2017, At: 02:37
Neotropical Biodiversity
ISSN: (Print) 2376-6808 (Online) Journal homepage: http://www.tandfonline.com/loi/tneo20
Biodiversity patterns and ecological processes
in Neotropical dry forest: the need to connect
research and management for long-term
conservation
Gema Escribano-Avila, Laura Cervera, Leonardo Ordóñez-Delgado, Andrea
Jara-Guerrero, Luis Amador, Bruno Paladines, Joahana Briceño, Violeta
Parés-Jiménez, Diego J. Lizcano, David H. Duncan & Carlos Iván Espinosa
To cite this article: Gema Escribano-Avila, Laura Cervera, Leonardo Ordóñez-Delgado, Andrea
Jara-Guerrero, Luis Amador, Bruno Paladines, Joahana Briceño, Violeta Parés-Jiménez, Diego
J. Lizcano, David H. Duncan & Carlos Iván Espinosa (2017) Biodiversity patterns and ecological
processes in Neotropical dry forest: the need to connect research and management for long-term
conservation, Neotropical Biodiversity, 3:1, 107-116
To link to this article: http://dx.doi.org/10.1080/23766808.2017.1298495
© 2017 The Author(s). Published by Informa
UK Limited, trading as Taylor & Francis
Group
Published online: 15 Mar 2017.
Submit your article to this journal
View related articles
View Crossmark data
Biodiversity patterns and ecological processes in Neotropical dry forest: the need to connect
research and management for long-term conservation
Gema Escribano-Avila
a
*, Laura Cervera
b
, Leonardo Ordóñez-Delgado
a
, Andrea Jara-Guerrero
a
,
Luis Amador
c,d
, Bruno Paladines
e
, Joahana Briceño
a
, Violeta Parés-Jiménez
b
, Diego J. Lizcano
b
,
David H. Duncan
a
and Carlos Iván Espinosa
a
a
Natural Science, Universidad Tecnica Particular de Loja, Loja, Ecuador;
b
Central Department of Research, Universidad Laica Eloy
Alfaro de Manabi, Manta, Ecuador;
c
Science Faculty, Universidad Austral de Chile, Valdivia, Chile;
d
Research Department,
Universidad Laica Vincente Rocafuerte de Guayaquil, Guayaquil, Ecuador;
e
Naturaleza y Cultura Internacional, Loja, Ecuador
(Received 26 April 2016; accepted 17 February 2017)
Neotropical dry forests are among the most diverse and threatened ecosystems worldwide. The extent and knowledge of
Neotropical dry forests are quite heterogeneous with forests located in the Ecuadorian province especially diverse, threat-
ened and poorly studied. In this work, we review patterns and conservation status of biodiversity, ecosystem processes
and human perception of tropical dry forest of the Ecuadorian province. We found that patterns of biodiversity, ende-
mism and conservation status are generally poorly studied. Overall, these forests provide habitat for at least 900 species
including trees, birds, mammals, amphibians and reptiles. On average 18% of these species (range 625%) are endemic
to the region and 25% (350%) are recognized as threatened. Little is known about groups such as invertebrates, sh,
fungi, or herbaceous plants, and as well as about processes generating and maintaining critical ecosystem functions.
Available literature points out the importance of positive ecological interactions such us plantfrugivore and plantplant
facilitation interactions in maintaining the regeneration dynamics of these forests. Faced by the formative state of
knowledge about basic biodiversity patterns and ecological functions, the implementation of ecosystem risk assessment
under the IUCN criteria for the Red List of Ecosystems may offer constructive means to organize, integrate and advance
existing knowledge and conservation priorities for dry forests of the Ecuadorian province. With examples of existing
conicts between people and protected areas, we emphasize the importance of consultation and involvement of local
communities in the development of conservation measures including new protected areas. Lastly, we reect on some
encouraging examples where ecosystem goods and services provided by these forests may be used in a sustainable
manner, contributing to local communitiesincome and preserving biodiversity. In this regard, we highlight how the
interaction between research and innovation together with local management may lead to sustainable development and,
thus, encourage these sectors to work together for the conservation of dry forests of the Ecuadorian province.
Keywords: Tumbesian region; Equatorial Pacic region; IUCN ecosystem risk assessment; endemism; sustainable
development
Introduction
Tropical dry forests (TDFs), perhaps the most threatened
terrestrial biome in the tropics [1,2], sustain a high diver-
sity of plants and animals and ecological functions that
supply a multitude of ecosystem goods and services to
local human populations [3]. These forests encompass a
variety of form and composition across three continents
[4] and have long resisted the imposition of a clear
denition [3,5]. However, two simple characteristics
dene TDFs globally: forests occurring in the tropics
(temperatures above 20º, complete absence of frosts) that
experience marked rainfall seasonality with several
months of drought (58 months) [6,7] generating a partic-
ular ecosystem structure and physiognomy that is shared
among TDFs [2,5]. For many of the same reasons, tropical
dry forests have typically coincided with areas of dense
human inhabitation and land conversion [2], which now
render this ecosystem extremely threatened [8,9].
Despite its more acute exposure to anthropogenic
threats, and great relevance for human settlements over
millennia, TDF remain poorly studied, especially in
comparison to their latitude neighbors, tropical wet and
rainforests [3,8]. From 1945 to 2004, for each research
paper published in the Web of Science portal about tropi-
cal dry forests, six were published on tropical wet forests
[10]. In a more recent review (19972014), this ratio has
slightly increased to one research article on tropical dry
forests for each 4.5 in rainforest, but still only 10% of
the research work performed in the tropics is about dry
ecosystems (Cayuela et al. unpublished results) Within
those articles concerning dry forests from the Neotropical
region, one also nds a stark imbalance. Two countries,
México and Costa Rica have attracted ~70% of the sci-
entic research (Figure 1), while other areas in Central
and South America are virtually unknown. For example,
TDFs in Bolivia are the subject of less than 5% of
*Corresponding author. Email: gema.escribano.avila@gmail.com,gescribano@imedea.uib-csic.es
© 2017 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits
unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Neotropical Biodiversity, 2017
Vol. 3, No. 1, 107116, http://dx.doi.org/10.1080/23766808.2017.1298495
research articles despite representing 25% of the overall
extension of Neotropical dry forests [10].
Around the border of Ecuador and Peru exists
another highly diverse and threatened area of tropical
dry forest, variously known as the Equatorial Pacic
Region, Tumbesian region, or Tumbes-Piura [1115].
Despite the great importance of such forests in terms of
biodiversity and endemism [11,14], these have been lar-
gely neglected in the literature on Neotropical dry forests
[2,11]. According to the above-mentioned analysis, the
area is represented by only 3% of tropical dry forest
studies (Figure 1). Part of this area is the focus of our
review. According to the most recent update on Neotrop-
ical bioregionalization, the area target of this review has
been denominated the Ecuadorian province [16].
Hereafter, we will refer to the target area and ecosystem
of this study as tropical dry forest of the Ecuadorian Pro-
vince (EP-TDF).
The EP-TDF conforms one the ecological sub-units
of the Tumbes-Chocó-Magdalena hotspot [17]. It covers
a narrow strip of coastal hinterland delimited by the
Pacic Ocean and the lower slopes of the Andes, extend-
ing over a distance of ~900 km from southeast Ecuador
to northeast of Peru (Figure 2). The TDF-EP experiences
a profound inuence of the cold and dry Humboldt cur-
rent and it is intermittently inuenced by the warm and
wet El Niño current, both determining the strong season-
ality encountered in this area [18].
The extent of the EP-TDF has been vastly deforested
to the present day, where only about 30% is thought to
remain relatively undisturbed (25 and 5% in Ecuador
and Peru respectively) and only 10% of it is under some
kind of protective land designation. These historical
facts, together with the current rate of deforestation,
510% for the period 20002012, [9,19] suggest a bleak
future for TDF-EP, unless signicant changes to land use
and development patterns actions are implemented. The
future wellbeing of the EP-TDF is tightly bound to the
livelihoods of the many people who live in and around
them. To satisfy the needs of people living in EP-TDF is
a necessary condition for the present and future conser-
vation of this ecosystem; this may be achieved only if
all stakeholders, people, managers and researches work
together.
The aim of this review is to identify gaps in knowl-
edge of biodiversity and ecosystem functioning of EP-
TDF. We also consider how the nearby human popula-
tions, historically the principal degrading agent of the
forests, currently perceive the forest and how they may
contribute to a more sustainable future. We think the
consideration of all this information together, may guide
future efforts to generate the necessary knowledge to
perform an ecosystem risk evaluation according to the
IUCN criteria for the Red List of Ecosystems. The topics
discussed herein resulted from a dynamic multidisci-
plinary workshop hosted by Universidad Técnica
Particular de Loja (Ecuador) in November 2015. The
workshop brought together researchers from various dis-
ciplines and conservationists, both concerned and enthu-
siastic about the future of the EP-TDF.
