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Taxonomy & Evolution
1 - Taxonomy
2 - Butterfly Families and subfamilies
3 - What is a species ?
4 - Evolution and Speciation
5 - Lepidoptera and the Evolutionary table
6
- Butterfly World Census
Butterfly
Families and Subfamilies
The 2
butterfly superfamilies Hesperioidea and Papilionoidea are positioned
systematically between the moth superfamilies
Geometroidea and
Drepanoidea, indicating that 'butterflies' are essentially a sub-group
or branch of moths, rather than being a separate group of organisms.
The Hesperioidea
comprises of a single family, the Hesperiidae. The Papilionoidea were
traditionally divided into 5 individual families - Papilionidae,
Pieridae, Lycaenidae, Riodinidae and Nymphalidae. In 2011 however
phylogeneticists discovered as a result of molecular analysis that the
moth family Hedylidae had more in common with the 5 'butterfly'
families than with other moths. Consequently they were transferred to
the Papilionoidea, and are now commonly known as butterfly-moths!
Hesperiidae
The
classification proposed by Evans has been shown by later
revisionists to be unreliable. In this treatment I follow the
arrangements of Mielke, Austin, Lamas and Warren. There are about
4130 known species of Hesperiidae worldwide. Members of this family
have a characteristic rapid darting flight, hence they are popularly
known as skippers or darts. Many of the South American species are
brightly coloured, but elsewhere most species are brownish and quite
drab in appearance. Compared to other butterflies, skippers have
stout bodies and large eyes. They also have widely separated
antennae that are usually tipped with a hooked club. The larvae of
most species live in shelters or tubes constructed from the foliage
of their foodplants. The pupae are usually formed loosely within a
frail silk cocoon.
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Pyrrhopyginae
-
Swifts, Sabre-wings
Jemadia menechmus
( Peru )
A subfamily
confined to South America and the part of Central America
south of the Rio Grande, Mexico. There are 168 known species.
Genera include
Myscelus,
Passova,
Zonia,
Pyrrhopyge,
Jemadia
&
Mysarbia.
The
butterflies are very stout-bodied, and average 40-60mm in
wingspan. They fly powerfully around the forest canopy, but
descend to feed at mud or bird-droppings.
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Pyrginae
-
Flats, Spreadwings
Pyrgus malvae
( England )
Pyrginae are represented in all temperate and tropical areas
of the world. They commonly bask on the ground or on foliage
with their wings held flat. Many tropical species habitually
rest under leaves with wings outspread. Species from temperate
regions often roost overnight on dead flower-heads. The males
of many genera have a small fold on the leading edge of the
forewings, containing androconial scales.
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Heteropterinae
-
Chequered Skippers
Dalla cypselus
( Peru )
The Heteropterinae are a fairly small sub-family comprised of
about 140 neotropical species, and 30 or so in the Holarctic
region. They are all small butterflies, averaging between
30-40mm in wingspan. Most have dark brown uppersides, dappled
with cream or orange spots. The larvae of all species feed on
grasses.
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Hesperiinae
-
Grass Skippers, Trapdoor Skippers
Ochlodes sylvanus
( England )
All Hesperiine larvae feed on monocotyledons. Adults of most
species have fuscous or chocolate wings, often marked with
pale spots or hyaline windows. When resting they hold their
wings erect, or adopt the posture illustrated here. Temperate
species commonly nectar at flowers, but those from tropical
areas feed at mud, rotting vegetation or bird droppings. The
former subfamily Megathyminae is now included in the
Hesperiinae.
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Trapezitinae
-
Australian Skippers
Trapezites symmomus ( Australia )
A group of about 60-70 smallish, heavy-bodied skippers, all of
which are confined to the Australian/Papuan region. Unlike the
Hesperiinae, males of Trapezitinae do not possess androconial
sex brands. There are 18 genera.
Trapezites is the largest, with between 15-18 species
depending on taxonomic interpretation. Each genus of the
Trapezitinae specializes on particular larval foodplants, e.g.
Trapezites all feed on spike rush
Lomandra.
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Coeliadinae
-
Policemen, Awls
The Policemen and Awls are a group of large robust skippers
with a fast whirring flight similar to that of hawkmoths. They
are active from dawn to early evening, and frequently visit
flowers for nectar. There about 75 species worldwide - 18 in
Africa, 12 in Australasia, and about 45 in the Oriental
region.
