JANUARY – JUNE 2018

SPECIES INVENTORY AND FOREST BIODIVERSITY

COMPARISION FOR THE AREA OF JALOVA,
TORTUGUERO NATIONAL PARK, COSTA RICA

PRODUCED BY:

Victoria Hawkins – Forest Biodiversity Project Leader

IN COLLABORATION WITH:

FIELD COORDINATOR: Cormac Healy

FIELD STAFF: Danny Guy, Thijs Claes, Stephanie Butera, Allen Bush, Emily Underhill, Dan
Sturgess, Amber Searcy.

GVI COSTA RICA COUNTRY DIRECTOR: Cynthia Arochi

 GVI Costa Rica

Jalova, Tortuguero National Park

The research for this project has been conducted by Global Vision International under the
investigation permit RESOLUCIÓN SINAC-ACTO-D-RES-018- 2018 EXPEDIENTE ACTO-PIN-
030-2016 The findings and recommendations made in this report are based upon analysis of
data collated during the period January – June 2018.

Cited as

Hawkins, V. 2018. Species inventory and biodiversity comparison for the area of Jalova,
Tortuguero National Park, Costa Rica.

Submitted in whole to

Global Vision International (GVI)

The Ministry of Environment, Energy of Costa Rica (MINAE)

 With thanks to all the staff and volunteers who contributed to the Incidentals Project
during the study period and since its commencement in 2010.

The research during the study period was conducted with the help of the following
volunteers:

Anna Bowland, Jessica Floyd, Nina Champion, Lindsey Agee, Lakesha Smith, Matilda Bray, Blaire
Borstad, Sophie Hood, Erica Bowker, Chris Beckingham, Emilie Andersen, Joanna Pittet, Elsa
Liljekvist, Ryan Collier, Thomas Cockell, Lindsay Gilbert, Tracey Hinton, Adam Grushcow, Rebecca
Higham, Charlotte Howard, Christine Davison, Katherine Andrews, Jordan Phoenix, Isobel Bond,
Laura Marschall, Eva Dahlqvist, Amelia Scott, Joshua Klindienst, Hana Akselrod, Jacqueline Esteban,
Alana Toms, Alberte Larsen, Hannah Venter, Rachel Ford, Steffanie Knoll, Neil Dittrich, Steffanie De
Ree, Vivien Lauenstein, Evelyn Salguero, Gloria Guerrero, Wessel Peters, Emelie Ahrendsen, Rosa
Ghoneim, Francesca Galietta, Alexander Langenus, Sarah Drijkoningen, Ben Ryan, Courtney Watkins,
Lynsey Whittaker, Simone Ganpat, Adele Green-Armytage, Ebba Eriksson, Tora Godow Bratt, Kalisyn
So, Finn Hampton, Justin Hedlund.

GVI Costa Rica (Jalova)

Email: costarica@gviworld.com

Web page: http://www.gvi.co.uk and http://www.gviusa.com

ABSTRACT

For the previous eight years, Global Vision International (GVI) has been working in
collaboration with the Ministry of Environment and Education (MINAE) to document the
vertebrate species found in the southern area of Tortuguero National Park (TNP), known as
Jalova. This work is formed by two studies. The first is the Incidental Project which has been
running since 2010. The aim of this project is to build a vertebrate species inventory to
increase the knowledge of biodiversity in this area. This report confirms that new species are
being consistently added year on year, in particular for the last six month study period January
to June 2018, a total of six new species were recorded. Along side the species, the type of
habitat it was recorded in was also noted. This allows for direct comparison of habitat types
and the sightings of species. In this study period the class of Aves were the most sighted and
associated more with the disturbed habitat of the coconut plantation. A more even
distribution of species is found in the forest habitat, particularly the swamp forest habitat.
The second project, the Forest Biodiversity project, incorporates a more balanced time effort
into the study with certain trails being surveyed for an equal amount of time over the study
period. The results from this project resemble the results from the incidental project where
again the Aves class dominates the disturbed habitat of the coconut plantation and the forest
habitat types have a more even distribution of species. Longer term trends are also examined
with a focus on some of the IUCN species of interest. Continuation of the Incidentals Project
and the Forest Biodiversity Project to collect long-term data and add to the species inventory
for the southern tip of the TNP would be beneficial.

INTRODUCTION

Costa Rica is a small country, making up less than 0.05% of the world’s terrestrial surface
(Wainwright & Arias, 2007), but it is listed in the top 20 countries for holding the most
biodiversity on Earth (Nielson-Munroz et al. 2012). Despite its small size, it holds almost 4%
of the world’s biodiversity (Nielson-Munroz et al. 2012). Over 900 species of birds (Garrigues
and Dean 2007), nearly 400 species of amphibian and reptiles (Leenders 2001) and at least
240 mammal species (Wainwright and Arias 2007) are found in Costa Rica. Central America is
well known to be a mega diverse ‘hotspot’ for biodiversity (Nielson-Munroz et al. 2012)
however only a fraction of the estimated amount of species have as yet been described
(SINAC 2009).

Central America acts as a natural land bridge between North and South America; as such
species from both continents are represented as well as endemic species (Baltensperger &
Brown 2015). The large number of life zones and ecoregions which provide a multitude of
habitat types explains the high biodiversity found in this region (Stiles and Skutch 1989;
Garrigues and Dean 2007). It is well documented that habitat change, fragmentation and
deterioration often at the hand of anthropogenic forces are key drivers in the risk of
biodiversity loss (e.g. Baltensberger & Brown 2015, Hooper 2008). Furthermore, the threat of
spreading non-native species coupled with these extant threats also poses a great risk to
biodiversity (Roy et al 2014). Therefore, it is crucial to understand habitat change, particularly
in regards to regeneration of secondary habitats and within plantations (Beirne et al. 2013).
Costa Rica has experienced high levels of deforestation with up to a 50% loss of forests since
1950 (Baltensberger & Brown 2015). As a result of this the landscape is fragmented and now
consists of a mosaic of habitats. One way to tackle this issue was to instigate a national
network of protected areas encompassing a multitude of habitat types (Wainwright & Arias
2007).

In the 1970’s as part of this national network, Torutugeuro National Park (TNP) was
established (Boza 1993). The park was originally established to protect a valuable nesting site
for green turtles (Chelonia mydas) but as a consequence of this new status the tropical forest
was also preserved, and former plantations were acquired and turned back into forest (Boza
1993). A high level of biodiversity was discovered in the park, increasing its value and ensuring
its protection. Almost 44% of the known biodiversity found in Costa Rica is found in TNP
(Bermudez 2006). Specifically, this includes 375 species of aves, 125 of mammalia, and 124 of
reptilia and amphibian. However, one consequence of a protected area network is that it can
result in the patch effect. Species are unable to move from one protected area to another
resulting in their increased isolation (Cabrera-Guzman and Reynoso 2012). It was therefore
vital to establish connectivity between these sites to ensure habitat continuity. The San Jan –
La Selva Biological Corridor which includes TNP was established to prevent fragmented
populations and secondary growth forests play a large role in conservation for this reason
(Hooper 2008).

The regeneration of forested areas can take up to 100-180 years to recover fully back to
primary status after anthropogenic disturbance (Chazdon et al. 2009) One way to monitor the
success of regrowth is to look at changes to the fauna species assemblages (e.g. Takano 2014).
As the forest develops and matures, species assemblages also change from generalist species
to ones which are more specialised (Devictor et al. 2008, Christian et al. 1998). However,
many studies of these systems are limited to young regenerating forests, and therefore it is
important to get more information on maturing secondary forest, and more long term studies
on species assemblages (Chazdon et al. 2009; Hooper 2008).

Knowing where specific species occur within a managed area is also useful for maximising
tourist experience whilst minimising interference with sensitive or vulnerable species (Bryan
2014). Species inventories can assist conservation by signifying the ecological status of an
area, particularly some assemblages of indicator species, which are indicators of good
ecological health and the condition of the habitat (Stotz et al. 1996). Some bird species are
often considered good indicator species because they can have specific requirements for
areas they live in i.e. nesting sites, certain types of fruit or seeds and canopy cover (Stotz et
al. 1996). Amphibians are also useful indicator species because of their high sensitivity to
pollution, and their need for complex microhabitats (Wilson and McCranie 2003).

The aims of the project undertaken by Global Vision International (GVI) at the southern end
of TNP are to build a species inventory of the vertebrate species found here. This inventory
will increase the knowledge and understanding of the species assemblages and therefore
inform management decisions. The Jalova research station is in the unique position of being
able to compare directly disturbed habitat types with secondary and primary forest habitat.
As a result, a further aim of the research undertaken here is to monitor the changes in species
assemblages and the habitat zones found here to gain understanding on how well the forest
is regenerating.
OBJECTIVES

INCIDENTALS PROJECT:

The Incidentals Project collects data on the presence of all mammal, aves, reptile, and
amphibian species found within the southern tip of TNP. The main aim of the project is to
construct a thorough species inventory for MINAE so that there is a record of every species
identified within the area and date of first sighting. A secondary aim of the project is recording
habitat types that species are encountered in to compare number of sightings in different
areas, and compose a database that can be used to compare trends over time.

FOREST BIODIVERSITY PROJECT:

The Forest Biodiversity Project’s main aim is to collect and compare information on the
vertebrate species richness and abundance within the different habitats found in the
southern tip of TNP, and look at richness and abundance between different trails that are
used by GVI expedition members. A secondary aim is to look at ecological information about
the species found such as locations in which species are sighted. In 2016 two new trails were
added within the coconut habitat to be able to compare the species diversity between the
forest habitats and the human maintained disturbed habitat. By adding these two trails to
our existing six forest trails we are able to compare an area significantly and continuously
impacted by humans to those less impacted. This also helps to reduce bias in the data when
comparing species richness between the forest and coco habitats.

STUDY AREA

The Jalova Biological Research Station is operated by Global Vision International and is located
in the southern tip of the Tortuguero National park. Tortuguero National Park is located on
the Caribbean coast of Costa Rica as shown in figure 1, includes 76 000 ha of protected land
and sea (26 000 ha land) and is classified as a tropical rainforest. The topography of the area
ranges from 0 to 311m above sea level, and the average rainfall is near 6000mm per year with
an average temperature of 26̊C (Lewis et al. 2010; Bermúdez and Hernández 2004; Bermúdez
2006). Within this area are several different habitat types including several different types of
swamp forest (some dominated by several palm species, some more species rich including
high canopy tree species), herbaceous marshes, coastal forest (has a more open canopy, often
dominated by sea grape - Coccoloba uvifera), dense forest (drier than the swamp forest areas
often dominated by Ficus species) coconut plantation and beach (Halewood et al. 2013; Lewis
et al. 2010; Ramsar 2006).

