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Butterflies of the Golfo Dulce Region, Costa Rica

June 2010

Publisher: Verein zur Förderung der Tropenstation La Gamba
ISBN: 978-3-200-01883-9

Authors:

Harald Krenn

University of Vienna

Martin Wiemers

Senckenberg Society for Nature Research

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This guide book focuses on the butterflies of the Golfo Dulce Region in Costa Rica. It introduces the reader to a representative selection of species from the region. The text and numerous color photographs provide essential information for the identification of the most common and conspicuous butterfly species. The booklet also includes a general introduction to butterflies and the natural history of the region.

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Content uploaded by Martin Wiemers

Content may be subject to copyright.

Butteries

of the

Golfo Dulce Region

Costa Rica

Corcovado National Park

Piedras Blancas National Park

‚Regenwald der Österreicher‘

Authors

Harald Krenn

Department of Evolutionary Biology

University of Vienna

Althanstraße 14, 1090 Vienna, Austria

harald.krenn@univie.ac.at

Martin Wiemers

Department of Animal Biodiversity

University of Vienna

Rennweg 14, 1030, Vienna, Austria

martin.wiemers@univie.ac.at

Werner Huber

Department of Structural and

Functional Botany

University of Vienna

Rennweg 14, 1030, Vienna, Austria

werner.huber@univie.ac.at

Roland Albert

Department of Chemical Ecology

and Ecosystem Research

University of Vienna

Rennweg 14, 1030, Vienna, Austria

roland.albert@univie.ac.at

Anton Weissenhofer

Department of Structural and

Functional Botany

University of Vienna

Rennweg 14, 1030, Vienna, Austria

anton.weissenhofer@univie.ac.at

Lisa Maurer Veronika Pemmer

Contents

The ‘Tropical Research Station La Gamba’ 4

The rainforests of the Golfo Dulce region 6

Butteries of the Golfo Dulce Region, Costa Rica 8

Papilionidae - Swallowtail Butteries 13

Pieridae - Sulphures and Whites 17

Nymphalidae - Brush Footed Butteries 21

Subfamily Danainae

Subfamily Ithomiinae

Subfamily Charaxinae

Subfamily Satyrinae

Subfamily Cyrestinae

Subfamily Biblidinae

Subfamily Nymphalinae

Subfamily Limenitidinae

Subfamily Heliconiinae 40

Riodinidae - Metalmarks 47

Lycaenidae - Blues 53

Hesperiidae - Skippers 57

Appendix- Checklist of species 61

Acknowledgements 74

References 74

Picture credits 75

Index 78

The ‘Tropical Research Station La Gamba’

Roland Albert

Secretary General of the ‘Society for the Preservation of the Tropical Research Station La Gamba’

Department of Chemical Ecology and Ecosystem Research, University of Vienna

In 1991, Michael Schnitzler, a distinguished

musician and former professor at the Univer-

sity of Music and Performing Arts in Vienna,

founded the ‘Association Rainforest of the Aus-

trians’ (Verein Regenwald der Österreicher). Over

many years, he collected contributions in Aus-

tria and forwarded the funds to the Costa Rican

government. With the money, the government

purchased large tracts of forest from landowners

and placed it under the control of the ‘Parque Na-

cional Piedras Blancas’, thereby preserving the

Esquinas Rainforest in southwestern Costa Rica.

It soon became apparent that this protected area

also provided ideal conditions for promoting

Austrian research and teaching programmes in

rainforests. To make the area more accessible, the

‘Association Rainforest of the Austrians’bought

an old farmhouse (a nca) in 1993. This formed

the nucleus for a tropical eld station, located di-

rectly on the border of the National Park near the

small village of La Gamba. With the support of

the ‘Association Rainforest of the Austrians’, the

Ministry of Science and Research and the Uni-

versity of Vienna, the ‘Tropical Research Station

La Gamba’ ourished and grew. It now compris-

es several buildings and includes comfortable

living and research facilities for more than 30

visitors as well as a botanical garden. In 2008 an

airconditioned laboratory was constructed with

the support of the Faculty of Life Sciences of the

University of Vienna. With its infrastructure, the

eld station is currently on the threshold of be-

coming an internationally established research

institution and education centre, focussing on

the exploration and conservation of Neotropical

rainforests. Since its foundation, many scientists

have carried out eld research in the Esquinas

rainforest and many visitors have admired its

beautiful surroundings. As well as support-

ing research and teaching activities, the station

helps the inhabitants of La Gamba by means of

a series of applied projects to improve their liv-

ing conditions. These projects are partly run in

The main building of the Tropical Research Station

Scientists at work in La Gamba

collaboration with the nearby ‘Esquinas Rainfor-

est Lodge’, another brainchild of Michael Schnit-

zler, which represents an Austrian example of

sustainable ecotourism. Indeed, the lodge was

awarded some years ago the ofcial Costa Ri-

can ‘Three-Leaf Certicate of Sustainable Tour-

ism’. We are particularly proud that the former

European Commissioner for Agriculture, Dr.

Franz Fischler, has been a patron of the ‘Society

for the Preservation of the Tropical Station La

Gamba’(Verein zur Förderung der Tropenstation La

Gamba) since 2006. Numerous scientic publica-

tions have resulted from research performed at

the Tropical Research Station La Gamba, includ-

ing about 70 doctoral and diploma theses. The

scientic work initially focussed on the ora and

vegetation of the Esquinas forest but it has now

broadened to include a wide range f other topics,

such as animal-plant interactions, herpetology,

ornithology, entomology (especially studies of

butteries), limnology, chemical eco-physiology,

bio-geochemistry, geography and sociology.

About 60 eld courses and excursions have been

organized, enabling students and scientists from

universities all over the world to visit the Piedras

Blancas National Park.

Our Society is particularly interested in pro-

viding all our visitors, and beyond that the

many friends and admirers of the tropics, with

information on the biology and ecology of the

local and regional rainforests and of the adja-

cent cultural land. Following the publication

of an internationally acclaimed ‘Field Guide of

Flowering Plants of the Golfo Dulce Rainforests’

in 2001, seven colourful brochures have to date

been published: ‘Amphibians and Reptiles of

the Golfo Dulce Region’, ‘The Birds of the Golfo

Dulce Region, Costa Rica’, ‘Das Leben hier und

dort – La vida aquí y allá’, ‘Ecosystem diversity

in the Piedras Blancas National Park and adja-

cent areas (Costa Rica)’, ‘Fruits in Costa Rican

Markets’, ‘Corbiculate Bees of the Golfo Dulce

Region, Costa Rica’, ‘Dragonies of the Golfo

Dulce Region, Costa Rica’. Adittionally, a scien-

tic monograph ‘Natural and Cultural History

of the Golfo Dulce Region, Costa Rica’ has been

published in 2008.

We are now proud to offer a further colorful

issue dealing with a group of animals - the but-

teries – which on the one hand belong to the

most eye-caching insect groups, and can be en-

countered on the other hand due to their behav-

iour usually more frequent by rain forest visitors

as some other insect groups. We hope that this

booklet – as all our former brochures and books

- will be a source of information and joy to lov-

ers of tropical life, to scholars und students, to

teachers and researchers at the University and

– nally - to all the friends of the ‘Rainforest of

the Austrians’ and the ‘Tropical Research Station

La Gamba’.

For further information see:

www.lagamba.at

and

www.regenwald.at

The ‘Comedor’ Martin Wiemers with students in the laboratory

The rainforests of the Golfo Dulce region

Werner Huber/Anton Weissenhofer

Department of Structural and Functional Botany, University of Vienna

The Golfo Dulce region is located in the

southern corner of the Pacic coast of Costa

Rica, near the border with Panama. Within this

region are the Corcovado and Piedras Blancas

national parks. The forests of the region are the

only moist or wet evergreen lowland forests that

still exist on the Pacic coast of Central America.

The elevations range from sea level to 745 m on

the Cerro Rincón, Peninsula de Osa; the annual

precipitation is up to 6,000 mm with a short or al-

most nonexistent dry season from December un-

til March; the relief is strongly structured at the

landscape level and contains many microhabitats

and niches; and the soil types are diverse and the

vegetation highly dynamic. All of these factors,

coupled with the biogeographical position of the

Golfo Dulce, have led to a very species-rich for-

est: about 2,700 species of higher plant have been

recorded from the area.

The tallest trees in the moist and wet ever-

green lowland forests grow up to 60 m high (e.g.

Morning clouds above the forest

Waterfall on the Quebrada Chorro

Climatic diagramm, La Gamba

Anacardium excelsum, Parkia pendula, Carapa guia-

nensis, Brosimum utile and Caryocar costaricense).

Most trees retain their foliage throughout the

year. Palms (about 45 species) such as Socratea

exorrhiza, Iriartea deltoidea, Wela regia and As-

terogyne martiana are typical features of the for-

est. Heliconia herbs are obvious near streams and

in gaps. Lianas (e.g. Bauhinia and Entada), vines

and epiphytes (Bromeliaceae, Orchidaceae) are

found on many trees. About 100 species of or-

chid (e.g. vanilla) and 40 species of bromeliad are

distributed in the region.

Hemi-epiphytic plants of the genus Clusia

are abundant in the very wet parts of the forest

while g trees (Ficus) are more frequent in the

drier regions (on the Corcovado plains). On the

coast, the coconut palm (Cocos nucifera) and the

‘‘tropical almond tree’ (Terminalia catappa) are the

most conspicuous plants; in the mangroves this

position is taken by red mangrove (Rhizophora

mangle) and mangle pinuela (Pelliciera rhizopho-

rae). In addition to the various types of primary

and secondary forest, the area also features other

types of habitat such as pastures, pools, rivers,

cultivated land and various kinds of plantation

(e.g. oil palm, teak and manioc). Bees are very

abundant and are found in all habitats.

Costa Rica’s geographic location on the

‘bridge’ between North and South America and

the fact that the country formed a kind of refuge

for tropical animals and plants during the last ice

age has led to remarkable biogeographical pat-

terns. Restricted-range plants and animals are

abundant and many reach their northern limits

in southern Costa Rica. The region is an excellent

place for naturalists to enjoy tropical nature. An

extensive system of trails in the Corcovado and

Piedras Blancas national parks (La Gamba) offers

wonderful insights into tropical rainforests.

Interior of the forest of the NP Piedras Blancas

The Rió Bonito in the NP Piedras Blancas

Butteries of the Golfo Dulce Region

Costa Rica

Introduction

The butteries of Costa Rica are extraordi-

narily diverse and rich in species. They are one

of the most thoroughly studied buttery faunas

of the Neotropics. At least 1,323 species of true

butteries and skippers (Papilionoidea and Hes-

perioidea) have been identied in the various

habitats of the country, according to the study of

DeVries (1983). It is estimated that about 5% of

the total number of buttery species of the world

live in Costa Rica (Wiemers & Fiedler 2008).

The preliminary checklist of the surround-

ings of the Tropical Research Station La Gamba

accounts for a total of 216 buttery species (202

Papilionoidea and 14 Hesperioidea, Appendix

1). The list summarizes the results of four stan-

dardized one-week-surveys which were carried

out from 2006 to 2010 (mainly in February), and

includes species, which were encountered out-

side of the time period of the studies. Additional

species were added from two longer surveys car-

ried out by Hellena Binz (December 2008 – Janu-

ary 2009 and November 2009 – February 2010)

who also employed fruit baiting for her second

study. The results of the rst two surveys in 2006

and 2007 were published by Wiemers & Fiedler

(2008), who list 144 buttery species (130 Papil-

ionoidea and 14 Hesperioidea) and estimate at

least 156-204 species of Papilionoidea for the area

of La Gamba within the Esquinas Rainforest. An

additional 64 species (49 Papilionoidea and 15

Hesperioidea) were reported from the Corco-

vado National Park. Thus, a total of 280 buttery

species (251 Papilionoidea and 29 Hesperioidea)

are currently known from the Golfo Dulce region

of the Costa Rican province Puntarenas.

The habitats surrounding the research station

can be divided into three types with varying de-

grees of land use: the natural forest, the secondary

forest and the intensively used land systems. The

natural forest consists of primary forests which

are located mainly in the Piedras Blancas Na-

tional Park. This habitat type contains the high-

est number of buttery species. It is followed by

the secondary forest habitat, which borders the

protected areas of the National Park. The habitat

type characterized by intensive land use consists

of pastures, oil palm plantations, roadsides, and

gardens and shows the least number of buttery

species (Wiemers & Fiedler 2008). Typical for the

natural forests are the nymphalid species Phila-

ethria dido, Heliconius sapho, H. cydno, Eueides lybia

and the lycaenid species, Arumecla galliena. In the

intensively used habitats, the nymphalids Her-

meuptychia hermes, Anartia jatrophae, A. fatima and

the pierid buttery Pyrisitia nisa are most com-

mon (Wiemers & Fiedler 2008).

Diurnal butteries, unlike moths, are active

during the daytime. They are presently classi-

ed into two superfamilies: Papilionoidea and

Hesperioidea. The Papilionoidea (or true but-

teries) are split into ve families: Papilionidae

(swallowtail butteries), Pieridae (sulphures and

whites), Nymphalidae (brush-footed butteries),

Lycaenidae (blues) and Riodinidae (metalmarks).

Of these, the Nymphalidae is the group richest

in species. The Hesperioidea includes only one

family, Hesperiidae (skippers).

This booklet includes the most frequent and

most conspicuous butteries in the examined re-

gion. A short description is given for each of the

selected species along with general information

about aspects of its natural history. In addition to

the scientic names of the butteries, we provide

common English and Spanish names, in as much

as they are available. Most species in the book-

let can be seen during the whole year; however,

the frequencies of the species will vary over the

seasons. Typical for tropical biodiversity is that

a high number of species may be present but the

number of individuals is usually low. Thus, on a

brief visit, one should not expect to be able to see

all the butteries mentioned in the book.

This guide book is a product of a eld course

in tropical biology for students of the University

of Vienna (Austria) during August and Septem-

ber 2009 and is based on results of similar stu-

dent courses of previous years. Determination

of the buttery species was based on the two

volumes of ‘The Butteries of Costa Rica and

their Natural History’ (DeVries 1987, 1997) and

the recent eld guide ‘Mariposas de Costa Rica

– Butteries and moths of Costa Rica (Chacón

Eggs of Heliconius melpomene

& Montero 2007). The nomenclature, systematic

classication and order in the book follow La-

mas (2004) and the preliminary checklist of the

Papilionoidea and Hesperioidea of the Tropical

Research Station La Gamba (Wiemers & Fiedler

2008). English common names were taken from

the internet site: www.neotropicalbutteries.

com. For comprehensive information on the bi-

ology of butteries, see DeVries (1987, 1997) and

Janzen (1983). Aspects of climate, habitats and

biodiversity are well summarized in ‘Natural

and cultural history of the Golfo Dulce Region,

Costa Rica’ (Weissenhofer et al. 2008).

The best time to observe butteries is from

morning to noon. In these hours, most butteries

are active and searching for food or mates. The

best opportunity to closely observe and photo-

graph these beautiful insects is while they are

occupied with feeding. Most buttery species

can be found on or near owering plants. Some

are attracted to rotting fruits and others take up

uid from wet ground. The photographs and the

text in this guidebook should be helpful in iden-

tifying some of the most frequent species and in

becoming more acquainted with their natural

history.

Life cycle of butteries

The life cycle of a buttery can be divided into

four stages: egg, larva (caterpillar), pupa (chrys-

alis) and adult (imago).

Adult females lay individual eggs or clusters

of eggs on surfaces of leaves, ower buds or stems

of the host plants. The eggs undergo embryonic

development until they hatch from the chorion

shell, after which begins the larval stage.

The larvae of butteries are known as cater-

pillars. Most are voracious herbivores and make

good use of their biting-chewing mouthparts.

They generally feed on more or less closely re-

lated host plants or exceptionally on a single

plant species. Some are able to feed from a rela-

tively wide variety of plant species. Caterpillars

increase their size over one hundredfold. In the

process, they molt four to seven times. The phas-

es between successive molts are called instars.

Many caterpillars assume different shapes and

appearances from instar to instar. Not all instars

are easy to nd in nature.

During the pupal stage, larval tissues are

broken down, and the new organs of the adult

become differentiated (i.e., metamorphosis). Fi-

Larva of Heliconius melpomene

Pupa of Heliconius melpomene

nal adjustments for adult life are made while the

buttery is still emerging from the pupal encas-

ing. The wings unfold, and the adult feeding or-

gan or proboscis becomes fully assembled.

As adult insects, butteries take ight, con-

sume liquid nutrients and reproduce. Many but-

teries display special courtship behavior. After

mating, the females search for host plants on

which to lay their eggs.