This review is articulated around several themes
dealing rst with the interaction of people and forests.
EP-TDF has been occupied and utilized by various peo-
ples for millennia [3]. Now that only 30% of the former
extent of EP-TDF still remains, the management of these
interactions is more important than ever. Researchers
should seek to generate the necessary knowledge to help
overcome such challenges [20]. More information about
the biological diversity in EP-TDF should help identify
and develop management options for conservation and
sustainable use. We discuss key advances and gaps in
knowledge about biodiversity patterns and ecological
functions and processes, and also we tackle some
possible synergies between ecological research and local
management that may contribute to the long-term conser-
vation of the EP-TDF.
Figure 1. Number of studies published in the Web of Science in the period 19002015 (search performed on December 2015) using
the words Tropical dry forestand country name*in the title, keywords and abstract. Numbers above each bar represent the
percentage of studies in each country.
108 G. Escribano-Avila et al.
Figure 2. Location of dry forest of the Ecuadorian province in Peru and Ecuador, highlighting protected areas.
Neotropical Biodiversity 109
Tropical dry forest of the Ecuadorian province,
protected areas and people
Tropical dry forests of the Ecuadorian province are
found mainly along the coastal hinterland (Figure 2).
Therein, the gentle topography, ease of access, fertile
soils and moderate climate have favored the presence
and settlement of human populations for around ve to
nine millennia [21]. The expansion of the human settle-
ments and activities such as natural resource extraction,
agriculture and extensive grazing have combined to ren-
der EP-TDF one of the most threatened ecosystems in
the world [2]. Activities involving deforestation in partic-
ular, have fragmented extant habitat directly impacting
species richness and biodiversity [22,23].
Ofcial protection of remnant EP-TDF habitat has
progressed through a variety of regional, national and
internationally recognized reserves. In Peru, ~10% of
remnant dry forest (2337 km
2
) is protected by means of
six protected areas (Figure 2), most of them included in
the UNESCO Northwest Biosphere Reserve (NBR),
which covers an area of 231 km
2
. In Ecuador, dry forests
occupy more than 6000 km
2
[9]. Until 2014, only 24%
was protected in conservation reserves [9,24]; however,
this percentage recently rose to an estimated 8% of the
extant tropical dry forests, following the declaration of
Bosque Seco Biosphere Reserve (BSBR). The BSBR
declaration was facilitated by the efforts of various
municipal authorities to dene zones within their juris-
diction critical for conservation and ecosystem connectiv-
ity. The Municipal Reserves that resulted from the
zoning process today largely comprise the core area of
the BSBR. The recognition of the conservation value
and objectives, and the legal status established by the
declaration were positive and necessary steps, and the
establishment of protected areas has contributed to slow
down deforestation rates. However, the designation of
protected areas does not in itself assure long-term biodi-
versity conservation (reviewed in [25]). The management
of protected areas in developing countries is often com-
plex because of poverty, population growth and the lack
of resources available for effectively protecting those
areas [25].
Machalilla National Park, the only Ecuadorian
national park that includes dry forests, exemplies the
conicting interests of local communities and national
park management objectives. The park was created in
1979 without much consultation with local communities.
Consequently, illegal activities have continued to occur
in the park in the face of decient law enforcement [26,
Cervera et al. unpublished results]. Engaging local com-
munities in environmental policies is vital for the success
of conservation initiatives [27,28]. A good example of
consultation and involvement can be found in the rem-
nant dry forests around Zapotillo county (Loja Province)
in the south of Ecuador. In 2000, Nature and Culture
International (NCI) began working to create a network
of private ecological reserves in collaboration with local
communities. In this area, owing to the remoteness and
climate, the dry forests had not been cleared like in most
other areas in the country. With virtually no infrastruc-
ture present, and a climate hostile to agricultural use, the
area consisted of a few owners of vast haciendas, whose
principal activity of extensive goat and cow farming
depended on the dry forest ecosystem for stock feed and
shelter. Two co-managed reserves of 18 km
2
La Ceiba,
and Cazaderos, were established and now form part of
the BSBR. Importantly, local people contributed to the
establishment of protection policies and regulations for
forest use. Given that farmers make a living from goats
and the animals in turn forage in the forest, farmers
easily realized the forest needed to be protected to assure
their livelihood [Nature and Culture International,
https://natureandculture.org/]
The perceptions of people living nearby La Ceiba,
one of the co-managed reserves in Zapotillo, were
recently compared with those of people living near Are-
nillas Ecological Reserve [29]. Arenillas Ecological
Reserve (REA) was a military base and exclusion zone
since the 1970s due to its strategic location on the Peru
border. In 2001, responsibility for this area was trans-
ferred to the environmental ministry and the ecological
reserve was established. The perception of the protected
areas was usually positive in the case of La Ceiba (pro-
vision of ecosystem services, economic income by tour-
ism) and negative in the case of Arenillas Ecological
Reserve (restrictions of use, indifference) [29]. Like
Machalilla National Park, it seems that the people around
REA lack of sense of attachment and ownership of the
objectives of the protected area. Precisely these elements
seem critical in the achievement of conservation and sus-
tainable management goals. Given that dry forests are a
highly threatened ecosystem in the region, and are
under-represented in protected areas at the global scale,
additional forest reserves may be required to establish a
more robust conservation network, both from govern-
ment and local initiative, such as the examples of the
Zapotillo area and the Dry Forest Biosphere Reserve.
However, given the tight relationship between human
settlements, livelihoods and the dry forests, initiatives to
identify and declare new reserves should follow a gen-
uine process of consultation of local stakeholders, to
assure that the parks may contribute to effective and last-
ing conservation of the ecosystem [27,28].
Biodiversity patterns in the dry forests of the
Ecuadorian province: what do we know?
To review the existing studies about biodiversity in the
region, we performed a search in the Web of Science
portal, with the key words tropical dry forestAND
Ecuador* OR Peru*. The search (performed 18 April
2016) yielded 30 records, of which most were about
plants (60%), being almost 50% of those articles about
woody plants. The remaining articles were about birds
110 G. Escribano-Avila et al.
(17%), mammals (13%), insects (10%) and reptiles (3%).
According to this evidence, further research on the biodi-
versity patterns in the EP-TDF region is required for all
groups, but particularly for herpetofauna, sh, fungi and
invertebrates, which would be expected to be the most
species rich of all. Here, we provide some insights about
the current knowledge on species numbers, endemism
and conservation status for groups of woody plants,
birds, mammals, reptiles and amphibians.
Overall, EP-TDF is home of at least ~900 species of
trees, birds, mammals, amphibians and reptiles with an
average of 18% being endemic (range 256%) as shown
in Table 1. This is a high level of endemism similar to
that reported for the whole Tumbes-Chocó-Magdalena
region and other Neotropical biodiversity hotspots [11].
Around 30% of these endemic species are under some
threatened IUCN category (Table 1). Amongst tree spe-
cies, the richest taxonomic group of those well docu-
mented, 20% of the species are endemic to EP-TDF
[13,30]. There are 318 recorded tree species belonging to
180 genera and 54 families. Five families (Leguminosae,
Malvaceae, Boraginaceae, Cactaceae and Moraceae),
constitute around 40% of all recorded species; of those,
the Leguminosae alone accounts for 22% of species.
Most families, however, are represented by a few species
and 13 families by a single species. Woody species rich-
ness, considering trees and shrubs (DBH 5 cm), is
around 600 trees ha
1
(538625), while when including
individuals over 10 cm of DBH (thus probably excluding
shrubs) density values become lower and much variable
with a range of 22524 trees ha
1
(Table 2).
Birds are the second richest taxonomic group with an
estimated 250300 species. Tropical dry forest of the
Ecuadorian Province (in the case of birds, better known
as Tumbesian region) sustains one of the highest
concentrations of species with restricted distribution in
South America [14] with 21% of species endemic to the
region, and 30% under some IUCN threatened category
[24,3133], (Table 1). Although with much lower levels
of endemism, the mammal community is represented by
200 species, with 12 considered endemic [34,35]. Two
squirrel species deserve special mention in this regard,
the Guayaquil squirrel (Simosciurus stramineus), which
is endemic to the coast of Ecuador and distributed from
Esmeraldas to Guayas, and the white-napped squirrel
(Simosciurus nebouxii), which is found in the dry forests
of Arenillas, El Oro and north Piura, in Peru [36].
Amphibians and reptiles are also important compo-
nents of biodiversity in EP-TDF, contributing almost 100
species of which around 20% are endemic (see Table 1).
