Explanation of Nymphalidae Butterflies

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International Academic
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International Academic Journal of Science and Engineering
Vol. 5, No. 4, 2018, pp. 24-47.
ISSN 2454-3896
24
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International Academic Institute
for Science and Technology
Explanation of Nymphalidae Butterflies
Vitthalrao Bhimasha Khyadea, Priti Madhukar Gaikwadb, Pranita Rajendra Vareb
a Department of Zoology, Shardabai Pawar Mahila Mahavidyalaya, Shardanagar Tal. Baramati Dist. Pune – 413115 Maharashtra,
India.
b Graduate Student (T. Y. B. Sc.), Shardabai Pawar Mahila Mahavidyalaya, Shardanagar Tal. Baramati Dist. Pune 413115
Maharashtra, India.
Abstract
The Nymphalidae is a family of several thousand species found in all zoogeographical regions of the
world. Most are medium or large in size, but the family is highly variable given that it also includes the
Satyrinae, a subfamily that has been designated as a family in its own right in earlier classifications. The
forelegs in both sexes are vestigial and useless for walking. In the male, there are typically only 2 tarsal
joints and the legs have a brush-like appearance, resulting in a common name for this family - the "brush-
footed butterflies". The female foreleg has 4 tarsal joints which, when compared with the male, provides a
mechanism of determining the sex of the adult. The midleg and hindleg are normal in both sexes and both
tibia and tarsus may have spines. This family is represented by the following subfamilies: Danainae;
Satyrinae; Morphinae; Heliconiinae; Limenitidinae; Apaturinae and Nymphalinae.
Keywords: Brush-Footed Butterflies; Danainae; Monarch; Gatekeeper

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Introduction:
Butterflies of family: Nymphalidae are with reduced forelegs. The reduction in the length of forelegs is
the most significant, which made them to appear as if with only two pairs of legs (mesothorasic and
metathorasic). Therefore, butterflies of family: nymphalidae are also called as “Four Footed Butterflies”.
Reduction of the first pair of legs (prothorasic legs) in butterflies of family: nymphalidae appears
particularly extensive in the femur. Remaining two pairs of legs (mesothorasic and metathorasic) in
nymphalidae are frequently hairy and resemble brushes. The “Feather legged butterflies”, alternative
name derives from the fact of hairy or feathery legs of these butterflies. The Nymphalidae is with about
six thousand species distributed throughout most of the world. These butterflies are usually medium-sized
to large butterflies. Most species of family: Nymphalidae have a reduced pair of forelegs and many hold
their colourful wings flat when resting. The butterflies of family: Nymphalidae are also called brush-
footed butterflies or four-footed butterflies, because they are known to stand on only four legs while the
other two are curled up; in some species, these forelegs have a brush-like set of hairs which gives this
family its other common name.
Many species of family: Nymphalidae are brightly coloured and include popular species such as
the emperors, monarch butterfly, admirals, tortoiseshells, and fritillaries. However, the under wings of the
members of family: Nymphalidae are, in contrast, often dull and in some species look remarkably like
dead leaves, or are much paler, producing a cryptic effect that helps the butterflies blend into their
surroundings. Rafinesque (1815) introduced the name Nymphalia as a subfamily name in diurnal
Lepidoptera. Rafinesque did not include Nymphalis among the listed genera, but Nymphalis was
unequivocally implied in the formation of the name (Code Article 11.7.1.1). The attribution of the
Nymphalidae to Rafinesque has now been widely adopted (Vane-Wright & de Jong, 1978). In the adult
butterflies, the first pair of legs is small or reduced, giving the family the other names of four-footed or
brush-footed butterflies. The caterpillars are hairy or spiky with projections on the head, and
the chrysalids have shiny spots. The forewings have the submedial vein (vein 1) unbranched and in one
subfamily forked near the base; the medial vein has three branches, veins 2, 3, and 4; veins 5 and 6 arise
from the points of junction of the discocellulars; the subcostal vein and its continuation beyond the apex
of cell, vein 7, has never more than four branches, veins 8–11; 8 and 9 always arise from vein 7, 10, and
11 sometimes from vein 7 but more often free, i.e., given off by the subcostal vein before apex of the cell
(Bingham, 1905). The hind wings have internal (1a) and precostal veins. The cell in both wings is closed
or open, often closed in the fore, open in the hind wing. The dorsal margin of the hind wing is channelled
to receive the abdomen in many of the forms (Bingham, 1905). The antennae always have two grooves on
the underside; the club is variable in shape. Throughout the family, the front pair of legs in the male, and
with three exceptions (Libythea, Pseudergolis, and Calinaga) in the female also, is reduced in size and
functionally impotent; in some, the atrophy of the forelegs is considerable, e.g.,
the Danainae and Satyrinae. In many of the forms of these subfamilies, the forelegs are kept pressed
against the underside of the thorax, and are in the male often very inconspicuous (Bingham, 1905).
The butterfly family Nymphalidae has been an important taxon for developing some of the mentioned
hypotheses. Nymphalidae contains around 6000 species. Several members of nymphalidae are considered
model organisms in evolutionary biology (Joron, et al, 2006; Willmott, et al, 2006; Brakefield, et al,
2009). The family most likely originated around 94 MYA in the mid Cretaceous. Diversification of the
group began in the Late Cretaceous and most major radiations (current subfamilies) appeared shortly after

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the Cretaceous-Paleogene (K-Pg) boundary (Heikkil, et al, 2012). Several attempts have used time-
calibrated phylogenies and diversification models to reconstruct the evolutionary history of the group to
identify patterns of accelerated or decelerated diversification of some Nymphalidae clades (Heikkil, et al,
2012; Elias, et al, 2009; Fordyce, et al, 2010; Wahlberg, et al, 2009). For example, Pena and Wahlberg
(2008) suggested that climate change in the Oligocene and the subsequent diversification of grasses has
led to diversification of the subfamily Satyrinae (Pena and Wahlberg, 2008) due to the abundance of
grasses over extensive geographic areas (“resource abundance-dependent diversity dynamics”
hypothesis): Fordyce (2010) found increased net diversification rates in some Nymphalidae lineages after
a major host plant shift, which appears to be in agreement with the “escape-and-radiate” model of
diversification (Ehrlich and Raven, 1964). Although it has been suggested that part of the great diversity
of Nymphalidae butterflies is a result of hostplant-insect dynamics, it is necessary to use modern
techniques to investigate whether the diversification patterns of Nymphalidae are in agreement with the
theoretical predictions. It is necessary to test whether the overall diversification pattern of Nymphalidae is
congruent with events of sudden diversification bursts due to hostplant shift or climatic events (Nylin, et
al, 2014; Ferrer-Paris, et al, 2013). In the study, Pena and Espeland (2015) used a time-calibrated genus-
level phylogenetic hypothesis for Nymphalidae butterflies (Wahlberg, et al, 2009) to investigate patterns
of diversification. Pena and Espeland (2015) applied the statistical method MEDUSA (Malware detection
using statistical analysis) of system (Alfaro, et al, 2009), to study the diversification pattern of
Nymphalidae butterflies. MEDUSA fits likelihood models of diversification into a time-calibrated tree
and tests whether allowing increases or decrease the speciation and extinction rates within the tree
produces better fit of the models. MEDUSA is able to take into account unsampled extant species
diversity during model fitting and it is normally applied to the maximum clade credibility phylogenetic
tree. Particularly, Pena and Espeland (2015) wanted to study the effects of phylogenetic uncertainty and
by using the extended MEDUSA method called MultiMEDUSA (Alfaro, et al, 2009). Pena and
Esplanade (2015) also tested whether hostplant association dynamics can explain the diversification
patterns of component Nymphalidae lineages by testing whether character states of host plant use affected
the diversification pattern of those lineages employing the binary speciation and extinction model
(BiSSE) as implemented in the R package diversitree (Fitz John, 2012).
Butterfly Migration
Biological migration is the relatively long-distance movement of individual animals, usually on a
seasonal basis. It is the most common form of migration in ecology. It is found in all major animal
groups, including birds, mammals, fish, reptiles, amphibians, insects, and crustaceans (Dingle, et al,
2007). To be counted as a true migration, and not just a local dispersal or irruption, the movement of the
animals should be an annual or seasonal occurrence, such as Northern Hemisphere birds migrating south
for the winter; wildebeest migrating annually for seasonal grazing; or a major habitat change as part of
their life, such as young Atlantic salmon or Sea lamprey leaving the river of their birth when they have
reached a few inches in size (Silva, et al, 2013). It's possible to find a few butterfly species straying into
city parks and gardens, investigating weedy areas of wasteland or flying in other 'unnatural' habitats, but
despite such apparently cosmopolitan lifestyles butterflies are extremely choosy about where they lay
their eggs. Butterflies have strict requirements in terms of minimum / maximum temperature tolerance.
The habitats they occupy are determined by where their larval food plants grow, and by the availability of