Morphology of adult butteries

Like all insects, the body of a buttery is com-

posed of three main parts: head, thorax and ab-

domen. The head contains the brain, the major

sensory organs, mouthparts (proboscis) and the

suction pump of the frontal alimentary tract.

The proboscis is a long coil that can be extended

to suck on uids such as nectar, the juice of rot-

ting organic matter or water. The large spherical

compound eyes form the visual organs. The two

elongate and club-tipped antennae on the head

serve primarily tactile and olfactory functions.

They are used to locate food and mates, as well

as the appropriate site on the host plant to lay

eggs. In addition, antennal movements help the

buttery keep its balance when ying. The thorax

bears the locomotory organs, i.e., three pairs of

legs and two pairs of wings. The fore and hind-

wings fold vertically over the body when the

buttery is at rest. The wings vary in color, size

and form in the different groups and species.

They are exceedingly thin structures, yet with an

immensely large surface area in comparison to

the actual body. Inside the wings is a network of

supporting veins. The hind part of the body, the

abdomen, contains the major organs for digestion,

excretion and reproduction. The morphology of

the male sex organs is complex. The genitalia are

thus not only useful in taxonomic studies but of-

ten they provide the distinguishing traits, which

unambiguously identify the species.

Wing pattern and defense mechanisms

Butteries are among the most popular in-

sects, mainly because of their beautifully colored

wing patterns. The wing surfaces are coated

with minute cuticle scales, each capable of a

single color. Some scales reect light and appear

silvery. When iridescent hues are present, it is

because the scales produce a particular blue or

green color by physical diffraction of light. Not

all patterns are visible to human eyes since we

are unable to see UV-reected or polarized light,

which many butteries can. The wing pattern

may serve multiple biological functions. In many

species, the underside of the wings is dark and

cryptically colored so that the buttery is hardly

Imago of Heliconius melpomene

detectable when sitting still with its wings closed.

The upperside of the wings is often colorful and

functions as a signal enabling the butteries to

recognize members of their own species as either

sexual partners or co-rivals. The wings of some

butteries display eyespot patterns, and others

sport a short process (tail) on the hindwing. It is

postulated that eyespots and tails steer the atten-

tion of an attacking bird away from the delicate

body to the wing margins thereby better enabling

a buttery to escape. In many cases, particularly

colorful wing patterns are aposematic and warn

potential predators of the toxicity or distasteful

properties of the buttery.

A great number of butteries defend them-

selves with chemical compounds, which they

acquire as larvae when feeding on toxic host

plants and which they transfer via the pupae to

the adults. Toxic or unpalatable butteries of-

ten have conspicuous patterns on the wings. A

bird which has eaten a poisonous buttery may

be forced to vomit it out. If it can remember the

color pattern, it may avoid eating similar look-

ing butteries. The success of such chemical de-

fenses has led many species to resemble closely

each other in behavior and wing color pattern. In

other words, they imitate each other. Two types

of mimicry are distinguished.

In Batesian mimicry, a non-toxic edible but-

tery looks like a toxic or distasteful species. The

non-toxic species prots because an experienced

predator will associate the conspicuous color

pattern with the distastefulness of the toxic but-

tery. However, if non-toxic mimics are too com-

mon, predators will not make the association.

Thus for Batesian mimicry to be maintained, it

is important that non-toxic mimics are present in

lower frequency in the habitat than toxic model

species.

Mimicry ring with Mechanitis lysimnia (top), Heliconius hecale

(bottom, left) and Lycorea halia (bottom, right)

Mimicry ring with Dismorphia theucharila (Pieridae) (top), Greta oto (Nymphalidae) (bottom, left) and Phanoptis vitrina (Notodon-

tidae) (bottom, right)

In the second type, Müllerian mimicry, all in-

volved butteries are more or less distasteful and

have evolved a similar color pattern of the wings.

They prot from the mimetic resemblance since

a predator will learn faster when the warning

color pattern is more frequent. This type of mim-

icry is much more common, and it is considered

to be a more stable evolutionary strategy.

Rainforest gap in the Piedras Blancas National Park. In natural rainforests, gaps are regularly produced by fallen trees. Many

buttery species such as those of the Swallowtail genus Parides are most easily observed in light gaps.

Papilio larva with everted osmaterium Mud puddling swallowtail (Heraclides thoas)

Papilionidae

Swallowtail Butteries

The Papilionidae include many large and impressive butteries. Some, but not all, have typical short

tails on their hindwings that lend the entire group the name swallowtails. One feature, which uniquely

characterizes all caterpillars of Papilionidae, is the osmaterium, a stink-producing organ that delivers

an odious smell when it is extruded, and probably serves as a defense mechanism against predators.

The host plants of the Costa Rican swallowtail caterpillars belong to the plant families Aristolochia-

ceae, Annonaceae, Rutaceae, Piperaceae and Umbelliferae. Swallowtails are conspicuous insects and

can be seen in all habitats of Costa Rica. They are fast iers and quite active in the sunshine. In many

species, the males and females hardly resemble each other in appearance. In such cases, their wing

patterns are described as being sexually dimorphic. Normally, both sexes visit owers to feed on nectar,

but males are often encountered on moist sand or around puddles. They use their proboscis to suck up

mineral substances from the water to supplement their diets. This behavior is called ‚mud puddling‘.

Parides crithalion (Variable Cattleheart)

Forewing length: 38 - 43 mm

The sexes of this buttery are conspicuous-

ly dimorphic. The males have a green band of

variable size on the forewing and they are dis-

tinguished by having very reduced white spots

on the forewing. Both sexes have large red ar-

eas on the hindwings. The species is restricted to

rainforest. Both sexes visit owers in the morn-

ing and remain active until late afternoon. Like

many other Parides species, these butteries ut-

ter and move their forewings while feeding nec-

tar on owers, yet at the same time, they keep

their hindwings open and motionless. The ow-

er visiting butteries of this genus are thus very

conspicuous, even from a great distance. Females

lay their eggs during midday on host plants of

the genus Aristolochia that may grow along for-

est edges and in light openings of the forest.

Three other species of Parides can be found in the

Piedras Blancas National Park. One of the most

common ones is Parides iphidamas, which is toler-

ant of disturbed habitats. The males of these spe-

cies can be distinguished from each other mainly

by the shape of the green band. However, the

females are extremely similar to each other and

often cannot be reliably identied.

Parides larva on Aristolochia

Parides erithalion male

Parides erithalion female

Heraclides thoas (Thoas Swallowtail)

Forewing length: 57 - 61 mm

Heraclides thoas is a large and conspicuous

swallowtail. The hindwing tail is rather long.

The black wings are marked with oblong yellow

spots. Heraclides thoas appears very similar to its

sibling species, Heraclides cresphontes, and can be

distinguished from it by a notch in the genital

region of the male and the yellow spot on his tail.

Swallowtails are very fast iers, which help them

avoid predation by birds. Both species of Hera-

clides are regularly seen in moist and wet forests,

such as forest edges, clearings or along streams

where sunlight penetrates the forest canopy.

They are frequently encountered in gardens and

parks to which these marvelous butteries are

attracted. There they nd owers rich in nectar,

such as Stachytarpheta and Lantana. The mineral

substances, which the males take up from wet

sand or muddy roads, are necessary for produc-

tion of the spermatophore, which is transferred

to the female genital tract during copulation. The

eggs are laid singly on pepper trees (Piper sp.).

The caterpillars resemble bird droppings, which

represents a very effective camouage and dis-

courages birds from eating them. If a caterpil-

lar is disturbed, the stink organ (osmaterium) is

turned inside out and projects like a fork shaped

orange antenna. It exudes a nasty smell, which is

presumed to act as a repellent against predatory

birds.

Heraclides thoas caterpillars on Piper leaf

Heraclides thoas

Pasture near the research station. Open habitats such as pastures are the preferred habitat for many Sulphures and Whites.

Some species of the genera Phoebis and Pyrisitia can reach high abundances at those places.

Pieridae

Sulphures and Whites

Pieridae are small to medium-sized butteries usually with white, yellow or orange wings.

Many species present a simple pattern consisting of few small black or brown dots, lines or

dark wing margins on the otherwise uniformly colored wing. In addition, the wing pattern of

many species reects UV-light, which is invisible to human eyes, but important for con-spe-

cic butteries in courtship display. All species of Pieridae are ower visitors. They can be

observed readily when they are nectar feeding. In many species, the color pattern on the un-

derside of the wings is cryptic. When they close their wings, they remarkably resemble a leaf.

Most important host plants for the larvae are species of Mimosaceae, Fabaceae, Caesalpiniace-

ae, Brassicaceae, Capparidaceae and Loranthaceae. Many of these plants contain chemical toxins,

which have been shown to be important cues for egg laying and feeding stimulus. The caterpil-

lars are smooth; many are green or brown and very inconspicuous when on a plant. For pupa-

tion, the caterpillars spin a silk girdle that attaches the pupa at a 45° angle to a twig or other part

of a plant. The pupae are cryptic, some look like young leaves, buds or even bird droppings.

Pyrisitia nise (= Eurema nise) (Mimosa Yellow)

Forewing length: 16 - 19 mm

Pyrisitia nise is a small buttery; the wings are

yellowish; black markings are present along the

front margin of the forewing, which is arched

from the costa to the tornus. The underside of the

wings is dull yellow and lacks the black mark-

ings along the margin. The butteries y low to

the ground and visit a variety of owers to feed

on nectar. This is one of the most common but-

teries in disturbed habitats near the station.

Although they are small, they are known as fast

iers and are not easily caught by predators. The

larvae feed on leaves of the Sensitive Plant (Mi-

mosa pudica).

Similar and related species from the genus

Pyrisitia inhabit open areas and are especially

common where human activity has created pas-

ture and farmland. All species of this genus are

easily recognized due to their small size and the

yellow, orange or white wing coloring with a

black margin. The males often are brighter and Pyrisitia nise

Phoebis sennae (Cloudless Sulphur)

Forewing length: 29 - 36 mm

Phoebis sennae is a medium-sized buttery

with yellow wings. Those of the male are clear

yellow with lighter yellow distal margins. Those

of the females are usually lighter yellow with

a black wing margin. On the underside of the

wings are brown dots and spots. The butter-

ies y rapidly in the sunlight and are regularly

seen along roads, in gardens, pastures, and open

elds. They have a long proboscis and can gain

access to the nectar in long tubular owers, such

as Stachytarpheta. Phoebis sennae, like all species of

the genus Phoebis, is a migratory buttery. They

regularly travel long distances to colonize new

habitats.

The larvae of Phoebis sennae are green and as-

toundingly cryptic. Their host plants belong to

the family Caesalpiniaceae. When not feeding,

the caterpillars often rest beneath leaf petioles.

Phoebis sennae male Phoebis caterpillar

may aggregate on wet ground to take up mineral

substances.

Dismorphia theucharila (Clearwing-Mimic White)

Forewing length: 25 - 27 mm

This buttery species is encountered in wet

natural forests where it ies in low densities.

Dismorphia theucharila is the only pierid but-

tery in Costa Rica that has partly transparent

wings. The species is difcult to separate from

some nymphalid butteries because of the close

resemblance in wing coloration and ight behav-

ior. The resemblance is an amazing example of

Batesian mimicry, which includes toxic model

species and edible mimicking butteries. Dis-

morphia theucharila is palatable to bird predators,

in contrast to the distasteful and/or toxic model

species of nymphalid butteries, such as Mecha-

nitis polymnia (mentioned below). Nonetheless,

Dismorphia theucharila, like all Pieridae, can be

distinguished from the nymphalids because it

stands and walks on six legs, while the nympha-

lids use only four legs.

Dismorphia theucharila

Rainforest creek near La Gamba. Creeks and rivers often produce gaps in the rainforest which are used for patrolling male but-

teries, e.g. the Longwings of the genus Heliconius, in search of food or females.

Nymphalidae

Brush Footed Butteries

Nymphalidae contain a large number of species exhibiting amazing life history traits.

The family includes species of different sizes from small to very large. The shape, color-

ation and pattern of the wings are highly diverse. Likewise, the feeding preferences of the

adults vary greatly. Some representatives of this group feed on oral nectar, others suck

on rotting fruits or tree sap. In several species, the adults have never been observed feed-

ing on owers or other plant sources, rather only on fresh dung, urine or decaying carrion.

Despite the great diversity of life histories and the unresolved phylogeny of these butteries,

all adult Nymphalidae can be recognized by the fact that they stand and walk on four legs only.

The greatly reduced forelegs bear sensory organs, which probably detect larval host plants. The

caterpillars, too, show a great variety of sizes, shapes and feeding preferences. Most pupae hang

with their head down, anchored with special hooks to a silk web, which was spun by the caterpillar.

Subfamily Danainae

Lycorea halia (= cleobaea) (Tiger Mimic-Queen)

Forewing length: 44 - 51 mm

Lycorea halia is a medium-sized buttery with

an orange to brown wing coloration and white

dots along the black wing margins. The color

pattern is named ‘tiger-stripe’. It is a conspicu-

ous warning coloration (aposematic) and strong-

ly resembles that in several distasteful and/or

toxic Neotropical butteries. All these co-mimic

species take advantage of imitating each other

since predatory birds learn to avoid unpalatable

species more rapidly if butteries displaying the

same warning coloration occur in greater num-

ber in a given habitat.

In comparison to other tiger-stripe butteries,

Lycorea halia can be identied by the rather short

yellow antennae. The butteries are found in the

forest and open areas where they feed on nectar

from a variety of owers or where males wait

for females. The males perch in the morning un-

der the subcanopy and chase away conspecic

males. Occasionally, they extrude the hair pen-

cils on the abdomen and send out pheromones

to attract females.

The preferred host plants of the larvae belong

to Papaya and Asclepias, both of which contain

milksap. The larvae cut a ring into the leaf tis-

sue or veins and wait for about an hour before

feeding. The behavior presumably prevents mo-

bilization of defensive plant compounds. Later

instars are white with black rings and have a

pair of movable laments behind the head. The

caterpillars are probably distasteful to birds due

to the toxicity of the host plants. The larvae store

the secondary chemical compounds and trans-

fer them to the adults via the pupal stage. The

substances are used by the adult butteries as a

means of chemical defense. Together with toxic

pyrrolizidine alkaloids, which the butteries

gather from other plants, the substances are es-

sential precursors for the production of the male

sex pheromone.

Pupa of Lycorea halia

Lycorea halia (right) and co-mimic Heliconius hecale (left) Lycorea halia larva

Danaus plexippus (Monarch)

Forewing length: 41 - 48 mm

The Monarch buttery has a large global dis-

tribution and is most famous for its extraordinary

migrations across North America. The butteries

of the last generation of the summer will travel

up to 3,600 km south to their overwintering sites.

The populations in eastern North American mi-

grate to the Central Mexican highlands. There

millions of individuals congregate among the

conifer trees. However in Costa Rica, the species

is found all year round in small numbers, usu-

ally in open areas. The caterpillars feed on poi-

sonous milkweeds (Asclepias sp.). They are able

to sequester cardiac glycosides from their food

plant, which renders them unpalatable for most

predators. This protection extends into the adult

stage, and the conspicuous orange color pattern

signals to potential predators that the butteries

are unsuitable as food.

Danaus plexippus

Danaus plexippus larva on Asclepias

Subfamily Ithomiinae

Mechanitis polymnia (Polymnia Tigerwing)

Forewing length: 34 - 38 mm

Mechanitis polymnia has wings that are large-

ly orange with black stripes and white dots. It

is another abundant member of the tiger-stripe

mimicry complex, which includes several unre-

lated distasteful and toxic butteries. Mechanitis

polymnia can be distinguished by the rather small

eyes and the round orange spot on the upperside

of the forewing margin. It is very common and

widespread in Costa Rica. It ies in sunny open

areas where it feeds on the owers. Males settle

in the shade of the forest and wait for females.

The males are known to use their proboscis to

acquire precursor substances to produce their

sex pheromones (pyrrolizidine alkaloids) from

the wilted leaves of particular species of Boragi-

naceae and Asteraceae. The females are attracted

by the male sex pheromone. During copulation,

the poisonous alkaloids are transferred via the

spermatophore of the male to the female. In ad-

dition to oral nectar, female butteries have

been recorded to feed from bird droppings, pre-

sumably as a source of nitrogen. The buttery

has received the name ‘ant buttery’ because of

its habit of following birds, which hunt insects

trying to escape the procession of army ants.

The larvae feed on the poisonous plant genus

Solanum, and they possess a chemical similarity

between their cuticular lipids and those of the

host plant which protects them from predatory

ants.