For reptiles, most species are snakes and lizards, one
species of blind snake, Amphisbaena occidentalis, and
one turtle (Rhinoclemmys annulata)[
37]. The conserva-
tion status of the herpetofauna, especially reptiles,
remains uncertain as the taxonomy is not well resolved.
For instance, the circumscription of several species of
snakes and lizards has recently been revised following
phylogenetic studies [38]. This is the case of the species
Lampropeltis micropholis,Erythrolamprus epinephelus,
and Alsophis elegans [39,40]. Taxonomic change is
rather common for tropical dry forest species, not just in
the case of the herpetofauna [41], but also for other
groups such as birds and mammals [42,43].
Aside from taxonomic revisions, the estimated ranges
of fauna from EP-TDF are also frequently modied as
eld studies continue to produce new records. Some of
these recent examples are new records for the lizard
Macropholidus ruthveni [44] and the Tumbes tyrant
Tumbezia salvini [45] both recently conrmed in the
Zapotillo area, Province of Loja in southeastern Ecuador.
These examples highlight that more research is required
to understand patterns of biodiversity in these forests,
and to understand the effects of deforestation and anthro-
pogenic activities on the distribution and species conser-
vation status [46]. As more work is conducted, often
with improved techniques to study the distribution and
abundance of species (especially cryptic and elusive
ones), it is to be expected that changes in the conserva-
tion status will arise. A better understanding of both tax-
onomy and conservation status is required, considering
that Neotropical dry forests constitute a hotspot of defau-
nation. A global phenomenon that is especially worri-
some in the tropics [47].
Ecosystem function and risk in tropical dry forests of
the Ecuadorian province: a gap of knowledge
Given the immature state of knowledge of some individ-
ual taxa in EP-TDF, and the urgency of their conserva-
tion, a dual focus on improving species knowledge, and
ecosystem level function and risk, should be pursued.
Based on the policy success of the IUCN red list of spe-
cies, an international movement to develop criteria for
the recognition of globally threatened ecosystems was
initiated in the early 2000s to create the IUCN Red List
of Ecosystems [48]. Importantly, an ecosystem level
approach underlines the fundamental structural and engi-
neering role that common species play in the mainte-
nance of ecosystems and the functions that they generate
[49], as well as gathering essential information about the
distribution and status of rare and threatened species. In
recent years, the ecosystem risk assessment criteria and
method of application have been improved [50,51] and
could now be considered for application in EP-TDF.
These criteria include: rates of ecosystem loss and
Table 1. Number of endemic species and their percentages.
Taxa Nº species Endemic (%) IUCN threaten (%)
Trees 318 67 (21) 11
Birds 250300 59 (21) 34
Mammals 200 12 (6) 33
Reptiles 64 12 (19) 17
Amphibians 32 8 (25) 38
Neotropical Biodiversity 111
decline at the distribution range level, restricted
distributions continuously declining or threatened, envi-
ronmental degradation, degradation of biotic processes
(e.g. mutualistic interactions) and a quantitative risk of
ecosystem collapse as integrative indicator of all the
above mentioned.
Recent work has established that from the processes
involved in forest regeneration, those more vulnerable to
human impact are pollination and seed dispersal medi-
ated by mutualistic interactions between plants and ani-
mals [52]. The disruption of mutualisms is considered as
a key indicator of the IUCN criteria biotic processes
degradation[50,51]. Accordingly, in this section we
highlight some insights on key ecological interactions,
such as endozoochorus seed dispersal and plant regenera-
tion dynamics, that may aid in the development of a risk
assessment approach for the ecosystem.
In spite of not being the most abundant dispersal
syndrome, zoochorous species account for around 50%
of woody plants in Neotropical dry forests [53,54], espe-
cially in well preserved patches [55]. This is also the
case of EP-TDF, where up to 54% of woody species pre-
sent a zoochorous dispersal syndrome [54]. Following
disturbance, zoochorous species tend to be the last ones
to recover [56], leading to impoverished plant communi-
ties and also reduced food resources for animals and less
nutrient supply to soils [57]. Despite this, relatively little
is currently known about seed dispersal performed by
animals in EP-TDF. A search of the subject performed in
the Web of Science portal (on 19 November 2015) with
the terms (frugivoryAND tropical dry forest)OR
(endozoochoryAND tropical dry forest)OR
(epizoochoryAND tropical dry forest) lead to 18
publications. Most were about primates, birds and bats,
with very few about reptiles or herbivorous mammals.
Surprisingly, there were no studies dealing with carnivo-
rous mammals as seed dispersers, despite the recognized
importance of this group as seed dispersal agents
(reviewed in [58]). An investigation of the importance of
the last herbivore-megafauna inhabiting EP-TDF, the
white tailed deer (Odocoileus virginianus peruvianus)
and also a mesocarnivore, the sechuran fox (Lycalopex
sechurae Thomas, 1900) is currently underway (Jara-
Guerrero et al., Escribano-Avila et al. unpublished data).
These two mammals, seem to be important dispersers as
they consume and deposit seeds of at least 15 woody
species, most of which have been shown to be viable
after passing through deer and fox gut. Birds, primates,
carnivores and bats are known as relevant dispersers in
mostly all kinds of ecosystems with eshy fruited spe-
cies. However, recent ndings on dispersal patterns in
other Neotropical dry forest areas highlight the relevance
of non-typical dispersers, such as iguanas [59,60] and
small terrestrial mammals [61,62]. Such work leads to
the expectation that these groups of animals may be also
relevant in the dispersal process of TDF-EC and thereby
warrant further research.
Once a seed is dispersed, many processes act to
inuence germination, early recruitment and further
stages until recruits reach reproductive adulthood [63].
Plant-plant interactions may play a key role in determin-
ing the spatial structure and diversity of plant communi-
ties in the EP dry forests [11]. Various authors propose
that certain individuals or species may facilitate the
recruitment of other species by modifying the surround-
ing micro-environmental conditions (e.g. [64,65]). This
hypothesis was tested in an Ecuadorian dry forest by
means of evaluating the behavior of species in relation
to their ability to attract or repel diversity, considered as
a facilitation or competition effect respectively. At short
distances (10 m), about 30% of species behaved as
accumulators of diversity, or more still (over 50%), when
the relationship was assessed between juveniles [66].
Some ongoing work in dry scrublands of the Ecuadorian
Province show higher recruitment success under the
fringe of the canopy of the dominant shrub compared
with open areas. Furthermore, the relationship was modi-
ed by environmental stress; seedlings were more depen-
dent on facilitation in more stressful habitats (Quintana
et al. unpublished data). Although this demonstration
pertains to scrubland rather than forest, and should be
further developed to test generality, these ndings
suggest that positive interactions between plants may be
relevant in the maintenance of biodiversity and regenera-
tion dynamics in EP-TDF. A greater understanding of
these dynamics and interactions in the dry forest ecosys-
tem should prove useful to guide management and strate-
gic documents, such as restoration plans.
Table 2. Diversity and density typical values of the dry forests of the Ecuadorian province. Note the minimum diameter threshold.
DBH
(cm)
Density trees
(Ha
1
)
Richness Spp
(Ha
1
)
Plot size
(Ha)
N
plots Country Locality Ref.
5 625 36 0,05 109 EC Zapotillo [69]
5 538 37 1 1 EC Machalilla NP [70]
5 588 4 9 1 EC Arenillas Ecol. Reserv. [73]
10 215 25 1 1 EC Machalilla NP [70]
10 22524 625 1 6 PE Cerros de Amotape
NP
[71]
10 31399 837 1 10 PE Cerros de Amotape
NP
[72]
112 G. Escribano-Avila et al.
Future challenges
Towards bi-national protection and management of the
tropical dry forest of the Ecuadorian province
Tropical dry forests of the Ecuadorian Province extend
over Ecuador and Peru. Whilst both countries have
established protected areas to preserve the forest, better
coordination could improve the connectivity of those
reserves, and consider the management of strategic areas
in the matrix between them. Both nations have declared
their intent to establish a Bi-National UNESCO reserve
that would protect 17000 km
2
including dry forests and
associated ecosystems (i.e. mangroves and wetlands).
This will allow the coordination of conservation aims
and management procedures. One of the proposed targets
of this bi-national reserve is to support the development
of a conservation culture, whereby local communities
may generate economic activities, such as ecotourism
(e.g. birdwatching, trekking or bicycle tours to enjoy the
Guayacan (Tabebuia spp) blooming). These options will
lead to widen the sources of economic income in the
local region and, at the same time, promote the conserva-
tion of dry forests. This initiative is already supported by
both countries and has been recently presented in the 4th
World Congress of Biosphere Reserves-UNESCO, held
in Lima, Peru on March 2016. We suggest that the work
being performed in this regard may be useful and inspir-
ing for both nations to collaborate towards an ecosystem
risk assessment process for EP-TDF. The process of doc-
umentation, and analysis of risks and opportunities for
the forests, should foster improved international coordi-
nation and collaboration in designing a sustainable future
for the region and its people.
