Passenger Pigeon

Passenger pigeon
The passenger pigeon or wild pigeon (Ectopistes migra- of the greatest and most senseless extinctions induced by
torius) is an extinct species of pigeon that was endemic humans.
to North America. Its common name is derived from
the French word passager, meaning passing by, due to
the migratory habits of the species. The scientic name 1 Taxonomy
also refers to its migratory characteristics. The morphologically similar mourning dove (Zenaida macroura) was
long thought to be its closest relative, and the two were
at times confused, but genetic analysis has shown that the
genus Patagioenas is more closely related to it than the
Zenaida doves.
The passenger pigeon was sexually dimorphic in size and
coloration. The male was 39 to 41 cm (15.4 to 16.1 in)
in length, mainly gray on the upperparts, lighter on the
underparts, with iridescent bronze feathers on the neck,
and black spots on the wings. The female was 38 to 40
cm (15.0 to 15.7 in), and was duller and browner than
the male overall. The juvenile was similar to the female, but without iridescence. It mainly inhabited the
deciduous forests of eastern North America and was also
recorded elsewhere, but bred primarily around the Great
Lakes. The pigeon migrated in enormous ocks, constantly searching for food, shelter, and breeding grounds,
and was once the most abundant bird in North America,
numbering around 3 to 5 billion at the height of its population. It was not always as abundant, and the population size uctuated rapidly over time. A very fast yer,
it could reach 100 km/h (62 mph). The bird fed mainly
on mast, as well as fruits and invertebrates. It practiced
communal roosting and communal breeding, and its extreme gregariousness may be linked with searching for
food and predator satiation.
Earliest published illustration of the species (a male), Mark

Catesby, 1731
Swedish naturalist Carl Linnaeus coined the binomial

name Columba macroura for both the mourning dove
and the passenger pigeon in the 1758 edition of his work
Systema Naturae (the starting point of biological nomenclature), wherein he appears to have considered the two
identical. This composite description cited accounts of
these birds in two pre-Linnean books. One of these
was Mark Catesby's description of the passenger pigeon,
which was published in his 17311743 work Natural History of Carolina, Florida and the Bahama Islands, which
referred to this bird as Palumbus migratorius, and was
accompanied by the earliest published illustration of the
species. Catesbys description was combined with the
1743 description of the mourning dove by George Edwards, who used the name C. macroura for that bird.
There is nothing to suggest Linnaeus ever saw specimens
of these birds himself, and his description is thought to
be fully derivative of these earlier accounts and their illustrations. In his 1766 edition of Systema Naturae, Linnaeus dropped the name C. macroura, and instead used
the name C. migratoria for the passenger pigeon, and C.
carolinensis for the mourning dove.[2][3][4] In the same
edition, Linnaeus also named C. canadensis, based on
Turtur canadensis, as used by Mathurin Jacques Brisson
in 1760. Brissons description was later shown to have
been based on a female passenger pigeon.[5]
Passenger pigeons were hunted by Native Americans, but

hunting intensied after the arrival of Europeans, particularly in the 19th century. Pigeon meat was commercialized as cheap food, resulting in hunting on a massive
scale for many decades. There were several other factors contributing to the decline and subsequent extinction
of the species, including shrinking of the large breeding
populations necessary for preservation of the species and
widespread deforestation which destroyed its habitat. A
slow decline between about 1800 and 1870 was followed
by a rapid decline between 1870 and 1890. The last conrmed wild bird is thought to have been shot in 1900.
The last captive birds were divided in three groups around
the turn of the 20th century, some of which were photographed alive. Martha, thought to be the last passenger pigeon, died on September 1, 1914, at the Cincinnati
Zoo. Eradication of the species has been described as one
1 TAXONOMY
In 1827 William John Swainson moved the passenger