Within the study area there are nine main trails that are used to survey the forest habitats, as
well as two trails within the coconut plantation that were added within the last year (figure
2). Trails are as follows;

Kingfisher: (600m) Dense Forest. Starts at the edge of coconut plantation and winds through
dense forest ending at the estuary. Dominated by high canopy trees with a reasonably open
mid canopy layer, and a lot of smaller palms and ferns at lower levels. It is well drained most
of the year with only a small amount of standing water present in the wetter seasons.

South Boundary: (580m) Swamp Forest. Starts in the coconut plantation near kingfisher and
heads north west, this trail is one of the wettest we have and most of the year there is
standing water and in the driest periods there is still thick mud. A lot of palm species are
present but with a variety of larger trees forming a high canopy in most places.

North Boundary A: (650m) Dense Forest. Starts in the coconut plantation and travels
perpendicular to the plantation. A dry trail at most times of the year with only one spot
subject to standing water in rainy periods. More open forest than in kingfisher with a sparser
mid canopy and less understory vegetation, at points gets very near to the plantation.

North Boundary B: (400m) Swamp forest. Dominated by raffia palms (Raphia taedigera), low
canopy for most of it, some parts are very open with a lot of herbaceous species such as peace
lilies (spathiphyllum friedrichsthalii). Most of the year is inundated and even in dry periods
has mud and a few pools of water.

North Trail A and North Trail B (Originally one trail of 1050m now divided into two different
trails of 700 m each; the habitat of these trails is very similar) Coastal Forest. Travels parallel
to the beach though coastal forest, a much drier trail than any of the others, canopy is
dominated by sea grape (Coccoloba uvifera) and tropical almond (Terminalia catappa), very
sparse understory. Very open and exposed, the widest and most used trails within the study
as it is often used by rangers on quad bikes and is maintained by rangers with strimmer’s,
machetes, and chainsaws.

Juana Lopez: (700m this trail was originally 990m, but the length of the survey was reduced
to be more comparable with other trails and to ensure only habitat consistent with dense
forest was included) Dense Forest. Located the furthest away from the base of all the trails
within the study it is the least used by GVI of all our trails. It starts 300 metres from the beach
at mile 15 and goes west though dense forest to end at the canal Cano Negro. This trail has
some of the oldest and least impacted forest that we survey, dominated by Figus spp. with a
dense canopy and mid canopy. At several points it travels though small herbaceous wetland
areas.

Cocos Back Trail: (650m) Coconut Plantation. This trail starts outside the base and is shielded
from the beach by the base and a thick hedge of vegetation. It travels through coconut
plantation that is bordered by dense forest. This is very open and dry habitat dominated by
coconut palms (Cocos nucifera) with a few other fruit trees interspersed, the grass is kept low
by a small herd of cows and manual slashing.

Cocos Front Trail: (700m) Coconut Plantation. Starting at the front gate to the base this trail
runs through the coconut plantation that boarders the beach, at points this trail is very close
to a mixed vegetation hedge that separates two areas of coconuts. Other than this the
dominant tree is coconut palms (Cocos nucifera) and it is a very open and dry habitat. The
grass is kept low by a small herd of cows and manual slashing. Because this trail is in full sight
of the beach at many points it will never be surveyed at night time during the turtle season
due to potential disturbance for nesting marine turtles.

METHODOLOGY
For full methodology please see: Hawkins, V. & Guy, D. 2018. Anteproyecto 2018 Inventario
de biodiversidad en el área de Jalova.
Data Analysis Methodology

The time and effort spent on each survey differed year on year, so it was important to use an
indices that negated this bias, which was the primary reason the Simpson’s Diversity Index
was used. This index compares species richness but also the evenness. Evenness is a measure
of relative abundance of the different species making up the richness of the area. The
Simpson’s Index of Diversity was used with the range 0 – 1, with the greater number
representing greater sample diversity. An average was determined for the trails, forest type
and study period and was used for direct comparisons, taking into account differing lengths
of trails and times the surveys were undertaken.

The abundance of species was calculated to correct for uneven sampling time and therefore
ensures more comparable data across the trails and forest habitat types. It was calculated by
taking the number of minutes spent per survey and dividing this by the number of species
seen over the equivalent time period.

A One-way ANOVA was used to examine species richness (using the Simpson’s Diversity Index)
against the different trails, time period (year) and class of species sighted for both the six
month comparison and the long-term comparison. Post hoc analyses were then conducted
following a statistically significant result on all relevant pairwise contrasts using a TUKEY post
hoc test.

Descriptive and visual statistics were carried out in Microsoft Excel and statistical testing in
Minitab 2017.

INCIDENTAL RESULTS AND DISCUSSION

During this study period (January – June 2018), six new species were added to the inventory
for the Jalova area of the National Park. This brings the total number of species recorded to
483. Of the six new species added, five were from the Aves class and one was from the
Mammalia class (Figure 1). All new species sighted were of least concern according to the
IUCN (2018).
 Common Name Scientific name IUCN status Population trend
Least Grebe Tachybaptus dominicus Least Concern Stable
Southern Lapwing Vanellus chilensis Least Concern Increasing
Buff-throated Foliage-gleaner Automolus ochrolaemus Least Concern Decreasing
Tree swallow Tachycineta bicolor Least Concern Stable
Dickcissel Spiza americana Least Concern Decreasing
Forest Spiny pocket mouse Heteromys desmarestianus Least Concern Stable

Figure 1. The six new species that have been added to the inventory during the period of January 2018- June
2018. IUCN status and population trends have also been included.

The primary objective of the Incidental project is to continue to build the species inventory
for the Jalova region of Tortuguero National Park, as such the addition of six new species in
this permit period demonstrates this aim is still being met and remains a valuable tool for
assessing the park.

Of the Automolus genus, the Buff-throated foliage-gleaner (Automolus ochrolaemus) is the

most common and widespread. It favours secondary growth forests and forages in the
understory. It has been assessed as Least Concern by the IUCN (2018) but the population
trend is decreasing due in large part to habitat loss. It is projected that 11.7 – 12.8% of
suitable habitat will be lost over the next three generations (12 years) (IUCN 2018). It is
susceptible to habitat fragmentation throughout its range so the sighting within the
National Park is encouraging.

The Tree swallow (Tachycineta bicolor) has an extremely large range and currently a large
population. Due to these factors it has been assessed as least concern with a stable
population (IUCN 2018). It is a non-breeding migrant to Costa Rica. The Dickcissel (Spiza
americana) is a non-breeding winter migrant to the Caribbean coast of Costa Rica. Its typical
habitat is open grasslands, but it has adapted well to secondary habitats. It is nomadic over
winter, often visiting several countries. However, it has a large population and large range
and therefore is considered stable and of Least Concern by the IUCN (2018).

The Forest Spiny pocket mouse (Heteromys desmarestianus) prefers areas of good second
growth and mature forests in the lowlands, as well as areas with abundant palms (IUCN
2018). Due to its prevalence in protected areas it is considered to be of least concern by the
IUCN (2018). It is encouraging for the health of the park that it has been sighted in Jalova
given its preferences for mature or good second growth forests.

The Least Grebe (Tachybaptus dominicus) is considered as Least concern by the IUCN and its
population trend is stable. During the dry season, this aquatic bird is known to inhabit slow
flowing large bodies of water. The Southern Lapwing (Vanellus chilensis) is found along
coastlines and near large bodies of open water. Their preferred substrate is open grasslands.
It is unlikely to be seen in closed canopy forests unless there are open areas with water
nearby. Common throughout their range from Costa Rica to South America, they are well
adapted to human disturbance and are increasing their range in response to deforestation
and cultivation. As such, the IUCN has assessed them as Least concern with their population
increasing.

There are several possible explanations behind the addition of these six new species to the
species inventory. All species are inside their natural range, so it is auspicious that they are
being recorded in the Jalova region of Tortuguero National Park. It is also possible that their
range has increased, which would be expected in secondary forest as it matures into more
complex habitat (Chazdon et al 2009; Baltensperger and Brown 2015). Despite these species
being of Least Concern (IUCN 2018) it is nevertheless encouraging that new species are
being recorded in the area.

A total of 7,738 sightings of species were recorded in this permit period. This is an average
of 50.57 sightings per day. This encompassed 266 individual species. Broken down further
by class (Figure 2), this equates to 5,677 Aves sightings, 1,029 Reptillia sightings, 562
Mammal sightings and 470 Amphibian sightings.

As expected the Aves class dominates the results, with the highest number of recorded
sightings coming from the Coconut Plantation (C), Base (A) and, of course, the Canals (R).
The reason that the canal zone (R) has a high number of bird sightings is due to a dedicated
project that actively surveys these areas with a consistent time effort in each permit period.
The target species are aquatic birds, but all sightings are recorded within the incidental
database.
The coconut plantations (C) and Base (A) are also dominated by the Aves class. There are
several reasons for this. Firstly, the areas are disturbed, the former being a plantation which
is maintained with the aid of grazing cattle and herbicides to control ‘weeds,’ and the latter
due to permanent human habitation. This disturbed habitat attracts more generalist bird
species such as flycatchers and grackles. The plantation is also a wide-open space which
lends itself more easily to spotting and identifying bird species with greater accuracy and
ease (Henderson, C. 2010). The combination of this and the habitat preferences of the
species encountered on a regular basis explains the higher number of sightings in these two
zones. Furthermore, for the Aves class, encounters can be recorded as either audio or visual.
As such their presence is easier to detect and could have resulted in the higher number of
sightings recorded. This technique is also relevant for amphibian species but as yet the
expertise to record audio does not exist within the team. Finally, because of the species
richness in the Aves class (where there are more species of bird found in Costa Rica than any
other class (Garrigues and Dean 2007)), it is unsurprising that the Aves class dominates the
recorded species sightings across all zones. As expected the three forest habitat types have
a more even distribution of recorded sightings across the classes. These habitats provide a
wide-ranging selection of niches for the four classes to occupy, which indicates that the
ecosystem is functioning as it should for a secondary regrowth forest and as the forest
continues to regenerate, the community will develop with it.
 Sightings per class
2500

2000
Number of sightings

1500

1000 Reptilia
Mammalia
500 Aves
Amphibia
0

Zone

Figure 2. Distribution of sightings across the zones broken down by class.