Joint projects of research, innovation and management
may lead to achieve sustainable development in
equatorial pacic dry forests
Protected areas are a pivotal instrument toward the con-
servation of EP-TDF, however human use of these for-
ests, even in protected areas, is the rule rather than the
exception. Thus, it is necessary to seek compromises that
permit sustainable development of economic activities by
local communities alongside biodiversity conservation.
Research and innovation may be of great help in this
regard. A good example is the Palo Santo project (this is
the common name given in Ecuador to Bursera grave-
olens (Burseraceae)), which proposes to manage one of
the most widespread and dominant trees in EP-TDF. A
valuable essential oil is obtained from B. graveolens,
typically from the trunk, and thus the exploitation of this
economic resource is at the expense of forest conserva-
tion. The chemistry department of Universidad Técnica
Particular de Loja (UTPL) developed a methodology to
extract the essential oil from the fruits instead of the
trunk; this was the rst step of the Palo Santo project
that brought together local communities, government
institutions and local NGOs, such as the aforementioned
NCI. The Palo Santo project resulted in the commercial-
ization of B. graveolens essential oil to the Brazilian
environmental enterprise Natura used in their natural
cosmetic products. Most project management is carried
out by the community association Bolívar Tello Cano.
This is probably the most remarkable example of how
the natural resources of dry forests may be used in a sus-
tainable manner, in addition to create social networking
and support local economies. Actually, in 2014 the
Bolívar Tello Cano community association was awarded
the United Nations Development Programmes Equator
Prize, in recognition of the Palo Santo projects
outstanding effectiveness in reducing poverty through
biodiversity conservation and sustainable business
practices [67].
Researchers could seek to play a similarly creative
and catalytic role in the development of sustainable
initiatives from the products and management of sting-
less native melaponid bees (Melipona spp). Ancestral
knowledge of the management of indigenous Melipona
bees has almost been lost due to habitat deterioration
and the introduction of the European bee (Apis
melifera). However, there are still some areas in which
meliponeculture still remains active, and these examples
may serve as a base to recover and expand this ances-
tral knowledge and the resources that Melipona bees
may provide. Researchers working together with local
people could assist in recovering and codifying ances-
tral knowledge for its dissemination to other interested
local communities, and at the same time increase scien-
tic information on the biology and conservation of
Melipona bees [68].
Concluding remarks
Tropical dry forests of the Ecuadorian Region are home
of at least 900 species of trees, birds, mammals, amphib-
ians and reptiles. Around one in ve of these species are
endemic to the region, and many of them are threatened
with extinction. These numbers, already startling, are
likely to be conservative, as much remains to be learned
about several groups such as mammals, sh, fungi and
especially invertebrates, the most diverse living group in
the world. It is possible that many species may go
extinct before being discovered owing to the ongoing
deforestation and fragmentation condition of EP-TDF.
However, the growing number of protected areas with a
strong focus on co-management with local people
together with initiatives of sustainable development are
encouraging and signaling the way forward. The IUCN
process for the Red Listing of Ecosystems should serve
as inspiration and direction for targeted research in EP-
TDF. This research should address identied gaps in
knowledge required for conservation, but also contribute
to the development of sustainable options for local
communities that are required to facilitate the long term
conservation of EP-TDF.
Neotropical Biodiversity 113
Author contributions
Gema Escribano-Ávila designed, led the manuscript and
performed most of scientic literature review; David
Duncan provided guidance with the manuscript structure,
content section and English language; Laura García
Cervera contributed in art work. All authors actively
collaborated in the literature revision of their eld of
expertise and manuscript writing (People-Forest interac-
tions: Joahana Briceoño, Bruno Paladines; Mammal
patterns: Laura García Cervera and Diego Lizcano; Bird
patterns: Leonardo Ordoñez; Herpetofauna patterns:
Luis Amador; Plant patterns and seed dispersal: Andrea
Jara-Guerrero, Carlos Iván Espinosa, Gema Escribano-
Ávila and Violeta Parés-Jimenez). All authors critically
reviewed the manuscript.
Associate Editor: Elisa Bonaccorso.
Disclosure Statement
No potential conict of interest was reported by the
authors.
Funding
This work was supported by the Research Program of
Universidad Técnica Particular de Loja through the project
CCNN-1054. Gema Escribano-Ávila was supported by the
Prometeo Project of the Secretary for Higher Education,
Science, Technology and Innovation of Ecuador (SENESCYT).
ORCID
Gema Escribano-Avila http://orcid.org/0000-0003-1362-8924
Leonardo Ordóñez-Delgado http://orcid.org/0000-0002-
4593-1728
Andrea Jara-Guerrero http://orcid.org/0000-0001-9394-2790
Luis Amador http://orcid.org/0000-0003-2638-4068
Joahana Briceño http://orcid.org/0000-0002-3056-4734
Violeta Parés-Jiménez http://orcid.org/0000-0001-5346-6566
Diego J. Lizcano http://orcid.org/0000-0002-9648-0576
David H. Duncan http://orcid.org/0000-0003-4411-8214
Carlos Iván Espinosa http://orcid.org/0000-0002-5330-4505
References
[1] Hoekstra J, Boucher T, Ricketts T, et al. Confronting a
biome crisis: global disparities of habitat loss and protec-
tion. Ecol Lett. 2005;8:2329.
[2] Miles L, Newton AC, DeFries RS, et al. A global over-
view of the conservation status of tropical dry forests. J
Biogeogr. 2006;33:491505.
[3] Murphy PG, Lugo AE. Ecology of tropical dry forest.
Annu Rev Ecol Evol Syst. 1986;17:6788.
[4] Pennington RT, Lavin M, Oliveira-Filho A. Woody plant
diversity, evolution, and ecology in the tropics: perspec-
tives from seasonally dry tropical forests. Annu Rev Ecol
Evol Syst. 2009; 40: 437457.
[5] Dexter KG, Smart B, Baldauf C, et al. Floristics and
biogeography of vegetation in seasonally dry tropical
regions. Int For Rev. 2015;17(S2):1032. DOI:10.1505/
146554815815834859
[6] Mooney HA, Bullock SH, Medina E. Introduction. In:
Bullock SH, Mooney HA, Medina E, editors. Seasonally
dry tropical forests. Cambridge: Cambridge University
Press; 1995.p.18.
[7] FAO. Global ecological zones for FAO forest reporting:
2010 update. Food and agriculture organization of the Uni-
ted Nations Forest resources assessment working paper
179. [cited 2016 Apr 20]. Available from: http://www.fao.
org/docrep/017/ap861e/ap861e00.pdf
[8] Janzen DH. Management of Habitat fragments in a
tropical dry forest: growth. Ann Mo Bot Gard. 1988;75:
105116.
[9] Portillo-Quintero CA, Sánchez-Azofeifa GA. Extent and
conservation of tropical dry forests in the Americas. Biol
Cons. 2010;143:144155.
[10] Sánchez-Azofeifa GA, Quesada M, Rodríguez JP, et al.
Research priorities for Neotropical dry forests. Biotropica.
2005;37:477485.
[11] Espinosa CI, De la Cruz M, Luzuriaga AL, et al. Bosques
tropicales secos de la región Pacíco Ecuatorial: diversi-
dad, estructura, funcionamiento e implicaciones para la
conservación. Ecosistemas. 2012;21:167179.
[12] Peralvo M, Sierra R, Young KR, et al. Identication of bio-
diversity conservation priorities using predictive modeling:
an application for the Equatorial Pacic region of South
America. Biodivers Conserv. 2007;16:26492675.
[13] Linares-Palomino R, Kvist LP, Aguirre-Mendoza Z, et al.
Diversity and endemism of woody plant species in the
Equatorial Pacic seasonally dry forests. Biodivers Con-
serv. 2010;19:169185.
[14] Best B, Kessler M. Biodiversity and conservation in
Tumbesian Ecuador and Peru (Vol. 218). Cambridge:
BirdLife International; 1995.
[15] Dinerstein E, Olson DM, Gram DJ, et al. Una Evaluación
del estado de conservación de las eco-regiones de America
Latina y Caribe. Washington, DC: Banco Internacional de
Reconstrucción y Fomento/Banco Mundial; 2011.
[16] Morrone JJ. Biogeographical regionalisation of the
Neotropical region. Zootaxa. 2014;3782(1):1110.
