This study examined the diversity and abundance of fruit-feeding butterflies across four habitat types in a Costa Rican cloud forest: primary forest, natural secondary regrowth forest, planted secondary regrowth forest, and pastureland. The researchers trapped 174 butterflies of 27 species over six weeks. They found that planted secondary regrowth forest had the highest species richness, diversity, and evenness, indicating reforestation efforts were improving diversity. Climate change may be causing butterflies to move to new elevations.

1. Duong and Junger, 2015
1
Frequency and Distribution of Fruit-Feeding Butterflies in a
Costa Rican Cloud Forest: From Grazeland to Primary Forest
Gabrielle
C.
Duong
&
Ashley
J.
Junger
Abstract
To determine whether the frequency and distribution of fruit-feeding Nymphalid
butterflies change through a habitat’s succession from pasture land to primary forest, as well as to
determine whether Cloudbridge’s reforestation efforts were expediting the growth of primary
forest butterfly populations, a community of fruit-feeding Nymphalid butterflies was sampled
daily for 6 weeks by trapping 174 individuals of 27 species in the understory of four habitat
types: primary forest, natural secondary regrowth forest, planted secondary regrowth forest, and
pastureland. We found the whole study area had a species evenness of 0.5, and a Simpson’s Index
of Diversity of 0.88. Planted regrowth had the highest species richness (20), diversity index (0.9),
and a relatively high evenness (0.58). Therefore, of the successional habitat types studied, planted
regrowth is the most diverse and rich, indicating that the process of planting climax species in
secondary forests improves community diversity and richness. This increase in community
diversity and richness may lead to higher diversity and richness in the climax community. We
also conclude that a large number of species are being found out of their natural elevation range,
which could indicate that butterflies in this area are experiencing the effects of climate change.
Key words: nymphalidae, butterfly, species abundance distribution, species richness,
environmental monitoring, habitat disturbance, tropical, conservation.
INTRODUCTION
Natural habitats in the tropics continue
to be globally threatened by habitat loss and
climate change, leading to the massive loss of
species. Remediation efforts are being
undertaken globally in an effort to reduce and
reverse the effects of habitat loss and climate
change. In order to focus these efforts and to
determine their effects on biological diversity,
reliable monitoring programs that assess
changes in biodiversity and ecosystem
function are needed. The choice of organism
investigated is crucial to this process due to
overall lack of funds and available expertise.
Trapping butterflies is often the
method of choice in tropical forests (Aduse-
Poku et al., 2012). Advantages of using
butterflies (Lepidoptera) as a target species
include: their relatively large size, colorful
appearance, relative ease of identification to
species level, their presence in all terrestrial
habitats, and sensitivity to microclimate
heterogeneity and disturbance (New, 1997;
DeVries et al., 1997). Butterflies are the best-
known group of insects (DeVries et al., 1997),
giving them great potential to provide
understanding of insect diversity and
conservation. The study of fruit feeding
butterflies has additional advantages,
including their ability to be vigorously
sampled with the use of fruit-baited traps.
Therefore, fruit-feeding butterflies provide a
standard means for comparing species
diversity within and among tropical insect
communities (DeVries et al., 2012).
Bait trapping is an inherently biased
method for assessing butterfly fauna. Issues
include: some butterflies are never captured in
traps, and some butterflies are more strongly
attracted than others, thus relative abundances
captured do not necessarily reflect the relative

2. Duong and Junger, 2015
2
abundances of fruit feeding species in the
region (Hughes et al., 1998). Despite these
drawbacks, bait trapping provides the most
effective and efficient method for monitoring
changes in species abundances, and measuring
diversity of tropical butterfly communities.