IUCN SPECIES OF INTEREST:

As part of the monitoring of the Jalova region of Tortuguero National Park, the IUCN species
of interest are of special importance to the project and for park management. These species
are good indicators for the continuing health of the forest and allow park management to
better understand how well protected the park is from outside threats. The numbers of
recorded sightings are used to assess their continued presence in this area of the park. This
data are not a reflection of the abundance of the focal species but rather an indication of
their continued presence in the park. It is also worth noting that 2018 is not a complete year
but a six month period and as such gives more of an indication on what is expected for the
year than a true representation of this. The Baird’s Tapir (Tapirus bairdii) has been recorded
eight times this year so far in comparison to the nine times it was recorded in the whole of
2017. The Great Green Macaw (Ara ambiguous) is also on track for being seen at least with
the same frequency as the previous year and more sightings have been recorded so far this
year than other preceding years 2010 – 2015. However, due to the way data are recorded,
all sightings have the potential to relate to the same individual, therefore we cannot
extrapolate further in terms of population numbers or trends based on this data. It is
propitious that the sightings are being recorded with more frequency which is a good
indication that at the very least Great Green Macaws are more active in this area than
previously. The remaining species listed (Figure 3) all follow similar trends which will be
examined in more detail when the full years data set is available for the end of year report.

Vulnerable 2010 2011 2012 2013 2014 2015 2016 2017 2018a
American Crocodile (Crocodylus
73 40 88 81 68 56 21 47 26
acutus)
Great Curassow (Crax rubra) 13 19 20 29 74 39 42 38 11
White-lipped Peccary (Tayassu pecari) 11 7 1 6 35 20 38 19 2
Endangered 2010 2011 2012 2013 2014 2015 2016 2017 2018a
Baird's Tapir (Tapirus bairdii) 0 3 3 16 36 19 25 9 8
Great Green Macaw (Ara ambiguus) 5 0 0 3 12 36 55 50 28
CA Spider Monkey (Ateles geoffroyi) 181 105 103 153 273 248 267 279 135

Figure 3. List of IUCN species that are Endangered or Vulnerable currently recorded in the Jalova region of
Tortuguero National Park. The data listed are the number of recorded sightings for each year since the
incidental project began. 2018a is of the first six months of the year only which is represented by the a.

These species all have in common their sensitivity to disturbance be it through hunting
pressure, habitat change/loss or their complex dietary requirements. It is therefore a
positive sign for the park and its management that they are present here and being seen
with increasing regularity in the majority of cases. It is a good indication that the forest is
regenerating and recovering well enough to supply these complex needs across the
keystone and umbrella species.

NON-NATIVE VERTEBRATE SPECIES:

One of the biggest threats to biodiversity is the spread of invasive species, particularly when
this interacts with other known drivers such as habitat loss (Roy et al. 2014). A crucial part
of managing and monitoring a national park is knowing and understanding the extant
threats to biodiversity including non-native species that have arrived in geographically
similar locations. As such, monitoring for invasive species is an important tool for assessing
the continuing health of the forest ecosystem at Jalova. Currently, of the vertebrate species,
there are only five invasive species listed in Costa Rica. Of the two amphibian species listed
as being present in Costa Rica, one (Osteopilus septentrionalis) occurs in Limon province,
which is geographically close enough to warrant monitoring. To date there has been no
recorded sighting of this species in Jalova. At present it appears that this species is
contained to highly disturbed areas, most likely residential (Savage 2002). Therefore, at the
moment, it is unlikely to spread into the national park in the immediate future.

Similarly, there are two known invasive lizard species, one of which has been recorded in
the park. The House gecko (Hemidactylus frenatus) is believed to have been introduced
post-1990 (Savage 2002) accidentally through shipping. It has been recorded incidentally
and only on the Jalova base. It resides exclusively in human dwellings (Savage 2002) and it
has never been recorded in the forest. It has adapted to feed on insects attracted to electric
lighting (Savage 2002) and consequently, it seems to pose no threat to the biodiversity of
the forest. However, it has been documented to out-compete the smaller native species of
Gecko that have potential to be found here (Rodder et al 2008).

The remaining known invasive vertebrate species do not occur in this region and
subsequently do not pose a threat to the biodiversity of the park. Monitoring for invasive
species will continue through the work of this project as potential threats to the ecosystem
are identified.

FOREST BIODIVERSITY RESULTS AND DISCUSSION

During the study period of January – June 8th 2018 a total of 198 surveys were conducted.
Of the 198 survey, 58 were conducted in Dense forest, 45 in the Coco Plantation, 49 in
Swamp Forest and 46 in Coastal Forest (Figure 4 illustrates the spread over time). Although
the aim is to have an even distribution of surveys across time and space, it is not always
possible (see methodology for how data are treated). The main reason for the uneven
sampling is that Kingfisher and Juana Lopez are not allowed to be surveyed at night due to
the potential for Jaguar presence and in consultation with partner organisations. The Cocos
Front trail was deemed to have too great a risk of disruption from white light to nesting sea
turtles. This trail is parallel to the beach and little to no vegetation separates the two. From
the sampling conducted, 4,639 individuals were recorded. These individuals can be broken
down by class into 2,060 Aves, 1,268 Reptillia, 942 Amphibia and 369 Mammalia.

Trail Forest Type Morning Afternoon Night

Cocos Back Coco Plantation 8 8 9
Cocos Front Coco Plantation 10 10 N/A
Juana Lopez Dense Forest 7 7 N/A
Kingfisher Dense Forest 9 9 N/A
North Boundary A Dense Forest 9 8 9
North Boundary B Swamp Forest 10 8 8
North Trail A Coastal Forest 8 8 10
North Trail B Coastal Forest 6 8 6
South Boundary Swamp Forest 8 7 8

Figure 4. List of all trails with the forest type and the number of times they were surveyed during the morning
(6.30am), afternoon (14.30) and at night (19.30).

TRAIL ANALYSIS

The community composition was assessed for all trails using an average diversity index
which was calculated from all the surveys undertaken during the study period (Figure 5).
Greater species richness was found on the Cocos Front trail (Coconut Plantation) with an
average index of 0.930, this was followed by South Boundary (Swamp Forest) with an
average across all surveys of 0.904 and Kingfisher (Dense Forest) 0.903. The three with the
lowest indices were North Trail (0.799), North Trail A (0.826) and North Trail B (0.787), all of
which are coastal forest habitat. The forest habitat types are assessed further later in this
report for a more detailed overview.
 Simpsons Diversity Index for Trails
1

0.95

0.9

0.85

0.8

0.75

0.7

0.65

0.6
Coco Back Coco Front Juana Kingfisher North North North Trail North Trail North Trail South
Lopez Boundary Boundary A B Boundary
A B

Figure 5. The average Biodiversity Index for the study period January – June 2018 for each survey undertaken
for each trail.

This interpretation of the data was confirmed by the results from an ANOVA with a Tukey
pairwise comparison test. Significant differences were found between the trails when the
average Diversity Index was used (p = 0.00). Cocos front and South Boundary were indeed
the highest trails for diversity with the North Trails having the lowest diversity index score.

Trail N Mean Grouping

Coco Front 65 0.93042 A
South Boundary 145 0.90424 A
Kingfisher 131 0.90357 A
Juana Lopez 96 0.88171 A B
North Boundary A 173 0.87985 A B
Coco Back 82 0.8697 A B
North Boundary B 152 0.8458 B C
North Trail A 84 0.8261 B C
North Trail 76 0.7998 C
North Trail B 72 0.7874 C

Figure 6. The average Biodiversity Index for each trail. Means that do not share a letter are significantly
different.
The trails were then examined in more detail assessing the distribution of class across them.
The data are taken from the raw observations made across all surveys and as such will differ
from the above results. Therefore, caution must be used when interpreting the raw data
because sampling effort and time effort vary and have not been accounted for hence why
no statistical analysis has been performed. Nevertheless, it is a useful visual clue as to the
community composition and its distribution across the trails (Figure 7) and the bearing this
has on previous results.

The Aves class again dominates the community, particularly so for the two Cocos trails as
found in the Incidental project. This form of biotic homogenisation is to be expected in a
disturbed open habitat. It has often been observed that species richness decreases whilst
bird abundance increases with disturbance (Garaffa 2009). Furthermore, the forest edge
surrounding the plantation may be facilitating a greater movement of birds in a fragmented
habitat and accommodates those bird species that are classified as edge dwellers better
than an enclosed forest setting (Ries et al. 2004, Garrigues & Dean 2007). However, as
previously stated, Aves class are the most biodiverse of all the classes in Costa Rica so it
should not be surprising that they dominate the class distribution sightings here. Of the 484
known species of bird in Costa Rica, 289 species are known to breed in the Caribbean
lowlands which is 59% of the total species (McClearn 2016). The variety of habitat types, the
general climate with high rainfall and lack of seasons guarantees wide ranging availability of
different food sources including fruits, insects and flowers (McClearn 2016). These factors
mean that species richness is high in the Caribbean lowlands, particularly for bird species.
Another factor to explain this higher diversity for this class is because we do record both
audio and visual which increases the effort of recording. Indeed, 332 audio recordings were
registered in the six month study period out of a total 2,071 recorded sightings of birds,
16.03% of all the records. If other classes were treated in the same manner, such as
amphibians, then it is suspected that other classes may increase in turn.

As seen from the indices, South Boundary has the highest diversity and now from the raw
data it appears to have the most even spread of richness across the four classes. The
presence of open areas, standing water areas and dense canopy found in swamp forest
provides a variety of niches for all classes to occupy (Chazdon et al. 2009) and could explain
why the species richness is more even here. There is also a higher presence of Amphibian
species on this trail (as well as on Juana Lopez) than the other trails. Amphibians need shade
and moisture to thrive (Savage 2002) which is provided in this habitat type. The swamp
forest habitat also provides plenty of suitable breeding habitats for Amphibians. The biotic
and climatic features of the Caribbean lowlands provide plenty of temporary ponds and
small forest pools perfect for lentic breeding species such as Dendrobatids and Hylids. The
high rainfall and absences of prolonged dry periods ensure that Craugastor species are
largely diverse because these factors suit there terrestrial breeding needs (McClearn 2016).
All these habitats are provided in the swamp forest areas and all are sensitive to change at a
local and global level. It is important therefore to the preserve these habitats, especially in
light of the increasing spread of infectious fungal diseases and the continuing decline found
elsewhere in Costa Rica and the world.