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Euschemoniinae
-
Regent Skipper
Euschemon rafflesia
( Australia )
This subfamily contains just one species -the Regent Skipper
Euschemon rafflesia, an
Australian insect in which the fore and hindwings are coupled
together in flight by a frenulum- a feature found in all
moths, and in the 'butterfly-moths' Hedylidae, but not in the
5 traditional butterfly families.
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Papilionidae
Here I follow the classification adopted by Tyler, Doubleday
and Lamas. There are about 590 known species of Papilionidae
worldwide. The majority are large colourful butterflies. Many
have short tail-like extensions on their hindwings, and hence
are popularly known as swallowtails. The caterpillars can be
either cryptically coloured or aposematic, depending on
subfamily and tribe. All Papilionid caterpillars possess an
extrusible organ called an osmaterium, which exudes foul
smelling scents to deter predators and parasitoids. The pupae
of all species are cryptically coloured, and are usually
formed head-upwards, attached by the cremaster and a silk
girdle to a leaf or stem.
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Papilioninae
-
Swallowtails, Birdwings, Dragontails
Papilio machaon gorganus
( France )
About 540 medium or large species, found throughout the
tropical, temperate and sub-arctic regions of the world. The
exquisite Dragontails of the Oriental region zip back and
forth very rapidly, and resemble dragonflies in flight. Most
others including the Swallowtails and the spectacular
Birdwings of New Guinea, have a fast fluttering flight, and
keep their wings constantly in motion when nectaring.
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Parnassiinae
-
Apollos
Parnassius apollo
( France )
About 50 species found in alpine and sub-arctic regions of the
northern hemisphere - 3 in North America, 3 in Europe, and the
remainder in mountainous regions of temperate Asia. All
species are poisonous to vertebrates. Most have
semi-transparent whitish wings, marked with prominent red or
orange ocelli.
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Baroniinae
Baronia brevicornis
( Mexico )
A sub-family
erected to accommodate a single species,
Baronia brevicornis,
a locally common endemic of the mountains of western Mexico.
It is a smallish butterfly, marked with pale yellow patches on
a brown ground colour, and is characterised by having
extremely short antennae.
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Pieridae
Here I retain the classification proposed by Klots.
The Pieridae
comprises 1036 described species worldwide and includes the
whites, yellows, orange tips, ithomiine-mimics and jezebels.
Their eggs are spindle-shaped and vertically ribbed. The
larvae of most genera are cryptically coloured, naked or
slightly hairy, and without appendages. The pupae are also
cryptically coloured, and attached head-upwards by the
cremaster and a silken girdle to the substrate.
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Coliadinae
-
Sulphurs, Yellows, Brimstones
Gonepteryx rhamni
( England )
The Coliadinae are found on all continents. Most species are
strongly migratory, including the Grass Yellows and Sulphurs,
which migrate seasonally along the routes of rivers, and
settle in vast numbers on the sandbanks of Amazonian
tributaries. Other species such as the Brimstones, are nomadic
by nature, and very adaptable to different environments.
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Pierinae
-
Whites, Orange-tips, Jezebels, Pereutes
Anthocharis cardamines
( England )
The Pierinae are very cosmopolitan of all butterflies, having
representatives occurring on all continents and in all
habitats including tropical rainforests, temperate woodlands,
alpine pastures, deserts, moorland and tundra. Most have a
white ground, often marked with black, yellow or orange. They
include the Orange tips, Whites, Arabs, Vagrants, Dotted
Borders, Flip-Flops and Jezebels.
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Dismorphiinae
-
Wood Whites, Mimics
Leptidea sinapis
( England )
The
name Dismorphiinae refers to the fact that many members of
this predominantly tropical group are sexually dimorphic -
i.e. the males and females differ markedly in appearance.
Many are also
mimetic, e.g.
Patia orise
from Brazil
is an almost perfect mimic of the Ithomiine Glasswing
Methona confusa.
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Lycaenidae
There are about 4500 described species of Lycaenidae
worldwide, although the number of known species is increasing
rapidly as a result of work by Robbins and others in the
neotropics. The adult butterflies are generally small and
brightly coloured, often with metallic blue or coppery
uppersides. The larvae are typically slug-shaped, cryptically
coloured, and often resemble leaf blisters or buds. In most
cases they live in association with ants, which milk their
honey glands to obtain a sugary secretion. The ants
coincidentally act as guards, protecting them from predatory
insects. In many species the relationship is complex, and can
involve the larvae living within ant nests, and feeding
carnivorously on ant grubs. Other species feed on aphids or
fungi cultivated by ants. The pupae do not possess a
cremaster. In some species they are formed loosely on the
surface of the ground. In others they are attached by a silk
girdle to a leaf or stem.