Mechanitis polymnia larva Mechanitis polymnia

Aeria eurimedia

(Black-and-Yellow Prestonian or Eurimedia Clearwing)

Forewing length: 22 - 25 mm

The wing pattern of this species is black and

yellow. A red stripe is evident behind the head.

This small buttery is commonly encountered

as a solitary individual ying in deeply shaded

forests close to the ground, usually near water-

courses. In primary forests, both sexes visit ow-

ers, such as Psychotria and Cephalis, during the

morning. The larvae feed on Prestonia portabellen-

sis. First, they cut veins in older leaves to stop the

ow of toxic milksap. After briey waiting, they

are able to feed on the leaves without becoming

poisoned.

Aeria eurimedia

Hyposcada virginiana (Virginia‘s Ticlear)

Forewing length: 32 - 35 mm

Hyposcada virginiana is widespread in Costa

Rica and one of the more common Ithomiines

in the rainforests around the station. It also in-

habits secondary forests. It usually ies in the

deep shade. The females

lay their large white eggs

on woody, hemiepiphytic

species of Drymonia, which

belong to the Gesneria-

ceae, a plant family that

is not used by other but-

teries in Costa Rica. The

caterpillars cut round

holes into the interior of

the leaf blades, a behav-

ior which is also found in

other Ithomiines, whereas

most caterpillars feed at

leaf margins. Several but-

teries from other sub-

families are very similar

in wing pattern and form

a mimicry ring. Among

them are Heliconius hecale Hyposcada virginiana

(Heliconiinae) and Eresia ithomioides (Nymphali-

nae) in its color form melaina. Hyposcada virgin-

iana can be distinguished by the rounded shape

of the outer forewing.

Ithomia celemia (Celemia Clearwing)

Forewing length: 30 - 33 mm

Ithomia celemia belongs

to the tiger-stripe mim-

icry complex and is most

similar to Hypothyris eu-

clea, a common Ithomiine,

as well as to the light color

morph of Eresia ithomioides

(Nymphalinae). Its range

extends from Costa Rica

southwards to Venezuela.

In Costa Rica, it rarely ap-

pears outside the Golfo

Dulce Region, and is only

found in rainforests on

the Pacic side. The larval

food plant is Witheringia

riparia, a member of the

family Solanaceae, which

is the major host plant

family for species of the

subfamily Ithomiinae. Ithomia celemia

Subfamily Charaxinae

Consul fabius (Tiger Leafwing)

Forewing length: 36 - 40 mm

Consul fabius is a member of the subfamily

Charaxinae, which is most diverse in the Afro-

tropical Region. At rest, this buttery looks like a

dead leaf. The upperside, however, is not cryptic

but bright orange and resembles members of the

tiger-stripe complex, such as Mechanitis polymnia

from the subfamily Ithomiinae. It is probably

one of the best examples of Batesian mimics in

Costa Rica. The butteries also imitate the slow

ight of their models, but when alarmed, they

can y very fast, like other members of the sub-

family. The butteries do not visit owers but

are attracted to rotten fruits. They are often ob-

served along forest trails where the males perch

in the subcanopy. The females lay their eggs on

numerous species of Piper, the larval host plant.

The mature caterpillars hide inside rolled tubes

at the end of the leaves. Consul fabius

Fountainea eurypyle (Pointed Leafwing)

Forewing length: 28 - 34 mm

Fountainea eurypyle belongs to a large group

of similar-looking species, which have previ-

ously been placed in the genus Memphis. All of

them have a cryptic underside resembling dead

leaves, and are therefore difcult to spot when

resting in the trees. The upperside, however, is

more colorful, ranging from orange brown (as

in Fountainea eurypyle) to an iridescent blue or

green. Like other Charaxinae, adult butteries

do not visit owers, but are attracted to fruits.

Most species occur in primary and secondary

rainforests. Fountainea eurypyle is one of the more

common species, which has been observed in

the garden of the station. Its caterpillars feed on

leaves of Croton (Euphorbiaceae).

Fountainea eurypyle

Subfamily Satyrinae

Morpho helenor (= peleides) (Peleides or Helenor Morpho)

Spanish name: Celeste común, morfo

Forewing length: 64 - 78 mm

This Morpho buttery is one of

the most famous in the rainforest. It

seems to be depicted in nearly ev-

ery book about the natural wildlife

of Costa Rica. It has iridescent blue

wings and ies with a low frequency

of wing beat. The large wings of the

males have a clear shiny blue hue and

black margins. In females, the blue

is reduced and the black margin is

broader. Morpho helenor is regularly

seen along forest trails and woodland

streams, but also in plantations. The

butteries are very popular in butter-

y gardens and nurseries since they

can be easily bred in captivity.

Although the appearance of the buttery is

very conspicuous, their fast and erratic ight

makes them difcult to catch, even for birds.

This behavior is an appropriate strategy, since

Morpho butteries do not store toxins in their

body as many other butteries.

Morpho helenor is peculiar since it does not

visit owers. Instead, the butteries feed on rot-

ting fruits, such as bananas and fruit peels. Also,

they drink tree sap, which ows from cuts in the

bark. Their proboscis is relatively short and has

long sensory structures near the tip. The shape of

these structures is important for sucking liquids

from moist surfaces, since they serve to acquire

liquids by adhesion at the tip of the proboscis.

When feeding, they always close their wings and

show their cryptic underside. A row of eyespots

along the wing margins probably serve to dis-

tract potential predators.

Morpho helenor, male

Morpho helenor, female Morpho helenor

Morpho helenor larva

Morpho menelaus (= amathonte) (Menelaus Morpho)

Forewing length: 73 - 87 mm

The sexes of this large

buttery are very dis-

tinct from each other in

appearance. The males

have an iridescent light

blue on the upperside of

the wings, while the fe-

males have wide brown

margins and two rows of

white spots. The under-

side of the wings in both

sexes is washed reddish

brown, and the eyespots

are reduced. The bright

blue of the upperside of

the wings in the males is

the result of light diffract-

ing from the wing scales.

Diffraction colors are

produced by a series of

parallel grooves or ridges

on the scales. The grooves

are separated from each

other by the distance of

the particular wavelength

of light. Light reecting

from such parallel struc-

tures interferes with the

reectance from neigh-

boring parallel structures.

Thus, at particular angles, the light of a given

wavelength will be either reinforced or cancelled

out. Flying male Morpho butteries are easily

identied by the blue reection of their wings,

even when they y through the dark forest with

their typical low frequency of wing beat. The

underside of the wings is cryptic, and when the

buttery rests, it closes its wings over the bark of

a tree and becomes nearly undetectable.

Morpho menelaus with wing damage

This buttery is regularly found in rainforest

habitats. The males actively patrol along forest

edges and rivers during the early morning soon

after sunrise. By late morning, they become less

active and feed on eshy fruits, as do the females.

At midday, the females search for oviposition

sites. Just before dusk, both sexes slowly utter

along the forest ridge tops and settle on the un-

dersurfaces of leaves to sleep for the night.

Caligo eurilochus (Owl Buttery or Forest Giant Owl)

Spanish name: Buhito pardo

Forewing length: 77 - 91 mm

The Caligo species are among the largest but-

teries in Costa Rica. Their wingspan can reach

up to 18 cm. The dark wings of the males show a

dull iridescent blue. The underside of the hind-

wing is cryptic and has a large eyespot, which

resembles a large pupil with a light-colored iris.

Probably, it distracts predators for a moment.

When a predator attacks, the conspicuous eye-

spot is often the target, and the buttery can es-

cape. Due to its large wings, the buttery is still

capable of ight, even if large parts of the wings

are missing. Caligo butteries are most active at

dusk and dawn. In the daytime, they rest with

closed wings on tree bark. Adults feed exclu-

sively on rotting fruits, tree sap and mammal

droppings.

The caterpillars are considered a pest in ba-

nana plantations. Late instars can measure about

10 cm in length. Astoundingly, a single caterpil-

lar can consume an entire banana leaf in a few

days. The natural host plants of the larvae in

Central America are species of the plant genus

Heliconia.

Caligo eurilochus caterpillars on a banana leaf

Caligo eurilochus

Caligo atreus (Yellow-Edged Giant-Owl or Arteus Owl)

Forewing length: 73 - 85 mm

Caligo atreus is an inhabitant of the Costa Rican

rainforests; the larvae feed on plants from genus

Heliconia, Musa and different genera of the fam-

ily Cyclanthaceae. Its wings are gray to brown

with large eyespots on the underside of the hind-

wings (similar to that of C. eurilochus, mentioned

above) and small eyespots on the forewings. The

buttery can be easily identied. When it is star-

tled, it reveals a conspicuous dull blue shimmer

on the upperside of the forewing that contrasts

to the yellow stripe of the hindwing. When in

ight, the huge size, low frequency of wing beat

and blue and yellow coloration on the upperside

of the wings are revealing of the species. How-

ever, the buttery is difcult to notice when rest-

ing on a tree trunk with closed wings because

of the cryptic coloration on the underside of the

wings. Caligo atreus

Catoblepia orgetorix (Orange-rimmed Owl-Buttery)

Forewing length: 48 - 55 mm

Apart from the Giant Owl-Butteries of the

genus Caligo, several smaller Owl-Butteries oc-

cur in Central America, most of which belong to

the genera Eryphanis, Opsiphanes and Catoblepia.

All are butteries of the forest understory and are

rarely observed due to their crepuscular behav-

ior. In contrast to their camouage wing pattern

on the underside, the upperside is often bright

orange or marked with iridescent blue. Catoblepia

orgetorix is a species which can been found in the

Piedras Blancas National Park. The upperside of

the wing is dark brown with an orange marginal

band. The underside bears prominent eye spots

and a pinkish hue. The butteries feed on fallen

fruits, the caterpillars feed on the leaves of vari-

ous species of palms (family Arecaceae).

Catoblepia orgetorix

Hermeuptychia hermes (= Cissia hermes) (Hermes Satyr)

Forewing length: 16 - 20 mm

Hermeuptychia hermes is one of the most com-

mon butteries in the garden of the station and

along the roads of La Gamba. The buttery

is small, and the wings have a grayish hue on

both sides. On the hindwings are several small

eyespots, three of which are darker. On the fore-

wings are three small spots, as well. The butter-

ies feed on fruits and dung; occasionally they

are seen on owers drinking nectar.

Hermeuptychia hermes can be distinguished

from similar looking species of the genus and

related genera Cissia and Magneuptychia, by its

smaller size, number and arrangement of the eye-

spots. Most of the other species are only found

singly along forest tracts. More common is Mag-

neuptychia libye, a large species with a forewing

length of 22-23 mm, which occurs in secondary

forests around the station.

The caterpillars of most species are nocturnal

and feed on various grasses from the family Poa-

ceae.

Hermeuptychia hermes Magneuptychia libye

Hermeuptychia hermes

Cithaerias pireta (= menander)

(Pink-Tipped Satyr)

Forewing length 30 - 33 mm

Cithaerias pireta occurs in wet rain forest habi-

tats. It ies close to the forest oor in deeply

shady areas. The wings of this medium-sized

buttery are transparent. The hindwings are

characterized by a reddish hue and an eyespot.

Male butteries have been observed over a peri-

od of several weeks to visit consistently the same

perching site in light openings of the rain forest.

Cithaerias pireta

Pareuptychia ocirrhoe

(= Cissia hesione) (White Satyr)

Forewing length: 18 - 21 mm

Unlike most related satyrines, this Central

American species has a pearly white upperside

and a striped underside. It is often encountered

in secondary bush and along waysides near the

station. The females lay their eggs on grasses of

the genus Eleusine at the base of trees.

Pareuptychia ocirrhoe

Pierella luna

(Quiet Diaph or Moon Pierella)

Forewing length: 33 - 39 mm

Pierella luna has a dark brown color on the

upperside of the wings. The hindwings are con-

spicuously large and have two black dots on the

front margin. One spot has a white dot in the

center that resembles an eyespot. The underside

of the wings is lighter.

The buttery is characteristic of primary forest

habitats. It is often encountered on forest trails

ying low to the ground. Because of its dark col-

oration, it is difcult to see in the low light of a

rainforest. Especially when it sits on the ground

with closed wings, the buttery seems to vanish

before our eyes. With some patience, male ter-

ritorial behavior can be observed in this species.

The males defend the boundaries of their terri-

tory against rival males by chasing them away.

Remarkably, the butteries have never been ob-

served to visit owers. Instead, they feed almost

entirely on matter exuded by fungi.

Pierella luna

Subfamily Cyrestinae

Marpesia merops (Merops Daggerwing)

Forewing length: 29 - 33 mm

The buttery is regularly encountered on un-

paved roads near the station, usually with its

proboscis in the wet sand (sucking in mineral

substances). The brown-gray wing coloration of

Marpesia merops blends in very well with ground

textures. The buttery is nearly impossible to see

from a distance. It resembles a brown swallow-

tail because of the long tail on each hind wing.

Near the tip, on the upperside of the forewing

are several small white spots. The underside of

the wings is grayish white with brown streaks.

Marpesia chiron is similar but bears a conspicuous

red mark near the tornus of the hindwing. More

different is Marpesia petreus which has a bright

orange upperside coloration. Marpesia species

have never been seen on owers.

Marpesia merops

Marpesia merops Marpesia chiron

Marpesia petreus

Subfamily Biblidinae

Dynamine tithia (=salpensa) (Tithian Sailor)

Forewing length: 16 - 18 mm

The genus Dynamine encompasses several

small-sized butteries, which have a similar or-

ange-banded pattern on their undersides, some-

times with silver spots. Among them, Dynamine

tithia is characterized by its reective blue upper-

side. In Costa Rica, this species is only found in

the southernmost part. It is most easily encoun-

tered at mud puddles along river banks. The fe-

males lay their eggs on the buds of Dalechampia

triphyllia (Euphorbiaceae), and the caterpillars

feed on the ower parts or bore into the develop-

ing ovary. Dynamine tithia

Hamadryas feronia and Hamadryas arinome (Cracker Butteries)

Forewing length: 35 - 38 mm

Hamadryas species of the feronia-group are

famous for the crackling noises that individuals

produce in interaction with other individuals.

The interactions are mostly male-to-male and

represent territorial ghts. Resident males im-

mediately y from their perch site in their ter-

ritory toward any intruding male to prevent it

from entering. The mechanism of sound produc-

tion involves movements of the wings, but is in-

completely understood.

Two species have been identied in the gar-

den of the station. The males can regularly be

seen perching on trees trunks with the head

characteristically in a downward position and

the wings spread open. Hamadryas feronia is dis-

tinguished from Hamadryas arinome by the grayer

color and by the black zigzag lines on the up-

perside of the wings. Hamadryas arinome can

be identied by the blue color of the wing pat-

tern and the white band on the forewing.

Hamadryas feronia

Hamadryas arinome

Subfamily Nymphalinae

Anartia fatima (Banded Peacock)

Spanish name: Cocinera

Forewing length: 27 - 30 mm

Anartia fatima is the most common buttery

in Costa Rica. It can be seen in the garden of

the station throughout the entire year where it

comes to feed on nectar from various owers.

The wings are dark brown except for a white or

yellow band. On the inside of their hindwings

are red dots, which enable one readily to rec-

ognize the species. Males start out with yellow

bands, but over time, the color of the bands fade

to white. The discoloration has allowed studies

to be conducted on age structure in various pop-

ulations. The adults store no defensive chemical

substances in the body or wings. Lacking chemi-

cal protection, the butteries are a welcome

meal for many predators, such as spiders, frogs,

lizards, birds and certain mammals. That some

individuals can escape and survive an attack is

occasionally evident in butteries with pieces of

the wings missing. Anartia fatima

Anartia jatrophae (White Peacock)

Forewing length: 28 - 30 mm

A second member of the genus, Anartia jat-

rophae is likewise very common and can be

observed daily on owers in the garden of the

station. It prefers to drink nectar from Lantana,

which is abundant in the surroundings of the

station. The wings of A. jatrophae are grayish-

white with brown streaks all over.

Actual ower visiting behavior of both Anar-

tia species can readily be observed on Lantana ca-

mara, for example. The buttery often sits in the

middle of the inorescence (i.e. a ower cluster).

It uncoils the proboscis and quickly begins to

probe for a ower opening. The tip of the pro-

boscis is inserted into a single ower by lower-

ing the entire proboscis. Sensory organs at the tip

help detect the presence of nectar in the corolla

of a ower. After taking up nectar, the buttery

withdraws from the ower by raising the pro-

boscis as a whole and turns to the next ower of

the inorescence. By repeating this sequence of

feeding movements, a buttery can exploit rap-

idly and effectively an inorescence with numer-

ous small owers, like that of Lantana camara.