Reforestation is an essential step in
restoring forest health. Monitoring and
assessment of the effects of reforestation on
communities within remediated areas is
essential to gaining an understanding of its
short- and long-term effects. Few studies
investigate the changes in species abundance
and diversity through various levels of
succession in tropical forests. Most studies in
this area focus on the differences between
disturbed and undisturbed habitats. Disturbed
habitats have been found to have higher
species richness and more unique species
(DeVries et al., 1997). Additionally, vertical
stratification is reduced in disturbed forests,
trapping canopy species in the understory,
leading to overestimates of species richness in
understory trapping in disturbed areas
(DeVries & Walla, 2001; Fermon et al., 2005;
Aduse-Poku et al., 2012). Rapid monitoring
programs will allow assessments of the
changes in diversity and abundance within
different stages of succession, allowing for
more effective and focused remediation
efforts.
Between the 1940s and 1980s
deforestation in Costa Rica was caused by
government-sponsored land colonization
schemes, expansion of the agricultural
frontier, cattle ranching to support the beef
industry, and both legal and illegal timbering
(Borges-Méndez, 2008). During this period
the national area covered with forests dropped
from about 70% to about 10% (United States
Agency for International Development
[USAID], 1996). In fact, between 1950 and
1994, the pace of deforestation in Costa Rica
was one of the fastest in the western
hemisphere, with a decrease of 40,000-50,000
hectares annually (Watson et al., 1998;
Borges-Méndez, 2008). Given this rapid and
drastic loss of forested areas throughout the
country, reforestation programs will be
essential to restoring the country’s former
biodiversity.
Costa Rican forests have sufficiently
diverse fruit-feeding butterfly fauna to warrant
their use as target organisms for monitoring
changes in biodiversity. There are
approximately 543 butterfly species present in
Costa Rica (DeVries, 1987). Of these, at least
40% feed exclusively upon rotting fruits as
adults (DeVries, 1987).
The aim of this study is to quantify the
differences in diversity and abundance of
fruit-feeding butterflies within habitats at
different stages of succession. We performed
a butterfly trapping study on the Cloudbridge
Nature reserve in Costa Rica during the onset
of the rainy season, studying four habitat types
(grazeland, natural regrowth, planted
regrowth, and primary forest). In each of these
habitat types three traps were established in
the understory.
MATERIALS AND METHODS
Study Site
This research was conducted at the
Chirripó Cloudbridge Nature Reserve, San
Isidro de General, south central Costa Rica.
Located in a cloud forest on one of the tallest
mountains of Central America, Cloudbridge is
a 700 acre nature reserve on the northern end
of one of the most important biological zones
of all Central America, and lies within a
designated “biological hot spot” on the Meso-
American Biological Corridor. Cloudbridge is
part of an area of forested land that includes
over a million hectares spanning northern
Costa Rica and southern Panama. Together
with the adjoining La Amistad International
Park, Chirripó National Park is comprised of
the largest unspoiled forest in the country.
Cloudbridge started off as privately
owned land in 2002, owned by Ian Giddy and
Genevieve (Jenny) Giddy, who made the first
of many subsequent purchases of cattle farms
bordering the Chirripó National Park to
impede the appalling denuding and erosive
effects that cattle grazing has had on the land.

3. Duong and Junger, 2015
3
Since then, their reserve has grown to
encompass 700 acres of reclaimed pasture
land and is used to re-build a corridor where
deforestation has left a gap between the large
Chirripó National Park, and the smaller nature
reserve of 4,000 acres on the other side of the
river. Our study was conducted within a
contiguous patch of the Cloudbridge reserve
that formed a disturbance gradient composed
of 3 contiguous habitat types: primary forest,
natural secondary regrowth forest, and planted
secondary regrowth forest, as well as the
pasture of a nearby cow farm.
Trap Sites
Each habitat type was fitted with 3
butterfly traps, whose locations were selected
based on elevation, walking distance, and
accessibility (Appendix B). In steeper hiking
areas, such as in the primary forest (PF) and
planted regrowth (PR) forest, traps were
spaced, on average, 117 meters apart in
elevation. In flatter and lower areas, such as
on the grazeland (GL) and along the natural
regrowth (NR) forest, traps were spaced, on
average, 14 meters apart in elevation. All
latitude, longitude, and elevation
measurements were taken using GIS with an
accuracy of ± 15 meters.
