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: , butterfly, species abundance distribution, species richness, environmental monitoring, habitat disturbance, tropical, conservation.

INTRODUCTION

Natural habitats in the tropics continue habitats, and sensitivity to microclimate to be globally threatened by habitat loss and heterogeneity and disturbance (New, 1997; climate change, leading to the massive loss of DeVries et al., 1997). Butterflies are the best- species. Remediation efforts are being known group of (DeVries et al., 1997), undertaken globally in an effort to reduce and giving them great potential to provide reverse the effects of habitat loss and climate understanding of diversity and change. In order to focus these efforts and to conservation. The study of fruit feeding determine their effects on biological diversity, butterflies has additional advantages, reliable monitoring programs that assess including their ability to be vigorously changes in biodiversity and ecosystem sampled with the use of fruit-baited traps. function are needed. The choice of organism Therefore, fruit-feeding butterflies provide a investigated is crucial to this process due to standard means for comparing species overall lack of funds and available expertise. diversity within and among tropical insect Trapping butterflies is often the communities (DeVries et al., 2012). method of choice in tropical forests (Aduse- Bait trapping is an inherently biased Poku et al., 2012). Advantages of using method for assessing butterfly fauna. Issues butterflies () as a target species include: some butterflies are never captured in include: their relatively large size, colorful traps, and some butterflies are more strongly appearance, relative ease of identification to attracted than others, thus relative abundances species level, their presence in all terrestrial captured do not necessarily reflect the relative

Duong and Junger, 2015 1 abundances of fruit feeding species in the industry, and both legal and illegal timbering region (Hughes et al., 1998). Despite these (Borges-Méndez, 2008). During this period drawbacks, bait trapping provides the most the national area covered with forests dropped effective and efficient method for monitoring from about 70% to about 10% (United States changes in species abundances, and measuring Agency for International Development diversity of tropical butterfly communities. [USAID], 1996). In fact, between 1950 and Reforestation is an essential step in 1994, the pace of deforestation in Costa Rica restoring forest health. Monitoring and was one of the fastest in the western assessment of the effects of reforestation on hemisphere, with a decrease of 40,000-50,000 communities within remediated areas is hectares annually (Watson et al., 1998; essential to gaining an understanding of its Borges-Méndez, 2008). Given this rapid and short- and long-term effects. Few studies drastic loss of forested areas throughout the investigate the changes in species abundance country, reforestation programs will be and diversity through various levels of essential to restoring the country’s former succession in tropical forests. Most studies in biodiversity. this area focus on the differences between Costa Rican forests have sufficiently disturbed and undisturbed habitats. Disturbed diverse fruit-feeding butterfly fauna to warrant habitats have been found to have higher their use as target organisms for monitoring species richness and more unique species changes in biodiversity. There are (DeVries et al., 1997). Additionally, vertical approximately 543 butterfly species present in stratification is reduced in disturbed forests, Costa Rica (DeVries, 1987). Of these, at least trapping canopy species in the understory, 40% feed exclusively upon rotting fruits as leading to overestimates of species richness in adults (DeVries, 1987). understory trapping in disturbed areas The aim of this study is to quantify the (DeVries & Walla, 2001; Fermon et al., 2005; differences in diversity and abundance of Aduse-Poku et al., 2012). Rapid monitoring fruit-feeding butterflies within habitats at programs will allow assessments of the different stages of succession. We performed changes in diversity and abundance within a butterfly trapping study on the Cloudbridge different stages of succession, allowing for Nature reserve in Costa Rica during the onset more effective and focused remediation of the rainy season, studying four habitat types efforts. (grazeland, natural regrowth, planted Between the 1940s and 1980s regrowth, and primary forest). In each of these deforestation in Costa Rica was caused by habitat types three traps were established in government-sponsored land colonization the understory. schemes, expansion of the agricultural frontier, cattle ranching to support the beef

MATERIALS AND METHODS

Study Site over a million hectares spanning northern This research was conducted at the Costa Rica and southern Panama. Together Chirripó Cloudbridge Nature Reserve, San with the adjoining La Amistad International Isidro de General, south central Costa Rica. Park, Chirripó National Park is comprised of Located in a cloud forest on one of the tallest the largest unspoiled forest in the country. mountains of Central America, Cloudbridge is Cloudbridge started off as privately a 700 acre nature reserve on the northern end owned land in 2002, owned by Ian Giddy and of one of the most important biological zones Genevieve (Jenny) Giddy, who made the first of all Central America, and lies within a of many subsequent purchases of cattle farms designated “biological hot spot” on the Meso- bordering the Chirripó National Park to American Biological Corridor. Cloudbridge is impede the appalling denuding and erosive part of an area of forested land that includes effects that cattle grazing has had on the land.