Kingfisher trail, the North trails and North Boundary A have a greater presence of reptiles
which is consistent with their ectothermic regulation behaviours (Hertz 1992). These trails
have wide open spaces with a sparser canopy covering allowing greater levels of sunshine to
penetrate to the forest floor. More recent studies have also shown that reptile richness was
positively related to higher humidity, closed canopy cover, large patch size and the presence
of leaf litter (Cabrera-Guzmán & Reynoso 2012). These factors help explain why they are the
second largest class and are present on all trails as all their ecological needs are catered for
by all the forest types found in the study area.
 Trail with Class Distributions
3000

2500

2000

1500 Reptilia

1000 Mammalia
Aves
500
Amphibia
0

Figure 7. Class breakdown across all trails using raw data collected from all observations made on all surveys.

FOREST HABITAT TYPE ANALYSIS

The diversity index was calculated to investigate the community composition of the forest
habitat types. It is expected that these will yield similar results to the trail but this will
further enforce the categories used for each trail so it is a useful exercise. The Coconut
Plantation has the highest average Simpson’s Diversity Index of all the forest habitat types
with an average of 0.896, this was followed by Dense Forest with 0.888, Swamp Forest with
0.874 and Coastal forest with 0.804 (Figure 8). Whilst the first three habitat types are all
fairly even, Coastal forest as a whole is significantly lower in terms of its diversity. When the
trails were analysed separately the three North Trails were seen to have the lowest diversity
index but this was possibly due to uneven sampling of these trails. They were changed after
a year from one long trail into two smaller ones therefore the average indices for them
would appear lower. However, by treating it as a whole habitat type instead of individual
trails the results were similar and Coastal forest remains to have the lowest Diversity index
when compared to other forest types. This was born out with the results from the statistical
analysis carried out where p = 0.000 and the Tukey post hoc test (Figure 9) confirmed that
Coastal Forest is significantly different to the other forest types investigated here.
 Diversity by Habitat Type
0.95

0.9

0.85

0.8

0.75

0.7
Coastal Forest Coco Plantation Dense Forest Swamp Forest

Figure 8. Average Diversity of each forest habitat type.

Forest Type N Mean Grouping

Coco Plantation 147 0.8966 A
Dense Forest 403 0.88802 A
Swamp Forest 295 0.87439 A
Coastal Forest 231 0.80499 B

Figure 9. Tukey Post Hoc results. Means that do not share a letter are significantly different.

Using the diversity index alongside the class distribution breakdown (Figure 10), it is again
apparent that the Aves class dominates the Coconut Plantation and the Dense Forest
habitat type. Reptiles dominate the coastal forest habitat type and a more even community
is seen on the swamp forest habitat type. This broadly matches the results found in the trail
analysis and as such implies that the trails have been accurately categorised into suitable
habitat types.
 Habitat types with Class distributions
8000
7000
6000
5000
4000
3000
2000
1000
0
Coastal Forest coco plantation Dense Forest Swamp forest

Amphibia Aves Mammalia Reptilia

Figure 10. Class distribution across all Forest Habitat Types, taken from the raw data.

Contrary to expectations, the disturbed habitat of the Coconut plantation has the highest
diversity index. However, this is largely due to the high number of Avian species
encountered in this habitat and of the 484 known species of bird, 118 of them prefer open
habitats with an additional 66 species being generalist species. The remaining 166 species
prefer old or good secondary growth forests (McClearn 2016). As the Aves class dominates
the coconut plantation, it is useful to see which most commonly occurring birds are
recorded and what their habitat preferences are. Taking the top 10 most recorded birds, six
of the species are open or disturbed habitat dwellers, with two being unclassified as yet and
the remaining two are synonymous with secondary growth (Figure 11). These results are
expected from a disturbed habitat and also an open habitat where observations are easier
to record.

Species No. Individuals Habitat Type

Montezuma Oropendola 382 Unclassified
Variable Seedeater 120 Disturbed
Black Vulture 110 Disturbed
Western SLaty Antshirke 97 Disturbed
White Collared Manakin 90 Disturbed
Turkey Vulture 89 Unclassified
Bay Wren 88 Secondary
Tropical Pewee 69 Disturbed
Great Kiskadee 53 Disturbed
Slaty-tailed Trogon 53 Secondary
Figure 11. List of the top ten bird species found in the Jalova area of Tortuguero National Park. The Habitat
classifications were taken from Styles & Skutch 1989.

Looking at diversity and community composition is a valuable tool for understanding what is
happening spatially. Temporal investigations are also vital to understand trends and the
drivers behind these. Over the last six months (Figure 12), the trends over time in both
Coastal habitat and Swamp habitat remain relatively constant, with swamp habitat having
low peaks as expected following rainy periods. The Dense forest habitats experience clearer
peaks over the previous six months which match the low peaks of the swamp forest. This
can be attributed to the known breeding cycles of the amphibian species encountered there
when temporary ponds form and during breeding season for terrestrial breeders as
discussed above (McClearn 2016, Savage 2002). These patterns most likely follow those of
the rainy periods which occur in this region. The Coconut Plantation has a consistently high
average abundance, with a clear peak in May. As ascertained, it is dominated by the Aves
class therefore the high average abundance can be ascribed to the high numbers of bird
species known to breed in the Caribbean lowlands at this time (McClearn 2016, Styles &
Skutch 1989).

Abundance over time for habitat types

45
40
35
30
25
20
15
10
5
0
Jan Feb Mar Apr May Jun

Coastal Coco Dense Swamp

Figure 12. The average abundance of species over the six month study period for all forest
habitat types in the Jalova region of Tortuguero National Park. The average abundance was
calculated by dividing the number species seen on each survey by the length of tie spent on
survey.

The same patterns were observed for the trails (Figure 13), with Cocos Front and Back
matching the peaks and overall higher abundance. However more patterns emerge form the
trails when separated out which can be better interpreted. The Juana Lopez trail starts off
with high abundance during the first quarter of the year and then tails off and evens out for
the rest. The main explanation for this peak again aligns with breeding season for
amphibians. Although a dense forest trail, Juana Lopez has consistently returned high
numbers of amphibian species, especially juveniles during the first months of rainy season
(Savage 2002).

Abundance over time for the trails

60

50 Cocos Back
Cocos Front
40 Juana Lopez
Axis Title

Kingfisher
30
North Boundary A
North Boundary B
20
North Trail A
North Trail B
10
South Boundary

0
Jan Feb Mar Apr May Jun

Figure 13. Average abundance over the six month study period by trail in the Jalova region
of Tortuguero National Park. The average abundance was calculated by dividing the number
species seen on each survey by the length of tie spent on survey.

LONG-TERM TREND IN ABUNDANCE

Finally, the overall yearly trend was examined to observe the patterns of total abundance
over the total time the Forest Biodiversity project has been running (Figure 14). Although no
significant difference was found statistically (p = 0.07) for an increase in abundance over the
years, it is nevertheless encouraging that there is a slight increase. With more years of data
sampling and with a continued upwards projection shown in this regression model, there
could be a significant increase in the long term. This is auspicious for the continuing
recovery of the ecosystem as more species move into the area an increase in abundance is
expected which in turn indicates that the ecosystem is developing into a more complex
system and is able to provide niches associated with high abundance and richness
(Baltensperger and Brown 2015).

Abundance over years

160 R² = 0.0056
140
120
100
80
60
40
20
0
0 100 200 300 400 500 600

Figure 14. The abundance for each survey conducted over the three years of available data.

SPIDER MONKEY CENSUS RESULTS

As requested by ACTo, a census was conducted to estimate the population of the Central
American Spider Monkey (Ateles geoffroyi). The rationale behind this census was that of all
the mammals found in the park, the Central American Spider Monkey is the one with the
highest recorded sightings. A total of 209 individuals were recorded during the study period
of January to June 2018 (Figure 15), from the Forest Biodiversity data. This is out of a total
of 369 recorded sightings of all mammals from the same period. The dip seen in June is due
to the end of the permit period. The cut off date is 8th June. Therefore, the data are only
collected for a one-week period compared to a full month period for the previous months.
 Mammals
80
70
60
50 CA Spider Monkey
Axis Title

40 R² = 0.0125 All mammals

30 Linear (CA Spider Monkey)

R² = 0.0077 Linear (All mammals)
20
10
0
Jan Feb Mar Apr May Jun

Figure 15. The recorded sightings of Mammals within the study area. Due to the higher numbers of CA
spider monkey compared to all other mammals, it has been separated out. June represents only one
week of data.

Sightings were recorded from the trails we regularly survey and as such the census was
conducted in the same way. The same established trails and methodology used in incidental
surveys was followed but with one significant difference. All trails were surveyed by teams
of four simultaneously to reduce the likelihood of error in duplicating counts. Individuals
were counted when encountered and where possible their gender and age was determined
(Table 1). In total, the pilot census documented 22 individuals throughout the afternoon and
covered the majority of the study area. The exception was that the Juana Lopez trail was not
surveyed during the census due to the distance away from base this is located. In the future
the census will be conducted on a monthly basis and it is hoped that by the end of the next
study period the average monthly census figures will yield a robust estimate of the
population.

Location Total Adult Adult Adult Independent Dependent Breeding

Male Female Unknown Juvenile Juvenile Female

North Trail A & B 7 2 2 0 0 2 1

Cocos Back 0 0 0 0 0 0 0

South Boundary 0 0 0 0 0 0 0

Kingfisher 10 0 2 2 0 2 0

South Trail 0 0 0 0 0 0 0
 North Boundary A 2 1 0 0 1 0 0

North Boundary B 3 2 1 0 0 0 0

Table 1. Data from the first Central American Spider Monkey census, with information on life history
stages. The data are not enough for further analysis at this stage.