Primary Forest
Also known as an old-growth forest, a
primary forest is one that has remained
essentially unmodified by human activity.
Additionally, they are generally comprised of
climax species, a composition achieved as a
result of unrestrained ecological processes.
We chose to place our traps along the
primary forest areas of the El Jilguero and El
Hectare Trails.
Natural Regrowth Forest
A natural regrowth, or secondary,
forest is defined here as one that has naturally
re-grown after a major disruption, natural or
man-made, such as the deforestation efforts in
Costa Rica between the 1940’s and 1980’s. A
secondary forest has regrown for a long
enough period of time such that the effects of
the disturbance are no longer evident. It is
distinguished from a primary forest by species
composition; a secondary regrowth forest has
not yet reached a climax community.
We chose to place our traps along the
natural regrowth areas of the River Trail. Trap
1 was located near the bench by the river, and
next to a very small, narrow stream that
crossed over the trail. Traps 2 and 3 along the
River Trail were farther from the river and did
not have any streams of water flowing past
them. Trap 2 was placed in an open pocket of
forest under a tree surrounded by plants that
produced fruit. Trap 3 was placed in a more
open section of forest under a tree; no fruit-
growing species were observed.
Planted Regrowth Forest
A Planted regrowth, or secondary,
forest is defined here as one that, in addition to
having naturally regrown after a major
disturbance, is replenished through
reforestation efforts. During reforestation,
climax species are manually planted to
facilitate the transition of a secondary forest
into what more closely resembles a primary
forest.
We chose to place our traps along the
planted regrowth areas of the El Jilguero Trail.
Grazeland
A grazeland is a grassy field suitable
for grazing by livestock. In our study, the
grazeland is the state of disturbance from
which secondary regrowth forests are
recovering. We chose to utilize farmer Marcos
Romero’s land, located just down the road
from the Cloudbridge Nature Reserve. The
grazeland shared one of its borders with a
coffee bean plantation; it is along this border
that some of our traps were located.
Trap number 1 was located by the
road, just within the gate that fenced off the
grazeland, and was suspended from a lime
tree. Trap number 2 was located farther into
the grazeland and was suspended from a tree
located just past a narrow stream that ran
through the land. Trap number 3 was
suspended from a tree located at the top of the
hill.
Study Community: Fruit-feeding Nymphalids
Adult butterflies can be divided into
two main feeding guilds. One guild obtains all
nutritional requirements by feeding on the
nectar of flowers; this guild includes most
species of Papilonidae, Pieridae, Lycaenidae,
Riodinidae, and some groups within

4. Duong and Junger, 2015
4
Nymphalidae. (DeVries, Murray, & Lande,
1997). The other guild meets all nutritional
requirements by feeding on the juices of
rotting fruits or plant sap; this guild is
comprised of certain subfamilies of the
Nymphalidae, such as Charaxinae, Morphinae,
Brassolinae, Satyrinae, and Nymphalinae
(DeVries et al, 1997). It is this second
butterfly guild, which we call fruit-feeding
nymphalids, that can be easily baited and
trapped by exploiting their feeding habits and
escape mechanism. For completeness, we note
that some species in the subfamily Ithomiinae,
Limenitidinae, and Apaturinae can
occasionally be found in fruit-traps, although
they typically feed on flower nectar.
Additionally, some species in the Hespieriidae
family of skippers can also be occasionally
found in fruit-traps; however, because they are
not strictly butterflies, they are excluded from
the data analyzed here.
Butterfly Trap Design and Construction
Loosely following the trap dimensions
and construction instructions outlined by
George Austin and Thomas Riley, 1995, we
constructed our traps to be approximately 80
cm tall and 13 inches in diameter, with the
base hanging 1 inch below the bottom of the
trap netting. Each trap was constructed using
the following materials and procedures:
• Two wire hoops: 13-inches in
diameter. Bend wire into circle and
connect the ends by hooking them
together. Clamp the loop shut with
pliers.