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adult food sources and roosting sites. They are unable to survive and breed unless these and numerous
other vital conditions are precisely met. Suitable habitats are often highly localised so consequently many
species have an extremely patchy distribution. Most species never stray more than a kilometre away from
their established breeding grounds - it would be wasteful of their short lives to ramble across barren
habitats where there were no suitable plants on which to lay their eggs. Nevertheless, all species whether
humans or butterflies are to a greater or lesser extent genetically programmed to 'leave home', dispersing
to explore new areas. It is important to understand the difference between dispersal and migration. The
term dispersal is used to describe random and aimless movement away from the site where a butterfly
emerges. Dispersing butterflies are easily diverted from their course by minor changes in wind direction
or obstacles in their path. They will for example fly around the edge of a block of forest rather than fly
through it or over it. When they encounter hostile habitats such as arable farmland, lakes, rivers, roads or
buildings they steer left or right to try to find a route around them.
The butterfly migration refers specifically to medium or long distance directional movements by the
butterflies. Migrating butterflies have a strong purposeful flight and are unaffected by obstacles or hostile
landscapes. If for example they encounter a building they will fly over it rather than take an easier route
around it. Their flight path is not affected by wind. The contribution of Carrington Bonsor Williams (7
October 1889 – 12 July 1981) in butterfly migration is well accredited. This name is particularly
associated with insect migration, statistical ecology and biogeography. He made uncountable observations
himself during his years in the Tropics and he had colleagues all over the world making new
observations. He analyzed and published the results in a long series of publications and became a world-
leading authority on the subject. Through his research, he was able to shed light on many of the problems,
which he had first formulated in his 1930 thesis. He published a much enhanced account on the subject in
1958 (Wigglsworth, 1982). He analysed 470 migrations in well-known colorful butterfly, painted lady,
Vanessa cardui (L) (Family: Nymphalidae) migrations in various parts of the world and found no
correlation between flight direction and wind direction. Other studies involving migrations of Catopsilia
florella and Eurema hecabe in Africa, Pieris brassicae in England and Ascia monuste in North America
have arrived at the same conclusion. Migrations usually involve mass movements - a flight of Vanessa
cardui in California in 1924 was estimated to contain about 3000 million butterflies. It has been estimated
that a swarm of migrating snout butterfly, Libytheana carinenta (L) in Texas in 1921 was passing at a rate
of over a million butterflies per minute across a 250-mile-wide front. Migrations often begin
spontaneously across a wide area, but once in the air the insects all head towards a common destination. It
is generally accepted that migration is a 2-way process e.g. butterflies migrating south in autumn will
return north again in the spring. In some cases, an individual butterfly will take part in both legs of the
journey, but in many species a 'parent' generation flies in one direction, and it is the offspring that
undertake the return journey. The migrant species such as Danaus plexippus, Vanessa cardui or Pieris
brassicae would have an extremely directional flight pattern and would fly over a hostile habitat to
maintain its compass direction.
Random dispersal in Butterflies:
Nothing in nature is constant. Habitats are continually changing. Woodlands become overgrown and
shade out herbaceous plants on which the caterpillars of many butterflies depend. Heathlands catch fire,
grasslands become overgrown with scrub, deserts expand, cliffs crumble away. Species at the edge of

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their natural distribution range tend to inhabit sites which are sheltered from bad weather. Pearl-bordered
Fritillaries for example occupy open meadows in mainland Europe, but in Britain such habitats are too
cold for them so they form colonies in sheltered woodland clearings instead. Unfortunately, these habitats
are ephemeral. They fast become overgrown and the violets on which the Fritillary caterpillars feed
rapidly get shaded out. Within 2-3 years the habitat can no longer support the species, so the adult
butterflies disperse soon after mating, radiating out along forest trails in search of more suitable breeding
areas. Many other factors can trigger dispersion e.g. after a particularly successful breeding season Marsh
Fritillary Euphydryas aurinia populations can reach explosive levels. The larvae scour their habitat,
devouring every available leaf of their food plant. Ultimately they sense imminent starvation, and
undergo a chemical change which switches them into a high activity phase causing them to swarm across
the surrounding countryside in search of food.
Daily Commuting in Buterflies:
In Britain adults of Clossiana euphrosyne nectar almost exclusively on bugle - Ajuga reptans. They lay
their eggs on dead bracken or dry grass stems, but their caterpillars feed on Viola. Fortunately for the
butterfly all of these plants can be found in the same habitat - recently cleared woodland, so it is here that
the butterflies spend their entire lives. Many other species however are not so lucky - their larval food
plants may grow in entirely different places from the adult food sources, so they need to commute
between breeding and feeding sites. Species such as Apatura iris which occur at low density in woodland
habitats can't easily locate the opposite sex, so they have evolved 'hill-topping' - a strategy whereby both
sexes fly to the highest point in the vicinity, typically a tall oak tree on a ridge, where courtship and
copulation take place. After mating the females disperse to lay their eggs on sallow bushes which
typically grow alongside ditches on lower ground. The males also disperse to low lying areas where they
feed by imbibing mineralised moisture from the paths or patches of mud. Next morning, they commute
back to the 'master' oak tree to mate with other females. Purple Emperor Apatura iris commutes daily
between feeding and courtship sites.
Montane butterflies:
In temperate regions mountainous areas such as the Alps, Pyrenees, Rockies and Tien Shan, land above
about 1800m is covered in snow for much of the year. In the Andes, the Himalaya and the mountain
ranges of New Guinea areas as low as 3000m are subject to seasonal snow cover. During the short
montane summer however, the meadows and pastures become carpeted in vast swathes of flowers,
attended by hordes of butterflies. Swarms of Blues, Fritillaries and Skippers aggregate to imbibe moisture
from patches of mud; while Coppers, Ringlets, Apollos and Heaths nectar avidly at the abundant flowers.
Most of these species are sedentary insects, forming highly localised breeding colonies. Because they are
'stay-at-home' species, their flight season is limited to 2-3 weeks in mid-summer, and they have to spend
several months of their lives hibernating as either eggs or caterpillars. Other species such as Clouded
Yellows, Whites and Swallowtails are nomadic, and migrate down to the lowlands in late summer to
breed. In the early spring their progeny produces a further brood in the lowlands, but the habitat there
becomes too hot and dry in summer, so they then return to the mountainsides where there is cooler air
and an abundance of flowers for nectaring. Another form of altitudinal movement takes place on a daily
basis. In alpine regions one side of a mountain may remain in the shade until late morning. Sunlight

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reaches the east facing slopes first, but may not reach steep west facing slopes until late afternoon.
Consequently, butterflies commute around the mountainsides to keep pace with the sun. Butterflies
commute altitudinally too, moving from peaks to valleys and back again, to areas where the temperature
is most suitable. Sometimes these journeys take them to mountain passes. Over the millennia these
become established routes by which species migrate seasonally from one valley to another.
Seasonal Butterfly Migration
Seasonal migration is an entirely different phenomenon from commuting or random dispersal. It tends to
occur spontaneously and involves the mass movement of hundreds, thousands or even millions of
butterflies. Long-distance migration can lower parasite prevalence if strenuous journeys remove infected
animals from wild populations. The authors of this study examined wild monarch butterflies to investigate
the potential impact of the protozoan parasite Ophryocystis elektroscirrha on migratory success. The
researchers collected monarchs from two wintering sites in central Mexico to compare infection status
with hydrogen isotope measurements, an indicator of latitude of origin at the start of fall migration.
On average, uninfected monarchs had lower hydrogen isotope values than parasitized butterflies,
indicating that uninfected butterflies originated from more northerly latitudes and travelled farther
distances to reach Mexico. Within the infected butterflies, monarchs with higher levels of infection
originated from more southerly latitudes, indicating that heavily infected monarchs originating from
farther north were less likely to reach Mexico. The authors used observations of infection levels during
the breeding season to show that southerly latitudes did not simply generate more infected monarchs prior
to the start of the migration.
Collectively, these results emphasize that seasonal migrations may help lower infection levels in wild
animal populations. The authors posit that these results combined with recent observations of sedentary,
winter-breeding monarch populations in the southern U.S. indicate that the shifts from migratory to
sedentary behavior may lead to greater infection for North American monarchs.
The chemical markers allowed us to estimate where the monarchs started and how far they travelled to
reach the wintering sites in Mexico, something that would not be possible using other currently available
methods. It is also found that monarchs with larger wings migrated farther distances on average than
those with smaller wings, something that's supported by cross-population comparisons but hasn't been
previously shown within North American migratory monarchs.
The Evolution of Migratory Behaviour in Butterflies:
When butterflies first evolved the present day continents were united to form the giant land mass
Pangaea. In the centre of such a massive continent there would have been huge seasonal extremes of
temperature and humidity. Consequently, butterflies would have had to find some way to survive when
their larval food plants and nectar sources wilted in summer, or when temperatures were too high or too
low for normal activity. One choice would be to remain in situ and diapause, i.e. to aestivate. In a
diapaused state large numbers would perish due to predation by insectivorous birds. A better alternative
would be to fly to another location e.g. close to a river, where food plants would continue to be available
long after the surrounding land had dried out. Further food plants would then be found by migrating up or
down river. Natural selection would ensure that butterflies which survived as a result of such migratory
actions would be able to pass on the trait genetically. It is perhaps no coincidence that so many species in