The caterpillars are known to feed on dif-

ferent herbs of the plant families Acanthaceae,

Scrophulariaceae and Verbenaceae.

Anartia jatrophae

The larvae of Anartia fatima feed on plants

of the family Acanthaceae. The caterpillars are

abundant yet very difcult to nd in nature.

Junonia evarete (Mangrove Buckeye)

Forewing length: 26 - 29 mm

This middle-sized buttery is distinguishable

by the orange wing pattern of the upperside, an

eyespot on each forewing and two on each hind-

wing. Junonia evarete occurs in disturbed habi-

tats, such as cattle pastures and along beaches.

The buttery visits various owers to feed on

nectar. Males occasionally alight on the ground

to obtain mineral substances from mud puddles.

Without these minerals, they cannot successfully

mate. The larvae feed on Acanthaceae, such as

Ruellia.

Junonia evarete

Siproeta stelenes (Malachite)

Forewing length: 45 - 48 mm

Siproeta stelenes is common in a variety of hab-

itats — forest edges, secondary forests and cul-

tivated lands. It is easy to recognize because of

the pale green on the upperside of the wings, the

broad black wing margins and the small tail on

the hindwing. It probably is a Batesian mimic of

the very similarly colored nymphalid buttery

Philaethria dido. Siproeta stelenes is frequently seen

in open areas and gardens, where they feed on

oral nectar or squashed fruits. The larvae feed

on plants of the family Acanthaceae.

Siproeta stelenes

Anthanassa frisia (= tulcis) (Cuban Crescent)

Forewing length: 16 - 19 mm

This is a small and inconspicuous buttery,

which can be recognized by the indented margin

of the forewing and the brown coloration with

light dots and lines on both wings. The upperside

of the wings is darker than the underside. It is a

very common buttery in elds, pastures, along

roadsides, and even in oil palm plantations. An-

thanassa frisia ies close to the ground and often

visits small owers in the low vegetation to feed

on nectar. Dicliptera and possibly other genera of

Acanthaceae are used as larval hostplants.

Anthanassa frisia

Chlosyne janais (Crimson Patch)

Forewing length: 21 - 30 mm

Chlosyne janais is a common buttery in Cen-

tral America and one of several similar looking

species. All are characterized by scattered white

spots on the black forewing and an orange patch

on the hindwings. Fertile females may be so

weighted down with eggs in their abdomen that

they can barely manage to take ight. Nonethe-

less, they lay eggs in large clusters on various

host plants from the families Asteraceae, Acan-

thaceae and Amaranthaceae. The caterpillars

bear many small black spines on the body and

occur in two color morphs (red and black). They

feed gregariously on the host plants.

Chlosyne janais

Chlosyne janais larva,

black and red morph

Chlosyne janais pupa

Colobura dirce (Small Beauty, Zebra Mosaic)

Forewing length: 35 - 37 mm

The wings of this buttery are black with a

large yellow band on the upperside of the fore-

wing; the hindwing is brown in color. The un-

derside is cryptic and has zebra-like markings.

The host plants of the larvae are Cecropia trees,

which live in symbiosis with Azteca ants. Al-

though the ants vigilantly protect against other

insects, the caterpillars manage to feed on this

plant. Females also frequently oviposit on plants

without ant colonies. The adults of Colobura dirce

feed on rotting fruits, carrion, dung and abstain

entirely from oral nectar. A very similar sibling

species, Colobura annulata, which was described

and separated from C. dirce only a few years ago,

has now also been found in La Gamba. It merely

differs in the form of the stripes on the forewing

underside. The third dark brown submarginal

line narrows in the direction of the costa in C.

dirce, whereas it is uniformly thick throughout

its length in C. annulata.Colobura dirce

Historis odius (Orion Cecropian)

Forewing length: 50 - 55 mm

This large species resembles members of the

subfamily Charaxinae, although it is not close-

ly related to them. The underside of the wings

resembles dead leaves; the upperside bears an

extensive orange patch. The buttery is distin-

guished from its congener, H. acheronta, by the

Historis odius feeding on rotten bananas

single white spot at the forewing apex. The but-

teries are attracted to rotten fruit. Although

they usually spend most of their time in the

canopy, they are easily observed at baits placed

near the ground.The caterpillars feed on Cecropia

(Moraceae).

Subfamily Limenitidinae

Adelpha cytherea (Smooth-Banded Sister)

Forewing length: 22 - 25 mm

The butteries of the genus Adelpha are medi-

um-sized and typically have dark brown wings

with an orange and white pattern. In Costa Rica,

the genus contains a number of very similar look-

ing species, some of which are difcult to identify

at the species level. Species of Adelpha occur in all

habitats from sea level to mountainous regions.

In most Adelpha butteries, the dark brown up-

perside of the forewing has a prominent orange

patch or band, and it is aligned with a continu-

ous white band on the underside of both wings.

Otherwise, the undersides have a light brown

color and white bands across both wings. Adel-

pha cytherea is the most common Adelpha species

in cultivated lands and secondary forests where

it can be observed to feed on owers and fruits.

Its larvae feed on the Wooly Woodvine (Sabi-

cea villosa). Other similar looking members of

the genus occur in the natural rainforest habi-

tats around the station. They are usually seen

perching on the top of the trees and hardly ever

come close to the ground. The larval food plants

of many Adelpha species are still unknown, but

those which are known belong to a wide range of

plant families. Adelpha boeotia feeds on Cecropia

(Moraceae), as well as Luhea seemani (Tiliaceae),

while the larval host plant of Adelpha salmoneus

belongs to the genus Sabicea (Rubiaceae).

Adelpha cytherea

Adelpha boeotia

Adelpha salmoneus Adelpha salmoneus

Subfamily Heliconiinae

Dryas iulia (Julia)

Forewing length: 41 - 45 mm

Dryas iulia is easily identied by its bright or-

ange color and elongate forewings. The buttery

is a graceful ier with its long wings and low fre-

quency of wing beat. It occurs at the edge of the

forest, in the forest canopy and openings along

forest roads and trails. The adults frequently vis-

it many owers for nectar such as Lantana. They

can be readily observed in all times of the year in

the garden of the station and along roads in the

vicinity. The caterpillars of Dryas iulia are light

brown with white bands and bear long spiny

extensions on each body segment. They feed on

passion vines (Passioraceae).

Dryas iuliaDryas iulia larva

Eueides lybia (Sharp-edged Longwing)

Forewing length: 29 - 31 mm

Eueides lybia is associated with rainforest

habitats. This species rarely ies in direct sun-

light and does not occur in secondary vegetation

growth. The color of the wings is orange with a

black forewing apex and broad black hindwing

margins. E. lybia is one of the buttery species

which sleep gregariously. The caterpillars feed

on Passiora vitifolia.

Eueides lybia

Philaethria dido (Green Longwing)

Forewing length: 49 - 54 mm

Philaethria dido is a common buttery in the

lowland primary forests around the station.

However, it is rarely seen because this butter-

y mainly ies above the forest canopy. The fe-

males visit light openings and the forest edges to

Philaethria dido

oviposit on their host plants of the plant genus

Passiora. P. dido is one of the few examples of

a buttery with a green color pattern on the up-

perside of the wing. It is mimicked by Siproeta

stelenes.

Heliconius (Longwing Butteries or Passion Vine Butteries)

In our region of Costa Rica, eight species

have been recorded from the genus Heliconius.

All are characterized by the elongate forewings,

the long and club-tipped antennae, their slow

ight and unique mode of pollen feeding. Most

species are very colorful. They have stimulated

considerable biological interest, which has led to

studies on mimicry, feeding ecology and evolu-

tion. The host plants of the caterpillars are pas-

sion vines (Passioraceae). The plants contain

toxic cyanogenic glycosides, which protect them

against many herbivore insects. However, the

caterpillars of Heliconius and other butteries

have evolved the ability to process the poisons

and put them to use for their own protection.

Extraordinary is the fact that Heliconius spe-

cies are able to feed on pollen, aside from nectar.

The butteries actively gather pollen from ow-

ers onto the shaft of the proboscis and mash it

together with saliva. Afterward, they ingest the

liquid, which is rich in extracted amino acids.

The special pollen feeding behavior is crucially

tied together with their remarkable longevity

(up to nine months!), the ability to constantly

produce eggs and to mate on multiple occasions.

Ultimately, the chemical defense mechanisms

of the adults benet from the extra nitrogen as

well.

A further remarkable feature of the life his-

tory of some Heliconius species is pupal mating.

Male butteries claim female pupae by sitting

on them. Mating occurs invariably in the early

morning immediately upon eclosion (i.e., emer-

gence from pupal case) or beforehand with the

pupa. One knows that female pupae are present,

when several males are seen ying in the low

vegetation, trying to chase each other away from

the female pupae.

Heliconius buttery processing pollen with its proboscis

Heliconius sapho (= hewitsoni) (White-Patched Longwing)

Forewing length: 34 - 41 mm

Heliconius sapho is one of the most abundant

butteries in the forests near the station. It has

black wings with yellow bands. Viewed from the

underside, remarkable red areas become visible

at the base of the wings. The buttery greatly re-

sembles another species that is common in the

region, Heliconius cydno. The species are mem-

bers of a Müllerian mimicry complex — in which

both species are distasteful and prot from look-

ing like each other, since predatory birds learn

quickly to avoid potentially unpalatable prey.

Heliconius sapho Heliconius sapho

Heliconius cydno (= pachinus) (Cydno Longwing)

Forewing length: 38 - 43 mm

In our region of Costa Rica, Heliconius cydno

is almost identical in appearance to H. sapho.

The two species can be distinguished by looking

closely at the yellow stripe on the forewing close

to the body. In Heliconius pachinus, ve wing

veins cross the stripe, whereas in H. sapho only

two to three veins do. The close resemblance of

the butteries probably serves to protect both. If

a predatory bird learns to associate the yellow-

black coloration with a distasteful prey, butter-

ies of both species will be attacked less often.

The butteries emit a pungent smell that arises

from the abdominal glands. The odor can be used

as a eld character for species identication.

Heliconius cydno

Heliconius hecale (Hecale Longwing)

Forewing length: 42 - 50 mm

Heliconius hecale is one of the most common

members of the tiger-stripe mimicry complex.

Other Heliconius species and numerous butter-

ies from various groups belong to this complex,

as well. They all have black, orange and yellow

wing coloration. In general, bright colors serve

as a warning to birds that prey may be unpalat-

able or toxic. Heliconius hecale occurs in all habi-

tats of Central and South America. It has been

subdivided into a number of different looking

subspecies. Both sexes obtain nectar from a vari-

ety of owers, but pollen collection is restricted

primarily to owers of Psychotria (Rubiaceae),

Psiguria and Gurania (both Cucurbitaceae). Some-

times individuals aggressively defend owers

against other butteries. Several individuals of

H. hecale spend the night in low forest vegetation

close to each other in a small group. Similar to H.

cydno, the larvae of H. hecale feed on a variety of

Passiora species.

Heliconius hecale Heliconius hecale

Heliconius sara (Sara Longwing)

Forewing length: 30 - 35 mm

Heliconius sara is a black buttery with white

or yellow wing bands and a blue shining hue. It

is most commonly found along forest edges or in

secondary vegetation. The species lives in close

association with its larval host plant, Passiora

auriculata. Host plant specicity is a major factor

limiting the distribution of H. sara, and is one of

the reasons why this Heliconius species is always

associated with rainforest habitats

Heliconius sara

Heliconius melpomene

(Postman Buttery, Melpomene Longwing)

Forewing length: 35 - 39 mm

Like many other species of the genus Heli-

conius, H. melpomene is subdivided into various

subspecies each with a different coloration, par-

ticularly in other regions of Central and South

America. In Costa Rica, Heliconius melpomene is

black with a red band across the forewings and a

longitudinal light yellow stripe on the hindwings.

The buttery is very conspicuous and single in-

dividuals can be encountered along shady forest

trails and in the garden of the station. Its co-mim-

icking species, H. erato, is rare in La Gamba, and

is usually encountered in the sunshine. The two

species can only be told apart by counting the

number of small red dots on the underside of the

forewing upon close examination.

Heliconius melpomene

Heliconius eratoLarva of Heliconius melpomene

Heliconius ismenius (Tiger-striped Longwing)

Forewing length: 38 - 48 mm

Heliconius ismenius is a member of the Tiger-

striped mimicry complex, but can be distin-

guished from similar species by its large eyes. In

wing pattern it looks almost identical to Melinaea

lilis from the subfamily Ithomiinae. It inhabits

rainforest habitats and prefers to feed on the

owers of Psiguria and Gurania, which are also

excellent pollen sources. The females lay their

eggs singly on seedlings of different Passiora

species.

Heliconius ismenius

Laparus (= Heliconius) doris (Rayed Longwing)

Forewing length: 34 - 45 mm

This Longwing was placed in a separate ge-

nus due to differences in chromosome numbers,

but recent molecular phylogenetic results do not

conrm this splitting from the genus Heliconius.

This is a variable species which can appear in dif-

ferent color forms but can be readily identied

by the ray pattern on the hindwings. It is found

in different forest habitats and regularly seen

around the station. Its larvae are highly gregari-

ous and feed on old leaves of Passiora ambigua.

Sometimes large numbers of pupae can be found

on the tree trunks which hosted the passion

vine.

Laparus doris

Gallery forest near the research station. Gallery forests are often the only remains after clearance of surrounding forests for

agricultural development. They can provide important habitats for some species, including Calephelis and other Metalmarks,

and might also function as corridors to connect remaining rainforest fragments.

Juditha molpe larva feeding on a Passiora vine

The larva of the Molpe Metalmark (Juditha

molpe) is polyphagous and can feed on many dif-

ferent plant families. However, it has obligate as-

sociations with the ant species Dolichoderus bispi-

nosus, which appears to mediate the hostplant

choice of the ovipositing female buttery.

Riodinidae

Metalmarks

The butteries of the family Riodinidae are small to medium-sized. The group is characteristic

throughout the Neotropics for its great diversity of species. The females, like most insects, walk on

all six legs, however the males use only four. Their forelegs are too small to serve for locomotion,

but are probably connected with mating. Their wing patterns show extraordinary diversity among

species, most of which mimic members of other buttery or moth families. The metalmarks differ

also in behavior from other butteries. Most species perch on the underside of leaves and are there-

fore easily mistaken for moths. Their peak time of activity is often in the early morning or in the

late afternoon, whereas most other butteries are most active during midday. In certain genera the

larvae live symbiotically with ants (myrmecophily). The ants are attracted to and feed on a glandular

secretion of the larvae. In return, the ants care for the buttery larvae by defending them against

parasites. In some species, the caterpillars produce a vibration sound by rubbing their head against

a special organ. The sounds are not audible to humans but serve as important signals to the ants.

Eurybia lycisca (Blue-Winged Sheenmark)

Forewing length: 24 - 27.5 mm

The proboscis of Eurybia lycisca is one of the

longest among all butteries. It measures up to

45 mm and is almost twice as long as the body.

The upperside of the forewing is brown and has

two eyespots with iridescent centers; the upper-

side of the hindwing is remarkably metallic blue

across the entire distal area. The sexes appear

similar. However, the blue hues are less conspic-

uous in females. When the buttery settles on a

plant, it opens the wings, and the blue coloration

is noticeable from a distance at particular angles.

Eurybia lycisca is a medium-sized buttery. It ies

Eurybia lyciscaEurybia lycisca

fast and erratically in the shade of vegetation. In

the garden of the station, it is common and can

be found near its preferred nectar and larval host

plant, Calathea (Maranthaceae). Nectar feeding

predominantly occurs in the morning; in the af-

ternoon, female butteries can be observed to lay

eggs frequently on the inorescences of Calathea

plants. The caterpillars feed on the inorescence

and live in association with ants.

Euselasia regipennis (Purple-topped Sombermark)

Forewing length: 16 - 20 mm

This local Central American species, which is

found singly in rainforest habitats, resembles sa-

tyrines of and related to the genus Cissia (family

Nymphalidae). The sexes are dimorphic: In the

males, the upperside of the wings is black with a

shining deep purple iridescence. The females oc-

cur in two color morphs. One of them possesses

a white band on the upperside, as in Pareupty-

chia ocirrhoe. The other morph is brown, as Cis-

sia, Hermeuptychia and Magneuptychia. The early

stages of this buttery are still unknown

Euselasia regipennis

Ancyluris jurgensenii (Costa-spotted Beautymark)

Forewing length: 22 - 25 mm

Ancyluris jurgensenii is distributed throughout

Central America but is uncommon in Costa Rica.