OVERALL DISCUSSION

The risk of the national park becoming an isolated forest is a real one especially with
widespread logging occurring in the buffer zone (McClearn 2016). Recent local discussions
about a potential road to the park have not alleviated these concerns (pers. Comm.). For
some species, the Great Green Macaw, Tapir and Peccary for example, rely heavily on
connected habitats to migrate seasonally with fruits and other available food sources
(Hooper 2008). The Baird’s tapir is a keystone browser and plays a crucial role seed dispersal
(Foerster & Vaughan 2015) and the white lipped peccary has begun to range further and
wider in search of key resources (Reyna-Hurtado et al. 2009). Similarly the Great Green
Macaw is an excellent indicator species which relies heavily on second growth corridors
linking habitat patches together. It suffered a great decline in the 1990s and it starting to
recover, but will only continue to do so with adequate connectivity as it follows its food
sources around seasonally (McClearn 2016). Early indications from this study show
encouraging results but connectivity to wider habitats and the enabling of movement
between them remain a key concern.

The main aim of the project was to gather and evaluate information on species richness and
abundance and this coupled with, direct comparison to the disturbed habitat has been
invaluable for achieving this aim. As the project progresses and the disturbed habitat
changes this project will produce increasingly valuable insights and offers great potential for
individual species studies in the future.

RECOMMENDATIONS
Continued monitoring of the forest habitats and trails is recommended in order to achieve a
long term study in the Jalova area of Tortuguero National Park. This long term monitoring
will enable the early detection of potential risks to the area such as the increase in
accidental tourists, hunting pressures and the spread of invasive species. The monitoring
project coupled now with more accurate weather recording abilities allow us to observe any
future affects from climate change over the long term period.

This year two incidences of tourism in the Jalova area have been encountered and it should
be made clear to all lodges that this area of the park is only for those with scientific permits
and not for general visitors. This increase since the park first banned tourism in this area
will be monitored and recorded going forwards to ascertain the need for further measures.

Vertebrate studies can only take us so far in understanding regeneration of the park. To
really assess the secondary growth it is recommended that a full vegetational study be
undertaken by an expert in the field. This data combined with the work on vertebrate
species and butterflies will give a full picture into the state of ecosystem. This will also allow
for the monitoring of the spread of invasive flora species too which in turn could help
horizon scan for invasive vertebrate species, especially amphibians as they are known to
spread fungus and could decimate extant populations.

It would also be beneficial to lengthen one of the trails here in the area. North Boundary B is
the shortest trail but it is a valuable habitat to research due to its biotic characteristics. The
swamp changes constantly and alongside this the amphibian population and the species
which prey on them do too. As found in this report, South Boundary is one of the most
diverse habitats but this cannot be said of North Boundary B because of the lack data
compared to the other trails despite it being the same habitat type. Having a more
comparable length of the trail will we believe alter the results from this survey transect and
bring it more in line with South Boundary and the results we see from that trail, being one
of the most biodiverse. Lengthening it using the old trail which used to connect it to South
Boundary would have little detrimental effect on the forest and would be low impact.

A previous report for the incidental project noted the fact that we rely on tracks for the
recording of cryptic mammal species and recommended the use of camera trapping to
increase our recording of them, but after consultation with our partner project Coastal
Jaguar Conservation it was concluded that it would be duplicating data and the use of more
camera traps within the park was not going to be beneficial so this was not carried out.
BUTTERFLY PROJECT
INTRODUCTION

Lepidoptera are a very diverse taxon occupying a large variety of niches. They are highly
sensitive to environmental change, including climate change, farmland abandonment and
habitat fragmentation (Thomas, 2005). As such, they are one of the best groups to study to
monitor change in biodiversity. Hundreds of butterfly species are reported on the Caribbean
slope of Costa Rica (DeVries, 1983). However, no previous effort has been made to record
the species found at Jalova. The start of the Butterfly Biodiversity project in January to June
2018 aims to build a species inventory for the area as a short-term objective.

Multiple studies have used butterfly and bird communities when comparing habitats with
varying degrees of disturbance (Ernst et al. 2017; Posa & Sodhi, 2006). To properly assess
the regrowth of the forest in Tortuguero National Park, monitoring both these groups is
essential. As a secondary and long-term objective, the first permit period of this project
aimed to build a database of recorded butterfly species abundance. Over time, trends will
become discernable and will lead to inferring on the status of the regeneration of the forest
around Jalova. Eventually, both the data obtained from the monitoring of bird and butterfly
diversity will be used to strengthen the assessment of the area.

METHODOLOGY

For full methodology please see: Hawkins, V. & Guy, D. 2017. Anteproyecto 2017 Inventario
de biodiversidad en el área de Jalova.

RESULTS AND DISCUSSION

In total, 23 sweep netting surveys averaging 40 minutes per survey were conducted during
the permit period of January to June 2018. 193 specimens were caught for identification.
142 of these were identified to the species level with an average of 71.8 % identification
success per survey. Every species caught was released successfully with handling time
averaging 43 seconds per specimen. From these surveys, a species inventory was made,
family abundances calculated and population trends were observed.
Species Inventory

During the first six months of the project, 18 butterfly species were identified in the survey
area (Appendix 2). The species belong to the families Nymphalidae, Pieridae and
Papilionidae. Due to a lack of butterfly identification resources, only the specimens
belonging to the Nymphalidae, Pieridae, Papilionidae and Riodinidae families could have
been identified to species level. No specimen was identified as being part of the Riodinidae
family but several others were identified as Hesperiidae and Lycaenidae. The photographs
of such specimens are being kept on file for future identification once the appropriate
identification resources are acquired. For this reason, diversity indices were not calculated
for this period but will be discussed in the annual report of December 2018. Species that
were caught on survey are reported with an average abundance for each of the trails on
which they were caught. Some specimens were caught incidentally during the training of
new volunteers and are also reported in the species list, without a calculated average
abundance.

As shown in the appendix, the abundance of every species caught on the trails surveyed was
obtained. Most of the species’ reported habitat preferences coincide with the trails on
which they were caught. For example, Cissia hermes and Eurema nise, associated with
pastures and disturbed habitats, were found in relatively high abundance on the trail in the
coconut plantation. The butterflies found on South Trail are all associated with wet,
disturbed or second growth forest which coincides perfectly with the type of habitat of that
surveyed area. South Trail is maintained as a trail used by motorized vehicles, surrounded by
a forest of regrowth and some temporal ponds. The specimens found on the beach trail are
also classified as disturbed and secondary growth habitat dwellers as the surveyed area
borders such habitats (the coconut plantation and part of the regenerating coastal forest).
The preliminary results obtained from the current monitoring project demonstrate the
efficiency of the use of butterfly species as bioindicators. The continuation of this project
would contribute in the monitoring of the forest as it regenerates. As sampling efforts
increase and different sampling methods are used, butterflies associated with primary
forest habitats could be caught. For example, Cissia alcinoe, a primary rainforest dweller,
was caught on the beach and in the coconut plantation. Its low abundance in these areas
indicates that the captures were possibly chance encounters of this species during localized
daily migrations. These captures, if they occur in increasing frequencies, would allow us to
infer that Tortuguero National Park is gradually regenerating into the pristine habitat it once
was.

Family Abundance

All butterfly specimens that were caught were identified to family level and further. Figure
16 shows the average abundance of every family in the three sample areas. Nymphalids
were found to be the most abundant group followed by Hesperiids and Pierids. The
Lycaenids’ and Papilionids’ relatively low abundance could be explained by the lack of
suitable vegetation in the sampled area or could be an example of population seasonality of
their representative species in the area. Continued sampling using this method might help
to display this seasonality. Another avenue that could be explored by this project is sampling
of the vegetation in the area as an explanatory factor for butterfly abundance and species
composition at Jalova. As the forest regenerates, new plant species will grow and different
butterfly groups will become more abundant.

The coconut plantation had the highest average abundance for all families which can be
explained by the ease of catching butterflies in more open areas compared to areas that are
restricted by dense vegetation such as South Trail. The trail located on the beach may not be
a suitable habitat for a lot of species in the area which would explain the low family
abundance when compared to the two other trails. The continuation of the current study
will allow for a great sample size to be obtained and stronger inferences to be made.

Population Trends

A long-term objective of the study is to monitor population trends over time. The first six
months of the study allowed us to observe peaks in abundance for two common species
found in the area (Anartia jatrophae and Cissia hermes). Figure 17 displays these trends. The
frequent peaks correspond with the typically short generation times exhibited by
butterflies. The end of May had an abnormally large peak for both species. Many
environmental factors may be used to explain this peak such as temperature, humidity,
rainfall or food source availability. All of these factors may be quantified and included in a
multivariate model as the project progresses.
RECOMMENDATIONS

The preliminary results of the Butterfly Biodiversity project display the strong potential of
such a study. The continuation of the project would allow for more species to be added to
the inventory. Trends in populations and species composition would also be observable
after a certain amount of time. To increase the amount of species in the inventory more
efficiently, bait traps used to attract fruit-feeding butterflies could be installed weekly in the
coconut plantation and on South Trail. Multiple benefits would come from such a sampling
method such as the addition of species recorded and trends in population, species
composition and assemblages. To know more about the benefits and protocol of a bait trap
study at Jalova, refer to the project proposal (Hawkins & Guy 2018).

Average Family Abundance per Trail

14 9.18

12

10
ABUNDANCE

4 2.27 2.14
1 1.09 1.36 0.8
2 0.86 1
0.29
0 0.09 0 0 0
0
Hesperiidae Lycanidae Nymphalidae Papilionidae Pieridae
-2
FAMILY

Coconut Plantation South Trail Beach

Figure 16. Average abundance of butterfly families in the 3 trails sampled with 95% confidence
intervals.
 25

20
Total Abundance

15

10

Anartia jatrophae Cissia hermes

Figure 17. Trends in abundance of Anartia jatrophae and Cissia hermes over the first 6-month
period of the study.