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the tropics still use rivers and streams as migration paths. In subsequent geological periods the migratory
behaviour would have been interrupted as tectonic activity caused mountain ranges and seas to appear,
thereby splitting up formerly contiguous areas of breeding terrain. These geological movements took
place over millions of years, during which butterflies probably continued to migrate along the same
routes, crossing mountain ranges via low passes, and hopping from island to island to cross seas and
oceans. Many species would have been unable to overcome the new natural barriers, particularly the ever
widening oceans. Their populations therefore became permanently divided, gradually taking on new
characteristics, and ultimately evolving into new species. Other species however managed to cross the
barriers regularly, and migratory behaviour became genetically imprinted in them.
Triggers of Butterfly Migration:
Individual migrations appear to be triggered primarily by climatic phenomena, often in association with
over-abundance, and depletion of larval food resources. C. B. Williams in his book 'Insect Migration'
quotes Skertchley, who in March 1869 witnessed the commencement of a migration of Vanessa cardui in
desert behind Suakin by the Red Sea: "He noticed that the whole mass of grass, through which they were
riding on camels, was in a state of violent agitation although there was no wind. When he dismounted he
found that the cause was the emergence from the chrysalis of myriads of Painted Lady butterflies, which
dried their wings and about half an hour later flew off together eastwards towards the sea". Often climatic
differences on opposite sides of a mountain range will cause butterflies to migrate between them,
ascending to cross over high alpine passes. Beebe's observations include accounts of over 250 butterfly
species migrating through the 1000m high Portacheulo Pass in Venezuela. 22 of these species were said
to have occurred in "thousands". In late May 1946 he and 4 colleagues used stop watches and counters to
try to estimate the numbers of Eunica monima migrating through the pass. In Beebe's own words they
"completely failed to keep up with fast enough estimates of numbers, but at a minimum clocked 1000 per
second going past in the face of a gentle breeze".
Route-Finding in Butterfly Migration:
The studies on Monarchs has shown that their annual migration from Canada to Mexico is controlled by a
'time-compensated sun compass' that depends on light receptors, and a circadian clock built into the
antennae. When scientists removed the antennae from one group of Monarchs they flew strongly but in
random directions, but a control group with their antennae intact all flew in the same direction - their
south-westerly migration route. In another experiment the antennae of some were painted with black
enamel, and these butterflies when placed in a flight simulator all flew together, but in the 'wrong'
direction compared to their normal migration route. Another group had their antennae painted with
transparent paint, and these all migrated together in the right direction. Research by Chapman suggests
that migratory butterflies and other insects are programmed to seek out 'wind highways' in the sky, which
they use to enable them to travel quickly during their migratory flights. This may be the case with certain
species, but it is well documented that species including Colias crocea, Vanessa cardui and Pieris rapae
fly very low over the sea when migrating from the European mainland to Britain. There is for example a
famous account by Rev. Harrison, who in 1868, from a cliff near Marazion, Cornwall, observed "a
yellow patch out at sea, which as it came nearer showed itself to be composed of thousands of Clouded
Yellows, which approached flying close over the water, rising and falling over every wave till they

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reached the cliffs". A team of neurobiologists led by Steven Reppert of the University of Massachusetts
spent several years investigating Monarchs. In 2010 they published a paper indicating that they had
discovered that Monarchs have 2 types of photoreceptor proteins which not only allow them to see UV
light but also enable them to detect the Earth's magnetic field. A series of experiments on Drosophila fruit
flies whose genomes were engineered using Monarch Cry1 and Cry2 cryptochrome proteins proved that
they respond to magnetic fields when under the influence of UV-A/blue light.
Expansion and Contraction of Range of Butterfly Migration:
The range is nothing but the overall area in which a species can be found. Within that range there will be
areas of unsuitable habitat so the distribution within a butterfly's range is always patchy. The range and
distribution are limited by climate, geology, aspect and altitude, all of which affect the type of larval food
plants and adult nectar sources that will grow in an area. The distribution of a species within its range is
also greatly affected by human intervention - urban expansion has the greatest impact, but governmental
policy on farming, forestry and road planning also has a very profound effect on the distribution and
abundance of butterflies. The best example is the High Brown Fritillary Argynnis adippe which was
widely distributed and common in England until the 1950's. It's range then contracted rapidly as a result
of habitat fragmentation, and a change from traditional coppice woodland management to the mass
planting of conifers in English forests. By the turn of the 21st century the butterfly had disappeared from
virtually all of its former habitats. Now in 2012 it clings to its existence at just a handful of sites in
western England. Most other forest butterflies have also declined as a result of being shaded out of the
darker and cooler modern woodlands. Only one British species Pararge aegeria has benefited, as it
survives better in shadier conditions. It has even been able to increase its formerly patchy distribution in
the UK to the point where it is currently widespread and common over almost its entire range.
Monarh Migration: The Example of Butterfly Migration:
The most recognized butterflies of North America are the Monarch butterflies (Danaus plexippus L.)
These milkweed varieties of butterflies come from the family of Danainae, a subfamily of Nymphalidae.
People have seen this species in Australia and New Zealand since 1871, where they call it "the wanderer."
It inhabits the Azores, Canary Islands and Madeira. Occasionally you can sight them in Western Europe
and rarely in the United Kingdom. You can easily identify these beautiful butterflies by their wings with
black and orange pattern, and a wingspan of 3½–4 in or 8.9–10.2 cm. Males are a bit bigger than female
monarchs are and have a spot known as the androconium in the middle of each rear wing. Wings of
female’s boast of darker veins. Monarchs are among a small group of butterflies, mostly related species
that make a two-way migration in one generation. And, of this unique few, the fall migration of monarchs
from Canada and the United States to overwintering sites in Mexico is both the longest known and most
spectacular. How the monarch is able to accomplish this amazing feat is the subject of much speculation
and research. Scientists are getting closer to understanding how monarchs sense changing environmental
signals in both fall and spring and how they modify their behavior based on these changes to get to their
destinations. When monarchs take off for the south bound leg of their journey, people might see many
hundred, even a thousand, in well-known viewing places like Point Pelee in Southern Ontario, Canada.
Several migration routes in central southern Canada lead down through the central U.S. Most monarchs
that migrate to Mexico originate from the northern breeding grounds east of the Rocky Mountains and