[17] Mittermeier RA, Turner WR, Larsen FW, et al. Global
biodiversity conservation: the critical role of hotspots in
biodiversity hotspots. Berlin: Springer; 2011.p.322.
[18] International Conservation. Biological diversity in
Tumbes-Chocó-Magdalena. 2011. [cited 2015 Dec 4].
http://www.eoearth.org/view/article/150631/
[19] Hansen MC, Potapov PV, Moore R, et al. High-resolution
global maps of 21st-century forest cover change. Science.
2013;342:850853.
[20] Sanchez-Azofeifa GA, Kalacska M, Quesada M, et al.
Need for integrated research for a sustainable future in
tropical dry forests. Conserv Biol. 2005;19:285286.
DOI:10.1111/j.1523-1739.2005.s01_1.x
[21] Piperno DR, Dillehay TD. Starch grains on human teeth
reveal early broad crop diet in northern Peru. Proc Natl
Acad Sci USA. 2008;105:1962219627.
[22] Gillespie TW, ONeill K, Keppel G, et al. Prioritizing con-
servation of tropical dry forests in the Pacic. Oryx.
2014;48:337344.
[23] Tapia-Armijos MF, Homeier J, Espinosa CI, et al. Deforesta-
tion and forest fragmentation in south Ecuador since the
1970s losing a hotspot of biodiversity. PLoS ONE.
2015;10(9):e0133701. DOI:10.1371/journal.pone.0133701
[24] Albuja L, Almendáriz R, Barriga LD, et al. Fauna de Ver-
tebrados del Ecuador. Quito: Escuela Politécnica Nacional,
Instituto de Ciencias Biológicas; 2012.
[25] Naughton-Treves L, Holland MB, Brandon K. The role of
protected areas in conserving biodiversity and sustaining
local livelihoods. Annu Rev Environ Resour. 2005;30:
219252.
114 G. Escribano-Avila et al.
[26] Fiallo EA, Jacobson SK. Local communities and protected
areas: attitudes of rural residents towards conservation and
Machalilla National Park, Ecuador. Environ Conserv.
1995;22:241249.
[27] Sheppard SR. Participatory decision support for sustain-
able forest management: a framework for planning with
local communities at the landscape level in Canada. Can J
For Res. 2005;35:15151526.
[28] Brooks JS, Waylen KA, Mulder MB. How national con-
text, project design, and local community characteristics
inuence success in community-based conservation pro-
jects. Proc Nat Acad Sci USA. 2012;109:2126521270.
[29] Briceño J, Iniguez-Gallador V, Ravera F. Factores que
inuyen en la apreciación de servicios eco-sistémicos de
los bosques secos del sur del Ecuador. Ecosistemas.
2016;25(2):000000.
[30] Leal-Pinedo JM, Linares-Palomino R. Los bosques secos
de la Reserva de Biosfera del Noroeste (Perú): diversidad
arbórea y estado de conservación. Caldasia. 2005;27(2):
195211.
[31] Freile JF, Carrión JM, Prieto-Albuja F, et al. La orni-
tología en Ecuador: un análisis del estado actual del cono-
cimiento y sugerencias para prioridades de investigación.
Ornitología Neotropical. 2006;17:183202.
[32] Ridgely R, Greeneld P. Aves del Ecuador. Volumen II.
Guía de Campo. Academia de Ciencias Naturales de Fila-
dela. Fundación de Conservación. Quito: Jocotoco; 2006.
[33] BirdLife International. Endemic Bird Area factsheet:
Tumbesian region. 2016. [cited 11/02/2016. Available
from: http://www.birdlife.org
[34] Pacheco V, Cadenillas R, Salas E, et al. Diversidad y
endemismo de los mamíferos del Perú. Revista peruana de
biología. 2009;16:532.
[35] Albuja VLH. Lista de mamíferos actuales del Ecuador.
Quito: Escuela Politécnica Nacional; 2011.
[36] de Vivo M, Carmignotto AP. Family Sciuridae G. Fischer,
1817. In: Patton JL, Pardiñas UFJ, DElía G, editors,
Mammals of South America, Volume 2: Rodents Vol. 2.
Chicago: University of Chicago Press; 2015.
[37] Venegas PJ. Herpetofauna del bosque seco ecuatorial de
Perú: Taxonomía, Ecología y Biogeografía. Zona Áridas.
2005;9:926.
[38] Pyron RA, Burbrink FT, Wiens JJ. A phylogeny and
revised classication of Squamata, including 4161 species
of lizards and snakes. BMC Evol Biol. 2013;13:253.
DOI:10.1186/1471-2148-13-93
[39] Ruane S, Bryson RW, Pyron RA, et al. Coalescent species
delimitation in milksnakes (Genus Lampropeltis) and
impacts on phylogenetic comparative analyses. Syst Biol.
2014;63:231250.
[40] Zaher H, Gobbi-Grazziotin F, Cadle JE, et al. Molecular
phylogeny of advanced snakes (Serpentes, Caenophidia)
with an emphasis on South American Xenodontines a
revised classication and descriptions of new taxa. Papéis
Avulsos de Zoologia. 2009;49:115153.
[41] Duellman WE, Barley AJ, Venegas PJ. Cryptic species
diversity in marsupial frogs (Anura: Hemiphractidae:
Gastrotheca) in the Andes of northern Peru. Zootaxa.
2014;3768:159177.
[42] Schulenberg TS, Stotz DF, Lane DF, et al. Birds of Peru.
Revised edition. Princeton, NJ: Princeton University Press;
2010.
[43] Molina M, Molinari J. Taxonomy of Venezuelan white-
tailed deer (Odocoileus, Cervidae, Mammalia), based on
cranial and mandibular traits. Canadian J Zool. 1999;77:
632645.
[44] Torres-Carvajal O, Gaona FP, Zaragoza C, et al. First
record of Macropholidus ruthveni Noble 1921 (Squamata:
Gymnophthalmidae) from Ecuador. Herpetol Notes. 1921;
2015(8):2526.
[45] Ordoñez-Delgado L, Tomas G, Espinosa CI. Nueva locali-
dad del Tirano de Tumbes Tumbezia salvini (Aves: Tyran-
nidae) en el suroeste del Ecuador. Avances en Ciencias e
Ingenierias. 2016;8:14.
[46] Loaiza CR. The Tumbesian center of endemism: biogeog-
raphy, diversity and conservation. Biogeografía. 2013;6:4.
[47] Dirzo R, Young HS, Galetti M, et al. Defaunation in the
Anthropocene. Science. 2014;345:401406.
[48] Rodriguez JP, Rodriguez-Clark K, David AK, et al. IUCN
Red List of Ecosystems, S.A.P.I.EN.S [Online], 5.2 |
2012, Online since 2012 Aug 12. Available from: http://
sapiens.revues.org/1286
[49] Keith DA. Assessing and managing risks to ecosystem
biodiversity. Aust Ecol. 2015;40:337346.
[50] Keith DA, Rodríguez JP, Rodríguez-Clark KM, et al. Sci-
entic foundations for an IUCN red list of ecosystems.
PLoS ONE. 2013;8:e62111.
[51] Rodriguez JP, Keith DA, Rodriguez-Clark, KM, et al. A
practical guide to the application of the IUCN red list of
ecosystems criteria. Philos Trans R Soc Lond B Biol Sci.
2015;370:20140003.
[52] Neuschulz EL, Mueller T, Schleuning M, et al. Pollination
and seed dispersal are the most threatened processes of
plant regeneration. Sci Rep. 2016;6:29839.
[53] López-Martínez JO, Sanaphre-Villanueva L, Dupuy JM,
et al. β-diversity of functional groups of Woody plants in
a tropical dry forest in Yucatan. PLoS ONE. 2013;8:
e73660.
[54] Hilje B, Calvo-Alvarado J, Jiménez-Rodríguez C, et al.
Tree species composition, breeding systems, and pollina-
tion and dispersal syndromes in three forest successional
stages in a tropical dry forest in Mesoamerica. Trop Con-
serv Science. 2015;8:7694.
[55] Jara-Guerrero A, De la Cruz M, Méndez M. Seed disper-
sal spectrum of woody species in south Ecuadorian dry
forests: environmental correlates and the effect of consid-
ering species abundance. Biotropica. 2011;43:722730.
[56] Ceccon E, Hernández P. Seed rain dynamics following
disturbance exclusion in a secondary tropical dry forest in
Morelos, Mexico. Revista de Biología Tropical. 2009;
57:257269.
[57] Patricia L, Morellato C. Nutrient cycling in two south-east
Brazilian forests. Litterfall and litter standing crop. Revista
de Biología Tropical. 1992;8:205215.