pigeon from the genus Columba to the new monotypic
genus Ectopistes, due in part to the length of the wings and
the wedge shape of the tail.[6] In 1906 Outram Bangs suggested that because Linnaeus had wholly copied Catesbys
text when coining C. macroura, this name should apply to
the passenger pigeon, as E. macroura.[7] In 1918 Harry
C. Oberholser suggested that C. canadensis should take
precedence over C. migratoria (as E. canadensis), as it
appeared on an earlier page in Linnaeus book.[5] In 1952
Francis Hemming proposed that the International Commission on Zoological Nomenclature (ICZN) secure the
specic name macroura for the mourning dove, and the
name migratorius for the passenger pigeon, since this was
the intended use by the authors on whose work Linnaeus
had based his description.[4] This was accepted by the
ICZN, which used its plenary powers to designate the Band-tailed pigeon, a species in the related genus Patagioenas
species for the respective names in 1955.[8]
1.1
Evolution
The physically similar mourning dove is not closely related.
in an ancient DNA analysis for the rst time (in a paper focusing mainly on the dodo), and it was found to
be the sister taxon of the cuckoo-dove genus Macropygia.
The Zenaida doves were instead shown to be related to
the quail-doves of the genus Geotrygon and the Leptotila
doves.[13][14][15]
Stued male passenger pigeon, Field Museum of Natural History
The passenger pigeon was a member of the pigeon and

dove family, Columbidae. Its closest living relatives were
long thought to be the Zenaida doves, based on morphological grounds, particularly the physically similar
mourning dove (now Z. macroura).[9][10] It was even suggested that the mourning dove belonged to the genus Ectopistes and was listed as E. carolinensis by some authors,
including Thomas Mayo Brewer.[11] The passenger pigeon was supposedly descended from Zenaida pigeons
that had adapted to the woodlands on the plains of central
North America.[12] The passenger pigeon diered from
the species in the genus Zenaida in being larger, lacking a facial stripe, being sexually dimorphic, and having
iridescent neck feathers and a smaller clutch. In a 2002
study by American geneticist Beth Shapiro et al., museum specimens of the passenger pigeon were included
A more extensive 2010 study instead showed that the passenger pigeon was most closely related to the New World
Patagioenas pigeons, including the band-tailed pigeon (P.
fasciata) of western North America, which are related
to the Southeast Asian species in the genera Turacoena,
Macropygia and Reinwardtoena. This clade is also related
to the Columba and Streptopelia doves of the Old World
(collectively termed the typical pigeons and doves).
The authors of the study suggested that the ancestors of
the passenger pigeon may have colonized the New World
from South East Asia by ying across the Pacic Ocean,
or perhaps across Beringia in the north.[15] In a 2012
study, the nuclear DNA of the passenger pigeon was analyzed for the rst time, and its relationship with the Patagioenas pigeons was conrmed. In contrast to the 2010
study, these authors suggested that their results could indicate that the ancestors of the passenger pigeon and its Old
World relatives may have originated in the Neotropical
region of the New World.[14]
3
The cladogram below follows the 2012 DNA study showing the position of the passenger pigeon among its closest
relatives:[14]
DNA in old museum specimens is often degraded and
fragmentary, and passenger pigeon specimens have been
used in various studies to discover improved methods of
analyzing and assembling genomes from such material.
DNA samples are often taken from the toe pads of bird
skins in museums, as this can be done without causing
signicant damage to valuable specimens.[16][17] The passenger pigeon had no known subspecies.[9] Hybridization
occurred between the passenger pigeon and the Barbary
dove (Streptopelia risoria) in the aviary of Charles
Otis Whitman (who owned many of the last captive
birds around the turn of the 20th century, and kept
them with other pigeon species) but the ospring were
infertile.[12][18]
1.2
Etymology
The genus name, Ectopistes, translates as moving about