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north of the central United States (e.g. Oklahoma). A smaller monarch population that breeds in areas
west of the Rockies overwinters at numerous sites scattered along the California coastline from San
Francisco to San Diego. Getting to Mexico isn’t easy. It’s not just a matter of flying south. If they only
flew south, monarchs would end up in the Gulf of Mexico. Rather to get to Mexico, they set a course, and
this course is different for different regions of the country. Monarchs passing through Washington, D.C.
are moving west-southwest, near Atlanta the direction is almost due west, and in Lawrence, Kansas about
south-southwest. How monarchs set these courses is unknown. Do they use celestial cues or magnetic
information, some combination of the two or are other factors involved? Scientists are working to solve
this puzzle. We do know that they use a combination of biological clocks, one set in the antennae and
another in the brain that are used to stay on course. Surviving the trip to Mexico involves elements of luck
as well as specific adaptations that improve the odds of reaching the overwintering sites. The butterflies
must avoid everything from vehicles to pesticides, spider webs, storms and the occasional bird that has
yet to learn that monarchs aren’t to be eaten due to their distasteful chemistry. In addition to their unique
ability to set a course that will take them to the right place, monarchs have flight adaptations that improve
the chances of arriving at the overwintering sites in good condition. Rather than using powered flight –
that is, constant flapping –monarchs use gliding and soaring to advance to the southwest. As the Sun
heats the ground, thermals form. As monarchs encounter these warm rising air masses, they set their
wings like hawks and spiral upward on the rising air. Once they reach the top of the thermal, they begin to
glide. The glide ratio is about 3.5 to 1, meaning that for every 3.5 feet forward they loose about a foot in
altitude. By flapping their wings 2-3 times every 20-30 feet, they can extend the glide until they reach
another thermal. Thermals frequently carry monarchs to 1,000 feet, and some have been sighted a mile or
more above the ground. This behavior conserves energy and saves their muscles and fragile wings. Most
monarchs arriving at the overwintering sites are in remarkably good condition despite flights of up to
2,000 miles. As monarchs move across the continent, they encounter relatively flat forests, farmlands and
grasslands, as well as mountain ranges and large lakes. They deal with differences in topography in a
variety of ways. When they encounter large lakes, if the weather isn’t favorable, monarchs may
accumulate for days waiting for favorable winds that will aid their passage. In mountainous areas they
often ride the thermals that rise along the ridges, frequently taking the same pathways as migrating
hawks. In relatively flat areas they move to the southwest in directions that are appropriate for the latitude
and longitude in which they find themselves. As the migration progresses, monarchs from as far east as
New Brunswick and Maine and as far west as the front range of the Rockies converge on Texas, in effect
funneling down to a 50-mile wide gap of cool river valleys between Eagle Pass, Texas, and Del Rio,
Texas, their last stops before moving into the mountains of northern Mexico. Once in Mexico, monarchs
tend to follow the mountains to the general area of the overwintering sites. Once they reach the
overwintering area at the end of October, they begin to cluster on the oyamel fir trees on the ridge tops.
With each day, as more monarchs arrive through November, the clusters in the oyamels become larger
while moving toward more protected sites below the ridges. Distinct clusters are called colonies and some
of these colonies can cover many acres with up to 25 million butterflies per acre. These colonies persist
from November to March with some monarchs beginning to move northward by the end of February. The
“Flight of the Butterflies” is about Dr. Fred Urquhart’s quest to discover where and how monarchs
overwintered. It is likely that what he found exceeded his expectations. In the film, he is seen with his
map, filled with known monarch migration paths. These lines represent paths that are known to contain

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33
thousands of butterflies, where there were numerous monarch sightings. Two frequently asked questions
about monarchs are “Why migrate?” and “Why migrate to these specific sites in Mexico?” Monarchs are
not winter-hardy. Their resistance to freezing is minimal. To survive from one season to the next, they
migrate to oyamel fir forests above 10,500 feet, a cool habitat where the daytime temperatures seldom
exceed 65°F and the nighttime lows are seldom below freezing. The forest is a blanket that protects the
monarchs. Monarchs remain relatively inactive through the winter, surviving by converting fats stored in
the fall to blood sugars necessary to keep their bodies functioning. The monarch colonies are located in a
relatively small area at 19.5 north latitude, but why this latitude (and longitude), and why these same
locations in the forest each year to form colonies? Scientists don’t have answers to these questions –but
there is no lack of speculation and interest. The beauty of monarch butterfly lies in it’s strict follow up in
the flight schedule. A migratory monarch probably maintain a modest flight schedule, flying three to five
5 hours and advancing 30-50 miles to the SW when weather conditions favour flight.
Table- 1: Flight Schedule of monarch butterfly, Danaus plexippus plexippus (L).
Day Time
Monarch Schedule
9:00 a.m. – 9:45 a.m.
Sun bath. Warmed by the Sun hitting the overnight roost site
9:00 a.m. – 9:45 a.m.
Visit to flower for nectar
10:00 a.m. – 4.30 p.m.
The migratory flight. Stopping periodically for 10-15 minute feeding
episodes
4:30 p.m. – 5:00 p.m.
Stops migratory flight and feeds for at least 30 minutes on flower nectar.
5:00 p.m. – 5:30 p.m.
Searches for overnight roosting site, preferably one that is well sheltered
with many other monarchs
5:00 p.m. – 6:00 p.m.
Settles on roost for the remainder of the night, converts sugars from last
nectar
feeding to fats and blood sugar needed for the next day's flight
6:30 p.m. – 9:00 a.m.
(Next Morning)
Sleep

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34
Fig.1: Adult Monarch butterfly, Danaus plexippus (L).
This scenario is based on the flight of a late season monarch that averaged 61 miles per day from
North Carolina to Austin, Texas, and probably required close to sixhours of flight per day. Monarchs
average about 11mph with powered flight –slower when gliding and soaring. The Monarch butterfly
migrates for 2 reasons. They can not withstand freezing weather in the northern and central continental
climates in the winter. Also, the larval food plants do not grow in their winter overwintering sites, so the
spring generation must fly back north to places where the plants are plentiful.
Nymphalidae Butterfly Diversity of Mayureshwar Wildlife Sanctuary of Baramati Tehsil in Pune
district (India):
The Nymphalidae is a family of several thousand species found in all zoogeographical regions of the
world. Most are medium or large in size, but the family is highly variable given that it also includes the
Satyrinae, a subfamily that has been designated as a family in its own right in earlier classifications. The
forelegs in both sexes are vestigial and useless for walking. In the male, there are typically only 2 tarsal
joints and the legs have a brush-like appearance, resulting in a common name for this family - the "brush-
footed butterflies". The female foreleg has 4 tarsal joints which, when compared with the male, provides a
mechanism of determining the sex of the adult. The midleg and hindleg are normal in both sexes and both
tibia and tarsus may have spines. Butterflies of Family: Nymphalidae recorded at Mayureshwar Wildlife
Sanctuary of Baramati Tehsil Dist. Pune (India) are listed in table-2 and Fig.2.
Table 2. Butterflies of Family: Nymphalidae recorded at Mayureshwar Wildlife Sanctuary of
Baramati Tehsil Dist. Pune (India).
Serial
No.
Common
Name
Scientific Name
Food Plants
preferred by
Larval Stages
Relative
Abundance
1.
Dark Blue
Tiger
Tirumala
septentrionis Butler.
Ageratum
conyzoides,
Uncommon

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35
Vallaris spp.
2.
Plain Tiger
Danaus chrysippus
L.
Frerea spp.,
Calotropis sp.
Very
Common
3.
Striped Tiger
Danaus genutia
Cramer.
Uncommon
4.
Glassy Tiger
Parantica aglea
Stoll.
Calotropis sp.
5.
Chocolate
Tiger
Parantica melaneus
Cramer.
Rare
6.
Striped Blue
Crow
Euploea mulciber
Cramer.
Ficus sp.
Uncommon
7.
Common Crow
Euploea core
Cramer.
Ficus sp., Nerium
sp.
Common
8.
Common
Duffer
Discophora sondaica
Boisduval.
Dendrocalamus
sp.
Common
9.
Common
Evening Brown
Melanitis leda L.
Panicum spp.,
Sorghum spp.
Very
Common
10.
Dark Evening
Brown
Melanitis phedima
Cramer.
Rare
11.
Common
palmfly
Elymnias
hypermnestra L.
Calamus spp.,
Areca spp.
Very
Common
12.
Common
Bushbrown
Mycalesis perseus
Fabricius.
Grasses
Very
Common
13.
Dark-Brand
Bushbrown
Mycalesis mineus L.
Grasses
Common
14.
Common
Fourring
Ypthima huebneri
Kirby.
Grasses
Common
15.
Common
Fivering
Ypthima baldus
Fabricius.
Common
16.
Vagrant
Vagrans egista
Cramer.
Uncommon
17.
Common
Leopard
Phalanta phalantha
Drury.
Flacourtia spp.
Very
Common
18.
Commander
Moduza procris
Cramer.
Mussaenda
frondosa
Uncommon
19.
Common
Sergeant
Athyma perius L.
Glochidion sp.
Common
20.
Colour
Sergeant
Athyma nefte
Cramer.
Glochidion sp.
Uncommon
21.
Common
Lascar
Pantoporia hordonia
Stoll.
Acacia spp.
Uncommon