[58] Escribano-Ávila G, Pías Couso B, Escudero Alcántara A,
et al. Ecological relevance of frugivorous mammals in the
regeneration dynamics of old elds in Mediterranean envi-
ronments. Ecosistemas. 2015;24:3542.
[59] Lasso E, Barrientos LS. Epizoochory in dry forest Green
iguana: an overlooked seed dispersal mechanism? Colom-
bia Forestal. 2015;18:151159.
[60] Traveset A, Nogales M, Vargas P, et al. Galápagos land
iguana (Conolophus subcristatus) as a seed disperser.
Integr Zool. 2016;11:207213.
[61] Lessa LG, Geise L, Costa FN. Effects of gut passage on
the germination of seeds ingested by didelphid marsupials
in a Neotropical savanna. Acta Botanica Brasilica.
2013;27:519525.
[62] Genrich CM, Mello MA, Silveira FA, et al. Duality of
interaction outcomes in a plant-frugivore multilayer net-
work. Oikos. 2016. DOI:10.1111/oik.03825
[63] Harper JL. Population biology of plants. London:
Academic Press; 1977.
[64] Espinosa CI, Luzuriaga AL, de la Cruz M, et al. Climate
and grazing control nurse effects in an Ecuadorian dry
shrubby community. J Trop Ecol. 2013;30:2332.
Neotropical Biodiversity 115
[65] Brooker RW, Maestre FT, Callaway RM, et al. Facilitation
in plant communities: the past, the present, and the future.
J Ecol. 2008;96:1834.
[66] Espinosa CI, de la Cruz M, Jara-Guerrero A, et al. The
effects of individual tree species on species diversity in a
tropical dry forest change throughout ontogeny. Ecogra-
phy. 2016;39:329337. DOI:10.1111/ecog.01328
[67] Nature and Culture. 2014. [cited 2016 Apr 20]. https://na
tureandculture.org/places/ecuador/the-palo-santo-project
[68] Martínez-Fortún MS. Desarrollo sostenible y conservación
etnoecológica a través de la meliponicultura, en el sur de
Ecuador. Tesis de Maester. Universidad Internacional de
Andalucía; 2015.
[69] Espinosa CI, Cabrera O, Luzuriaga AL, Escudero A. What
Factors Affect Diversity and Species Composition of
Endangered Tumbesian Dry Forests in Southern Ecuador?
Biotropica. 2011;43(1):1522. 2
[70] Josse C, Balslev H. The composition and structure of a
dry, semideciduous forest in western Ecuador. Nordic
Journal of Botany. 1994;14(4):425434.
[71] Linares-Palomino R, Ponce-Alvarez SI. Tree community
patterns in seasonally dry tropical forests in the Cerros de
Amotape Cordillera, Tumbes. Peru. Forest Ecology and
Management. 2005;209:261272.
[72] Leal-Pinedo JM, Linares-Palomino R. Los bosques secos
de la Reserva de Biosfera del Noroeste (Perú): Diversidad
arbórea y estado de conservación. Caldasia. 2005;27(2):
195211.
[73] Jara-Guerrero A, De la Cruz M, Espinosa CI, Méndez M,
Escudero A. Does spatial heterogeneity blur the signature
of dispersal syndromes on spatial patterns of woody
species? A test in a tropical dry forest. Oikos. 2015;124
(10):13601366.
116 G. Escribano-Avila et al.
... Seasonally dry forests in the Pacific-Equatorial exhibit reduced species diversity, but have a pronounced degree of endemism, notably among vertebrates and flora (Chvala et al. 1972;Cracraft 1985;Cueva-Ortiz et al. 2019). It is estimated that one in every five species within these ecosystems is endemic (Escribano-Avila et al. 2017). Furthermore, distinct sections of this region are recognized as one of the 25 global biodiversity hotspots. ...
... In this biogeographic zone, few studies have been conducted on the richness, distribution, and endemism of various groups, especially arthropods, including horseflies (Buestán 1980;Lattke et al. 2016;Escribano-Avila et al. 2017;Padrón et al. 2023). This implies that our understanding of the biological diversity of one of the most medically and ecologically relevant groups of blood-feeding insects in one of the least studied and highly threatened biogeographic zones in Ecuador and South America is limited. ...
Article
The Equatorial Pacific Region (EPR) in Ecuador is characterized by high rates of vegetation diversity, and rapid loss of vegetation cover due to anthropogenic pressures. In this study, general ecological aspects of the Tabanidae family, including richness, endemism, and seasonality, were evaluated. Analyses reveal that approximately 42% of the species recorded for Ecuador are present in the EPR, and out of the 84 species cataloged in the EPR, 6 are endemic, representing an endemism of 7.14%. Furthermore, it was established that tabanid populations in a coastal dry forest significantly increased their population density during the dry season, while decreasing during the wet season.
... It is estimated that only 12.6% of TDF are protected worldwide, well below other types of forest such as the temperate broadleaf evergreen forest (34.2%), the tropical montane forest (26.1%) and the tropical semi-evergreen moist broadleaf forest (26.4%), and this protection is below the global forest average of 15.9% (Schmitt et al. 2009). It is also estimated that for each paper related to TDF that is published, there are 4.5 on evergreen forests, signifying that only 10% of the total scientific production related to tropical forests concerns TDF (Escribano-Avila et al. 2017), thus making it the least studied ecosystem (Blackie et al. 2014). ...
... It is, therefore, necessary to study the fragmentation and connectivity of SATDF (Escribano-Avila et al. 2017). The aim of this study was to assess changes in the fragmentation and connectivity of SATDF between 1992 and 2020 with the following specific objectives: (i) to assess changes in dry forest cover in South America at country scales; (ii) to evaluate changes in fragmentation patterns during three different periods (1992-2000, 2000-2010 and 2010-2020), and (iii) to study the evolution of the functional and structural connectivity of SATDF. ...
Article
Full-text available
Tropical dry forests are the most threatened of all the major tropical forest types and less than 25% of TDF currently remain worldwide. They are located mostly in South America. Parameters such as habitat loss, fragmentation and loss of connectivity have been identified as the main threats to biodiversity. This research aims to discover the forests changes, the evolution of connectivity and fragmentation of the South American tropical dry forest between 1992 and 2020. Land uses layers provided by the Copernicus program were employed, and land uses change, fragmentation and structural connectivity were analyzed in GIS systems. To calculate the functional connectivity, the Graphab software was used. The results showed a loss of forest surface, where fragmentation increased and a loss of functional connectivity between 25% and 49% occurs depended on the parameter analysed. On the other hand, some areas were recovered. Brazil is the country that recovered the most forest area and Argentina, Paraguay and Bolivia those that lost the most area. Only 31% of the area was not altered between 1992 and 2020. Human activities such as deforestation, agriculture expansion, and urbanization have led these forests to become increasingly fragmented and worse connected impacting on both ecological and socio-economic aspects. Supranational measures must be taken to mitigate the negative impacts of fragmentation and the loss of connectivity.
... Conversely, isolation by the environment is associated with greater genetic similarity between populations in similar environmental conditions (Alberto et al., 2013;Sexton et al., 2014). Research suggests that climatic factors might play a role in driving biogeographical patterns, as closely related species' distributional boundaries match climate regimen boundaries at the transition zone between the two regions in Western Ecuador (Albuja et al., 2012;Amador et al., 2019;Escribano-Avila et al., 2017;Morrone, 2006;Prieto-Torres et al., 2019). ...
... STRUCTURE when K = 4. DAPC identified K = 4 as the most likely number of clusters, while STRUCTURE regarded it as the third most likely. Employing K = 4 not only enables a finer classification but also resulted in genetic clusters with geographical boundaries that correspond closely to those of biogeographical regions that have been previously reported(Amador et al., 2019;Escribano-Avila et al., 2017;Morrone, 2006;Prieto-Torres et al., 2019) enabling us to explore potential barriers within Western Ecuador. ...