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APPENDICES
APPENDIX 1

The full list of all vertebrate species recorded in the Jalova area of Tortuguero National Park
from 2010 until 8th June 2018 (the end of the permit period).
Species Name Scientific Name IUCN status
Acadian Flycatcher Empidonax virescens Least concern
Agami Heron Agamia agami Vulnerable (A3c)
Alder Flycatcher Empidonax alnorum Least concern
Allen's Coralsnake Micrurus alleni Least concern
Alston's Mouse Opossum Micoureus alstoni Least concern
Amazon Kingfisher Chloroceryle amazona Least concern
American Coot Fulica americana Least concern
American Crocodile Crocodylus acutus Vulnerable (A2cd)
American Golden-Plover Pluvialis dominica Least concern
American Pygmy Kingfisher Chloroceryle aenea Least concern
American Redstart Setophaga ruticilla Least concern
Anhinga Anhinga anhinga Least concern
Annulated Tree Boa Corallus annulatus Not Yet assessed
Armored Rat Hoplomys gymnurus Least concern
Baird's Tapir Tapirus bairdii Endangered (A2abcd, 3bce)
Baltimore Oriole Icterus galbula Least concern
Bananaquit Coereba flaveola Least concern
Band-backed Wren Campylorhynchus zonatus Least concern
Band-tailed Barbthroat Threnetes ruckeri Least concern
Band-tailed Pigeon Patagioenas fasciata Least concern
Bank Swallow Riparia riparia Least concern
Bare-crowned Antbird Gymnocichla nudiceps Least concern
Bare-throated Tiger-Heron Tigrisoma mexicanum Least concern
Barn Swallow Hirundo rustica Least concern
Barred Antshrike Thamnophilus doliatus Least concern
Bat Falcon Falco rufigularis Least concern
Bay Wren Thryothorus nigricapillus Least concern
Bay-breasted Warbler Dendroica castanea Least concern
Belted Kingfisher Megaceryle alcyon Least concern
Bicolored Hawk Accipiter bicolor Least concern
Bicoloured Antbird Gymnopithys leucaspis Least concern
Black Myotis Myotis nigricans Least concern
Black Rat Rattus rattus Least concern
Black River Turtle Rhinoclemmys funerea Lower risk/ near threatened
Black Skimmer Rynchops niger Least concern
Black Tern Chlidonias niger Least concern
Black Vulture Coragyps atratus Least concern
Black-and-white Owl Ciccaba nigrolineata Least concern
Black-and-White Warbler Mniotilta varia Least concern
Black-bellied Plover Pluvialis squatarola Least concern
Black-bellied Whistling-duck Dendrocygna autumnalis Least concern
Blackburnian Warbler Dendroica fusca Least concern
Black-capped Pygmy-Tyrant Myiornis atricapillus Least concern
Black-cheeked Woodpecker Melanerpes pucherani Least concern
Black-collared Hawk Busarellus nigricollis Least concern
Black-cowled Oriole Icterus prosthemelas Least concern
Black-crowned Night-Heron Nycticorax nycticorax Least concern
Black-crowned Tityra Tityra inquisitor Least concern
Black-faced Grosbeak Caryothraustes poliogaster Least concern
Black-headed Saltator Saltator atriceps Least concern
Black-headed Tody-flycatcher Todirostrum nigriceps Least concern
Black-necked Stilt Himantopus mexicanus Least concern
Blackpoll Warbler Setophaga striata Least concern
Black-striped Sparrow Arremonops conirostris Least concern
Black-throated Trogon Trogon rufus Least concern
Blue Dacnis Dacnis cayana Least concern
Blue Ground-dove Claravis pretiosa Least concern
Blue-black Grassquit Volatinia jacarina Least concern
Blue-black Grosbeak Cyanocompsa cyanoides Least concern
Blue-chested Hummingbird Amazilia amabilis Least concern
Blue-gray Tanager Thraupis episcopus Least concern
Blue-throated Goldentail Hylocharis eliciae Least concern
Blue-winged Teal Anas discors Least concern
Blue-Winged Warbler Vermivora pinus Least concern
Boa Constrictor Boa constrictor Not Yet assessed
Boat-billed Flycatcher Megarynchus pitangua Least concern
Boat-billed Heron Cochlearius cochlearius Least concern
Bottlenose Dolphin Tursiops truncatus Least concern
Boulenger's Snouted Treefrog Scinax boulengeri Least concern
Bransfords Litterfrog Craugastor bransfordii Least concern
Bright-rumped Attila Attila spadiceus Least concern
Broad-billed Motmot Electron platyrhynchum Least concern
Broad-winged Hawk Buteo platypterus Least concern
Bronze Parrotsnake Leptophis mexicanus Least concern
Bronze-Backed Climbing-Skink Mabuya unimarginata Least concern
Bronzed Cowbird Molothrus aeneus Least concern
Bronze-tailed Plumeleteer Chalybura urochrysia Least concern
Bronzy Hermit Glaucis aeneus Least concern
Brown Booby Sula leucogaster Least concern
Brown Four-eyed Opossum Metachirus nudicaudatus Least concern
Brown Jay Psilorhinus morio Least concern
Brown Noddy Anous stolidus Least concern
Brown Pelican Pelecanus occidentalis Least concern
Brown Three-toed Sloth Bradypus variegatus Least concern
Brown Vinesnake Oxybelis aeneus Not Yet assessed
Brown Wood Turtle Rhinoclemmys annulata Lower risk/ near threatened
Brown-capped Tyrannulet Ornithion brunneicapillus Least concern
Brown-hooded Parrot pyrilia haematotis Least concern
Buff-throated Foliage-gleaner Automolus ochrolaemus Least Concern
Buff-throated Saltator Saltator maximus Least concern
Canada Warbler Wilsonia canadensis Least concern
Cape May Warbler Dendroica tigrina Least concern
Carpenter's Anole Anolis carpenteri Least concern
Casque-headed Lizard Corytophanes cristatus Not Yet assessed
Cattle Egret Bubulcus ibis Least concern
Central American Agouti Dasyprocta punctata Least concern
Central American Coralsnake Micrurus nigrocinctus Least concern
Central American Spider Monkey Ateles geoffroyi Endangered (A2c)
Central American Whiptail Holcosus festivus Least concern
Central American Wooly Opossum Caluromys derbianus Least concern
Checker-throated Antwren Epinecrophylla fulviventris Least concern
Chestnut-backed Antbird Myrmeciza exsul Least concern
Chestnut-coloured Woodpecker Celeus castaneus Least concern
Chestnut-headed Oropendola Psarocolius wagleri Least concern
Chestnut-mandibled Toucan Ramphastos ambiguus near threatened
Chestnut-sided Warbler Dendroica pensylvanica Least concern
Chimney Swift Chaetura pelagica near threatened
Chuck-will's-widow Antrostomus carolinensis Least concern
Cinnamon Becard Pachyramphus cinnamomeus Least concern
Cinnamon Woodpecker Celeus loricatus Least concern
Clay-colored Rainfrog Pristimantis cerasinus Least concern
Clay-colored Robin Turdus grayi Least concern
Cliff Swallow Petrochelidon pyrrhonota Least concern
Cloudy Slugeater Sibon nebulatus Not Yet assessed
Cocoa Woodcreeper Xiphorhynchus susurrans Least concern
Collared Aracari Pteroglossus torquatus Least concern
Collared Forest-Falcon Micrastur semitorquatus Least concern
Collared Peccary Pecari tajacu Least concern
Collared Plover Charadrius collaris Least concern
Common Black-Hawk Buteogallus anthracinus Least concern
Common Blunthead Imantodes cenchoa Not Yet assessed
Common Mexican Treefrog Smilisca baudinii Least concern
Common Nighthawk Chordeiles minor Least concern
Common Opossum Didelphis marsupialis Least concern
Common Pauraque Nyctidromus albicollis Least concern
Common Potoo Nyctibius griseus Least concern
Common Tent-making Bat Uroderma bilobatum Least concern
Common Tern Sterna hirundo Least concern
Common Tody-Flycatcher Todirostrum cinereum Least concern
Common Yellowthroat Geothlypis trichas Least concern
Costa Rican Coralsnake Micrurus mosquitensis Least concern
Crane Hawk Geranospiza caerulescens Least concern
Crested Caracara Caracara cheriway Least concern
Crested Eagle Morphnus guianensis near threatened
Crested Guan Penelope purpurascens Least concern
Crested Owl Lophostrix cristata Least concern
Crimson-fronted Parakeet Aratinga finschi Least concern
Dickcissel Spiza americana Least Concern
Dot-winged Antwren Microrhopias quixensis Least concern
Double-toothed Kite Harpagus bidentatus Least concern
Drab Treefrog Smilisca sordida Least concern
Dusky Antbird Cercomacra tyrannina Least concern
Dusky Rice Rat Melanomys caliginosus Least concern
Dusky-capped Flycatcher Myiarchus tuberculifer Least concern
Dusky-faced Tanager Mitrospingus cassinii Least concern
Eastern Kingbird Tyrannus tyrannus Least concern
Eastern Wood-pewee Contopus virens Least concern
Elegant Littersnake Rhadinaea decorata Not Yet assessed
Eyelash Palm Pitviper Bothriechis schlegelii Not Yet assessed
Fer-de-Lance Bothrops asper Not Yet assessed
Fitzinger's Rain Frog Craugastor fitzingeri Least concern
Forest Spiny Pocket Mouse Heteromys demarestianus Least concern
Four-lined Whiptail Holcosus quadrilineatus Least concern
Fringe-toed Foamfrog Leptodactylus melanonotus Least concern
Giant Cowbird Molothrus oryzivorus Least concern
Golden spectacled lizard Gymnophthalmus speciosus Least concern
Golden-hooded Tanager Tangara larvata Least concern
Gray-cheeked Thrush Catharus minimus Least concern
Gray-crowned Yellowthroat Geothlypis poliocephala Least concern
Gray-headed Chachalaca Ortalis cinereiceps Least concern
Great Black-Hawk Buteogallus urubitinga Least concern
Great Blue Heron Ardea herodias Least concern
Great Curassow Crax rubra Vulnerable (A2cd, 3cd,4cd)
Great Egret Ardea alba Least concern
Great Green Macaw Ara ambiguus Endangered (A2cd,3cd,4cd)
Great Kiskadee Pitangus sulphuratus Least concern
Great Potoo Nyctibius grandis Least concern
Great Tinamou Tinamus major near threatened
Great-crested Flycatcher Myiarchus crinitus Least concern
Greater Ani Crotophaga major Least concern
Greater Grison Galictis vittata Least concern
Greater White-Lined Bat Saccopteryx bilineata Least concern
Greater Yellowlegs
Great-tailed Grackle
Green Basilisk
Green Climbing Toad
Green Heron
Green Honeycreeper
Green Ibis
Green Iguana
Green Kingfisher
Green Parrotsnake
Green Tree Anole
Green Turtle
Green Vinesnake
Green-and-Rufous Kingfisher
Green-breasted Mango
Grey Catbird
Grey Four-eyed Opossum
Grey Kingbird
Grey-breasted Martin
Grey-capped Flycatcher
Grey-chested Dove
Grey-headed Kite
Greyish Saltator
Grey-necked Wood-rail
Grey-rumped Swift
Groove-billed Ani
Ground Anole
Gull-billed Tern
Halloween Snake
Hawksbill
Herring Gull
Hoffman´s Two-toed Sloth
Hooded Warbler
Hook billed Kite
Hourglass Treefrog
House Gecko
House Wren
Jaguar
Keel-billed Toucan
Kentucky Warbler
Killdeer
King Vulture
Kinkajou
Laughing Falcon
Laughing Gull
Tringa melanoleuca Least concern
Quiscalus mexicanus Least concern
Basiliscus plumifrons Least concern
Incilius coniferus Least concern
Butorides virescens Least concern
Chlorophanes spiza Least concern
Mesembrinibis cayennensis Least concern
Iguana iguana Not Yet assessed
Chloroceryle americana Least concern
Leptophis ahaetulla Not Yet assessed
Anolis biporcatus Not Yet assessed
Chelonia mydas Endangered (A2bd)
Oxybelis fulgidus Not Yet assessed
Chloroceryle inda Least concern
Anthracothorax prevostii Least concern
Dumetella carolinensis Least concern
Philander opossum Least concern
Tyrannus dominicensis Least concern
Progne chalybea Least concern
Myiozetetes granadensis Least concern
Leptotila cassini Least concern
Leptodon cayanensis Least concern
Saltator coerulescens Least concern
Aramides cajanea Least concern
Chaetura cinereiventris Least concern
Crotophaga sulcirostris Least concern
Anolis humilis Not Yet assessed
Gelochelidon nilotica Least concern
Pliocercus euryzonus Least concern
Eretmochelys imbricata Critically endangered (A2bd)
Larus argentatus Least concern
Choloepus hoffmanni Least concern
Wilsonia citrina Least concern
Chondrohierax uncinatus least concern
Dendropsophus ebraccatus Least concern
Hemidactylus frenatus Least concern
Troglodytes aedon Least concern
Panthera onca near threatened
Ramphastos sulfuratus Least concern
Oporornis formosus Least concern
Charadrius vociferus Least concern
Sarcoramphus papa Least concern
Potos flavus Least concern
Herpetotheres cachinnans Least concern
Larus atricilla Least concern
Leaf-breeding Rainfrog Pristimantis caryophyllaceus near threatened
Least Bittern Ixobrychus exilis Least concern
Least Flycatcher Empidonax minimus Least concern
Least Grebe Tachybaptus dominicus Least concern
Least Sandpiper Calidris minutilla Least concern
Least Tern Sternula antillarum Least concern
Leatherback Dermochelys coriacea Vulnerable (A2bd)
Lemur Anole Norops lemurinus Not Yet assessed
Lesser Greenlet Hylophilus decurtatus Least concern
Lesser Nighthawk Chordeiles acutipennis Least concern
Lesser Swallow-tailed Swift Panyptila cayennensis Least concern
Lesser Yellowlegs Tringa flavipes Least concern
Lichen Anole Anolis pentaprion Not Yet assessed
Lichen Coloured Snaileater Sibon longifrenis least concern
Limpkin Aramus guarauna Least concern
Lineated Woodpecker Dryocopus lineatus Least concern
Litter Gecko Lepidoblepharis xanthostigma Least concern
Litter Skink Sphenomorphus cherriei Least concern
Little Blue Heron Egretta caerulea Least concern
Little Tinamou Crypturellus soui Least concern
Loggerhead Caretta caretta Vulnerable (A2b)
Long-billed Gnatwren Ramphocaenus melanurus Least concern
Long-billed Hermit Phaethornis longirostris Least concern
Long-billed Starthroat Heliomaster longirostris Least concern
Long-tailed Tyrant Colonia colonus Least concern