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36
22.
Common Sailer
Neptis hylas L.
Bombax sp.
Very
Common
23.
Short-Banded
Sailer
Phaedyma columella
Cramer.
Dalbergia sp.
Uncommon
24.
Clipper
Parthenos sylvia
Cramer.
Very Rare
25.
Common
Baron
Euthalia aconthea
Cramer.
Mangifera indica
L.
Uncommon
26.
Plain Earl
Tanaecia jahnu
Moore.
Rare
27.
Archduke
Lexias pardalis
Moore.
Garcinia sp.
Very Rare
28.
Common
Castor
Ariadne merione
Cramer.
Ricinus
communis
Common
29.
Grey Pansy
Junonia atlites L.
Barleria sp.
Very
Common
30.
Peacock Pansy
Junonia almanac L.
Barleria sp.
Very
Common
31.
Yellow Pansy
Junonia hierta
Fabricius.
Barleria sp.
Very
Common
32.
Lemon Pansy
Junonia lemonias L.
Barleria sp.
Common
33.
Chocolate
Pansy
Junonia iphita
Cramer.
Uncommon
34.
Great Eggfly
Hypolimnas bolina
L.
Hibiscus sp.
Common
Fig. / Plate – 2 : Butterflies of Family: Nymphalidae recorded at Mayureshwar Wildlife Sanctuary
of Baramati Tehsil Dist. Pune (India).
Dark Blue Tiger,
Tirumala septentrionis
Butler.
Plain Tiger, Danaus
chrysippus L.
Striped Tiger, Danaus
genutia Cramer.
Glassy Tiger,
Parantica aglea Stoll.

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37
C
hocolate Tiger,
Parantica melaneus
Cramer.
Striped Blue Crow,
Euploea mulciber
Cramer.
Common Crow,
Euploea core Cramer.
Common Duffer,
Discophora sondaica
Boisduval.
Common Evening
Brown, Melanitis leda
L.
Dark Evening Brown,
Melanitis phedima
Cramer.
Common palmfly,
Elymnias
hypermnestra L.
Dark-Brand
Bushbrown, Mycalesis
mineus L.
Common Fourring,
Ypthima huebneri
Kirby.
Common Fivering,
Ypthima baldus
Fabricius.
Vagrant, Vagrans
egista Cramer.
Common Leopard,
Phalanta phalantha
Drury.
Commander, Moduza
procris Cramer.
Common Sergeant,
Athyma perius L.
Colour Sergeant,
Athyma nefte Cramer.

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38
Common Lascar,
Pantoporia hordonia
Stoll.
Common Sailer, Neptis
hylas L.
Short-Banded Sailer,
Phaedyma columella
Cramer.
Clipper, Parthenos
sylvia Cramer.
Common Baron,
Euthalia aconthea
Cramer.
Plain Earl, Tanaecia
jahnu Moore.
Archduke, Lexias
pardalis Moore.
Common Castor
butterfly, Ariadne
merione L.
Grey Pansy, Junonia
atlites L.
Peacock Pansy,
Junonia almanac L.
Yellow Pansy,
Junonia hierta
Fabricius.
Lemon Pansy,
Junonia lemonias L.
This is a large and diverse family that includes the typical Nymphalinae (brush-footed butterflies,
admirals, checkerspots), Libytheinae (snout butterflies), Satyrinae (wood nymphs, ringlets), Heliconiinae
(long wings, fritillaries), Morphinae (morpho and owl butterflies), and Danainae (milkweed and
glasswing butterflies). All possess three longitudinal ridges (carinae) on the ventral surface of the
antennae that are unique in Lepidoptera, and the forelegs of males are reduced or modified (less so in
Libytheinae), usually lacking claws and nonfunctional for walking. Adults are small to very large (FW
usually 10–50 mm, ranging to 75 mm in tropical Morpho and Caligo), mostly broad winged except in
Heliconinae, and usually brightly colored, often with orange, black, and white dominating, but mostly
brown and tan in Satyrinae. Many tropical nymphalids are involved in mimicry complexes, either as
models (Danainae, Heliconiinae) or as mimics of them or other distasteful butterflies and moths and/or
they benefit in both roles. Glasswing butterflies (Ithomiini) live primarily in deep shade of tropical forests
and have sparsely scaled areas or transparent wings, with subtle color patterns, while owl butterflies fly at

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39
dusk. They, morphos, and satyrines have conspicuous eyelike spots near the margins of the wing
undersides, presumably confusing would-be predators or diverting their attacks from the body. The larvae
are cylindrical caterpillars with full complement of abdominal prolegs, but there are diverse
modifications, e.g., densely spinose or with dorsal projections (verrucae) that are spinose (Nymphalinae),
smooth with filaments (Danainae), smooth with bifid caudal segment (Satyrinae), pubescent with hair
tufts and usually bifid caudally (Morphinae). The pupa hangs head downward, attached by a cremaster,
without a silken girdle. The larvae feed on a diverse array of flowering plants, with considerable
specialization within subfamilies: Morphinae and Satyrinae almost exclusively on monocots, including
Arecaceae, Bromeliadaceae, Helioconiacae, and Musaceae (a few species are pests on bananas) in the
tropics, mostly Poaceae and Cyperaceae in the Holarctic, with 2 genera on Selaginellaceae; other
nymphalids eat mostly dicot angiosperms, often specializing on plants with toxic chemicals (e.g.,
Heliconiinae on Flacourtiaceae, Passifloraceae, Urticaceae, Violaceae) or latex-producing plants
(Danainae on Apocynaceae, Asclepiadaceae, Moraceae).
Keith Brown suggested that feeding on Solanaceae was an important event in the diversification of
Ithomiini butterflies (Brown, 1987). Ithomiini butterflies are exclusively Neotropical and most species
feed on Solanaceae hostplants during larval stage (Willmott and Freitas, 2006). Optimizations of the
evolution of hostplant use on phylogenies evidence a probable shift from Apocynaceae to Solanaceae in
the ancestor of the tribe (Willmott and Freitas, 2006; Brower, et al, 2006). Fordyce (2010) found that the
Gamma statistics, a LTT plot of an Ithomiini phylogeny and the fit of the density-dependent model of
diversification are consistent with a burst of diversification in Ithomiini following the shift from
Apocynaceae to Solanaceae.
The present attempt on butterflies of Family: Lymphalidae recorded at Mayureshwar Wildlife Sanctuary
of Baramati Tehsil Dist. Pune (India) investigated whether the strong signal for an increase in net
diversification rate for Ithomiini (found by MEDUSA) can be explained due to the use of Solanaceae
plants as hosts during larval stage. For this, we used a Bayesian approach (FitzJohn, et al, 2009) to test
whether the trait “feeding on Solanaceae” had any effect on the diversification of the group. The BiSSE
analysis through the attempt on butterflies of Family: Lymphalidae recorded at Mayureshwar Wildlife
Sanctuary of Baramati Tehsil Dist. Pune (India) extended to take into account missing taxa and
phylogenetic uncertainty, shows a significantly higher net diversification rate for Ithomiini taxa, which
can be attributed to the trait “feeding on Solanaceae host plants”. This result is in agreement with
previous findings using other statistical methods (Fordyce, 2010). Due to the fact that Ithomiini are
virtually the only nymphalids using Solanaceae as host plants, it is possible that the trait responsible for a
higher diversification of Ithomiini might not be the host plant character. As noted by Maddison et al.
(2007), the responsible trait might be a co-distributed character such as a trait related to the ability to
digest secondary metabolites. Solanaceae plants contain chemical compounds and it has been suggested
that the high diversity of Ithomiini is consistent with the “escape-and-radiate scenario” due to a shift onto
Solanaceae (Fordyce, 2010) and radiation scenarios among chemically different lineages of Solanaceae
plants (Fordyce, 2010; Brown, 1987). According to this theory, the shift from Apocynaceae to Solanaceae
allowed Ithomiini to invade newly available resources due to a possible key innovation that allowed them
cope with secondary metabolites of the new hosts. Additional studies are needed to identify the actual
enzymes that Ithomiini species might be using for detoxification of ingested food as they have been found
in other butterfly groups (Wheat, et al, 2007). The increase in diversification rate inferred by MEDUSA