This treatment follows Robbins' revision of the Theclinae. The
remaining sub-families follow the classification used by Lamas
et al.
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Theclinae
-
Hairstreaks, Sunstreaks, Oakblues etc.
Thecla betulae
( England )
The Theclinae comprises about 2000 species worldwide, over
1100 of which occur in the neotropics. They are commonly known
as hairstreaks, a name derived from the white hair-line
streaks found on the undersides of most species. Another
common characteristic is the presence on most species of tails
on each hindwing. The butterflies are noted for their fast and
very erratic flight around bushes and trees.
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Polyommatinae
-
Blues, Arguses, Caeruleans
Polyommatus icarus
( England )
Polyommatinae means "many-spotted", and refers to the pattern
of small dark spots or blotches on the undersides of most
species. Males generally have blue uppersides, while females
tend to be brownish, often with orange or red sub-marginal
lunules. The butterflies primarily breed in grassland or
sub-alpine habitats, and visit flowers for nectar.
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Lycaeninae
-
Coppers, Sapphires, Silverlines
Lycaena phlaeas
( England )
The Lycaeninae are found mainly in grassland, moorland and
marshland areas of the Holarctic region, although a few
species occur in the tropics and in New Zealand. Most species
have fiery copper uppersides, often with dark borders and a
pattern of dark spots. Males typically set up territories,
voraciously defending them against any intruding insect.
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Poritiinae
-
Gems
Poritia phormedon
( Borneo )
The Poritiinae are exclusively Asian, with about 30 species
found from Assam to Malaysia, the Philippines and Sulawesi.
All are very rare, and found in rainforests and cloud-forests
up to an elevation of about 2000m. All Poritiinae species are
sexually dimorphic. Males are marked with iridescent blue or
green, while females are marked with small patches of white,
orange or red, depending on species.
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Miletinae
-
Brownies, Darkies
Allotinus horsfieldi
( West Malaysia )
A fascinating group of small delicate butterflies, mottled or
stippled with brown on a whitish ground colour. There are
about 80 species in south-east Asia, 45 in Africa, and one in
North America. The larvae of all species are carnivorous,
feeding on aphids and other Homoptera, although one Sumatran
species lives in ant nests and feeds on the grubs.
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Curetinae
-
Sunbeams
Curetis santana
( Singapore )
This sub-family
comprises of a single genus,
Curetis,
with 15 known species, found in rainforest, tropical dry
forest and coastal habitats in the Oriental region. Males have
fiery iridescent copper uppersides, but the females are brown,
with large yellowish or white patches, and can be mistaken for
small Pierids when in flight.
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Lipteninae -
African Liptenids
Epitola posthumus
( Ghana )
An exclusively African subfamily of 522 species, found mainly
in the rainforests of Ghana, Cameroon, Gabon and the Congo.
The larvae feed on lichens. The butterflies are small and
often brilliantly coloured. They are sometimes seen flying
around the trunks of fig trees, or settled on dry twigs. If
disturbed by ants, they remain on the twigs, slowly fanning
their wings.
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Liphyrinae
-
Moth butterflies
Liphyra brassolis
( Sumatra )
A small sub-family
comprising of 22 small stout-bodied African species, and 3
Liphyra species including the
"moth butterfly" Liphyra brassolis.
Its larvae live during the latter part of their lives in nests
of the weaver ant
Oecophylla smaragdina.
They are smooth, elliptical and flattened, with a hard shell
which protects them from being attacked by the ants. The adult
butterflies lack a proboscis and cannot feed.
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Riodinidae
This family was in the past known as the Eryciinidae or
Nemeobiidae and is regarded by some as being a sub-family of
the Lycaenidae. In this treatment I follow Stichel, Harvey,
Hall, d'Abrera, Lamas, Willmott and Callaghan, who regard it
as a family in it's own right.
There are
about 1500 species of Riodinidae, the vast majority of which
are confined to the neotropical region. They are generally
small in size, but very diverse in shape, colour and pattern.
Many have metallic green, blue or orange areas on the
uppersides, hence they are popularly known as metalmarks. Some
species have long tails on the hindwings and resemble
miniature swallowtails. Metalmark larvae are similar to those
of the Lycaenidae, but do not normally associate with ants. In
many neotropical species the larvae are adorned with prominent
tubercules or other protuberances.
As with the Lycaenidae, the pupae do not possess a cremaster,
and according to species are either formed loosely on the
surface of the ground, or are attached by a silk girdle to a
leaf or stem.