Article
Full-text available
Environmental gradients have the potential to influence genetic differentiation among populations ultimately leading to allopatric speciation. However, environmental gradients can also facilitate hybridization between closely related taxa. We investigated a putative hybrid zone in western Ecuador, involving two polytypic wren species (Aves: Troglodytidae), Campylorhynchus zonatus and C. fasciatus. Our study addressed two primary questions: (1) Is there evidence of population structure and genetic admixture between these taxa in western Ecuador? and (2) What are the relative contributions of isolation by distance and isolation by the environment to the observed genetic differentiation along the environmental gradient in this region? We analyzed 4409 single‐nucleotide polymorphisms (SNPs) from 112 blood samples sequenced using ddRadSeq and a de novo assembly. The optimum number of genetic clusters ranged from 2 to 4, aligning with geographic origins, known phylogenetics, and physical or ecological constraints. We observed notable transitions in admixture proportions along the environmental gradient in western Ecuador between C. z. brevirostris and the northern and southern genetic clusters of C. f. pallescens. Genetic differentiation between the two C. f. pallescens populations could be attributed to an unreported potential physical barrier in central western Ecuador, where the proximity of the Andes to the coastline restricts lowland habitats, limiting dispersal and gene flow, especially among dry‐habitat specialists. The observed admixture in C. f. pallescens suggests that this subspecies may be a hybrid between C. z. brevirostris and C. fasciatus, with varying degrees of admixture in western Ecuador and northwestern Peru. We found evidence of isolation by distance, while isolation by the environment was less pronounced but still significant for annual mean precipitation and precipitation seasonality. This study enhances our understanding of avian population genomics in tropical regions.
... The observed changes in vegetation have a direct effect on the availability of resources for frugivorous species, which in the medium term can generate a cascading effect on the whole forest ecosystem 8,59 . Although zoochory and plant regeneration dynamics are recognized as key processes to ecosystem functionality, there is need to aid them in the development of a risk assessment approach for the ecosystem 60 . Understanding the broader implications of these functional shifts is vital for informing management practices. ...
Article
Full-text available
Chronic disturbance is a major driver of biodiversity loss in seasonally dry tropical forests (SDTFs). However, its consequences on the functional dimension of diversity, particularly dispersal traits, remain unclear. We evaluated the effects of chronic disturbance on the functional traits of woody plant community and its potential effects for frugivores. We characterized eight traits related to seed dispersal and calculated the community weighted means and functional diversities for trees and whole woody community. We used GLMs to evaluate the effects of the chronic disturbance on these functional metrics, including the abundance and diversity of fruits as resources for frugivorous group. Chronic disturbance filtered traits related to dispersal, reducing the proportion of species with fleshy and heavier fruits, and producing more seeds per fruit. Functional richness and dispersion declined with disturbance. We found a general pattern of reduction in the availability and richness of fruits. Our results suggest that the changes in species richness and abundance are not random but result from environmental filtering on traits related to dispersal costs and stress tolerance. Changes in vegetation directly affected resources availability for frugivorous group, which in the medium term can affect the woody species persistence and catalyze the woody species loss.
... caudiscutatus), which have geographical distributions restricted to the northwestern part of the continent, which are also affected by cat predation. These findings illustrate the potential interaction between cats and locally adapted species, which can carry consequences for the function of ecosystems (Kitts-Morgan 2015;Escribano-Avila et al. 2017). A recent meta-analysis reported 89 native species as prey of cats for the South American continent (Lepczyk et al. 2023), while we recorded 30 native taxa (21 native species) solely in the study area, which has an area of approximately 3,000 km 2 . ...
Article
Full-text available
Domestic cats pose a latent threat to wildlife that lives within the remnants of natural vegetation in urban ecosystems. Both intrinsic (e.g., age, weight, sterilization status) and extrinsic factors (e.g., night confinement, interaction time with owners at home) can influence the number of prey items caught by cats. We assessed the fauna predation by domestic cats in three cities on the coast of Ecuador. We aimed to: (i) evaluate the composition of the prey brought home by cats, counting the taxa number and their capture frequency, as well as their conservation status, and (ii) identify the intrinsic and extrinsic factors that influence the quantity of prey brought home by cats (henceforth referred to as ‘prey captured’). A citizen science approach was employed to gather information about wildlife taxa caught and brought home by 100 cats in 50 households between March and October 2023. Cats captured 132 prey items, of which 53.8% were invertebrates, 27.3% reptiles, 8.3% birds, 6.8% small mammals, and 3.8% amphibians. These prey items belonged to 53 taxa, 56.6% native and 15.1% non-native. Non-native reptiles Hemidactylus sp. and Anolis sagrei were the most frequently captured taxa, and ten native taxa were among the most commonly captured, particularly odonates. This is the first study to register predation of cats on amphibians in northwestern South America. The capture by cats of Coniophanes dromiciformis, a vulnerable and probably endemic snake, is noteworthy. Three factors—age, nocturnal confinement, and the presence of toys in their homes—were the most important factors that contributed to predation events. We recommend controlling these factors to reduce the potential impacts caused by domestic cats on wildlife.
Article
Full-text available
Croton anomalus was described by Henri Pittier in 1930 from a collection made in Estado Lara, Venezuela, and the use of the name has so far been restricted to several states in this country. A reevaluation of the species has led to its recircumscription and recognition in several other countries, including Bolivia, Brazil, Colombia, French Guiana, Guyana, Mexico, and Suriname. Previously, it was confused with species referred to here as the “Croton anomalus group”, namely C . acapulcensis , C. blanchetianus , C . chiapensis , C . jacobinensis (= C . sonderianus ), and C. stahelianus . We integrated morphological, phylogenetic, and ecological evidence to understand species limits and relationships within the Croton anomalus group. We first studied ca. 650 herbarium specimens covering the geographic range of the group, and we inferred species phylogenetic relationships using DNA sequences from the nuclear and plastid regions (ITS and trnL-F ). We also used ecological niche modeling to infer potential suitable areas for the occurrence of the studied species and to determine the variables that most contribute to their distribution model. Both morphological and phylogenetic data provide evidence for the synonymization of C . acapulcensis , C. chiapensis , and C . stahelianus under C . anomalus . On the other hand, our results support the recognition of C. blanchetianus and C. jacobinensis as two independent lineages, both distinct from C. anomalus . An emended description of C. anomalus is provided, as well as the designation of lectotypes, illustrations, updates of distribution data, and morphological comparisons with closely related species. Regarding niche modeling, annual precipitation and the precipitation of the warmest quarter were the most important variables explaining species distributions. Croton anomalus showed suitable areas in most seasonally dry tropical forests in the Neotropics, while C. blanchetianus and C. jacobinensis had their most suitable areas restricted to the Caatinga Dry Forest (Brazil), and Caatinga + northern South America, respectively. Our study shows the importance of taxonomic revisions using integrative approaches to disentangling species boundaries and to elucidate their biogeography and conservation status.
Article
Full-text available
Los bosques secos tropicales, son ecosistemas únicos y diversos, pese a su importancia biológica se encuentran entre los más amenazados del mundo, las presiones sociales y económicas han provocado una reducción significativa de su cobertura, lo que ha afectado su capacidad de proporcionar servicios ecosistémicos. El objetivo de este estudio fue conocer los cambios de contenido de carbono en la biomasa aérea en dos pisos altitudinales 600 y 1200 m s.n.m. en la Reserva Natural Laipuna de Naturaleza y Cultura Internacional. Se registró el diámetro a la altura del pecho (DAP) y altura total de todos los individuos arbóreos con DAP ≥ 10 cm en seis parcelas permanentes de una hectárea. Para la estimación del carbono en la biomasa aérea se usó la ecuación de Chave, considerando el DAP (cm) y la altura de los árboles (m), así como también la densidad de madera (g/cm3) de cada una de las especies presentes dentro de las parcelas. No se encontraron diferencias estadísticas entre los contenidos de carbono dentro del área evaluada estas reservas fueron entre 35,6 Mg C ha-1 y 43,2 Mg C ha-1 a 600 m s.n.m. y 1 200 m s.n.m. respectivamente, demostrando así que la altitud es un factor que no influye en el almacenamiento de carbono aéreo. Se considera que este tipo de bosque representa una opción para contrarrestar el aumento de CO2 atmosférico, siendo este un justificativo importante para su conservación, más aún cuando el bosque se encuentra bajo procesos dinámicos de crecimiento.
Article
Full-text available
Understanding the spatial distribution of animal species, which involves integrating species occurrence with environmental data, provides crucial information for conservation planning, especially for threatened species. In this study, we used niche-based species distribution models to create a potential distribution map of Leptonycteris curasoae, a vulnerable fruit- and nectar-feeding bat. This model incorporated occurrence data from field sampling, mammal collections, scientific literature, and environmental variables. Additionally, we mapped the threats faced by L. curasoae and overlaid this data with protected area boundaries to identify priority conservation regions. Our results indicate that the current and potential distribution of this species is considerably smaller (~9 %) than the area previously considered according to the IUCN Red List. The potential distribution of L. curasoae is environmentally restricted to arid and semiarid areas and dry forests in Aruba, Bonaire, Curaçao, Northern Colombia, and Venezuela, characterized by high temperatures, low precipitation, and seasonality in temperature and precipitation. Approximately 22 % of the suitable areas for this species are within protected areas, and we observe differences in the magnitude of the area under protection and the impact of identified threats among countries. Roost vandalism is the most critical threat in Curaçao and Venezuela, while mining, tourism, and wind farms are more frequent in Colombia. Expanding or creating protected areas and roosts, jointly with establishing conservation corridors and connected private reserves across political boundaries, are high-priority conservation actions needed to guarantee safe mating and maternity roosts, long-distance movements, and connectivity of L. curasoae populations along its entire geographic distribution.