Louisiana Waterthrush Parkesia motacilla Least concern
Lowland Rainfrog Craugastor ranoides Critically endangered (A2ace)
Magnificent Frigatebird Fregata magnificens Least concern
Magnolia Warbler Dendroica magnolia Least concern
Mangrove Cuckoo Coccyzus minor Least concern
Mangrove Swallow Tachycineta albilinea Least concern
Mantled Howler Monkey Alouatta palliata Least concern
Many-banded Coralsnake Micrurus mipartitus Not Yet assessed
Marine Toad Rhinella marina Least concern
Masked Tityra Tityra semifasciata Least concern
Masked Treefrog Smilisca phaeota Least concern
Mealy Parrot Amazona farinosa near threatened
Melodious Blackbird Dives dives Least concern
Mexican Prehensile-tailed Porcupine Sphiggurus mexicanus Least concern
Mississippi Kite Ictinia mississippiensis Least concern
Montezuma Oropendola Psarocolius montezuma Least concern
Mourning Warbler Oporornis philadelphia Least concern
Muscovy Duck Cairina moschata Least concern
Narrow-bridged Mud-turtle Kinosternon angustipons Vulnerable (B1, 2c)
Neotropical Chameleon Polychrus gutturosus Not Yet assessed
Neotropical Cormorant Phalacrocorax brasilianus Least concern
Neotropical River Otter Lutra longicaudis near threatened
Nicaraguan Seed-Finch Oryzoborus nuttingi Least concern
Nine-banded Armadillo Dasypus novemcinctus Least concern
Noble's Rainfrog Craugastor noblei Least concern
Northern Barred Woodcreeper Dendrocolaptes sanctithomae Least concern
Northern Bird-eating Snake Pseustes poecilonotus Least concern
Northern Cateye Leptodeira septentrionalis Not Yet assessed
Northern Jacana Jacana spinosa Least concern
Northern Masked Rainfrog Craugastor mimus Least concern
Northern Parula Parula americana Least concern
Northern Raccoon Procyon lotor Least concern
Northern Rough-winged Swallow Stelgidopteryx serripennis Least concern
Northern Tamandua Tamandua mexicana Least concern
Northern Waterthrush Parkesia noveboracensis Least concern
Ocelot Leopardus pardalis Least concern
Ochre-bellied Flycatcher Mionectes oleagineus Least concern
Olingo Bassaricyon gabbii Least concern
Olive Snouted Treefrog Scinax elaeochrous Least concern
Olive-backed Euphonia Euphonia gouldi Least concern
Olive-backed Quail-Dove Geotrygon veraguensis Least concern
Olive-crowned Yellowthroat Geothlypis semiflava Least concern
Olive-sided Flycatcher Contopus cooperi near threatened
Olive-throated Parakeet Aratinga nana Not Yet assessed
Orange-bellied Littersnake Urotheca guentheri Least concern
Orange-Bellied Swamp Snake Tretanorhinus nigroluteus Not Yet assessed
Orange-billed Sparrow Arremon aurantiirostris Least concern
Orange-chinned Parakeet Brotogeris jugularis Least concern
Orchard Oriole Icterus spurius Least concern
Osprey Pandion haliaetus Least concern
Ovenbird Seiurus aurocapilla Least concern
Paca Cuniculus paca Least concern
Pale-billed Woodpecker Campephilus guatemalensis Least concern
Pale-vented Pigeon Patagioenas cayennensis Least concern
Palm Tanager Thraupis palmarum Least concern
Paltry Tyrannulet Zimmerius vilissimus Least concern
Passerini's Tanager Ramphocelus passerinii Least concern
Pectoral Sandpiper Calidris melanotos Least concern
Peregrine Falcon Falco peregrinus Least concern
Pied-billed Grebe Podilymbus podiceps Least concern
Piratic Flycatcher Legatus leucophaius Least concern
Plain Wren Thryothorus modestus Least concern
Plain-brown Woodcreeper Dendrocincla fuliginosa Least concern
Plain-coloured Tanager Tangara inornata Least concern
Prothonotary Warbler Protonotaria citrea Least concern
Pug-nosed Anole Anolis capito Not Yet assessed
Purple Caecilian Gymnopis multiplicata Least concern
Purple Gallinule Porphyrio martinicus Least concern
Purple Martin Progne subis Least concern
Purple-crowned Fairy Heliothryx barroti Least concern
Purple-throated Fruitcrow Querula purpurata Least concern
Pygmy Rainfrog Pristimantis ridens Least concern
Red Brocket Deer Mazama americana data deficient
Red Coffeesnake Ninia sebae Least concern
Red-billed Pigeon Patagioenas flavirostris Least concern
Red-capped Manakin Pipra mentalis Least concern
Red-eyed Treefrog Agalychnis callidryas Least concern
Red-eyed Vireo Vireo olivaceus Least concern
Red-lored Parrot Amazona autumnalis Least concern
Red-ringed Snaileater Sibon annulatus Least concern
Red-tailed Squirrel Sciurus granatensis Least concern
Red-throated Ant-Tanager Habia fuscicauda Least concern
Resplendent Quetzal Pharomachrus mocinno near threatened
Reticulated Crowned Snake Tantilla reticulata Least concern
Reticulated Sheepfrog Gastrophryne pictiventris Least concern
Ringed Kingfisher Megaceryle torquata Least concern
Ring-necked Duck Aythya collaris Least concern
Roadside Hawk Rupornis magnirostris Least concern
Roseate Spoonbill Platalea ajaja Least concern
Rose-breasted Grosbeak Pheucticus ludovicianus Least concern
Rose-throated Becard Pachyramphus aglaiae Least Concern
Royal Flycatcher Onychorhynchus coronatus Least concern
Royal Tern Thalasseus maximus Least concern
Ruddy Ground-Dove Columbina talpacoti Least concern
Ruddy Quail Dove Geotrygon montana Least concern
Ruddy Turnstone Arenaria interpres Least concern
Ruddy-tailed Flycatcher Terenotriccus erythrurus Least concern
Rufescent Tiger-Heron Tigrisoma lineatum Least concern
Rufous Mourner Rhytipterna holerythra Least concern
Rufous Piha Lipaugus unirufus Least concern
Rufous-tailed Hummingbird Amazilia tzacatl Least concern
Rufous-Winged Woodpecker Piculus simplex Least concern
Rugose littersnake Nothopsis rugosus Least concern
Salmon-bellied Racer Mastigodryas melanolomus Least concern
San Carlos Treefrog Dendropsophus phlebodes Least concern
Sanderling Calidris alba Least concern
Sandwich Tern Thalasseus sandvicensis Least concern
Satany Parrotsnake Leptophis depressirostris Not Yet assessed
Savages Thin-toed Frog Leptodactylus pentadactylus Least concern
Scarlet Tanager Piranga olivacea Least concern
Scarlet-webbed Tree Frog Hypsiboas rufitelus Least concern
Semipalmated Plover Charadrius semipalmatus Least concern
Semipalmated Sandpiper Calidris pusilla near threatened
Semiplumbeous Hawk Leucopternis semiplumbeus Least concern
Shining Honeycreeper Cyanerpes lucidus Least concern
Shiny Cowbird Molothrus bonariensis Least concern
Short-billed Dowitcher Limnodromus griseus Least concern
Short-billed Pigeon Patagioenas nigrirostris Least concern
Short-tailed Hawk Buteo brachyurus Least concern
Short-tailed Nighthawk Lurocalis semitorquatus Least concern
Slaty-backed Forest-Falcon Micrastur mirandollei Least concern
Slaty-headed tody-flycatcher Poecilotriccus sylvia Least concern
Slaty-tailed Trogon Trogon massena Least concern
Slender Anole Anolis limifrons Not Yet assessed
Slender Caecilian Dermophis parviceps Least concern
Smokey-brown Woodpecker Leuconotopicus fumigatus Least concern
Snowy Cotinga Carpodectes nitidus Least concern
Snowy Egret Egretta thula Least concern
Snowy Plover Charadrius nivosus near threatened
Social Flycatcher Myiozetetes similis Least concern
Song Wren Cyphorhinus phaeocephalus Least concern
South Polar Skua Catharacta maccormicki Least concern
Southern Rough-winged Swallow Stelgidopteryx ruficollis Least concern
Speckled Racer Drymobius margaritiferus Not Yet assessed
Spectacled Caiman Caiman crocodilus lower risk/ least concern
Spectacled Owl Pulsatrix perspicillata Least concern
Spix's Disc-winged Bat Thyroptera tricolor Least concern
Spotted Antbird Hylophylax naevioides Least concern
Spotted Sandpiper Actitis macularius Least concern
Spotted Woodcreeper Xiphorhynchus erythropygius Least concern
Squirrel Cuckoo Piaya cayana Least concern
Streak-crowned Antvireo Dysithamnus striaticeps Least concern
Streak-headed Woodcreeper Lepidocolaptes souleyetii Least concern
Stream Anole Anolis oxylophus Not Yet assessed
Stripe-breasted Wren Thryothorus thoracicus Least concern
Striped Basilisk Basiliscus vittatus Not Yet assessed
Stripe-headed Sparrow Peucaea ruficauda Least concern
Stripe-throated Hermit Phaethornis striigularis Least concern
Sulphur-bellied Flycatcher Myiodynastes luteiventris Least concern
Summer Tanager Piranga rubra Least concern
Sunbittern Eurypyga helias Least concern
Sungrebe Heliornis fulica Least concern
Swainson's Hawk Buteo swainsoni Least concern
Swainson's Thrush Catharus ustulatus Least concern
Swallow-tailed Kite Elanoides forficatus Least concern
Talamancan Galliwasp Diploglossus bilobatus Least concern
Talamancan Rain Frog Craugastor talamancae Least concern
Tawny Treefrog Smilisca puma Least concern
Tawny-chested Flycatcher Aphanotriccus capitalis vulnerable (B1abc - i,ii,iii,v)
Tawny-faced Gnatwren Microbates cinereiventris Least concern
Taylor's Leopard Frog Lithobates taylori Least concern
Tayra Eira barbara Least concern
Tennessee Warbler Vermivora peregrina Least concern
Thick-Billed Seed-Finch Oryzoborus funereus Least concern
Tiger Ratsnake Spilotes pullatus Not Yet assessed
Tink Frog Diasporus diastema Least concern
Tiny Hawk Accipiter superciliosus Least concern
Tome's spiny rat Proechimys semispinosus Least concern
Traill's Flycatcher Empidonax sp Least concern
Tree swallow Tachycineta bicolor Least Concern
Tricolored Heron Egretta tricolor Least concern
Tropical Gnatcatcher Polioptila plumbea Least concern
Tropical Kingbird Tyrannus melancholicus Least concern
Tropical Night Lizard Lepidophyma flavimaculatum Least concern
Tropical Pewee Contopus cinereus Least concern
Tropical seep snake Hydromorphus concolor Least concern
Tropical Slider Chrysemys ornata unknown
Turkey Vulture Cathartes aura Least concern
Turnip-tailed Gecko Thecadactylus rapicauda Not Yet assessed
Uniform Crake Amaurolimnas concolor Least concern
Vaillant's Frog Lithobates vaillanti Least concern
Variable Seedeater Sporophila corvina Least concern
Variegated Squirrel Sciurus variegatoides Least concern
Vesper Rat Nyctomys sumichrasti Least concern
Violaceous Trogon Trogon violaceus Least concern
Violet-crowned Woodnymph Thalurania colombica Least concern
Watsons Climbing Rat Tylomys watsoni Least concern
West Indian Manatee Trichechus manatus vulnerable (C1)
Western Sandpiper Calidris mauri Least concern
Western Slaty Antshrike Thamnophilus atrinucha Least concern
Western Wood-pewee Contopus sordidulus Least concern
Wet Forest Toad Incilius melanochlorus Least concern
Whimbrel Numenius phaeopus Least concern
White Hawk Pseudastur albicollis Least concern
White-breasted Wood-wren Henicorhina leucosticta Least concern
White-collared Manakin Manacus candei Least concern
White-collared Swift Streptoprocne zonaris Least concern
White-crowned Parrot Pionus senilis Least concern
White-flanked Antwren Myrmotherula axillaris Least concern
White-lined Tanager Tachyphonus rufus Least concern
White-lipped Mud-turtle Kinosternon leucostomum Not Yet assessed
White-lipped Peccary Tayassu pecari Vulnerable (A2bcde 3bcde)
White-necked Jacobin Florisuga mellivora Least concern
White-necked Puffbird Notharchus hyperrhynchus Least concern
White-nosed Coati Nasua narica Least concern
White-ringed Flycatcher Conopias albovittatus Least concern
White-Tailed Deer Odocoileus virginianus Least concern
White-throated Capuchin Cebus capucinus Not Yet assessed
White-throated Crake Laterallus albigularis Least concern
White-tipped Dove Leptotila verreauxi Least concern
White-Vented Euphonia Euphonia minuta least concern
White-whiskered Puffbird Malacoptila panamensis Least concern
White-winged Becard Pachyramphus polychopterus Least concern
Willet Tringa semipalmata Least concern
Willow Flycatcher Empidonax traillii Least concern
Wilson's Plover Charadrius wilsonia Least concern
Wilson's Storm-Petrel Oceanites oceanicus Least concern
Wilson's Warbler Cardellina pusilla Least concern
Wood Thrush Hylocichla mustelina near threatened
Worm-eating Warbler Helmitheros vermivorum Least concern
Yellow Blunt-Headed Vinesnake Imantodes inornatus Least concern
Yellow Tyrannulet Capsiempis flaveola Least concern
Yellow Warbler Dendroica petechia Least concern
Yellow-bellied Elaenia Elaenia flavogaster Least concern
Yellow-bellied flycatcher Empidonax flaviventris Least concern
Yellow-bellied Sapsucker Sphyrapicus varius Least concern
Yellow-billed Cuckoo Coccyzus americanus Least concern
Yellow-crowned Euphonia Euphonia luteicapilla Least concern
Yellow-crowned Night-heron Nyctanassa violacea Least concern
Yellow-headed Caracara Milvago chimachima Least concern
Yellow-headed Gecko Gonatodes albogularis Not Yet assessed
Yellow-lipped Parrotsnake Leptophis nebulosus Least concern
Yellow-margined Flycatcher Tolmomyias assimilis Least concern
Yellow-olive Flycatcher Tolmomyias sulphurescens Least concern
Yellow-rumped Warbler Dendroica coronata Least concern
Yellow-tailed Dwarf-gecko Sphaerodactylus homolepis Least concern
Yellow-tailed Oriole Icterus mesomelas Least concern
Yellow-throated Euphonia Euphonia hirundinacea Least concern
Yellow-throated vireo Vireo flavifrons Least Concern
Yellow-throated Warbler Setophaga dominica Least concern
Zone-tailed Hawk Buteo albonotatus Least concern
APPENDIX 2