Single specimens have also been observed along

forest tracts near the station. In the early morn-

ing, males perch on the underside of leaves, and

chase any approaching buttery. The females are

more active during midday. The wing patters of

Ancyluris jurgensenii

the sexes are dimorphic. The males have a red

band; the females have a white band and their

hindwings lack the beautiful blue iridescence

along the margins. The caterpillars are known to

feed on Hyeronima oblonga which belongs to the

family Euphorbiaceae.

Calephelis laverna (Small Scintillant)

Forewing length: 9 - 10 mm

Calephelis laverna is one of

many similar looking spe-

cies, but is distinguished by

the red-brown underside and

its small size. In Costa Rica,

it is only found in the South

of the Osa Peninsula but its

range extends to Brazil. The

butteries can be seen in

the gallery forests close to

the station. The life history

of this small but pretty spe-

cies in Costa Rica is still un-

known, but in Trinidad the

larvae feed on Asteraceae

(e.g. Chromolaena and Eupa-

torium).

Calephelis laverna

Charis anius (= auius) (Anius Metalmark)

Forewing length: 12 - 14 mm

The genus Charis com-

prises several similar look-

ing species which can be

abundant in open areas

along forest edges or along

streams where they y close

to the ground and are there-

fore easy to observe. Little is

known about their life his-

tories, but most species ap-

pear to be detrivorous, feed-

ing on dead leaves on the

ground. This is the case for

Charis anius, where egg-lay-

ing was observed on dead

leaves. The caterpillars are

extremely hirsute, which is

very unusual in butteries

and much more typical for Charis anius

Nymphidium ascolia (Creamy Metalmark)

Forewing length: 19 - 20 mm

Nymphidium ascolia is a medi-

um-sized buttery with white col-

oration of the wing and body. Both

wings have broad brown margin

with a blue zigzag band. On the

hindwing are orange markings.

The buttery visits owers, for ex-

ample Lantana, to ll up on nectar.

It is common at the biological sta-

tion, on the surrounding trails and

in forest gaps. Morning is the best

time to search for the species. Le-

gumes of the genus Inga are used

as larval food plants. The caterpil-

lars are attended by different ants

of the subfamilies Formicinae and

Myrmicinae. Nymphidium ascolia

Mesenopsis melanochlora (Orange-striped Metalmark)

Forewing length: 15 - 18 mm

Mesenopsis melanochlora is an

excellent mimic of day-ying

moths from different families, for

example, Josia ligata (Notodonti-

dae) and various Josiomorpha spe-

cies (Arctiidae). However, due to

its slightly club-tipped antennae it

can be differentiated from them. It

is encountered along forest tracts

in the Piedras Blancas National

Park, where the males perch sev-

eral meters above the ground on

the upperside of leaves during

the morning. The butteries visit

owers of various Asteraceae and

the females deposit their eggs sin-

gly in the leaf axils or in the bark

of the tree Miconia argentea (Melas-

tomataceae). The caterpillars usu-

ally feed on sapling plants where

they fold the edge of the leaf into

a tube. Mesenopsis melanochlora

Light gap in the Esquinas rainforest. Many rainforest butteries y in the canopy and are therefore difcult to observe. Natural

gaps provide the best opportunity to nd such species, including many Hairstreaks.

Lycaenidae

Blues

The Lycaenidae are predominantly small-sized butteries. In number of species, they are the sec-

ond largest buttery family with about 6,000 species worldwide. Three main groups are rec-

ognized: the blues, coppers and hairstreaks. Almost all species from Central and South America

are hairstreaks; many of them possess small tails on the hindwings. Most hairstreaks are depen-

dent on trees as larval food plants. The butteries spend most of the time in the tree canopy and

therefore are difcult to observe. Little is known about their biology. Like metalmarks, the cater-

pillars of many blues live in symbiotic relationship with ants (myrmecophily). The caterpillars offer

the ants a honeydew secretion. In return, the ants protect the caterpillars from parasites. Caterpil-

lars of some species are even transported by the ants into the ant nests, where they are cared for

and fed like ant brood. In a few species, the caterpillars become predators and attack ant larvae.

Arawacus lincoides

Forewing length: 15 - 18 mm

Arawacus lincoides is a species of

hairstreak with an unusual striped

pattern on its underside reminiscent

of small satyrines (e.g. Pareuptychia

ocirrhoe). It has often been confused

with its sibling Arawacus togarna,

which occurs on the Atlantic slope of

Costa Rica and further north to Mex-

ico, whereas A. lincoides is only found

on the Pacic slope of Costa Rica and

further south to Panamá and Colum-

bia (Robbins 2010). A. lincoides differs

from its sibling by genitalia charac-

ters and a white marginal spot at the

distal end of the hindwing underside

between its two tails (Robbins, pers.

comm.). The tails of hairstreaks are

thought to distract potential preda-

tors from essential body parts (false

head hypothesis). If a bird or lizard

bites off the tail, the buttery sur-

vives essentially unharmed. This

hairstreak can be seen quite often in

the garden of the station and along

waysides usually sitting on small bush-

es. The caterpillars feed on plants of the

family Solanaceae.

Arawacus lincoides

Arumecla galliena (Red-Based Groundstreak)

Forewing length: 12 - 15 mm

Arumecla galliena is found in Cen-

tral and northern South America. It is

one of the most common hairstreaks

in the primary and secondary rain-

forests of the Golfo Dulce region.

It is usually encountered in small

groups, ying around the treetops.

The butteries are most easily ob-

served in forest gaps around smaller

trees. Males are territorial and spend

their time chasing away other males.

Arumecla galliena belongs to a group

of hairstreaks which is detrivorous.

The females lay their eggs on dead

plant material and the caterpillars

then feed on leaf litter. Arumecla galliena

Hemiargus hanno (Hanno Blue)

Forewing length: 9 - 11 mm

Not all tropical butteries

are large and colorful. This tiny

blue is among the smallest but-

teries in Costa Rica. It is most

common on pastures with

woody legumes, such as plants

of the genera Cassia or Aca-

cia, which constitute the food

plants of the caterpillars. They

are quite polyphagous and also

feed on plants of the families

Mimosaceae and Oxalidaceae.

Like many other blues, the cat-

erpillars possess three types of

myrmecophilous organs (the

pore cupola, dorsal nectary and

tentacle organ) to entice ants,

which in turn guard the cater-

pillars. The buttery-ant asso-

ciation is known from Brazil,

where the ants were identied

as belonging to the formicine

genus Brachymyrmex.

Hemiargus hanno

Strymon megarus (= Tmolus basilides)

(Scrub-Hairstreak, Pineapple Fruit Borer)

Spanish name: Taladro de la Piña

Forewing length: 11 - 14.5 mm

This hairstreak is rare in its

natural habitat, but can become

a serious pest in pineapple

plantations. The females lay

their eggs on pineapple ow-

ers and the larvae bore into

the fruits. In its natural habi-

tat, various other bromeliads

and heliconias are used as food

plants. In the garden of the

station, females have been ob-

served laying eggs on the ow-

ers of Heliconia.

Strymon megarus

Secondary forest near the research station. The larvae of many Skippers feed on grasses which grow better in lighter places than

in the deep shade of primary rainforests. Therefore secondary forests can be a suitable habitat for many species of this family.

However, only a small fraction of rainforest species can survive in those degraded forests.

Hesperiidae

Skippers

Skippers are characterized by a large head, a compact body, and relatively small wings compared

to their body, which helps them to attain their quick and darting ight pattern. The short anten-

nae are club-tipped, but unlike other butteries the clubs are hooked backwards like a crochet

hook. The family is probably the sister group to the remaining ‘true’ butteries, but their phylo-

genetic position is still debated. Many species have a resting position which is very unusual for

butteries: only the hindwings are spread out at, whereas the forewings are angled upward,

producing a ‘v‘ shape. Other skippers (e.g. from the genus Urbanus) usually fold their wings over

the body giving it a triangular shape. These butteries are frequent ower visitors. They pos-

sess remarkably long mouthparts in comparison to their rather small body so that they are able

to extract nectar even from long tubed owers like Stachytarpheta. Many skippers are also fond of

dung or carrion, or can be observed taking up minerals at mud puddles. Unfortunately, this spe-

cies-rich family is the least well known among all buttery families and there is no identication

guide available for species of Costa Rica. The many species are usually dull brown or gray and

very similar to each other. Genitalia examination is often necessary for reliable identication.

Skipper in the typical resting position Skipper feeding on owers of Stachytarpheta

Urbanus teleus (Teleus Longtail)

Urbanus teleus is widespread throughout

Central and South America. The butteries are

regularly seen in the garden of the station. The

brown and medium-sized buttery is distin-

guished by the elongate hindwings which are

reminiscent of the tails in swallowtail butteries.

The females lay their eggs on the food plant, i.e.

different kinds of grasses. The young caterpillars

construct shelters of folded leaf blades.

Urbanus teleus

Pyrrhopyge thericles (= pseudophidias) (Thericles Firetip)

Unlike most other groups of skippers, the ge-

nus Pyrrhopyge contains many beautiful species

with an entirely black color or often with red or

white wing patterns. One of them, Pyrrhopyge

thericles, is distributed throughout tropical South

America. Some authors, however, consider the

Central American populations to represent a

different species (P. pseudophidias). P. thericles

has been observed ying in gallery forests along

the river near the station. Nothing appears to be

known about its life history.

Pyrrhopyge thericles

Appendix

Preliminary checklist of the Butteries of the Golfo Dulce

Region (Papilionoidea & Hesperioidea)

The table includes our own data (La Gamba) as well as published data from Corcova-

do National Park (Corcovado). The checklist from DeVries (1978) excludes six species

which appear to represent misidentications because they are not listed in DeVries (1987).

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995)

or Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Hesperiidae:

Achlyodes mithridates (Fabricius, 1793) •

Antigonus nearchus (LatreiLLe, 1817) •

Autochton bipunctatus (GmeLin, 1790) •

Autochton neis (Geyer, 1832) • •

Celaenorrhinus monartus (PLötz, 1884) • •

Celaenorrhinus stallingsi Freeman, 1946 •

Cogia calchas (HerricH-scHäFFer, 1869) •

Helias cama eVans, 1953 •

Heliopetes arsalte (Linnaeus, 1758) • •

Hylephila phyleus (Drury, 1773) •

Mnasilus allubita (butLer, 1877) • •

Molo mango (Guenée, 1865) •

Nisoniades rubescens (möscHLer, 1877) •

Nyctelius nyctelius (LatreiLLe, 1824) •

Ouleus fridericus (Geyer, 1832) Ouleus panna • •

Panoquina ocola (eDwarDs, 1863) • •

Pompeius pompeius (LatreiLLe, 1824) • • •

Pyrgus oileus (Linnaeus, 1767) •

Pyrgus orcus (stoLL, 1780) •

Pyrrhopyge thericles (mabiLLe, 1891) Pyrrhopyge pseudophidias • •

Pythonides jovianus (stoLL, 1782) •

Remella vopiscus (HerricH-scHäFFer, 1869) •

Timochares trifasciata (Hewitson, 1868) •

Urbanus dorantes (stoLL, 1790) •

Urbanus procne (PLötz, 1880) •

Urbanus proteus (Linnaeus, 1758) •

Urbanus simplicius (stoLL, 1790) •

Urbanus teleus (Hübner, 1821) •

Xenophanes tryxus (stoLL, 1780) • •

Lycaenidae:

Arawacus lincoides (Draudt, 1917) Arawacus togarna • • • •

Arumecla galliena (Hewitson, 1877) Thecla galliena • • •

Brangas caranus (stoLL, 1780) • •

Calycopis atnius (HerricH-scHäFFer, 1853) Thecla atrius • •

Calycopis demonassa (Hewitson, 1868) Thecla demonassa • •

Calycopis isobeon (butLer & Druce, 1872) Thecla beon • • •

Calycopis pisis (GoDman & saLVin, 1887) Thecla pisis • • •

Calycopis trebula (Hewitson, 1868) Thecla trebula • • •

Cupido comyntas (GoDart, 1824) Everes comyntas • • •

Eumaeus godartii (boisDuVaL, 1870) • •

Hemiargus hanno (stoLL, 1790) Hemiargus ceraunus • • •

Iaspis talayra (Hewitson, 1868) Thecla talayra • •

Ocaria thales (Fabricius, 1793) Thecla thales • •

Panthiades bathildis (FeLDer & FeLDer, 1865) •

Panthiades phaleros (Linnaeus, 1767) Cydno phaleros •

Pseudolycaena damo (Druce, 1875) •

Pseudolycaena marsyas (Linnaeus, 1758) • •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995)

or Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Hesperiidae:

Achlyodes mithridates (Fabricius, 1793) •

Antigonus nearchus (LatreiLLe, 1817) •

Autochton bipunctatus (GmeLin, 1790) •

Autochton neis (Geyer, 1832) • •

Celaenorrhinus monartus (PLötz, 1884) • •

Celaenorrhinus stallingsi Freeman, 1946 •

Cogia calchas (HerricH-scHäFFer, 1869) •

Helias cama eVans, 1953 •

Heliopetes arsalte (Linnaeus, 1758) • •

Hylephila phyleus (Drury, 1773) •

Mnasilus allubita (butLer, 1877) • •

Molo mango (Guenée, 1865) •

Nisoniades rubescens (möscHLer, 1877) •

Nyctelius nyctelius (LatreiLLe, 1824) •

Ouleus fridericus (Geyer, 1832) Ouleus panna • •

Panoquina ocola (eDwarDs, 1863) • •

Pompeius pompeius (LatreiLLe, 1824) • • •

Pyrgus oileus (Linnaeus, 1767) •

Pyrgus orcus (stoLL, 1780) •

Pyrrhopyge thericles (mabiLLe, 1891) Pyrrhopyge pseudophidias • •

Pythonides jovianus (stoLL, 1782) •

Remella vopiscus (HerricH-scHäFFer, 1869) •

Timochares trifasciata (Hewitson, 1868) •

Urbanus dorantes (stoLL, 1790) •

Urbanus procne (PLötz, 1880) •

Urbanus proteus (Linnaeus, 1758) •

Urbanus simplicius (stoLL, 1790) •

Urbanus teleus (Hübner, 1821) •

Xenophanes tryxus (stoLL, 1780) • •

Lycaenidae:

Arawacus lincoides (Draudt, 1917) Arawacus togarna • • • •

Arumecla galliena (Hewitson, 1877) Thecla galliena • • •

Brangas caranus (stoLL, 1780) • •

Calycopis atnius (HerricH-scHäFFer, 1853) Thecla atrius • •

Calycopis demonassa (Hewitson, 1868) Thecla demonassa • •

Calycopis isobeon (butLer & Druce, 1872) Thecla beon • • •

Calycopis pisis (GoDman & saLVin, 1887) Thecla pisis • • •

Calycopis trebula (Hewitson, 1868) Thecla trebula • • •

Cupido comyntas (GoDart, 1824) Everes comyntas • • •

Eumaeus godartii (boisDuVaL, 1870) • •

Hemiargus hanno (stoLL, 1790) Hemiargus ceraunus • • •

Iaspis talayra (Hewitson, 1868) Thecla talayra • •

Ocaria thales (Fabricius, 1793) Thecla thales • •

Panthiades bathildis (FeLDer & FeLDer, 1865) •

Panthiades phaleros (Linnaeus, 1767) Cydno phaleros •

Pseudolycaena damo (Druce, 1875) •

Pseudolycaena marsyas (Linnaeus, 1758) • •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Siderus leucophaeus (Hübner, 1813) Thecla leucophaeus • •

Strymon megarus (GoDart, 1824) • • •

Theorema eumenia Hewitson, 1865 • •

Theritas hemon (cramer, 1775) • •

Theritas lisus (stoLL, 1790) Thecla hisbon • •

Theritas mavors Hübner, 1818 •

Ziegleria syllis (GoDman & saLVin, 1887) Strymon syllis • •

Nymphalidae:

Actinote lapitha (stauDinGer, 1885) •

Adelpha basiloides (bates, 1865) •

Adelpha boeotia (FeLDer & FeLDer, 1867) • • •

Adelpha cocala (cramer, 1779) • • •

Adelpha cytherea (Linnaeus, 1758) • • • • • •

Adelpha heraclea (FeLDer & FeLDer, 1867) • •

Adelpha iphiclus (Linnaeus, 1758) • •

Adelpha justina (FeLDer & FeLDer, 1861) • •

Adelpha salmoneus (butLer, 1866) • •

Adelpha serpa (boisDuVaL, 1836) •

Aeria eurimedia (cramer, 1777) • • • •

Agraulis vanillae (Linnaeus, 1758) • • •

Anartia fatima (Fabricius, 1793) • • • • • •

Anartia jatrophae (Linnaeus, 1763) • • • • •

Anthanassa frisia (Poey, 1832) Anthanassa tulcis • • • • •

Antirrhea philoctetes (Linnaeus, 1758) Antirrhea tomasia • •

Archaeoprepona demophon (Linnaeus, 1758) • • •

Archaeoprepona demophoon (Hübner, 1814) •

Caligo atreus (KoLLar, 1850) • • •

Caligo eurilochus (cramer, 1775) • • •

Caligo telamonius (FeLDer & FeLDer, 1862) Caligo memnon • •

Callicore lyca (DoubLeDay, 1847) • •

Callicore texa (Hewitson, 1855) •

Callicore tolima (Hewitson, 1852) Callicore pacica •

Callithomia hezia (Hewitson, 1854) • • •

Castilia eranites (Hewitson, 1857) • • •

Catoblepia orgetorix (Hewitson, 1870) • •

Catonephele numilia (cramer, 1775) • • •

Catonephele nyctimus (westwooD, 1850) Catonephele mexicana • • •

Ceratinia tutia (Hewitson, 1852) • • •

Caerois gerdrudtus (Fabricius, 1793) • •

Chloreuptychia arnaca (Fabricius, 1776) Chloreuptychia arnaea • • • •

Chlosyne hippodrome (Geyer, 1837) •

Chlosyne janais (Drury, 1782) •

Chlosyne lacinia (Geyer, 1837) • •

Chlosyne theona (ménétriés, 1855) Thessalia ezra • • • • •

Cissia confusa (stauDinGer, 1887) • • •

Cissia pompilia (FeLDer & FeLDer, 1867) Cissia usitata • •

Cissia pseudoconfusa sinGer, DeVries & eHrLicH, 1983 •

Cithaerias pireta (stoLL, 1780) Cithaerias menander • • •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Siderus leucophaeus (Hübner, 1813) Thecla leucophaeus • •

Strymon megarus (GoDart, 1824) • • •

Theorema eumenia Hewitson, 1865 • •

Theritas hemon (cramer, 1775) • •

Theritas lisus (stoLL, 1790) Thecla hisbon • •

Theritas mavors Hübner, 1818 •

Ziegleria syllis (GoDman & saLVin, 1887) Strymon syllis • •

Nymphalidae:

Actinote lapitha (stauDinGer, 1885) •

Adelpha basiloides (bates, 1865) •

Adelpha boeotia (FeLDer & FeLDer, 1867) • • •

Adelpha cocala (cramer, 1779) • • •

Adelpha cytherea (Linnaeus, 1758) • • • • • •

Adelpha heraclea (FeLDer & FeLDer, 1867) • •

Adelpha iphiclus (Linnaeus, 1758) • •

Adelpha justina (FeLDer & FeLDer, 1861) • •

Adelpha salmoneus (butLer, 1866) • •

Adelpha serpa (boisDuVaL, 1836) •

Aeria eurimedia (cramer, 1777) • • • •

Agraulis vanillae (Linnaeus, 1758) • • •

Anartia fatima (Fabricius, 1793) • • • • • •

Anartia jatrophae (Linnaeus, 1763) • • • • •

Anthanassa frisia (Poey, 1832) Anthanassa tulcis • • • • •

Antirrhea philoctetes (Linnaeus, 1758) Antirrhea tomasia • •

Archaeoprepona demophon (Linnaeus, 1758) • • •

Archaeoprepona demophoon (Hübner, 1814) •

Caligo atreus (KoLLar, 1850) • • •

Caligo eurilochus (cramer, 1775) • • •

Caligo telamonius (FeLDer & FeLDer, 1862) Caligo memnon • •

Callicore lyca (DoubLeDay, 1847) • •

Callicore texa (Hewitson, 1855) •

Callicore tolima (Hewitson, 1852) Callicore pacica •

Callithomia hezia (Hewitson, 1854) • • •

Castilia eranites (Hewitson, 1857) • • •

Catoblepia orgetorix (Hewitson, 1870) • •

Catonephele numilia (cramer, 1775) • • •

Catonephele nyctimus (westwooD, 1850) Catonephele mexicana • • •

Ceratinia tutia (Hewitson, 1852) • • •

Caerois gerdrudtus (Fabricius, 1793) • •

Chloreuptychia arnaca (Fabricius, 1776) Chloreuptychia arnaea • • • •

Chlosyne hippodrome (Geyer, 1837) •

Chlosyne janais (Drury, 1782) •

Chlosyne lacinia (Geyer, 1837) • •

Chlosyne theona (ménétriés, 1855) Thessalia ezra • • • • •

Cissia confusa (stauDinGer, 1887) • • •

Cissia pompilia (FeLDer & FeLDer, 1867) Cissia usitata • •

Cissia pseudoconfusa sinGer, DeVries & eHrLicH, 1983 •

Cithaerias pireta (stoLL, 1780) Cithaerias menander • • •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Colobura annulata wiLLmott, constantino & HaLL, 2001 previously confused with Colobura dirce • • •

Colobura dirce (Linnaeus, 1758) • • •

Consul fabius (cramer, 1776) • • • •

Danaus gilippus (cramer, 1775) • • • •

Danaus plexippus (Linnaeus, 1758) • • •

Diaethria astala (Guérin-méneViLLe, 1844) •

Diaethria clymena (cramer, 1775) Diaethria marchalli •

Dione juno (cramer, 1779) • • • •

Dione moneta (Hübner, 1825) • •

Dircenna dero (Hübner, 1823) •

Dryadula phaetusa (Linnaeus, 1758) • • • •

Dryas iulia (Fabricius, 1775) • • • • • •

Dynamine agacles (DaLman, 1823) • • •

Dynamine tithia (Hübner, 1823) Dynamine salpensa • •

Eresia clio (Linnaeus, 1758) Eresia clara • •

Eresia eunice (Hübner, 1807) Eresia mechanitis • • •

Eresia ithomioides Hewitson, 1864 Eresia eutropia & Eresia melaina • • •

Eryphanis automedon (cramer, 1775) Eryphanis polyxena •

Eueides aliphera (GoDart, 1819) • •

Eueides isabella (stoLL, 1781) • •

Eueides lineata saLVin & GoDman, 1868 • •

Eueides lybia (Fabricius, 1775) • • • •

Eunica alpais (GoDart, 1824) Eunica excelsa • • • •

Eunica chlororhoa saLVin, 1869 Eunica mira • •

Eunica sydonia (GoDart, 1824) Eunica caresa • •

Eunica volumna (GoDart, 1824) Eunica venusia •

Euptoieta hegesia (Cramer, 1779) • • • •

Euptychia insolata butLer & Druce, 1872 • • • •

Euptychia jesia butLer, 1869 • • • •

Euptychia westwoodi butLer, 1867 Euptychia mollis • • •

Fountainea eurypyle (FeLDer & FeLDer, 1862) Memphis eurypyle • •

Godyris zavaleta (Hewitson, 1855) Godyris zygia •

Hamadryas amphinome (Linnaeus, 1767) •

Hamadryas arinome (Lucas, 1853) • •

Hamadryas feronia (Linnaeus, 1758) • • •

Hamadryas guatemalena (bates, 1864) • • •

Heliconius charithonia (Linnaeus, 1767) Heliconius charitonius • • •

Heliconius cydno (DoubLeDay, 1847) Heliconius pachinus • • • • •

Heliconius erato (Linnaeus, 1758) • •

Heliconius hecale (Fabricius, 1776) • • • • •

Heliconius hecalesia (Hewitson, 1854) • •

Heliconius ismenius LatreiLLe, 1817 • • • •

Heliconius melpomene (Linnaeus, 1758) • • • • •

Heliconius sapho (Drury, 1782) Heliconius hewitsoni • • • • •

Heliconius sara (Fabricius, 1793) • • • • • •

Hermeuptychia hermes (Fabricius, 1775) Cissia hermes • • • • •

Historis acheronta (Fabricius, 1775) •

Historis odius (Fabricius, 1775) • • •

Hypoleria lavinia (Hewitson, 1855) Hypoleria cassotis • •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Colobura annulata wiLLmott, constantino & HaLL, 2001 previously confused with Colobura dirce • • •

Colobura dirce (Linnaeus, 1758) • • •

Consul fabius (cramer, 1776) • • • •

Danaus gilippus (cramer, 1775) • • • •

Danaus plexippus (Linnaeus, 1758) • • •

Diaethria astala (Guérin-méneViLLe, 1844) •

Diaethria clymena (cramer, 1775) Diaethria marchalli •

Dione juno (cramer, 1779) • • • •

Dione moneta (Hübner, 1825) • •

Dircenna dero (Hübner, 1823) •

Dryadula phaetusa (Linnaeus, 1758) • • • •

Dryas iulia (Fabricius, 1775) • • • • • •

Dynamine agacles (DaLman, 1823) • • •

Dynamine tithia (Hübner, 1823) Dynamine salpensa • •

Eresia clio (Linnaeus, 1758) Eresia clara • •

Eresia eunice (Hübner, 1807) Eresia mechanitis • • •

Eresia ithomioides Hewitson, 1864 Eresia eutropia & Eresia melaina • • •

Eryphanis automedon (cramer, 1775) Eryphanis polyxena •

Eueides aliphera (GoDart, 1819) • •

Eueides isabella (stoLL, 1781) • •

Eueides lineata saLVin & GoDman, 1868 • •

Eueides lybia (Fabricius, 1775) • • • •

Eunica alpais (GoDart, 1824) Eunica excelsa • • • •

Eunica chlororhoa saLVin, 1869 Eunica mira • •

Eunica sydonia (GoDart, 1824) Eunica caresa • •

Eunica volumna (GoDart, 1824) Eunica venusia •

Euptoieta hegesia (Cramer, 1779) • • • •

Euptychia insolata butLer & Druce, 1872 • • • •

Euptychia jesia butLer, 1869 • • • •

Euptychia westwoodi butLer, 1867 Euptychia mollis • • •

Fountainea eurypyle (FeLDer & FeLDer, 1862) Memphis eurypyle • •

Godyris zavaleta (Hewitson, 1855) Godyris zygia •

Hamadryas amphinome (Linnaeus, 1767) •

Hamadryas arinome (Lucas, 1853) • •

Hamadryas feronia (Linnaeus, 1758) • • •

Hamadryas guatemalena (bates, 1864) • • •

Heliconius charithonia (Linnaeus, 1767) Heliconius charitonius • • •

Heliconius cydno (DoubLeDay, 1847) Heliconius pachinus • • • • •

Heliconius erato (Linnaeus, 1758) • •

Heliconius hecale (Fabricius, 1776) • • • • •

Heliconius hecalesia (Hewitson, 1854) • •

Heliconius ismenius LatreiLLe, 1817 • • • •

Heliconius melpomene (Linnaeus, 1758) • • • • •

Heliconius sapho (Drury, 1782) Heliconius hewitsoni • • • • •

Heliconius sara (Fabricius, 1793) • • • • • •

Hermeuptychia hermes (Fabricius, 1775) Cissia hermes • • • • •

Historis acheronta (Fabricius, 1775) •

Historis odius (Fabricius, 1775) • • •

Hypoleria lavinia (Hewitson, 1855) Hypoleria cassotis • •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Hyposcada virginiana (Hewitson, 1855) • • • •

Hypothyris euclea (GoDart, 1819) • •

Ithomia celemia Hewitson, 1854 • •

Ithomia patilla Hewitson, 1852 • • •

Janatella leucodesma (FeLDer & FeLDer, 1861) •

Junonia evarete (cramer, 1779) • • •

Laparus doris (Linnaeus, 1771) Heliconius doris • • •

Lycorea halia (Hübner, 1816) Lycorea cleobaea • • •

Magneuptychia gomezi (sinGer, DeVries & eHrLicH, 1983) Cissia gomezi • • •

Magneuptychia libye (Linnaeus, 1767) Cissia libye • • • • •

Marpesia berania (Hewitson, 1852) • • •

Marpesia chiron (Fabricius, 1775) • • •

Marpesia furcula (Fabricius, 1793) Marpesia iole • • •

Marpesia merops (Doyère, 1840) • • • • •

Marpesia petreus (cramer, 1776) • • •

Mechanitis lysimnia (Fabricius, 1793) • • • •

Mechanitis polymnia (Linnaeus, 1758) • • • • •

Megeuptychia antonoe (cramer, 1775) • • •

Melinaea lilis (DoubLeDay, 1847) Melinaea scylax • • • •

Memphis forreri (GoDman & saLVin, 1884) •

Memphis glauce (FeLDer & FeLDer, 1862) Memphis centralis • •

Memphis oenomais (boisDuVaL, 1870) • • •

Memphis proserpina (saLVin, 1869) • •

Memphis xenocles (westwooD, 1850) • •

Morpho cypris westwooD, 1851 •

Morpho helenor (cramer, 1776) Morpho peleides • • •

Morpho menelaus (Linnaeus, 1758) Morpho amathonte • • •

Morpho theseus DeyroLLe, 1860 •

Nessaea aglaura (DoubLeDay, 1848) • • •

Nica avilla (GoDart, 1824) • •

Oleria paula (weymer, 1883) • •

Oleria rubescens (butLer & Druce, 1872) •

Opsiphanes cassina FeLDer & FeLDer, 1862 • •

Opsiphanes invirae (Hübner, 1808) • •

Opsiphanes tamarindi FeLDer & FeLDer, 1861 • • •

Pareuptychia metaleuca (boisDuVaL, 1870) • • •

Pareuptychia ocirrhoe (Fabricius, 1776) Cissia hesione • • • •

Perophthalma lasus westwooD, 1851 • •

Philaethria dido (Linnaeus, 1763) • • • •

Pierella helvina (Hewitson, 1859) Pierella helvetia • • •

Pierella luna (Fabricius, 1793) • • • •

Posttaygetis penelea (cramer, 1777) Taygetis penelea • •

Prepona laertes (Hübner, 1811) Prepona omphale • •

Pteronymia alcmena (GoDman & saLVin, 1877) Eunica alcmena •

Pteronymia aletta (Hewitson, 1855) Pteronymia agalla •

Pyrrhogyra crameri auriViLLius, 1882 Pyrrogyra crameri • • •

Pyrrhogyra otolais bates, 1864 •

Siproeta stelenes (Linnaeus, 1758) • • • • •

Taygetis laches Fabricius, 1793 Taygetis andromeda •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Hyposcada virginiana (Hewitson, 1855) • • • •

Hypothyris euclea (GoDart, 1819) • •

Ithomia celemia Hewitson, 1854 • •

Ithomia patilla Hewitson, 1852 • • •

Janatella leucodesma (FeLDer & FeLDer, 1861) •

Junonia evarete (cramer, 1779) • • •

Laparus doris (Linnaeus, 1771) Heliconius doris • • •

Lycorea halia (Hübner, 1816) Lycorea cleobaea • • •

Magneuptychia gomezi (sinGer, DeVries & eHrLicH, 1983) Cissia gomezi • • •

Magneuptychia libye (Linnaeus, 1767) Cissia libye • • • • •

Marpesia berania (Hewitson, 1852) • • •

Marpesia chiron (Fabricius, 1775) • • •

Marpesia furcula (Fabricius, 1793) Marpesia iole • • •

Marpesia merops (Doyère, 1840) • • • • •

Marpesia petreus (cramer, 1776) • • •

Mechanitis lysimnia (Fabricius, 1793) • • • •

Mechanitis polymnia (Linnaeus, 1758) • • • • •

Megeuptychia antonoe (cramer, 1775) • • •

Melinaea lilis (DoubLeDay, 1847) Melinaea scylax • • • •

Memphis forreri (GoDman & saLVin, 1884) •

Memphis glauce (FeLDer & FeLDer, 1862) Memphis centralis • •

Memphis oenomais (boisDuVaL, 1870) • • •

Memphis proserpina (saLVin, 1869) • •

Memphis xenocles (westwooD, 1850) • •

Morpho cypris westwooD, 1851 •

Morpho helenor (cramer, 1776) Morpho peleides • • •

Morpho menelaus (Linnaeus, 1758) Morpho amathonte • • •

Morpho theseus DeyroLLe, 1860 •

Nessaea aglaura (DoubLeDay, 1848) • • •

Nica avilla (GoDart, 1824) • •

Oleria paula (weymer, 1883) • •

Oleria rubescens (butLer & Druce, 1872) •

Opsiphanes cassina FeLDer & FeLDer, 1862 • •

Opsiphanes invirae (Hübner, 1808) • •

Opsiphanes tamarindi FeLDer & FeLDer, 1861 • • •

Pareuptychia metaleuca (boisDuVaL, 1870) • • •

Pareuptychia ocirrhoe (Fabricius, 1776) Cissia hesione • • • •

Perophthalma lasus westwooD, 1851 • •

Philaethria dido (Linnaeus, 1763) • • • •

Pierella helvina (Hewitson, 1859) Pierella helvetia • • •

Pierella luna (Fabricius, 1793) • • • •

Posttaygetis penelea (cramer, 1777) Taygetis penelea • •

Prepona laertes (Hübner, 1811) Prepona omphale • •

Pteronymia alcmena (GoDman & saLVin, 1877) Eunica alcmena •

Pteronymia aletta (Hewitson, 1855) Pteronymia agalla •

Pyrrhogyra crameri auriViLLius, 1882 Pyrrogyra crameri • • •

Pyrrhogyra otolais bates, 1864 •

Siproeta stelenes (Linnaeus, 1758) • • • • •

Taygetis laches Fabricius, 1793 Taygetis andromeda •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Taygetis virgilia (cramer, 1776) • •