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

Duong and Junger, 2015 3 Nymphalidae. (DeVries, Murray, & Lande, 1997). The other guild meets all nutritional A 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, , 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, B 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 C 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 D 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)

Illustration 1. Trap design and construction

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

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

Duong and Junger, 2015 6 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 and Their Natural History: Volume I. All Of the 27 species captured 10 (37%) butterflies found out of their range were found were found at least once between 30 m and above their specified range; 72.5% were found 819 m out of their elevation range as specified out of their elevation range by 100 m or more by DeVries in The Butterflies of Costa Rica at least once.

Duong and Junger, 2015 7 The number of individuals captured line with this, as the study continued, the steadily increased as the study continued. 56% number of unique species captured increased of all individuals captured were captured in (Figure 5). the last two weeks of the study (Figure 4). In

Figure 4. Number of individuals captured per week Figure 5. Species accumulation

DISCUSSION

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

Duong and Junger, 2015 8 migrations (DeVries & Walla, 2001), which with 72.5% found out of their range by 100 m could have caused more species to be present or more at least once. This large shift in at low numbers as they travel, (ii) the short species to higher habitats could indicate that study period was not sufficient enough to climate change is affecting tropical butterfly capture representative numbers of individuals species’ ranges. Warming of habitats, along from each species, and with a continuation of with the affects of shifts in cloud density and the study relative abundance could increase. precipitation patterns may be forcing butterfly Despite their low relative abundance, species into higher elevation habitats to the species that we did capture appear to combat these changes. This shift represents a provide a good representation of the large concern for tropical insect communities, Nymphalidae species. 44.4% of the species as habitat types typically change drastically captured were from the subfamily Satyrinae. with increases in elevation. Butterflies may The Satyrinae subfamily represents nearly half find they are rapidly displaced from suitable of Nymphalidae diversity. Therefore, the habitat, and higher elevation habitats that are species we captured appear to be in proportion suitably wet and cool do not contain suitable to the overall species present. plant species. This shift has the potential to This study was conducted at the close cause a drastic decrease in the abundance and of the dry season and the onset of the rainy diversity of butterfly populations in tropical season. The first three weeks of the study were areas. relatively dry, receiving little rainfall From this study we can conclude that sporadically. The second three weeks saw the of the successional habitat types studied, onset of the rainy season, with heavy rains planted regrowth is the most diverse and rich, almost every day. This change in weather indicating that the process of planting forests patterns correlates to a drastic increase in the improves community diversity and richness. number and species of butterflies This increase in community diversity and captured. 70.11% of all individuals captured richness may lead to higher diversity and were trapped in the last three weeks of the richness in the climax community. We can study, with 56% of all individuals captured also conclude that a large number of species trapped in the last two weeks alone. This is are being found out of their natural elevation consistent with the idea that a seasonal range, which could indicate that butterflies in correlation with rainfall is typical of tropical this area are experiencing the affects of insect communities (Wolda, 1978, 1992; Kato climate change, and that, if remediation efforts et al., 1995; Novoty & Basset, 1998). This are not undertaken soon, then a drastic seasonal shift in population size has two decrease in butterfly populations and diversity possible explanations: (i) butterflies typically may ensue. undergo seasonal, multi-species migrations at Improvements and Further Studies this time of year (DeVries & Walla, 2001), An area this study needs improvement causing more species to be present than would in is studying butterflies on the grazeland. normally be found at other times of year, and Because grazeland traps were more exposed to (ii) the availability of natural fruit sources may the sun, rain, and wind, we frequently found cause differential attraction of butterflies to these traps and/or the bait in an ineffective banana-baited traps (DeVries & Walla, 2001), state: dried out, diluted, or dumped out by the causing butterflies to be less picky about their wind. We suggest weighing down the base of food source and more likely to feed from the the trap with a handful of gravel to prevent the traps. Further studies that span both the rainy wind from tipping the bait out, or using season and dry season and specifically heavier material for the base of the trap. investigate the seasonal components of Perhaps stirring in a bit of water to dried out biodiversity should be undertaken. bait may restore the bait to a more suitable We found that a large number of the condition, and carrying extra bait may be species captured (37%) were found at least worthwhile in case of diluted bait or an empty once higher than their natural elevation range, bait dish. Further studies may also consider