Species Name Date First Coconut South Trail Beach Average Habitat
Observed Plantation Average Abundance Preferences
Average Abundance (Devries, 1987)
Abundance

Adelpha celerio 08-06-2018 0.09 N/A N/A All forest habitats

Adelpha cytherea 22-02-2018 N/A 0.17 N/A All forest

marcia habitats;
secondary
growth

Aeria eurimedia 22-02-2018 N/A 0.5 N/A Wet forest

agna habitat

Anartia fatima 21-05-2018 0.64 N/A N/A All disturbed

habitat

Anartia jatrophae 28-01-2018 3.91 N/A N/A Coastal lowland

rain forest

Cissia alcinoe 28-01-2018 0.3 N/A 0.2 Primary

rainforest habitat

Cissia hermes 28-01-2018 3.82 0.5 0.6 All habitats; most

common in
pastures & open
areas.

Cissia usitata 09-06-2018 N/A 0.17 N/A Rain forest and

wet forest

Colobura dirce 19-02-2018 N/A N/A N/A All forest types;

especially
second-growth

Consul fabius 22-02-2018 N/A 0.17 N/A All forested

cecrops habitats

Dryas iulia 27-03-2018 0.18 N/A N/A Open areas and

forest canopy

Eurema nise 20-03-2018 1.27 N/A 0.6 Disturbed habitat

Heliconius cydno 22-02-2018 N/A 0.33 N/A Forest habitats

galanthus

Heliconius erato 30-01-2018 0.18 0.5 0.2 Disturbed habitat

petiverana & secondary
growth

Heliconius sara 30-01-2018 N/A N/A N/A Rain forest

fulgidus habitat;
 secondary
growth

Historis odius 19-02-2018 N/A N/A N/A All habitat types

Papilio 08-06-2018 N/A N/A N/A Disturbed

anchisiades habitats
idaeus

Phoebis argante 26-03-2018 0.09 N/A N/A Disturbed

habitats