• For body of the trap, cut a piece of
mosquito netting 42-inches wide and
34-inches tall; sew ends together
along the 42-inch edges to produce a
“tube” of netting. Then sew one end
of the tube such that this end is 2
inches narrower in diameter with
respect to the wire hoops you just
made. (Illustration 1.B)
• For the top of the trap, cut a sheet of
plastic tarp approximately 4 inches
larger in diameter than the wire hoops
you made. Wrap and tape this over the
wire hoop. (Illustration 1.B)
A
B
C
D
Illustration 1. Trap design and construction

5. Duong and Junger, 2015
5
• For the top of the trap, cut a sheet of
plastic tarp approximately 4 inches
larger in diameter than the wire hoops
you made. Wrap and tape this over the
wire hoop. (Illustration 1.B)
• Fit the plastic covered hoop into the
narrow end of the tube and secure in
place by placing two 30-inch lengths
of wire in an “x” pattern under the
plastic-covered hoop; the wires should
support the plastic-covered hoop, and
the plastic-covered hoop should
support the body of the netting. The
ends of the wire should meet near the
center of the upper side of the plastic-
covered hoop. Fashion the ends into a
hoop or hook, to which you will tie
the suspending rope. Plastic across the
top should be flat. (Illustration 1.C)
• For trap base, sew the second wire
hoop directly onto the bottom end of
the tube. When hung, hoops should
lay parallel to one another and to the
ground. (Illustration 1.A)
• Attach four hooks fashioned out of
metal wire to the base, leaving
approximately 1 inch of space
between the base and the bottom edge
of trap netting (Illustration 1.A).
• Punch four holes into a plastic plate.
This will serve as the trap base.
• Tape a bait container in the center of
the trap base, and hang the base to the
trap via the four hooks. For bait
containers, we repurposed cream
cheese containers and tuna cans.
Field Methods
Within the study areas, each of the
four habitat types was fitted with three traps,
providing a total of twelve traps. These
understory traps were suspended such that the
bottom of the traps were approximately 1.3
meters above the ground, with the exception
of the pastureland traps, whose bottoms were
approximately 1.8 meters above the ground to
keep the curious residential cows from
destroying them. Traps in the pastureland were
suspended from thin ropes run over branches
of an emergent tree, such that the traps could
be raised and lowered from the ground without
disturbance. The free end of the rope was
fastened to a branch at least 2 meters high with
all excess rope either wrapped around the
branch or tucked away to prevent the cows
from chewing through the ropes. All other
traps were suspended from lower tree
branches and could be serviced directly.
Each Monday morning, traps were baited
with rotting bananas obtained free-of-charge
from the local village store. Bananas were
sprinkled with 1 teaspoon of dry yeast and 1
teaspoon of sugar, mixed and mashed, and
fermented for 24 hours in one large container
prior to use. On the last day of the weekly
four-day sampling period, bait was removed
from all traps, and traps were tied shut over
the weekend. New bait was made prior to the
subsequent sampling interval, and this
protocol repeated throughout the study, which
extended from 15 June 2015 to 24 July 2015.
When checking each trap, we first
cinched the middle section shut with string to
prevent any butterflies from escaping. Trapped
butterflies were then individually extracted via
a plastic bag and photographed within the bag.
Then, butterflies were handled so they could
be photographed outside of the bag; both
dorsal and ventral sides were photographed for
identification. Butterflies were then released to
the area in which they were found.
Information was first recorded in a field
notebook, and later transferred to a
spreadsheet to perform data analyses.
All butterflies were identified using
the DeVries butterfly field book, which
follows the more conservative estimates of
Ackery, is based upon the work of Ehrlich,
and represents a widely used, functional
classification of nymphalid subfamilies
(DeVries et al, 1997).