or wandering, while the specic name, migratorius, indicates its migratory habits.[19] The full binomial can thus
be translated as migratory wanderer. The English common name passenger pigeon derives from the French
word passager, which means to pass by in a eeting
manner.[20][21] While the pigeon was extant, the name
passenger pigeon was used interchangeably with wild
pigeon.[22] The bird also gained some less-frequently
used names, including blue pigeon, merne rouck pigeon,
wandering long-tailed dove, and wood pigeon. In the 18th
century, the passenger pigeon was known as tourte in New
France (in modern Canada), but to the French in Europe
it was known as tourtre. In modern French, the bird is
known as tourte voyageuse or pigeon migrateur, among
other names.[23] In the Native American Algonquian languages, the pigeon was called amimi by the Lenape,
omiimii by the Ojibwe, and mimia by the Kaskaskia
Illinois.[24][25][26] Other names in indigenous American
languages include ori'te in Mohawk, and putchee nashoba,
or lost dove, in Choctaw.[27] The Seneca people called
the pigeon jahgowa, meaning big bread, as it was a
source of food for their tribes.[28] Chief Simon Pokagon
of the Potawatomi stated that his people called the pigeon
O-me-me-wog, and that the Europeans did not adopt native names for the bird, as it reminded them of their domesticated pigeons, instead calling them wild pigeons,
as they called the native peoples wild men.[29]
Turnaround video of an adult male specimen at Naturalis Biodiversity Center
hindneck. On the sides of the neck and the upper mantle were iridescent display feathers that have variously
been described as being a bright bronze, violet or goldengreen, depending on the angle of the light. The upper
back and wings were a pale or slate gray tinged with olive
brown, that turned into grayish-brown on the lower wings.
The lower back and rump were a dark blue-gray that
became grayish-brown on the upper tail-covert feathers.
The greater and median wing-covert feathers were pale
gray, with a small number of irregular black spots near
the end. The primary and secondary feathers of the wing
were a blackish-brown with a narrow white edge on the
outer side of the secondaries. The two central tail feathers were brownish gray, and the rest were white.[20][31]
The tail pattern was distinctive as it had white outer edges
with blackish spots that were prominently displayed in
ight.[31] The lower throat and breast were richly pinkishrufous, grading into a paler pink further down, and into
white on the abdomen and undertail covert feathers. The
undertail coverts also had a few black spots. The bill was
black, while the feet and legs were a bright coral red. It
had a carmine-red iris surrounded by a narrow purplishred eye-ring.[31] The wing of the male measured 196215
mm (7.78.5 in), the tail 175210 mm (6.98.3 in), the
bill 1518 mm (0.590.71 in), and the tarsus was 2628
mm (1.01.1 in).[20]
Description
The passenger pigeon was sexually dimorphic in size and Turnaround video of an adult female specimen at Naturalis
coloration. It weighed between 260 and 340 g (9 and 12
oz).[30] The adult male was about 39 to 41 cm (15.4 to The adult female passenger pigeon was slightly smaller
16.1 in) in length.[31] It had a bluish-gray head, nape, and than the male at 38 to 40 cm (15.0 to 15.7 in) in length.
4
It was duller than the male overall, and was a grayishbrown on the forehead, crown, and nape down to the
scapulars, and the feathers on the sides of the neck had
less iridescence than those of the male. The lower throat
and breast were a bu-gray that developed into white on
the belly and undertail-coverts. It was browner on the
upperparts and paler bu brown and less rufous on the
underparts than the male. The wings, back, and tail were
similar in appearance to those of the male except that the
outer edges of the primary feathers were edged in bu
or rufous bu.[20][31] The wings had more spotting than
those of the male.[30] The tail was shorter than that of the
male, and the legs and feet were a paler red. The iris was
orange red, with a grayish blue, naked orbital ring. The
wing of the female was 180210 mm (7.18.3 in), the tail
150200 mm (5.97.9 in), the bill 1518 mm (0.590.71
in), and the tarsus was 2528 mm (0.981.10 in).[20]
Turnaround video of a juvenile female specimen at Naturalis
The juvenile passenger pigeon was similar in plumage to

the adult female, but lacked the spotting on the wings,
and was a darker brownish-gray on the head, neck, and
breast. The feathers on the wings had pale gray fringes
(also described as white tips), giving it a scaled look. The
secondaries were brownish-black with pale edges, and the
tertial feathers had a rufous wash. The primaries were
also edged with a rufous-brown color. The neck feathers had no iridescence. The legs and feet were dull red,
and the iris was brownish, and surrounded by a narrow
carmine ring.[20][31] The plumage of the sexes was similar during their rst year.[32]
Of the hundreds of surviving skins, only one appears to be
aberrant in color an adult female from the collection
of Walter Rothschild, Natural History Museum at Tring.
It is a washed brown on the upper parts, wing covert, secondary feathers, and tail (where it would otherwise have
been gray), and white on the primary feathers and underparts. The normally black spots are brown, and it is
pale gray on the head, lower back, and upper-tail covert
feathers, yet the iridescence is unaected. The brown
mutation is a result of a reduction in eumelanin, due to incomplete synthesis (oxidation) of this pigment. This sexlinked mutation is common in female wild birds, but it
is thought the white feathers of this specimen are instead
the result of bleaching due to exposure to sunlight.[33]
DESCRIPTION
Skeleton of a male bird, 1914
The passenger pigeon was physically adapted for speed,