Tegosa anieta (Hewitson, 1864) • •

Temenis laothoe (cramer, 1777) •

Temenis pulchra (Hewitson, 1861) • •

Thyridia psidii (Linnaeus, 1758) • • •

Tigridia acesta (Linnaeus, 1758) • •

Tithorea tarricina Hewitson, 1858 • • • •

Yphthimoides renata (stoLL, 1780) Cissia renata •

Zaretis ellops (ménétries, 1855) • •

Papilionidae:

Battus lycidas (cramer, 1777) • •

Battus polydamas (Linnaeus, 1758) •

Eurytides orabilis (butLer, 1872) •

Heraclides androgeus (cramer, 1775) Papilio androgeus •

Heraclides cresphontes (cramer, 1777) Papilio cresphontes •

Heraclides thoas (Linnaeus, 1771) Papilio thoas • • •

Mimoides ilus (Fabricius, 1793) Eurytides ilus • •

Parides childrenae (Gray, 1832) • • • •

Parides erithalion (boisDuVaL, 1836) • •

Parides eurimedes (stoLL, 1782) Parides arcas •

Parides iphidamas (Fabricius, 1793) • • •

Parides panares (Gray, 1853) Parides lycimenes • •

Protesilaus protesilaus (Linnaeus, 1758) Eurytides protesilaus •

Protographium calliste (bates, 1864) Eurytides calliste •

Protographium thyastes (Drury, 1782) Eurytides marchandi •

Pterourus menatius (Hübner, 1819) Papilio cleotas & Papilio victorinus •

Pieridae:

Anteos clorinde (GoDart, 1824) •

Aphrissa boisduvalii (FeLDer & FeLDer, 1861) •

Aphrissa statira (cramer, 1777) • • • • •

Archonias brassolis (Fabricius, 1776) Archonias tereas • • •

Ascia monuste (Linnaeus, 1764) • • • •

Dismorphia theucharila (DoubLeDay, 1848) • • •

Enantia melite (Linnaeus, 1763) Enantia licinia • •

Eurema albula (cramer, 1775) • • • •

Eurema daira (GoDart, 1819) • • • •

Ganyra phaloe (GoDart, 1819) Ascia limona • •

Glutophrissa drusilla (cramer, 1777) Appias drusilla • • •

Perrhybris pamela (stoLL, 1780) Perrhybris pyrrha •

Phoebis argante (Fabricius, 1775) • • • •

Phoebis neocypris (Hübner, 1823) Phoebis rurina •

Phoebis philea (Linnaeus, 1763) • • •

Phoebis sennae (Linnaeus, 1758) • • • •

Pyrisitia lisa (boisDuVaL & Le conte, 1830) Eurema lisa •

Pyrisitia nise (cramer, 1775) Eurema nise • • •

Pyrisitia proterpia (Fabricius, 1775) Eurema proterpia •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Taygetis virgilia (cramer, 1776) • •

Tegosa anieta (Hewitson, 1864) • •

Temenis laothoe (cramer, 1777) •

Temenis pulchra (Hewitson, 1861) • •

Thyridia psidii (Linnaeus, 1758) • • •

Tigridia acesta (Linnaeus, 1758) • •

Tithorea tarricina Hewitson, 1858 • • • •

Yphthimoides renata (stoLL, 1780) Cissia renata •

Zaretis ellops (ménétries, 1855) • •

Papilionidae:

Battus lycidas (cramer, 1777) • •

Battus polydamas (Linnaeus, 1758) •

Eurytides orabilis (butLer, 1872) •

Heraclides androgeus (cramer, 1775) Papilio androgeus •

Heraclides cresphontes (cramer, 1777) Papilio cresphontes •

Heraclides thoas (Linnaeus, 1771) Papilio thoas • • •

Mimoides ilus (Fabricius, 1793) Eurytides ilus • •

Parides childrenae (Gray, 1832) • • • •

Parides erithalion (boisDuVaL, 1836) • •

Parides eurimedes (stoLL, 1782) Parides arcas •

Parides iphidamas (Fabricius, 1793) • • •

Parides panares (Gray, 1853) Parides lycimenes • •

Protesilaus protesilaus (Linnaeus, 1758) Eurytides protesilaus •

Protographium calliste (bates, 1864) Eurytides calliste •

Protographium thyastes (Drury, 1782) Eurytides marchandi •

Pterourus menatius (Hübner, 1819) Papilio cleotas & Papilio victorinus •

Pieridae:

Anteos clorinde (GoDart, 1824) •

Aphrissa boisduvalii (FeLDer & FeLDer, 1861) •

Aphrissa statira (cramer, 1777) • • • • •

Archonias brassolis (Fabricius, 1776) Archonias tereas • • •

Ascia monuste (Linnaeus, 1764) • • • •

Dismorphia theucharila (DoubLeDay, 1848) • • •

Enantia melite (Linnaeus, 1763) Enantia licinia • •

Eurema albula (cramer, 1775) • • • •

Eurema daira (GoDart, 1819) • • • •

Ganyra phaloe (GoDart, 1819) Ascia limona • •

Glutophrissa drusilla (cramer, 1777) Appias drusilla • • •

Perrhybris pamela (stoLL, 1780) Perrhybris pyrrha •

Phoebis argante (Fabricius, 1775) • • • •

Phoebis neocypris (Hübner, 1823) Phoebis rurina •

Phoebis philea (Linnaeus, 1763) • • •

Phoebis sennae (Linnaeus, 1758) • • • •

Pyrisitia lisa (boisDuVaL & Le conte, 1830) Eurema lisa •

Pyrisitia nise (cramer, 1775) Eurema nise • • •

Pyrisitia proterpia (Fabricius, 1775) Eurema proterpia •

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Riodinidae:

Ancyluris jurgensenii (saunDers, 1850) • •

Anteros allectus westwooD, 1851 • •

Anteros chrysoprasta Hewitson, 1867 Anteros chrysoprastus • •

Anteros formosus (cramer, 1777) • •

Calephelis browni McAlpine, 1971 • •

Calephelis iris (stauDinGer, 1876) Charis iris • •

Calephelis laverna (GoDman & saLVin, 1886) • • •

Calospila cilissa (Hewitson, 1863) • •

Charis anius (cramer, 1776) Charis auius • • •

Chimastrum argentea (bates, 1866) Chimastrum argenteum •

Comphotis ignicauda (GoDman & saLVin, 1878) Phaenochitonia ignicauda • •

Detritivora gynaea (GoDart, 1824) Charis gynaea • •

Esthemopsis clonia FeLDer & FeLDer 1865 • • •

Eurybia elvina sticHeL, 1910 • • •

Eurybia lycisca westwooD, 1851 • • • •

Eurybia patrona weymer, 1875 • •

Eurybia unxia GoDman & saLVin, 1885 • •

Euselasia aurantia (butLer & Druce, 1872) • •

Euselasia aurantiaca (saLVin & GoDman, 1868) • •

Euselasia procula (saLVin & GoDman, 1885) • •

Euselasia regipennis (butLer & Druce, 1872) • •

Juditha molpe (Hübner, 1808) • • •

Leucochimona lepida (GoDman & saLVin, 1885) • •

Menander pretus (cramer, 1777) • •

Mesene phareus (cramer, 1777) • • •

Mesene viz.phareus (cramer, 1777) • •

Mesenopsis melanochlora (GoDman & saLVin, 1878) • •

Mesosemia asa Hewitson, 1869 • • •

Mesosemia hesperina butLer, 1874 • •

Mesosemia telegone (boisDuVaL, 1836) • •

Mesosemia zonalis GoDman & saLVin, 1885 • • •

Napaea eucharila (bates, 1867) • •

Nymphidium ascolia Hewitson, 1853 • • •

Perophthalma lasus westwooD, 1851 • •

Perophthalma tullius (Fabricius, 1787) • •

Pheles strigosa (stauDinGer, 1876) Lepricornis strigosa • •

Pirascca arbuscula (möscHLer, 1883) Stichelia arbuscula • •

Pseudonymphidia clearista (butLer, 1871) • •

Sarota chrysus (stoLL, 1781) Sarota dematira • •

Sarota gyas (cramer, 1775) • •

Setabis cleomedes (Hewitson, 1870) • •

Theope acosma sticHeL, 1910 • •

Theope thestias Hewitson, 1860 • •

Theope virgilius (Fabricius, 1793) • •

Thisbe lycorias (Hewitson, 1853) • •

•

Total number of species 73 84 128 216 115 51

Species (Lamas 2004) Differing taxon names according to

DeVries (1987 & 1997) D’abrera (1995) or

Warren (in litt.)

La Gamba Corcovado

Intensive

land use

Secondary

forest

Natural

forest

ToTaL DeVries (1978) KunTe (2008)

Riodinidae:

Ancyluris jurgensenii (saunDers, 1850) • •

Anteros allectus westwooD, 1851 • •

Anteros chrysoprasta Hewitson, 1867 Anteros chrysoprastus • •

Anteros formosus (cramer, 1777) • •

Calephelis browni McAlpine, 1971 • •

Calephelis iris (stauDinGer, 1876) Charis iris • •

Calephelis laverna (GoDman & saLVin, 1886) • • •

Calospila cilissa (Hewitson, 1863) • •

Charis anius (cramer, 1776) Charis auius • • •

Chimastrum argentea (bates, 1866) Chimastrum argenteum •

Comphotis ignicauda (GoDman & saLVin, 1878) Phaenochitonia ignicauda • •

Detritivora gynaea (GoDart, 1824) Charis gynaea • •

Esthemopsis clonia FeLDer & FeLDer 1865 • • •

Eurybia elvina sticHeL, 1910 • • •

Eurybia lycisca westwooD, 1851 • • • •

Eurybia patrona weymer, 1875 • •

Eurybia unxia GoDman & saLVin, 1885 • •

Euselasia aurantia (butLer & Druce, 1872) • •

Euselasia aurantiaca (saLVin & GoDman, 1868) • •

Euselasia procula (saLVin & GoDman, 1885) • •

Euselasia regipennis (butLer & Druce, 1872) • •

Juditha molpe (Hübner, 1808) • • •

Leucochimona lepida (GoDman & saLVin, 1885) • •

Menander pretus (cramer, 1777) • •

Mesene phareus (cramer, 1777) • • •

Mesene viz.phareus (cramer, 1777) • •

Mesenopsis melanochlora (GoDman & saLVin, 1878) • •

Mesosemia asa Hewitson, 1869 • • •

Mesosemia hesperina butLer, 1874 • •

Mesosemia telegone (boisDuVaL, 1836) • •

Mesosemia zonalis GoDman & saLVin, 1885 • • •

Napaea eucharila (bates, 1867) • •

Nymphidium ascolia Hewitson, 1853 • • •

Perophthalma lasus westwooD, 1851 • •

Perophthalma tullius (Fabricius, 1787) • •

Pheles strigosa (stauDinGer, 1876) Lepricornis strigosa • •

Pirascca arbuscula (möscHLer, 1883) Stichelia arbuscula • •

Pseudonymphidia clearista (butLer, 1871) • •

Sarota chrysus (stoLL, 1781) Sarota dematira • •

Sarota gyas (cramer, 1775) • •

Setabis cleomedes (Hewitson, 1870) • •

Theope acosma sticHeL, 1910 • •

Theope thestias Hewitson, 1860 • •

Theope virgilius (Fabricius, 1793) • •

Thisbe lycorias (Hewitson, 1853) • •

•

Total number of species 73 84 128 216 115 51

References

Chacón I., Montero J. 2007. Mariposas de Costa Rica. Butteries and moths of Costa Rica. Instituto

Nacional de Biodiversidad. Santo Domingo de Heredia: INBio

D’Abrera B. 1995. Butteries of the Neotropical region. Vol. 7. Lycaenidae. Victoria: Black Rock.

DeVries P.J. 1978. An annotated checklist of the butteries of Parque Nacional Corcovado during the

dry season. – Brenesia 14-15: 47-56.

DeVries P.J. 1983. Checklist of Butteries. In: Janzen D.H. (Ed.), Costa Rican Natural History. Chi-

cago: University of Chicago Press.

DeVries P.J. 1987. The Butteries of Costa Rica, Volume I: Papilionidae, Pieridae, Nymphalidae.

Chichester: Princeton University Press.

DeVries P.J. 1997. The Butteries of Costa Rica, Volume II: Riodinidae. Chichester: Princeton Uni-

versity Press.

Janzen D.H. 1983. Costa Rican Natural History. Chicago: University of Chicago Press.

Kunte K. 2008. Competition and species diversity: removal of dominant species increases diversity in

Costa Rican buttery communities. – Oikos 117: 69-76.

Lamas G. 2004. Atlas of Neotropical Butteries. Checklist: Part 4A. Hesperioidea – Papilionoidea.

Association for Tropical Lepidoptera, Scientic Publishers, Gainsville.

Robbins R.K. 2010. Four commonly confused hairstreaks (Lycaenidae, Theclinae, Eumaeini): three

need names, one does not. - J. Lepid. Soc. 64(1): 1-13.

Weissenhofer A., Huber W., Mayer V., Pamperl S., Weber A., Aubrecht G. 2008. Natural and cultural

history of the Golfo. Stapa 88, Biologiezentum der Oberösterreichischen Landesmuseen.

Wiemers M., Fiedler K. 2008. Buttery diversity of the Piedras Blancas National Park and its vicinity

– A preliminary assessment (Lepidoptera: Papilionoidea & Hesperioidea). In: Weissenhofer A.,

Huber W., Mayer V., Pamperl S., Weber A., Aubrecht G., (Eds.): Natural and cultural history of

the Golfo Dulce Region, Costa Rica. Stapa 88, Biologiezentum der Oberösterreichischen Landes-

museen, pp. 277-294.

Acknowledgements

We thank Hellena Binz for contributing unpublished data from her diploma thesis (currently in

preparation), Robert K. Robbins and Jim Miller for help in Species determination, and Konrad Fiedler

for providing information on the biology of several species.