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Riodininae
-
Metalmarks, Underleafs, Doctors etc.
Hamearis lucina
( England )
The Duke of Burgundy illustrated here is the only European
member of the Riodinidae. In South and Central America there
are 1150 known species in the sub-family Riodininae. Most are
roughly similar in size to the Duke of Burgundy, but many are
marked with metallic green, blue, purple, copper and red
scales, and are commonly known as metalmarks.
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Euselasiinae
-
Euselasias
Euselasia gelanor
( Peru )
This sub-family is confined entirely to the neotropical
region. There are 172 known species. The butterflies range
between about 25 - 40mm in wingspan. They habitually hide
under leaves, with their wings held erect, occasionally
darting out to investigate other butterflies. The undersides
are generally marked with vertical bands in shades of brown,
white and orange, but the uppersides are typically black or
chocolate,
with large patches of bright red or iridescent blue.
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Nymphalidae
Many of the
sub-families described below were previously considered to be
families in their own right. In this treatment I follow
Wahlberg's 2008 classification
in which the 6000
Nymphalid species are divided into 12 subfamilies. These each
have their own distinctive characteristics but are united by
one common feature - the forelegs of the adults are greatly
reduced in size and are useless for walking. In males they are
reduced to such a degree that they are little more than hairy
stumps, hence the popular term 'brush-footed' butterflies.
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Libytheinae
-
Beaks
Libythea myrrha
( West Malaysia )
A very small sub-family with only 19 species worldwide, but
represented on all continents and on many islands. Some
taxonomists still consider Libytheidae to be a full family.
The butterflies are characterised by having very long
beak-like labial palpi. When at rest the jagged brown wings
take on the appearance of dead leaves, with the "beak"
resembling a leaf stalk.
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Nymphalinae
-
Aristocrats, Checkerspots etc
Inachis io
( England )
The Nymphalinae
includes many of the world's most familiar species, including
the Peacock, Red Admiral & Painted Lady. Other representatives
include the genera
Junonia,
Siproeta,
Anartia,
Kallima,
Hypolimnas
and the smaller Fritillaries in the genera
Euphydryas,
Chlosyne,
Melitaea
and
Mellicta,
all of which were formerly included in the now defunct
sub-family Melitaeinae.
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Heliconiinae
-
Longwings, Fritillaries etc
Argynnis paphia
( England )
A contentious
combination of taxa including about 70 species of Heliconiini
- neotropical rainforest species marked with patches of cream,
red and blue; and the Argynnini which comprises of about 70
black-spotted temperate woodland butterflies, typified by the
Silver-washed Fritillary illustrated here.
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Acraeinae
-
Acraeas, Altinotes etc
Acraea bonasia
( Ghana )
Some workers
regard the African Acraea and
South American Altinote /
Actinote /
Abananote genera as being members of the tribe
Acraeini, which they place in the subfamily Heliconiinae.
Recent DNA research however indicates that they are distinct
from the Heliconiinae, thus I treat them here as members of
the subfamily Acraeinae.
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Limenitidinae
-
Gliders, Sailors, Sisters, Foresters
Adelpha capucinus
( Peru )
In 2003 Wahlberg
reclassified the Limenitidinae, excluding many genera which
are now reassigned to the Biblidinae. The revised arrangement
is more rational, including only a few very obviously related
genera e.g.
Adelpha,
Limenitis,
Neptis,
Athyma
and
Moduza. All
are forest dwelling species noted for their graceful flight
and distinctive uppersides, banded with white and / or orange.
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Biblidinae
-
Crackers, Numberwings
Diaethria clymena
( Peru )
The Biblidinae are
a large group of very colourful forest butterflies which to
the layman have few obvious features in common. The vast
majority of genera such as
Diaethria,
Nessaea,
Hamadryas &
Panacea
are
restricted to the neotropics although a few genera including
Ariadne
&
Neptidopsis
are found
in Africa.
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Cyrestinae
-
Daggerwings, Mapwings, Maplets
Marpesia petreus
( Peru )
The subfamily
Cyrestinae include the spectacular
Marpesia
Daggerwings
of the neotropics, which roost overnight in clusters of up to
50, hanging beneath leaves. Also included are the oriental
Cyrestis
Mapwings &
Chersonesia
Maplets,
which commonly rest under leaves, with their wings fully
outspread.