Bait Recipes
Different butterfly species use
different kinds of food sources to obtain the
nutrients they require to survive. Some species
are attracted to what is called stinky bait,
which includes rotting fish and other carrion.
Nymphalids, however, are attracted to sweet
bait, which includes overripe or rotting
bananas, mangos, and other fruit. More
specifically, Nymphalid butterflies are
attracted to the alcohol in sweet baits. We tried

6. Duong and Junger, 2015
6
two different sweet bait recipes: (1) Beer bait
and (2) Yeast bait.
1. Beer Bait
o 4 overripe/rotting bananas,
peeled and mashed
o Add 1 tablespoon sugar
o Add ⅓ cup beer
o Mix well
o Let ferment for 7 days
2. Yeast Bait
o 4 overripe/rotting bananas,
peeled and mashed
o Add 1 teaspoon sugar
o Add 1 teaspoon yeast
o Mix well
o Let ferment for 24 hours
On June 26th, 11 days into our study,
we switched from using the beer bait to using
the yeast bait for a few reasons: it was easier
to prepare, cheaper, and took less time to
ferment. We also found that the yeast bait
attracted slightly more butterflies than the beer
bait, as we started finding butterflies in traps
in which we had never before found
butterflies.
RESULTS
Over the course of 27 data collection
days, 174 individual butterflies were collected
(Figure 1). These individuals were represented
by 27 different species. All species captured
were members of the Nympalidae family;
44.4% of species captured belonged to the
Satyrinae subfamily, followed by the
subfamily Charaxinae (22.2%). Other
subfamilies captured include: Brassolinae
(14.81%), Nymphaline (7.4%), Ithomiinae
(3.7%), Opoptera (3.7%), and Pycina (3.7%).
Of the Satyrinae butterflies captured, 36.6%
were Cissia satyrina, which represented
25.28% of all individuals captured.
Planted Regrowth had a species
evenness of 0.58, and a Simpson’s Index of
Diversity of 0.9. Of the total individuals
captured 44.8% were found in planted
regrowth, and 14.81% of species captured
were found there exclusively (Figure 2).
Primary Forest had a species
evenness of 0.58, and a Simpson’s Index of
Diversity of 0.87. Of the total number of
individuals captured 25.9% were found in
primary forest, and 7.4% of species captured
were found there exclusively (Figure 2).
Natural Regrowth had a species
evenness of 0.48, and a Simpson’s Index of
Diversity of 0.76. Of the total number of
individuals captured 26.4% were found in
natural regrowth, and 11.11% of species
captured were found there exclusively (Figure
2).
Grazeland had a species evenness of
0.82, and a Simpson’s Index of Diversity of
0.9. Of the total number of individuals
captured 2.9% were found on grazeland. No
species were found there exclusively (Figure
2).
The whole study area had a species
evenness of 0.5, and a Simpson’s Index of
Diversity of 0.88. With respect to relative
abundance, 33.3% of species were represented
by 1 individual, and 70.37% of species were
represented by 5 individuals or fewer (Figure
3). Of the total species captured 11.11% were
found in all habitat types (Figure 2).
Figure 1. Distribution of individuals captured in each habitat
type

7. Duong and Junger, 2015
7
Of the 27 species captured 10 (37%)
were found at least once between 30 m and
819 m out of their elevation range as specified
by DeVries in The Butterflies of Costa Rica
and Their Natural History: Volume I. All
butterflies found out of their range were found
above their specified range; 72.5% were found
out of their elevation range by 100 m or more
at least once.
Figure 2. A Venn diagram showing the overlap of species in the habitat types
Figure 3. Relative abundance (A) Overall (B) Planted Regrowth (C) Primary Forest (D) Natural Regrowth (E) Grazeland

8. Duong and Junger, 2015
8
The number of individuals captured
steadily increased as the study continued. 56%
of all individuals captured were captured in
the last two weeks of the study (Figure 4). In
line with this, as the study continued, the
number of unique species captured increased
(Figure 5).