Picture credits

Werner Huber: The main building of the Tropical Research Station, Scientists at work in La Gamba,

The ‘Comedor’, Martin Wiemers with students in the laboratory, Waterfall on the Quebrada Chorro,

Interior of the forest of the NP Piedras Blancas, The Rió Bonito in the NP Piedras Blancas

Harald Krenn: Eggs of Heliconius melpomene, Larva of Heliconius melpomene, Pupa of Heliconius mel-

pomene, Imago of Heliconius melpomene, Papilio larva with everted osmaterium, Mud puddling swal-

lowtail (Heraclides thoas), Parides iphidamas female, Heraclides thoas caterpillars on Piper leaf, Pyrisitia

nise, Phoebis sennae male, Lycorea halia (right) and co-mimic Heliconius hecale (left), Lycorea halia larva,

Morpho helenor, female, Caligo eurilochus caterpillars on a banana leaf, Catoblepia orgetorix, Hermeupty-

chia hermes (bottom), Marpesia merops (top), Marpesia chiron, Hamadryas arinome, Anartia fatima, Junonia

evarete, Siproeta stelenes, Colobura dirce, Dryas iulia, Dryas iulia larva, Heliconius buttery processing

pollen with its proboscis, Heliconius sapho (both), Heliconius cydno, Heliconius hecale (both), Heliconius

sara, Heliconius melpomene, Heliconius erato, Eurybia lycisca (both), Nymphidium ascolia, Skipper feeding

on owers of Stachytarpheta

Peter Weish: Morning clouds above the forest

Martin Wiemers: Mechanitis lysimnia, arctiid moth, Rainforest gap, Parides erithalion male, Parides

larva on Aristolochia, Heraclides thoas, Pasture, Phoebis caterpillar, Dismorphia theucharila, Rainforest

creek, Pupa of Lycorea halia, Pupa of Lycorea halia, Danaus plexippus larva on Asclepias, Mechanitis

polymnia, Mechanitis polymnia larva, Aeria eurimedia, Hyposcada virginiana, Ithomia celemia, Consul fa-

bius, Fountainea eurypyle, Morpho helenor, male, Morpho helenor, Morpho helenor larva, Morpho menelaus,

Caligo eurilochus, Caligo atreus, Hermeuptychia hermes (top), Magneuptychia libye, Pareuptychia ocirrhoe,

Pierella luna, Cithaerias pireta, Marpesia merops (bottom), Marpesia petreus, Dynamine tithia, Hamadryas

feronia, Anartia jatrophae, Anthanassa frisia, Chlosyne janais (both), Chlosyne janais larva (black morph),

Chlosyne janais larva (red morph), Historis odius feeding on rotten bananas, Adelpha cytherea, Adelpha

boeotia, Adelpha salmoneus (both), Eueides lybia, Philaethria dido, Larva of Heliconius melpomene, Gal-

lery forest, Riodinid caterpillar, Euselasia regipennis, Ancyluris jurgensenii, Calephelis laverna, Charis an-

ius, Mesenopsis melanochlora, Light gap in the rainforest, Arawacus togarna, Arumecla galliena, Strymon

megarus, Hemiargus hanno, Secondary forest , Skipper in the typical resting position, Urbanus teleus,

Pyrrhopyge thericles

Adelpha boeotia 39

Adelpha cytherea 39

Adelpha salmoneus 39

Aeria eurimedia 24

Anartia fatima 35

Anartia jatrophae 35

Ancyluris jurgensenii 49

Anius Metalmark 50

Anthanassa frisia 36

Anthanassa tulcis 36

Arawacus lincoides 54

Arteus Owl 30

Arumecla galliena 54

Banded Peacock 35

Black-and-Yellow Prestonian Clearwing 24

Blue-Winged Sheenmark 48

Blues 53ff.

Brush Footed Butteries 21ff.

Calephelis laverna 50

Caligo atreus 30

Caligo eurilochus 29

Catoblepia orgetorix 30

Celemia Clearwing 25

Charaxinae 26ff.

Charis anius 50

Charis auius 50

Chlosyne janais 37

Cissia hermes 31

Cissia hesione 32

Cithaerias menander 32

Cithaerias pireta 32

Clearwing-Mimic White 19

Cloudless Sulphur 18

Colobura dirce 38

Consul fabius 26

Costa-spotted Beautymark 49

Cracker Butteries 34ff.

Creamy Metalmark 51

Crimson Patch 37

Cuban Crescent 36

Cydno Longwing 42

Cyrestinae 33ff.

Danainae 22ff

Danaus plexippus 23

Dismorphia theucharila 11, 19

Dryas iulia 40

Dynamine salpensa 34

Dynamine tithia 34

Eueides lybia 40

Eurema nise 18

Eurimedia Clearwing 24

Eurybia lycisca 48

Euselasia regipennis 48

Forest Giant Owl 29

Fountainea eurypyle 26

Green Longwing 41

Greta oto 11

Hamadryas arinome 34

Hamadryas feronia 34

Hanno Blue 55

Hecale Longwing 43

Helenor Morpho 27

Heliconiinae 40ff.

Heliconius 41ff.

Heliconius cydno 42

Heliconius doris 45

Heliconius erato 44

Heliconius hecale 11, 22, 43

Heliconius hewitsoni 42

Heliconius ismenius 45

Heliconius melpomene 44

Heliconius pachinus 42

Index

Heliconius sapho 42

Heliconius sara 43

Hemiargus hanno 55

Heraclides thoas 12, 15

Hermes Satyr 31

Hermeuptychia hermes 31

Hesperiidae 57ff.

Historis odius 38

Hyposcada virginiana 25

Ithomia celemia 25

Ithomiinae 24ff.

Juditha molpe 46

Julia 40

Junonia evarete 36

Laparus doris 45

Longwing Butteries 41ff.

Lycaenidae 53ff.

Lycorea cleobaea 22

Lycorea halia 11, 22

Magneuptychia libye 31

Malachite 36

Mangrove Buckeye 36

Marpesia chiron 33

Marpesia merops 33

Marpesia petreus 33

Mechanitis lysimnia 11

Mechanitis polymnia 24

Melpomene Longwing 44

Menelaus Morpho 28

Merops Daggerwing 33

Mesenopsis melanochlora 51

Metalmarks 47ff.

Mimosa Yellow 18

Molpe Metalmark 46

Monarch 23

Moon Pierella 32

Morpho amathonte 28

Morpho helenor 27

Morpho menelaus 28

Morpho peleides 27

Nymphalidae 21ff.

Nymphalinae 35ff.

Nymphidium ascolia 51

Orange-rimmed Owl-Buttery 30

Orange-striped Metalmark 51

Orion Cecropian 38

Owl Buttery 29

Papilionidae 13ff.

Pareuptychia ocirrhoe 32

Parides crithalion 14

Passion Vine Butteries 41ff.

Peleides Morpho 27

Phanoptis vitrina 11

Philaethria dido 41

Phoebis sennae 18

Pierella luna 32

Pieridae 17ff.

Pineapple Fruit Borer 55

Pink-Tipped Satyr 32

Pointed Leafwing 26

Polymnia Tigerwing 24

Postman Buttery 44

Purple-topped Sombermark 48

Pyrisitia nise 18

Pyrrhopyge pseudophidias 58

Pyrrhopyge thericles 58

Quiet Diaph 32

Rayed Longwing 45

Red-Based Groundstreak 54

Riodinidae 47ff.

Sara Longwing 43

Satyrinae 27ff.

Scrub-Hairstreak 55

Sharp-edged Longwing 40

Siproeta stelenes 36

Skippers 57ff.

Small Beauty 38

Small Scintillant 50

Smooth-Banded Sister 39

Strymon megarus 55

Sulphures 17ff.

Swallowtail Butteries 13ff.

Teleus Longtail 58

Thericles Firetip 58

Thoas Swallowtail 15

Tiger-striped Longwing 45

Tiger Leafwing 26

Tiger Mimic-Queen 22

Tithian Sailor 34

Tmolus basilides 55

Urbanus teleus 58

Variable Cattleheart 14

Virginia‘s Ticlear 25

White-Patched Longwing 42

White Peacock 35

Whites 17ff.

White Satyr 32

Yellow-Edged Giant-Owl 30

Zebra Mosaic 38

Krenn etal 2010 Butterflies of the Golfo Dulce Region Costa Rica - Sample

Data

April 2015

Harald Krenn · Martin Wiemers · Lisa Maurer · Veronika Pemmer · Anton Weissenhofer

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Manejo forestal sostenible del bosque y monitoreo ecológico en dos bosques muy húmedos tropicales de Zona Norte de Costa Rica

Article

Full-text available

Jun 2019

Se realizó un monitoreo ecológico en dos propiedades sometidas a manejo forestal sostenible (MFS), ubicadas en la Zona Norte de Costa Rica. Para el monitoreo se eligió como indicador de filtro grueso el área basal y como indicador de filtro fino la composición, y diversidad de mariposas diurnas de las familias Nymphalidae, Papilionidae y Pieridae. Los resultados indican que los bosques sometidos a MFS, que además fueron afectadas por el paso del Huracán Otto, sufrieron una pérdida importante de área basal, lo que implica ajustes en el nuevo ciclo de corta que aseguren la recuperación de las áreas de bosque. Se comprobó que a menor cobertura forestal mayor abundancia, riqueza y composición de mariposas. Para lograr utilizar los monitoreos de mariposas como indicador de la sostenibilidad se requiere de un esfuerzo mayor para lograr construir líneas de referencia para este indicador.

Functional morphology of the feeding apparatus and evolution of proboscis length in metalmark butterflies (Lepidoptera: Riodinidae)

Article

Full-text available

Oct 2013

An assessment of the anatomical costs of extremely long proboscid mouthparts can contribute to the understanding of the evolution of form and function in the context of insect feeding behaviour. An integrative analysis of expenses relating to an exceptionally long proboscis in butterflies includes all organs involved in fluid feeding, such as the proboscis plus its musculature, sensilla, and food canal, as well as organs for proboscis movements and the suction pump for fluid uptake. In the present study, we report a morphometric comparison of derived long-tongued (proboscis approximately twice as long as the body) and short-tongued Riodinidae (proboscis half as long as the body), which reveals the non-linear scaling relationships of an extremely long proboscis. We found no elongation of the tip region, low numbers of proboscis sensilla, short sensilla styloconica, and no increase of galeal musculature in relation to galeal volume, but a larger food canal, as well as larger head musculature in relation to the head capsule. The results indicate the relatively low extra expense on the proboscis musculature and sensilla equipment but significant anatomical costs, such as reinforced haemolymph and suction pump musculature, as well as thick cuticular proboscis walls, which are functionally related to feeding performance in species possessing an extremely long proboscis.

Análisis del perfil de coloración de alas de lepidópteros diurnos (Lepidoptera: Papilionoidea), en dos bosques secos de Guatemala (Parque Regional Municipal Los Cerritos, Baja Verapaz; Reserva Natural Heloderma, Zacapa)

Thesis

Nov 2023

Rocio Leon

Butterflies present a great diversity of colors and morphology on their wings, thereby being good materials for multiple biological investigations. Several authors have reported seasonal polyphenisms and changes in color patterns which were physiologically induced by thermal and stress shocks (e.g. Brakefield & Larsen, 1984; Hiyama, et al., 2012; Kertész, et al., 2017; Burg & Reed, 2021). In a seasonally dry forest in Guatemala, field observations have detected that the prevalent colorations of diurnal lepidopteran wings in the rainy season differ from the colorations prevalent in the dry season (J. Yoshimoto, personal observation). In the present study I examined whether there is a seasonal effect on wing coloration of diurnal lepidopterans collected from 2011 to 2021 in two dry forests of Guatemala: Reserva Natural Heloderma in Cabañas, Zacapa and Los Cerritos Regional Municipal Park in Salamá, Baja Verapaz. The R Colordistance package (Weller & Westneat, 2019) was used to quantify a color profile in butterfly wings by processing calibrated images. The results of the cluster analysis and the hypothesis tests based on the Analysis of Similarity (ANOSIM) showed that there is no significant difference in the colorations of butterfly wings between the dry and rainy seasons, although significant differences in coloration were detected among the different months of sampling. Interestingly, species with less conspicuous colors (browns and grays) in the subfamily Satyrinae of the family Nymphalidae occurred more in the dry season, suggesting that the drastic change in vegetation cover during the dry season favors the prevalence of butterflies with more cryptic colors.

Butterfly diversity of the Piedras Blancas National Park and its vicinity – a preliminary assessment (Lepidoptera: Papilionoidea & Hesperioidea)

Article

Full-text available

Jan 2008

Two short-term surveys were carried out in 2006 and 2007 to study butterfly diversity in different habitats around the Tropical Field Station La Gamba, adjacent to Piedras Blancas National Park. Three different land use types were examined: cul-tivated land (oil palm plantations, pastures, garden and roadside verges), secondary forest (regeneration forest and gallery wood-land), and primary forest (near-natural ridge and riverine forest). As expected, species richness was lowest in habitats with inten-sive land use. Forests are more species-rich than habitats more affected by human interventions, but secondary forests are surpris-ingly similar in butterfly species composition to cultivated land, due to the dominance of some widespread and abundant species of the open countryside. Butterfly assemblages of primary forests are significantly different from, and more heterogeneous than, those of disturbed habitats. Differences in butterfly community composition appear to be due mainly to larval host plant affilia-tions and are less strongly governed by bionomic traits related to adult resource use. Despite their limited extent, short-term as-sessments of adult butterflies appear to be suitable for inferring habitat quality for butterfly species and communities. True forest butterflies were rarely observed in secondary or gallery forests. It is therefore recommended that in order to facilitate the exchange of individuals at the landscape scale, forest corridor plantations should be broad enough and contain a high variety of tree species.

Four Commonly Confused Hairstreaks (Lycaenidae, Theclinae, Eumaeini): Three Need Names, One Does Not

Article

Apr 2010

Robert Robbins

The taxonomy of four relatively common Neotropical eumaeine hairstreak species has been confused. Newly described are Iaspis andersoni Robbins, new species, differentiated from J. talayra (Hewitson), 7. castitas (Druce), and I. exiguus (Druce); Michaelas Joseph Robbins, new species, differentiated from M. ira (Hewitson); and Ignota caldas Robbins, new species, differentiated from J. gadira (Hewitson). Iaspis andersoni is unnamed because of a taxonomic misidentification made more than a century ago. The latter two are undescribed because the types of M. ira and I. gadira are not the species that they had been thought to be. Populations in the Arawacus togarna (Hewitson) species complex from Mexico and Costa Rica have been treated as two distinct species, but new data on geographical variation of wing pattern and male genitalia suggests that this classification is incorrect. A lectotype for Thecla exiguus Druce, 1907 is designated because taxonomy of the Iaspis talayra group in the Amazon Region is unresolved. A lectotype for Thecla togarna Hewitson, 1867 is designated because an incorrect type locality has engendered confusion.

The Butterflies of Costa Rica and Their Natural History: Volume I

Article

Jan 1987

Philip J Devries

Competition and species diversity: Removal of dominant species increases diversity in Costa Rican butterfly communities

Article

Jan 2008
OIKOS

Krushnamegh Kunte

Biological communities are usually dominated by a few species and show characteristically skewed species abundance distributions. Although niche apportionment and resource competition are sometimes implicated in such patterns, few experimental studies have shown direct links between resource limitation, competition with dominant species and their impacts on the overall diversity and composition of large natural communities. Here I report the results of an experiment in which I first studied species diversity and composition in two Costa Rican nectar-feeding butterfly communities numerically dominated by two species of Anartia butterflies. Then I removed Anartia from these communities to study changes in resource availability, species abundance relationships, community diversity and composition as an outcome of the removal of the dominant competitors. In the face of competition with Anartia, nectar was scarce, species abundance distributions were highly skewed, and species diversity was low in both communities. Within two weeks after the removal of Anartia, there were parallel changes in both communities: competition for nectar reduced and the nectar quantity increased substantially, which facilitated increase in community diversity and resulted in significantly less skewed species abundance distributions. Higher nectar quantity also enabled the distribution of body size and proboscis length of constituent species in the communities to expand at both ends. This study thus experimentally showed that resource competition with the dominant species was excluding many species from the communities, lowering their diversity and skewing relative species abundance relationships. These findings are of fundamental importance for competition theory and community ecology because they indicate ways in which diverse communities may be affected by and recover from competition with dominant species.

Butterflies and moths of Costa Rica. Instituto Nacional de Biodiversidad

Jan 2007

I Chacón
J Montero

Chacón I., Montero J. 2007. Mariposas de Costa Rica. Butterflies and moths of Costa Rica. Instituto Nacional de Biodiversidad. Santo Domingo de Heredia: INBio

Butterflies of the Neotropical region

Jan 1995

B D'abrera

D'Abrera B. 1995. Butterflies of the Neotropical region. Vol. 7. Lycaenidae. Victoria: Black Rock.

An annotated checklist of the butterflies of Parque Nacional Corcovado during the dry season

Jan 1978
47-56

P J Devries

DeVries P.J. 1978. An annotated checklist of the butterflies of Parque Nacional Corcovado during the dry season. -Brenesia 14-15: 47-56.

Checklist of Butterflies

Jan 1983

P J Devries

DeVries P.J. 1983. Checklist of Butterflies. In: Janzen D.H. (Ed.), Costa Rican Natural History. Chicago: University of Chicago Press.

The Butterflies of Costa Rica

Jan 1997

P J Devries

DeVries P.J. 1997. The Butterflies of Costa Rica, Volume II: Riodinidae. Chichester: Princeton University Press.

Atlas of Neotropical Butterflies. Checklist: Part 4A. Hesperioidea -Papilionoidea. Association for Tropical Lepidoptera

Jan 2004

G Lamas

Lamas G. 2004. Atlas of Neotropical Butterflies. Checklist: Part 4A. Hesperioidea -Papilionoidea. Association for Tropical Lepidoptera, Scientific Publishers, Gainsville.

Butterflies of the Golfo Dulce Region, Costa Rica

Abstract

Supplementary resource (1)

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