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Apaturinae
-
Emperors, Princes, Doxocopas
Apatura iris
( England )
There are about 50
species of Apaturinae worldwide, all forest dwelling
butterflies noted for their with powerful flight. Males obtain
their sustenance from minerals dissolved in mud, sap, rotting
fruit, carrion or dung, and transfer the proteins to females
during copulation. Neither sex visit flowers. Males of
Apatura
and
Doxocopa
display a
brilliant purple, blue or green sheen when viewed at certain
angles.
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Charaxinae
-
Leafwings, Pashas, Preponas, Agrias
Archaeoprepona amphimachus
( Peru )
Only one member of
the Charaxinae is found in Europe,
Charaxes jasius.
The majority of species are restricted to tropical /
sub-tropical areas, with the greatest concentrations in Africa
and South America. They are powerful flyers, found primarily
in rainforest and tropical dry forest habitats. Many have
convincing "dead-leaf" undersides, but some such as
Agrias have intricate patterns in
red, cream and blue. The uppersides usually have bright blue
or red bands or patches.
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Satyrinae
-
Browns, Phantoms, Palmflies
Melanargia galathea
( England )
Satyrine larvae
feed on grasses, palms or bamboos. In the Holarctic region the
subfamily is represented by the Browns, Heaths, Ringlets,
Marbled Whites & Graylings. Their African and tropical Asian
counterparts include ringlet-like genera such as
Ypthima
&
Mycalesis.
South America has by far the greatest diversity, with the
transparent
Cithaerias,
skulking
Pierella,
and
Euptychia
ringlets
being well known examples.
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Morphinae
-
Morphos, Owls, Jungle Glories
Caligo oileus
( Peru )
The Morphinae
includes
some of the largest & most dazzling butterflies on Earth - the
enormous iridescent blue Morpho
butterflies which flap slowly along river courses in Central &
South America. Other members of the subfamily include the
Jungle Glories of Asia ( Amathusiini ), and the neotropical
Caligo Owl butterflies (
Brassolini ). The huge "owl" eyespots of the the latter serve
a dual role - acting as a threat when intimidated, and as
decoys in the event of attack. Recent molecular study strongly
suggests that the Morphinae are a tribe of Satyrinae.
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Danainae
( Danaini ) -
Monarchs, Tigers, Crows
Danaus genutia
( West Malaysia )
There are about
200 species of Danaini. Only 2 species have been recorded in
Europe -
Danaus plexippus
and
D.
chrysippus.
The larvae of all species assimilate toxins from their
foodplants, and pass them on to the adult butterflies,
rendering them unpalatable to birds. This unpalatability is
advertised by the bright orange and black patterning on the
wings, which in turn is mimicked by various other toxic and
non-toxic species from other sub-families.
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Danainae
( Ithomiini ) -
Glasswings, Tiger-mimics
Ithomia terra
( Peru )
Ithomiines fall (
unscientifically ) into 2 groups - the orange and black banded
Tiger-mimics, and the transparent Glasswings. They occur
exclusively in neotropical rainforests and cloudforests. The
adult males feed at
Heliotropium
and
Eupatorium
to obtain
pyrrolizidine alkaloids, which they chemically convert into
pheromones. They gather at "leks" to release their pheromones
to attract mates.
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Danainae
( Tellervini ) -
Hamadryads
Tellervo assarica
( New Guinea )
The Tellervini was originally
thought to comprise of a single Australasian species
Tellervo zoilus,
with 9 subspecies. Recent research however has led to all of
these being elevated to full species status. The butterflies
are allied to the Danaini and Ithomiini, but differ
morphologically from both in several respects. They are found
in Queensland, Papua and the Solomon Islands.
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Hedylidae
The 35 species of the family Hedylidae are nocturnal and
moth-like in appearance.
The early stages however have many butterfly-like
characteristics. The eggs for example are structurally closer
to those of Pieridae and Nymphalidae than to moth eggs. The
caterpillars have horn-like processes like Apaturinae, a bifid
tail as found in Satyrinae, and secondary setae as found in
Pieridae. They also have an "anal comb" used for expelling
droppings - a characteristic of the Hesperiinae. The pupae
likewise have structural characteristics more representative
of butterfly pupae, and are secured to the substrate with a
silken girdle, just like those of the Pieridae and
Papilionidae.
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Hedylidae
-
Butterfly-moths
Macrosoma heliconaria
( Peru )
The Hedylidae have not yet
been assigned any subfamilies or tribes.
All Hedylidae
are placed in the same genus - Macrosoma.
There are 35 species distributed variously from Mexico to
Bolivia. The greatest concentration is in southern Peru where
at least 26 species occur.
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