DISCUSSION
As with any trapping study, sampling
bias because of microenvironmental variance
among traps, as well as because of variance
among species in attraction to baits, may have
been a source of error. Pooling replicate traps
can reduce individual trap variance; however,
species attraction to bait can be addressed only
by intensive mark/recapture studies and/or by
natural history observations (DeVries et al.,
1997).
We found that the grazeland had the
highest species evenness and the highest
diversity index; however, it has the lowest
species richness. This is because few species
were found in the grazeland, but these species
were each represented by only 1 or 2
individuals, making it very even and diverse.
We can conclude that the grazeland provides
habitat for a small number of highly varied
species. This suggests that butterflies may use
grazeland as a transitory habitat and that it
may act as a habitat corridor. Thereby,
grazelands allow butterflies to travel between
more suitable fragmented habitats.
The planted regrowth had the highest
species richness, highest diversity index, and a
relatively high evenness rating. This suggests
that of the habitat types studied the planted
regrowth forests host the highest number of
individuals and the most diverse butterfly
population. There are a few explanations as to
why planted regrowth is more diverse and rich
than natural regrowth. Firstly, planted
regrowth is cleared of invasive species, such
as bracken, which shade out and compete with
other secondary species as part of the planting
process. This clearing may provide more
habitat for butterfly host plants that would not
have been able to grow otherwise. Secondly,
the clearing of the planted regrowth habitat
reduces the number of plants shading the
habitat. Butterflies often prefer areas that are
sunny and warm, so this may explain the
butterfly’s higher affinity to planted regrowth
habitats. Thirdly, the elimination of bushy and
invasive plant species may reduce habitat for
organisms that compete with or parasitize
butterfly species, giving butterflies in the
planted regrowth area a competitive
advantage. From the data collected during this
study we can conclude that the process of
planting trees in secondary forests encourages
an increase in butterfly richness and diversity.
This suggests that planting secondary forests
not only aids in speeding up the reforestation
process, but also improves the insect diversity
in the area. Further studies are needed to
pinpoint exactly how the process of planting
improves richness and diversity.
All habitat types showed relatively
low abundance, as 70.37% of species are
represented by 5 individuals or fewer. This
low abundance has two possible explanations:
(i) many butterfly species undergo seasonal
Figure 4. Number of individuals captured per week Figure 5. Species accumulation

9. Duong and Junger, 2015
9
migrations (DeVries & Walla, 2001), which
could have caused more species to be present
at low numbers as they travel, (ii) the short
study period was not sufficient enough to
capture representative numbers of individuals
from each species, and with a continuation of
the study relative abundance could increase.
Despite their low relative abundance,
the species that we did capture appear to
provide a good representation of the
Nymphalidae species. 44.4% of the species
captured were from the subfamily Satyrinae.
The Satyrinae subfamily represents nearly half
of Nymphalidae diversity. Therefore, the
species we captured appear to be in proportion
to the overall species present.
This study was conducted at the close
of the dry season and the onset of the rainy
season. The first three weeks of the study were
relatively dry, receiving little rainfall
sporadically. The second three weeks saw the
onset of the rainy season, with heavy rains
almost every day. This change in weather
patterns correlates to a drastic increase in the
number and species of butterflies
captured. 70.11% of all individuals captured
were trapped in the last three weeks of the
study, with 56% of all individuals captured
trapped in the last two weeks alone. This is
consistent with the idea that a seasonal
correlation with rainfall is typical of tropical
insect communities (Wolda, 1978, 1992; Kato
et al., 1995; Novoty & Basset, 1998). This
seasonal shift in population size has two
possible explanations: (i) butterflies typically
undergo seasonal, multi-species migrations at
this time of year (DeVries & Walla, 2001),
causing more species to be present than would
normally be found at other times of year, and
(ii) the availability of natural fruit sources may
cause differential attraction of butterflies to
banana-baited traps (DeVries & Walla, 2001),
causing butterflies to be less picky about their
food source and more likely to feed from the
traps. Further studies that span both the rainy
season and dry season and specifically
investigate the seasonal components of
biodiversity should be undertaken.