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40
occurred after the probable shift from Apocynaceae to Solanaceae, as the Solanaceae feeders in the sub-
tribes Melinaeina and Mechanitina are not included in the diversification shift. The apparent conflicting
results from MEDUSA and BiSSE can be explained by the low species-richness of the subtribes
Melinaeina and Mechanitina compared to the other subtribes included in the shift (fifty-two versus two
hundred seventy-two species). It can be that MEDUSA is more conservative than BiSSE and is not
including Melinaeina and Mechanitina in the shift due to low species numbers. Although the Solanaceae
genera used by the Ithomiini clades are well known (Willmott and Freitas, 2006), the present attempt does
not have any understanding on the physiological routes involved in the detoxification of Solanaceae
compounds by the several lineages of Ithomiini. The present attempt can speculate that older lineages
exploiting a novel toxic resource (Willmott and Freitas, 2006; Wahlberg, et al, 2009) may be inefficient
in metabolizing plant toxins and that younger lineages are able to deal with toxins more efficiently, so
that host switching events within Solanaceae are possible, which can lead to higher diversification.
Studies in Papilio species have reported that detoxification enzymes can become more efficient in
metabolizing toxins than ancestral configurations of the proteins, providing more opportunities for
hostplant switches (Li, et al, 2009). This might be the reason why the basal Ithomiini subtribes
Melinaeina and Mechanitina are so species-poor and restricted to few Solanaceae hosts (Willmott and
Freitas, 2006), while recent subtribes are species-rich and have expanded their host range into several
Solanaceae lineages (Willmott and Freitas, 2006). It might be that the switch to feeding in Solanaceae
was an important event in the evolutionary history of Ithomiini, but the actual radiation occurred after
critical physiological changes (a probable key innovation) allowed efficient detoxification of Solanaceae
toxins.
The hypothesis entitled, “Diffuse Co-speciation” predicts almost identical ages of insects and their host
plants. The hypothesis entitled, “Resource Abundance-Dependent Diversity” and the hypothesis entitled,
“Escape-And-Radiate” postulate that, insects diversify after their host plants (Nyman, et al, 2012; Ehrlich
and Raven, 1964). Wheat et al. (2007) found strong evidence for a model of speciation congruent with
Ehrlich and Raven’s hypothesis in Pieridae butterflies due to, in addition to the identification of a key
innovation, a burst of diversification in glucosinolate-feeding taxa shortly afterwards (with a lag of circa
10 MY). According to a recent dated phylogeny of the Angiosperms (Bell, et al, 2010), the family
Solanaceae split from its sister group about 59 (49–68) MYA and diversification started (crown group
age) around 37 (29–47) MYA. Wahlberg et al. (2009) give the ages for origin and diversification for
Ithomiini at 45 (39–53) and 37 (32–43) MYA, respectively. Thus, current evidence shows that Solanaceae
and Ithomiini might have diversified around the same time, during the Late Eocene and Oligocene, and
this would be congruent with the diffuse co-speciation hypothesis.
Butterflies of Danaini Tribe of Nymphalidae
The Danaini are a tribe of brush-footed butterflies (family Nymphalidae). Their type genus Danaus
contains the well-known monarch butterfly (D. plexippus) and is also the type genus of the tribe's
subfamily, the milkweed butterflies (Danainae). The Danaini do not have a fixed colloquial name for the
entire tribe, but in particular for subtribe Danaina the term tiger butterflies is occasionally used in
reference to the numerous species in several genera. MultiMEDUSA approach in present attempt showed
a significant slowdown in net diversification rate in the sub-tribe Danaina of the Danini. Both Danaina
and the sister clade Euploeina feed mainly on Apocynaceae and thus a host plant shift should not be

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41
responsible for the observed slowdown of diversification in the Danaina. As expected, BiSSE analysis of
Apocynaceae feeders shows that there is no effect of feeding on this plant family on the net
diversification rates of Nymphalidae lineages. Many of the Danaina are large, strong fliers, highly
migratory and involved in mimicry rings, including the best-known migratory butterfly, the monarch
(Danaus plexippus). The causes for a lower net diversification rate in the Danaina remains to be
investigated, but their great dispersal power might be involved in preventing allopatric speciation. It has
been found in highly vagile species in the nymphalid genus Vanessa that dispersal has homogenized
populations due to gene flow, as vagile species seem to be genetically homogeneous among populations
(Wahlberg and Rubinoff, 2011).
Butterflies of Satyrinae Tribe of Nymphalidae
The Satyrinae, the satyrines or satyrids, commonly known as the browns, are a subfamily of the
Nymphalidae (brush-footed butterflies). They were formerly considered a distinct family, Satyridae. This
group contains nearly half of the known diversity of brush-footed butterflies. The true number of the
Satyrinae species is estimated to exceed 2400. They are generally weak fliers and often shun bright
sunlight, preferring moist and semishaded habitats. The caterpillars feed chiefly on monocotyledonous
plants such as palms, grasses, and bamboos. The Morphinae are sometimes united with this group. The
taxonomy and systematics of the subfamily are under heavy revision. Much of the early pioneering work
of L. D. Miller (Savela, Markku, 2007) has helped significantly by creating some sort of order. Dyndirus
(Capronnier, 1874) is a satyrid incertae sedis. Other than this genus, according to the latest studies on the
classification of Nymphalidae, all satyrines have been assigned to one of the tribes, at least preliminarily
(Savela, Markku, 2007). Lineages in the diverse family Satyrinae radiated simultaneously with the
radiation of their main hostplant, grasses, during the climatic cooling in the Oligocene (Pena and
Wahlberg, 2008). Thus, it is somewhat surprising that part of Satyrinae were found to have accelerated
diversification in only 13% of the trees from the posterior distribution. Although this can be attributed to
low phylogenetic signal, the clade Satyrini is very robust and MEDUSA failed to identify any significant
accelerated net diversification rate for Satyrini (Wahlberg, et al, 2009). It appears that the radiation of
Satyrini as a whole was not remarkably fast and therefore not detected by MEDUSA, although it
estimated a diversification shift for Satyrini + its sister clade. This should be expected if the
diversification of Satyrini occurred in a stepwise manner, with pulses or bursts of diversification for
certain lineages but unlikely for the tribe Satyrini as a whole.
The present attempt on butterflies of Family: Lymphalidae recorded at Mayureshwar Wildlife Sanctuary
of Baramati Tehsil Dist. Pune (India) found that, even though MEDUSA estimated several diversification
shifts in the maximum clade credibility tree of Nymphalidae, only a few of these shifts were found in
more than 90% of the trees from the posterior distribution. In the literature, it is common practice that
conclusions are based on the shifts estimated on the maximum clade credibility tree. However, by using a
MultiMEDUSA approach, the attempt found that for this Nymphalidae dataset some of these shifts might
be greatly affected by phylogenetic uncertainty. Moreover, some of these shifts can be recovered either as
increases or decreases in net diversification rate depending on the tree from the posterior distribution that
was used for analysis. The method MEDUSA appears to be sensitive to the number of nodes with high
posterior probability and width of age confidence intervals. For data of present attempt, it would be
necessary to obtain a posterior distribution of trees with no conflicting topology, and very similar