Preprint
Full-text available
Although chronic disturbance is widely recognized as a main driver of biodiversity loss in tropical dry forests, their consequences beyond the taxonomic loss perspective (i.e the functional dimension of diversity) still need to be clarified, especially in those plant traits associated with dispersal. Here, we evaluated the effects of chronic disturbance on the functional diversity of a seasonally dry tropical forest, and their potential effects on the frugivores guild. We characterized eight plant traits related to seed dispersal and calculated the community weighted means and functional diversities for trees and the whole woody community. We used generalized linear models to evaluate the effects of the disturbance on these functional estimates in relation with the abundance and diversity of fruits as resources for wildlife. Our results revealed that, the dominance of plants with costly fruiting species was reduced with disturbance. Functional richness and divergence were reduced with the disturbance, mainly in the qualitative traits. Finally, the availability of resources was slightly different between groups of dispersers, observing a general pattern of reduction in the availability and richness of fruits with disturbance. Our results suggest that the changes in species richness and abundance are not random but the result of filtering on traits related to dispersal costs and their subsequent ability to withstand environmental stress. The observed changes in vegetation have a direct effect on the availability of resources for frugivorous species, which in the medium term can affect the woody species persistence and catalyze the woody species loss.
Preprint
Full-text available
Domestic cats pose a latent threat to wildlife in natural and urban ecosystems. Both intrinsic (age, weight, sterilization status, etc.) and extrinsic factors (night confinement, interaction time with owners at home, etc.) can influence the number of prey items caught by cats. We address the fauna predation by domestic cats in three cities on the coast of Ecuador. Two objectives were established: (i) determine the richness, composition, abundance, and conservation status of the prey captured by domestic cats, and (ii) identify the intrinsic and extrinsic factors that influence the quantity of prey captured. A citizen science approach was employed to gather information about wildlife species caught and brought home by 100 cats in 50 households between March and October 2023. Cats captured 132 prey items, of which 53.8% were invertebrates, 27.3% reptiles, 8.3% birds, 6.8% mammals, and 3.8% amphibians. These prey items belonged to 53 taxa, 56.6% native and 15.1% non-native. Non-native reptiles Hemidactylus sp. and Anolis sagrei were the most frequently captured taxa, and ten native taxa were among the most commonly captured, particularly odonates. This is the first study to register predation of cats on amphibians in northwestern South America. The capture by cats of Coniophanes dromiciformis, a vulnerable and probably endemic snake, is noteworthy. Three factors—age, nocturnal confinement, and the presence of toys in their homes—were the most important factors that contributed to predation events. We recommend these factors be considered to reduce the impacts by owned cats on wildlife.
Article
Full-text available
Reportamos la especie Tumbezia salvini (Aves: Tyrannidae) en el sector El Oro de Pilares, cantón Zapotillo, provincia de Loja, Ecuador. Esta localidad se ubica en los bosques secos del suroccidente del Ecuador y se ubica a 16 km de la única localidad previamente conocida en el país para la especie.
Article
Full-text available
Deforestation and fragmentation are major components of global change; both are contributing to the rapid loss of tropical forest area with important implications for ecosystem functioning and biodiversity conservation. The forests of South Ecuador are a biological 'hotspot' due to their high diversity and endemism levels. We examined the deforestation and fragmentation patterns in this area of high conservation value using aerial photographs and Aster satellite scenes. The registered annual deforestation rates of 0.75% (1976-1989) and 2.86% (1989-2008) for two consecutive survey periods, the decreasing mean patch size and the increasing isolation of the forest fragments show that the area is under severe threat. Approximately 46% of South Ecuador's original forest cover had been converted by 2008 into pastures and other anthropogenic land cover types. We found that deforestation is more intense at lower elevations (premontane evergreen forest and shrubland) and that the deforestation front currently moves in upslope direction. Improved awareness of the spatial extent, dynamics and patterns of deforestation and forest fragmentation is urgently needed in biologically diverse areas like South Ecuador.
Article
Full-text available
The importance of the ecosystem services provided by dry tropical forest is better understood when analyzing peoples' perceptions of such services. Particularly south ecuadorian dry forests have been scarcely studied from social sciences, though they are the most populated and threathened forests in the world. Therefore, little is known about factors that may influence the perception of ecosystem services provided by such forests. Based on a study case research in the Reserva Ecológica Arenillas and by using a qualitative and quantitative approach, we discuss how the history of the Reserve and factors such as participants' residence, frequency of visits, gender and age play a key role in the social perception and value given to the ecosystem services provided by the protected area.
Article
Full-text available
Plant regeneration is essential for maintaining forest biodiversity and ecosystem functioning, which are globally threatened by human disturbance. Here we present the first integrative meta-analysis on how forest disturbance affects multiple ecological processes of plant regeneration including pollination, seed dispersal, seed predation, recruitment and herbivory. We analysed 408 pairwise comparisons of these processes between near-natural and disturbed forests. Human impacts overall reduced plant regeneration. Importantly, only processes early in the regeneration cycle that often depend on plant-animal interactions, i.e. pollination and seed dispersal, were negatively affected. Later processes, i.e. seed predation, recruitment and herbivory, showed overall no significant response to human disturbance. Conserving pollination and seed dispersal, including the animals that provide these services to plants, should become a priority in forest conservation efforts globally.
Article
Full-text available
In plant–animal interactions, species are commonly labeled as either mutualists or antagonists, based on the most common, most studied, or most easily observed outcome. Nevertheless, evidence from simple systems comprising 2–4 species suggests that those labels are an oversimplification: individual species often function in both roles, either simultaneously or at different places or times. We include both mutualistic and antagonistic interactions between mammals and seeds in a multilayer network, to explore for the first time the community-level consequences of the dual roles played by some species. We tested whether negative and positive interactions within a plant-frugivore network are separated into different modules, or whether they overlap due to the presence of frugivores that both kill and disperse seeds. The frugivorous diets of nonvolant small mammals were studied at one dry tropical forest site in southeastern Brazil by analyzing fecal samples from individuals captured in live traps. Seed viability was assessed with a tetrazolium test to determine the outcome of those interactions, as estimated by whether or not seeds survived gut passage. Interactions were analyzed as a weighted multilayer network, subdivided into one potentially mutualistic (live seeds deposited) and one antagonistic (dead seeds deposited) layer. The two layers had similar structure with high overlap between them. Some mammal species exhibited highly central, dual roles, acting both as antagonists and mutualists, in many cases of the same plant species. Dispersal service by most of these small mammals is accompanied by seed destruction, suggesting that the selective pressures exerted by those animals on the plants is much more complex than often assumed. Our results demonstrate that the complexity of plant–frugivore networks cannot be fully understood without proper incorporating measures of seed fate following gut passage. This article is protected by copyright. All rights reserved.
Article
Full-text available
Deforestation and fragmentation are major components of global change; both are contributing to the rapid loss of tropical forest area with important implications for ecosystem functioning and biodiversity conservation. The forests of South Ecuador are a biological 'hotspot' due to their high diversity and endemism levels. We examined the deforestation and fragmentation patterns in this area of high conservation value using aerial photographs and Aster satellite scenes. The registered annual deforestation rates of 0.75% (1976–1989) and 2.86% (1989–2008) for two consecutive survey periods, the decreasing mean patch size and the increasing isolation of the forest fragments show that the area is under severe threat. Approximately 46% of South Ecuador's original forest cover had been converted by 2008 into pastures and other anthropogenic land cover types. We found that deforestation is more intense at lower elevations (premontane evergreen forest and shrubland) and that the deforestation front currently moves in upslope direction. Improved awareness of the spatial extent, dynamics and patterns of deforestation and forest fragmentation is urgently needed in biologically diverse areas like South Ecuador.
Article
Article
Seed dispersal, together with pollination, are two key services provided by animals to plants. On oceanic islands, where strong isolation limits the arrival of medium and large sized mammals (Gorman 1979), tortoises, iguanas or lizards often undertake an important ecological role as dispersers (Olesen & Valido 2003). Furthermore, the reported niche expansion or interaction release of island vertebrates, which tend to occupy underexplored ecological niches and adopt super-generalized diets, magnifies the ecological importance of insular native fauna (MacArthur et al. 1972; Cox & Ricklefs 1977; Traveset et al. 2015). This article is protected by copyright. All rights reserved.