We found that a large number of the
species captured (37%) were found at least
once higher than their natural elevation range,
with 72.5% found out of their range by 100 m
or more at least once. This large shift in
species to higher habitats could indicate that
climate change is affecting tropical butterfly
species’ ranges. Warming of habitats, along
with the affects of shifts in cloud density and
precipitation patterns may be forcing butterfly
species into higher elevation habitats to
combat these changes. This shift represents a
large concern for tropical insect communities,
as habitat types typically change drastically
with increases in elevation. Butterflies may
find they are rapidly displaced from suitable
habitat, and higher elevation habitats that are
suitably wet and cool do not contain suitable
plant species. This shift has the potential to
cause a drastic decrease in the abundance and
diversity of butterfly populations in tropical
areas.
From this study we can conclude that
of the successional habitat types studied,
planted regrowth is the most diverse and rich,
indicating that the process of planting forests
improves community diversity and richness.
This increase in community diversity and
richness may lead to higher diversity and
richness in the climax community. We can
also conclude that a large number of species
are being found out of their natural elevation
range, which could indicate that butterflies in
this area are experiencing the affects of
climate change, and that, if remediation efforts
are not undertaken soon, then a drastic
decrease in butterfly populations and diversity
may ensue.
Improvements and Further Studies
An area this study needs improvement
in is studying butterflies on the grazeland.
Because grazeland traps were more exposed to
the sun, rain, and wind, we frequently found
these traps and/or the bait in an ineffective
state: dried out, diluted, or dumped out by the
wind. We suggest weighing down the base of
the trap with a handful of gravel to prevent the
wind from tipping the bait out, or using
heavier material for the base of the trap.
Perhaps stirring in a bit of water to dried out
bait may restore the bait to a more suitable
condition, and carrying extra bait may be
worthwhile in case of diluted bait or an empty
bait dish. Further studies may also consider

10. Duong and Junger, 2015
10
studying butterfly frequency and diversity on
farm land instead of grazeland, as farm land
lacks animals that tamper with traps and has
more shade to prevent desiccation of bait.
Our study methods provide estimates
of species abundance using only the numbers
of adult butterflies caught in our traps, but
provides no information on the distribution of
host plants, roosting areas, courtship sites, or
other life history components. Further studies
should investigate life history components in
correlation with diversity and
frequency. Additionally, the type of bait we
selected mostly excludes adult butterflies that
do not belong to the family Nymphalidae, as
we used a sweet bait. Further studies may
consider incorporating different types of baits
to attract a wider variety of butterfly families.
A number of butterflies, particularly
of the larger variety, were excluded from our
data because they escaped from our trap and
were thus unable to be identified. Many times,
these larger butterflies were found sitting
directly on the bait dish and, when
approached, flew down and out of the trap.
One possible modification to the trap, such
that the base can be pulled up and the bottom
of the trap sealed from a distance, can prevent
such losses. Or, perhaps narrowing the
distance between the bottom of the trap, and
the base and bait dish of the trap, from 1 inch
to less than 1 inch may also help to prevent the
loss of specimens; however, such a
modification may also inhibit larger butterflies
from entering the trap entirely. Further studies
should consider the trade off between
including larger butterfly species and the rate
of butterfly escape from the traps.
This study focused on butterfly
diversity and abundance in habitat
understories. Vertical stratification is essential
to community structure (DeVries & Walla,
2001). Therefore, further studies should
consider investigating the vertical
stratification of communities by incorporating
canopy traps into their study.