Duong and Junger, 2015 9 studying butterfly frequency and diversity on directly on the bait dish and, when farm land instead of grazeland, as farm land approached, flew down and out of the trap. lacks that tamper with traps and has One possible modification to the trap, such more shade to prevent desiccation of bait. that the base can be pulled up and the bottom Our study methods provide estimates of the trap sealed from a distance, can prevent of species abundance using only the numbers such losses. Or, perhaps narrowing the of adult butterflies caught in our traps, but distance between the bottom of the trap, and provides no information on the distribution of the base and bait dish of the trap, from 1 inch host plants, roosting areas, courtship sites, or to less than 1 inch may also help to prevent the other life history components. Further studies loss of specimens; however, such a should investigate life history components in modification may also inhibit larger butterflies correlation with diversity and from entering the trap entirely. Further studies frequency. Additionally, the type of bait we should consider the trade off between selected mostly excludes adult butterflies that including larger butterfly species and the rate do not belong to the family Nymphalidae, as of butterfly escape from the traps. we used a sweet bait. Further studies may This study focused on butterfly consider incorporating different types of baits diversity and abundance in habitat to attract a wider variety of butterfly families. understories. Vertical stratification is essential A number of butterflies, particularly to community structure (DeVries & Walla, of the larger variety, were excluded from our 2001). Therefore, further studies should data because they escaped from our trap and consider investigating the vertical were thus unable to be identified. Many times, stratification of communities by incorporating these larger butterflies were found sitting 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.

Duong and Junger, 2015 10 References

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Duong and Junger, 2015 11 APPENDIX Appendix A- Cloudbridge Map with Trap Locations Appendix B- Site and species information

APPENDIX A - CLOUDBRIDGE Map with Trap Locations

Duong and Junger, 2015 12 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 PF 3

Catonephele chromis godmani Archaeoprepona demophon centralis Cissia gigas Cissia renata Cissia satyrina Cissia satyrina Cyllopsis argentella Cyllopsis argentella Dioriste tauropolis Dioriste tauropolis Drucina leonata Drucina leonata Pedaliodes dejecta

Memphis arginussa eubaena Opoptera staudingeri PR 1 Opsiphanes cassina chiriquensis Opsiphanes cassina fabricii Caligo eurilochus sulanus Cissia satyrina PF 2 Consul electra Dioriste tauropolis Catonephele chromis godmani Drucina leonata Cissia satyrina Megeuptychia antonoe Drucina leonata Memphis beatrix Pycina zamba zelys Opsiphanes cassina chiriquensis

Duong and Junger, 2015 13 Smyrna blomfildia datis NR 1

PR 2 Cissia satyrina Dioriste cothonides Dioriste tauropolis Memphis beatrix Archaeoprepona demophon centralis Smyrna blomfildia datis Cissia gigas Cissia satyrina NR 2 Cissia similis Dioriste tauropolis Cissia satyrina Drucina leonata Cyllopsis argentella Memphis beatrix Memphis pithyusa Memphis glycerium Opsiphanes bogotanus Opsiphanes cassina chiriquensis Smyrna blomfildia datis Pedaliodes dejecta Pronophila timanthes NR 3

PR 3 Cyllopsis argentella Dioriste tauropolis Drucina leonata Greta polissena umbrana Archaeoprepona demophon centralis Opsiphanes cassina chiriquensis Cissia gigas Pronophila timanthes Cissia renata Cissia satyrina GL 1 Cissia satyrina Dioriste tauropolis Drucina leonata Drucina leonata Memphis glycerium Memphis arginussa eubaena Opsiphanes cassina chiriquensis Memphis beatrix Smyrna blomfildia datis Memphis pithyusa

Opsiphanes cassina chiriquensis GL 2: zero Opsiphanes cassina fabricii GL 3: zero Pedaliodes dejecta Pedaliodes perperna

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