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42
estimated ages for nodes in order to consistently recover most of the diversification shifts on the posterior
distribution of trees that were inferred by MEDUSA on the MCC tree.
Conclusion
In a nut shell, let us conclude the journey. The family is known as the Brush-Footed butterflies because
the forelegs of the adults are small and hairy, resembling tiny brushes, and are not used for walking. At
least 150 species of Nymphalids can be found in North America, with 47 occurring in Idaho. These
include the Fritillaries (Subfamily Heliconiinae), the true Brush-Foots (Subfamily Nymphalinae), and the
Admirals and Relatives (Subfamily Limenitidinae). The butterflies in this family vary considerably in
their appearance, but generally can be characterized by the following, in addition to their reduced
forelegs:
1. Medium to large in size and brightly and/or uniquely marked.
2. The pattern of wing veins of the forewing is unique.
3. The rigid antennae are tipped with little knobs, called clubs.
Interesting traits demonstrated by some members of this family include lengthy migrations, territoriality,
and the ability to overwinter as adults. Eggs vary in shape and in their arrangement on the plant.
Caterpillars vary considerably in their appearance, but are often hairy or spiny. Pupae have a cremaster
from which they are suspended upside down, but have no silk girdle and form no cocoon. Caterpillars are
the typical overwintering stage, although the adults of some species may overwinter as well.
Utility of DNA barcoding may have a excellent dimension of diversity of butterflies in the Indian
sanctuaries (like Mayureshwar Wildlife Sanctuary of Baramati Tehsil Dist. Pune India). This method
should be explored for application in various fields including biodiversity, conservation biology, ecology,
population studies, phylogeographic, etc. It is more a novel and a rapid method for species description
and identification put forward based on the DNA sequences and it undoubtedly serves as a
complementary approach for the existing traditional taxonomical practices. Although the conventional
morphology-based method of species identification is available, it is on a slow track due to many reasons
such as inaccessibility of important type material for comparison, unavailability of old literature,
decreasing number of specialists and very importantly, negligible number of students entering in this
field.
Conclusion
With around 6,000 species, this Nymphalidae is the largest family of butterflies. Many species, like the
monarch butterfly, are very colorful. Butterflies in this family are called brush-foots because of their tiny
forelegs. Their front two legs look more like brushes than feet and are not used for walking. Many species
in this family have some brown or orange on their wings and veins on their forewings. Most species have
long antennae with rounded clubs at the end. Most of the caterpillars in this family are colorful and
covered with spines or hair. Traditionally the family Nymphalidae as now defined has been treated as a
number of families, including the satyrs (Satyridae), monarchs (Danaidae), heliconians (Heliconiidae),
and snouts (Libytheidae). More recent concepts of phylogenetic analysis and classification have resulted
in these groups being treated as subfamilies of the Nymphalidae. By this revised definition, the
Nymphalidae contains about 220 species in North America and 102 in Canada.The forelegs of
nymphalids are reduced in size, usually in both sexes (not in Libytheinae females), and covered with long

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43
hairs, so as to resemble a brush. They are so small as to be practically useless; nymphalids perch and walk
using four legs only, with the front pair held up under the "face".
Unlike the other families, nymphalids vary greatly in appearance in larval, pupal, and adult stages. Adults
range from small to large, with most being of medium size. In North America, the most common colour is
orange brown but there are many exceptions to this. Some species are powerful fliers (Polygonia) or
migrants (Vanessa), while others (Euphydryas) are very weak fliers living in small, highly localized
colonies.
Acknowledgements
The support received from Department of Zoology, Savitribai Phule Pune University, Pune Maharashtra ,
India and International Science Community Association, Indore deserve appreciations and exert a grand
salutary influence.
References
Bingham CT (1907) The Fauna of British India, including Ceylon and Burma, Butterflies Vol. 2. Taylor
and Francis, London, 480 pp. https://doi.org/10.1038/115115c0
Bingham, C.T. (1905). The Fauna of British India including Ceylon and Burma – Butterflies (Vol 1).
London: Taylor and Francis. p. 519. Retrieved 7 November 2010.
Bingham, C.T. (1907). The Fauna of British India, Including Ceylon and Burma. II (1st ed.).
London: Taylor and Francis, Ltd.
Vane‐Wright, R.I. 1978. Ecological and behavioural origins of diversity in butterflies. In: Diversity of
Insect Faunas. Symposia of the Royal Entomological Society of London: No. 9 (L. A. Mound &
N. Waloff, eds), pp. 56–70. Blackwell Scientific Publications, Oxford.
Brakefield PM, Beldade P, Zwaan BJ. The African butterfly Bicyclus anynana: a model for evolutionary
genetics and evolutionary developmental biology. Cold Spring Harb Prot. 2009;2009(5): pdb–
emo122.
Brower AVZ, Freitas AVL, Lee MM, Silva-Brandão KL, Whinnett A, Willmott KR. Phylogenetic
relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitochondrial
and two nuclear gene regions. Syst Entomol. 2006 Mar;31(2): 288–301.
Brower AVZ, Wahlberg N, Ogawa JR, Boppré M, Vane-Wright RI. Phylogenetic relationships among
genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA
sequences. Syst Biodivers. 2010;8(1): 75–89.
Brower, Andrew V. Z. 2007. Riodinidae Grote 1895. Metalmarks, Erycinidae Swainson 1827 (see
nomenclature section). Version 19 May 2007 [4] in The Tree of Life Web Project, [5]
Brower, Andrew V. Z. 2007. Riodinidae Grote 1895. Metalmarks, Erycinidae Swainson 1827 (see
nomenclature section). Version 19 May 2007 [4] in The Tree of Life Web Project,

International Academic Journal of Science and Engineering,
Vol. 5, No. 4, pp. 24-47.
44
Brower, Andrew V. Z. 2008. Lycaenidae. Version 25 April 2008 (temporary). [2] in The Tree of Life
Web Project, [3]
Brower, Andrew V.Z. & Warren, Andrew (2006): The higher classification of the Hesperiidae
(Lepidoptera: Hesperioidea) Full Article. Retrieved 2012-OCT-26.
Brower, Andrew V.Z. & Warren, Andrew (2008): Tree of Life Web Project – Hesperiidae. Version of
2008-APR-07. Retrieved 2009-DEC-24.
Brower, Andrew V.Z. & Warren, Andrew (2009): Tree of Life Web Project – Euschemoninae. Version of
2009-JUN-11. Retrieved 2009-DEC-24.
Brown KS Jr. Chemistry at the Solanaceae/Ithomiinae Interface. Ann Mo Bot Gard. 1987;74(2): 359–397.
Brown KS, Freitas AVL (2000) Atlantic forest butterflies: indicators for landscape conservation.
Biotropica 32: 934–956
Brown KS. The conservation of insects and their habitats, In: Conservation of Neotropical Environment s:
Insects as Indicators 15th Symposium of the Royal Entomological Society 1991, 350-403.
Brown, J. W., H. G. Real, and D. K. Faulkner. 1992. Butterflies of Baja California. Faunal Survey,
Natural History, Conservation Biology. The Lepidoptera Research Foundation, Inc.
Bell CD, Soltis DE, Soltis PS. The age and diversification of the angiosperms re-revisited. Am J Bot.
2010 Aug;97(8): 1296–1303. pmid:21616882
Alfaro ME, Santini F, Brock C, Alamillo H, Dornburg A, Rabosky DL, et al. Nine exceptional radiations
plus high turnover explain species diversity in jawed vertebrates. P Natl Acad Sci USA. 2009
Aug;106(32): 13410–13414.
Dingle, Hugh; Drake, V. Alistair (2007). "What is migration?". BioScience. 57 (2): 113–121.
doi:10.1641/B570206.
Ehrlich PR, Raven PH. Butterflies and plants: A study in coevolution. Evolution. 1964;18: 586–608.
Ehrlich, P. R. 1958. The comparative morphology, phylogeny and higher classification of the butterflies
(Lepidoptera: Papilionoidea). Univ. Kans. Sci. Bull. 39: 306-370.
Ehrlich, P.R. & Raven, P.H. 1965. Butterflies and plants: a study in coevolution. Evolution 18: 586–608.
Elias M, Joron M, Willmott KR, Silva-Brandão KL, Kaiser V, Arias CF, et al. Out of the Andes: Patterns
of diversification in clearwing butterflies. Mol Ecol. 2009 Apr;18(8): 1716–1729.
pmid:19386035.
Fordyce JA. Host shifts and evolutionary radiations of butterflies. P Roy Soc B. 2010 Dec;277(1701):
3735–3743.
FitzJohn RG. Diversitree: comparative phylogenetic analyses of diversification in R. Method Ecol Evol.
2012 Dec;3(6): 1084–1092.
FitzJohn RG, Maddison WP, Otto SP. Estimating trait-dependent speciation and extinction rates from
incompletely resolved phylogenies. Syst Biol. 2009 Dec;58(6): 595–611. pmid:20525612