Acknowledgements
This research internship trip was funded by the Hubbard Center for Student Engagement of
DePauw University, whose grant funds were used in the purchase of necessary field equipment
and supplies. Additionally, this research project was conducted on and made possible by
Cloudbridge Nature Reserve, as well as on the pastureland owned by the farmer Marcos Romero,
who opened his land for our research and helped to maintain the butterfly traps set on his land.
We would also like to thank Cloudbridge manager Frank Spooner for advising us on this study.

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APPENDIX
Appendix A- Cloudbridge Map with Trap Locations
Appendix B- Site and species information
APPENDIX A - CLOUDBRIDGE Map with Trap Locations

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APPENDIX B - SITES AND SPECIES INFORMATION
This is a list of all the sites where research has been conducted, including a list of species found at
each site.
PF = Primary Forest (El Jilguero/Hectare)
PR = Planted Regrowth Secondary Forest (El Jilguero)
NR = Natural Regrowth Secondary Forst (Rio)
GL = Grazeland (Marco’s pasture)
Habitat Type Trap Number Latitude Longitude Elevation (m)
PF 1 09°28'05.1 083°34'27.6 1857
PF 2 09°27'59.0 083°34'17.7 1970
PF 3 09°27'57.1 083°34'12.9 2030
PR 1 09°28'11.9 083°34'39.0 1519
PR 2 09°28'14.6 083°34'44.1 1662
PR 3 09°28'07.8 083°34'32.5 1813
NR 1 09°28'30.9 083°34'06.8 1672
NR 2 09°28'27.6 083°34'11.8 1681
NR 3 09°28'22.8 083°34'15.4 1704
GL 1 09°28'16.9 083°34'53.8 1536
GL 2 09°28'15.9 083°34'53.4 1563
GL 3 09°28'13.9 083°34'52.9 1561
PF 1
Archaeoprepona demophon centralis
Cissia renata
Cissia satyrina
Cyllopsis argentella
Dioriste tauropolis
Drucina leonata
Memphis arginussa eubaena
Opoptera staudingeri
Opsiphanes cassina chiriquensis
Opsiphanes cassina fabricii
PF 2
Catonephele chromis godmani
Cissia satyrina
Drucina leonata
Pycina zamba zelys
PF 3
Catonephele chromis godmani
Cissia gigas
Cissia satyrina
Cyllopsis argentella
Dioriste tauropolis
Drucina leonata
Pedaliodes dejecta
PR 1
Caligo eurilochus sulanus
Cissia satyrina
Consul electra
Dioriste tauropolis
Drucina leonata
Megeuptychia antonoe
Memphis beatrix
Opsiphanes cassina chiriquensis

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Smyrna blomfildia datis
PR 2
Archaeoprepona demophon centralis
Cissia gigas
Cissia satyrina
Cissia similis
Dioriste tauropolis
Drucina leonata
Memphis beatrix
Memphis glycerium
Opsiphanes cassina chiriquensis
Pedaliodes dejecta
Pronophila timanthes
PR 3
Archaeoprepona demophon centralis
Cissia gigas
Cissia renata
Cissia satyrina
Cissia satyrina
Dioriste tauropolis
Drucina leonata
Memphis arginussa eubaena
Memphis beatrix
Memphis pithyusa
Opsiphanes cassina chiriquensis
Opsiphanes cassina fabricii
Pedaliodes dejecta
Pedaliodes perperna
NR 1
Cissia satyrina
Dioriste cothonides
Dioriste tauropolis
Memphis beatrix
Smyrna blomfildia datis
NR 2
Cissia satyrina
Cyllopsis argentella
Memphis pithyusa
Opsiphanes bogotanus
Smyrna blomfildia datis
NR 3
Cyllopsis argentella
Dioriste tauropolis
Drucina leonata
Greta polissena umbrana
Opsiphanes cassina chiriquensis
Pronophila timanthes
GL 1
Drucina leonata
Memphis glycerium
Opsiphanes cassina chiriquensis
Smyrna blomfildia datis
GL 2: zero
GL 3: zero