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Taxonomy & Evolution
1 - Taxonomy
2 - Butterfly Families and
subfamilies
3 - What is a species / subspecies ?
4 - Evolution and Speciation
5 - Lepidoptera and the Evolutionary
table
Butterfly
Families and Subfamilies
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,
Jemadia,
Mysarbia
&
Pyrrhopyge.
Most species are
stout-bodied, and average 40-60mm in wingspan. They fly rapidly,
high in 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. 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,
Darters, Swifts
Ochlodes venata
( England )
The
larvae of all Hesperiine
skippers feed on palms, grasses or bamboos. Most species have fuscous or chocolate wings,
often marked
with pale spots or hyaline windows. When basking, many
adopt the posture illustrated here. The
feeding habits of the adults varies considerably - species
from temperate zones commonly nectar at flowers, but those from
tropical areas feed at mud, rotting vegetation or bird droppings.
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Megathyminae -
Giant Skippers, Trapdoor Skippers
Stallingsia maculosus
( USA - Texas )
32 large powerful
skippers found in the
deserts, woodlands and grasslands of Mexico and southern USA. The larvae feed on Yucca, Aloe or
Agave. When older, some species bore into roots. Others feed within the fleshy leaves
and construct trap doors from which the adult butterflies can later
emerge.
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Trapezitinae -
Australian Skippers
A group of about 60-70
smallish, heavy-bodied skippers, all of which are confined to the
Australian region ( including Papua ). Unlike 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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Papilionidae
In this treatment 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
This treatment retains the classification proposed by Klots, and recognised by Lamas.
The Pieridae
comprise 1036 known species worldwide, and include the whites,
yellows, orange tips, ithomiine-mimics and jezebels. Their eggs
are usually spindle-shaped and vertically ribbed. The caterpillars
of most genera are green or cryptically coloured, naked or
slightly hairy, and without appendages or protuberances.
The pupae are also cryptically coloured, and are 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 species of Lycaenidae worldwide. The
butterflies are generally small and brightly coloured, often with
metallic blue or coppery uppersides. The caterpillars 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. In return the ants act as "guards" to
protect them from predatory insects. In many species the
relationship is very complex, and can involve the caterpillars
living within ant nests, feeding either on the ant grubs or on
aphids or fungi cultivated by the ants. The pupae do not possess a
cremaster. In some species they are formed loosely on
the surface of the ground, while 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 1060 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 has in the past been
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
1430 known species of Riodinidae, the vast majority of which are
confined to the neotropical region. The butterflies are generally
small in size, but extremely diverse in wing 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 possess honey glands or 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. Some taxonomists
still adhere to the earlier system of classification as handed
down by the fathers of entomology, but most agree that the
former Libytheidae,
Morphidae,
Acraeidae, Heliconiidae,
Danaidae,
Ithomiidae & Satyridae should be relegated
to become subfamilies or tribes of the Nymphalidae. In this
treatment I follow Wahlberg's 2008 higher classification of
subfamilies, tribes and genera, full details of which can be found
here.
The
Nymphalidae as classified by Wahlberg comprises about 6000 known species, divided into 12
sub-families. These
subfamilies each have their own distinctive characteristics, but
are united by one common feature that is instantly recognisable -
the forelegs of the adult butterflies 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, Acraeas, 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; about 200 semi-transparent African
Acraea;
about
60 weak-flying neotropical cloudforest
Actinote
and
Altinote
species, and the Argynnini - a group of 70 black-spotted temperate
woodland butterflies, typified by the Silver-washed Fritillary
illustrated here.
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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.
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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
zoilus
( Australia )
The Tellervini exists to accommodate a single Australasian species
Tellervo zoilus,
but recent research has led to some authorities elevating it's
9 subspecies 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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