International Academic Journal of Science and Engineering,
Vol. 5, No. 4, pp. 24-47.
45
Ferrer-Paris JR, Sánchez-Mercado A, Viloria ÁL, Donaldson J. Congruence and Diversity of Butterfly-
Host Plant Associations at Higher Taxonomic Levels. PLoS ONE. 2013;8(5): e63570.
pmid:23717448
Heikkil¨a M, Kaila L, Mutanen M, Peña C, Wahlberg N. Cretaceous origin and repeated tertiary
diversification of the redefined butterflies. P Roy Soc B. 2012 Mar;279(1731): 1093–9.
Heikkilä M, Kaila L, Mutanen M, Peña C, Wahlberg N (2011) Cretaceous origin and repeated tertiary
diversification of the redefined butterflies. Proc R Soc B Biol Sci 279: 1093–1099
Heikkilä, M., Kaila, L., Mutanen, M., Peña, C., & Wahlberg, N. (2012). Cretaceous origin and repeated
tertiary diversification of the redefined butterflies. Proceedings of the Royal Society B:
Biological Sciences, 279(1731), 1093-1099.
Heikkilä, M.; Kaila, L.; Mutanen, M.; Peña, C.; Wahlberg, N. (2012). "Cretaceous Origin and Repeated
Tertiary Diversification of the Redefined Butterflies". Proceedings of the Royal Society B:
Biological Sciences. 279 (1731): 1093–1099. doi:10.1098/rspb.2011.1430.
Joron M, Jiggins C, Papanicolaou A, McMillan W. Heliconius wing patterns: an evo-devo model for
understanding phenotypic diversity. Heredity. 2006;97(3): 157–167. pmid:16835591
Li W, Schuler MA, Berenbaum MR. Diversification of furanocoumarin-metabolizing cytochrome P450
monooxygenases in two papilionids: Specificity and substrate encounter rate. P Natl Acad Sci
USA. 2003;100(Suppl 2): 14593–14598.
Maddison DR, Schulz KS. The Tree of Life Web Project; 2007. Accessed 16 June 2014. Available:
http://tolweb.org.
Maddison WP, Maddison DR (2001) Mesquite, a modular system for evolutionary analysis, Available:
http://mesquiteproject.org.
Maddison WP, Maddison DR. 2011. Mesquite 2.75: A modular system for evolutionary analysis.
Maddison WP, Midford PE, Otto SP. Estimating a binary character’s effect on speciation and extinction.
Syst Biol. 2007 Oct;56(5): 701–710. pmid:17849325
Maddison, W.P. & Maddison, D.R. 1999. MacClade: Analysis of Phylogeny and Character Evolution,
version 3.08a for Macintosh. Sinauer Associates, Sunderland, MA.
Nylin S, Slove J, Janz N. Host plant utilization, host range oscillations and diversification in Nymphalid
butterflies: A phylogenetic investigation. Evolution. 2014;68(1): 105–124. pmid:24372598
Nyman T, Linder HP, Peña C, Malm T, Wahlberg N. Climate-driven diversity dynamics in plants and
plant-feeding insects. Ecol Lett. 2012 Aug;15(8): 889–898
Peña C, Espeland M (2015) Diversity Dynamics in Nymphalidae Butterflies: Effect of Phylogenetic
Uncertainty on Diversification Rate Shift Estimates. PLoS ONE 10(4): e0120928.
https://doi.org/10.1371/journal.pone.0120928

International Academic Journal of Science and Engineering,
Vol. 5, No. 4, pp. 24-47.
46
Peña C, Nylin S, Freitas AVL, Wahlberg N. Biogeographic history of the butterfly subtribe Euptychiina
(Lepidoptera, Nymphalidae, Satyrinae). Zool Scr. 2010 May;39(3): 243–258.
Peña C, Nylin S, Wahlberg N. The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge
for phylogenetic methods. Zool J Linn Soc. 2011 Jan;161(1): 64–87.
Peña C, Wahlberg N, Weingartner E, Kodandaramaiah U, Nylin S, et al. (2006) Higher level phylogeny
of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Mol
Phylogenet Evol 40: 29–49
Peña C, Wahlberg N, Weingartner E, Kodandaramaiah U, Nylin S, Freitas AVL, et al. Higher level
phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data.
Mol Phylogenet Evol. 2006 Jul;40(1): 29–49. pmid:16563805
Peña C, Wahlberg N. Prehistorical climate change increased diversification of a group of butterflies. Biol
Lett. 2008 Jun;4(3): 274–278. pmid:18364308
Peña, L.E. & Ugarté, A.J. 1996. Las Mariposas de Chile. Editorial Universitaria, Santiago.
Silva, S., Servia, M. J., Vieira-Lanero, R., & Cobo, F. (2013). Downstream migration and hematophagous
feeding of newly metamorphosed sea lampreys (Petromyzon marinus Linnaeus, 1758).
Hydrobiologia, 700(1), 277–286.
Silva, S., Servia, M. J., Vieira-Lanero, R., Barca, S., & Cobo, F. (2013). Life cycle of the sea lamprey
Petromyzon marinus: duration of and growth in the marine life stage. Aquatic Biology, 18(1),
59–62.
Silva-Brandão KL, Wahlberg N, Francini RB, Azeredo-Espin AML, Brown KS Jr, Paluch M, et al.
Phylogenetic relationships of butterflies of the tribe Acraeini (Lepidoptera, Nymphalidae,
Heliconiinae) and the evolution of host plant use. Mol Phylogenet Evol. 2008 Feb;46(2): 515–
31. pmid:18191590
Vitthalrao B. Khyade & Sharad G. Jagtap (2017). Butterflies - Diversity in Mayureshwar Wildlife
Sanctuary of Baramati Tehsil in Pune district (India). Journal of Entomology and Zoology
Studies 2017; 5(2): 443-457.E-ISSN: 2320-7078 P-ISSN: 2349-6800 www.entomoljournal.com
Vitthalrao B. Khyade (2017). A Study of Review on Wildlife. International Academic Journal of
Innovative Research Vol. 4, No. 1, 2017, pp. 32-40.ISSN 2454-390X www.iaiest.com
Vitthalrao B. Khyade and Sharad G. Jagtap (2016).Diversity of butterflies (Order: Lepidoptera) in
MayureshwarWildlife Sanctuary of Baramati Tehsil Dist. Pune(India).International Academic
Journal of Innovative Research. Vol.3 No. 11:40 – 67. ISSN 2454-390X. www.iaiest.com
Vitthalrao B. Khyade(2016). Camouflage: the Dead – Leaf Butterfly, Kallimainachus(L). International
Academic Journal of Innovative Research Vol. 3, No. 12, 2016, pp. 13-21. ISSN 2454-2474.
http://iaiest.com/journals/international-academic-journal-of-innovative-research/volume-3-issue-
12-december-2016/

International Academic Journal of Science and Engineering,
Vol. 5, No. 4, pp. 24-47.
47
Wahlberg N, Leneveu J, Kodandaramaiah U, Peña C, Nylin S, Freitas AVL, et al. Nymphalid butterflies
diversify following near demise at the Cretaceous/Tertiary boundary. P Roy Soc B. 2009
Sep;276(1677): 4295–4302.
Wahlberg N, Rubinoff D. Vagility across Vanessa (Lepidoptera: Nymphalidae): mobility in butterfly
species does not inhibit the formation and persistence of isolated sister taxa. Syst Entomol. 2011
Apr;36(2): 362–370.
Wahlberg N, Wheat CW, Peña C (2013) Timing and patterns in the taxonomic diversification of
Lepidoptera (butterflies and moths). PLoS ONE 8: e80875
Wahlberg, N. 2001. The phylogenetics and biochemistry of host‐plant specialization in Melitaeine
butterflies (Lepidoptera: Nymphalidae). Evolution 55: 522–537.
Wahlberg, N., Braby, M.F., Brower, A.V.Z., De Jong, R., Lee, M.‐M., Nylin, S., Pierce, N.E., Sperling,
F.A.H., Vila, R., Warren, A.D. & Zakharov, E. 2005. Synergistic effects of combining
morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proc.
R. Soc. Lond. B 272: 1577–1586.
Wheat CW, Vogel H, Wittstock U, Braby MB, Underwood D, Mitchell-Olds T. The genetic basis of a
plant-insect coevolutionary key innovation. P Natl Acad Sci USA. 2007;104: 20427–20431.
Wigglesworth, V. (1982). "Carrington Bonsor Williams. 7 October 1889 – 12 July 1981". Biographical
Memoirs of Fellows of the Royal Society. 28: 666. doi:10.1098/rsbm.1982.0025.
JSTOR 769914.
Willmott KR, Freitas AVL. Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae):
classification, patterns of larval hostplant colonization and diversification. Cladistics. 2006
Aug;22(4): 297–368.
Sonia Altizer, Keith A. Hobson, Andrew K. Davis, Jacobus C. De Roode, Leonard I. Wassenaar. Do
Healthy Monarchs Migrate Farther? Tracking Natal Origins of Parasitized vs. Uninfected
Monarch Butterflies Overwintering in Mexico. PLOS ONE, 2015; 10 (11): e0141371 DOI:
10.1371/journal.pone.0141371 DOI: 10.1371/journal.pone.0141371.