endurance, and maneuverability in ight, and has been
described as having a streamlined version of the typical
pigeon shape, such as that of the generalized rock dove
(Columba livia). The wings were very long and pointed,
and measured 220 mm (8.7 in) from the wing-chord to the
primary feathers, and 120 mm (4.7 in) to the secondaries.
The tail, which accounted for much of its overall length,
was long and wedge-shaped (or graduated), with two central feathers longer than the rest. The body was slender
and narrow, and the head and neck were small.[20][34][35]
The internal anatomy of the passenger pigeon has rarely
been described. Robert W. Shufeldt found little to differentiate the birds osteology from that of other pigeons
when examining a male skeleton in 1914, but Julian
P. Hume noted several distinct features in a more detailed 2015 description. The pigeon had particularly large
breast muscles that indicate powerful ight (musculus
pectoralis major for downstroke and the smaller musculus
supracoracoideus for upstroke). The coracoid bone
(which connects the scapula, furcula, and sternum) was
large relative to the size of the bird, 33.4 mm (1.31
in), with straighter shafts and more robust articular ends
than in other pigeons. The furcula had a sharper Vshape and was more robust, with expanded articular ends.
The scapula was long, straight, and robust, and its distal
end was enlarged. The sternum was very large and robust compared to that of other pigeons; its keel was 25
mm (0.98 in) deep. The overlapping uncinate processes,
which stien the ribcage, were very well developed. The
wing bones (humerus, radius, ulna, carpometacarpus)
were short but robust compared to other pigeons. The leg
bones were similar to those of other pigeons.[35][36][37]
2.1 Vocalizations
The noise produced by ocks of passenger pigeons was
described as deafening, audible for miles away, and the
birds voice as loud, harsh, and unmusical. It was also described by some as clucks, twittering and cooing, and as
a series of low notes instead of actual song. The birds apparently made croaking noises when building nests, and
bell-like sounds when mating. During feeding, some indi-
Musical notes documenting male vocalizations, compiled by

Wallace Craig, 1911
Specimen in ying pose, Academy of Natural Sciences of Drexel
viduals would give alarm calls when facing a threat, and University
the rest of the ock would join the sound while taking
o.[20][38][39]
In 1911 American behavioral scientist Wallace Craig
published an account of the gestures and sounds of this
species as a series of descriptions and musical notations,
based on observation of C. O. Whitmans captive passenger pigeons in 1903. Craig compiled these records
to assist in identifying potential survivors in the wild (as
the physically similar mourning doves could otherwise be
mistaken for passenger pigeons), while noting this meager information was likely all that would be left on the
subject. According to Craig, one call was a simple harsh
keck that could be given twice in succession with a
pause in between. This was said to be used to attract the
attention of another pigeon. Another call was a more frequent and variable scolding. This sound was described as
kee-kee-kee-kee or tete! tete! tete!", and was used
to call either to its mate or towards other creatures it
considered to be enemies. One variant of this call, described as a long, drawn-out tweet, could be used to
call down a ock of passenger pigeons passing overhead,
which would then land in a nearby tree. Keeho was a
soft cooing that, while followed by louder keck notes
or scolding, was directed at the birds mate. A nesting
passenger pigeon would also give o a stream of at least
eight mixed notes that were both high and low in tone and
ended with keeho. Overall, female passenger pigeons
were quieter and called infrequently. Craig suggested that
the loud, strident voice and degenerated musicality was
the result of living in populous colonies where only the
loudest sounds could be heard.[38][40]
Distribution and habitat
The passenger pigeon was found across most of North

America east of the Rocky Mountains, from the Great
Plains to the Atlantic coast in the east, to the south of
Canada in the north, and the north of Mississippi in the
southern United States, coinciding with its primary habitat, the eastern deciduous forests. Within this range, it
constantly migrated in search of food and shelter. It is unclear if the birds favored particular trees and terrain, but
they were possibly not restricted to one type, as long as

their numbers could be supported.[20][30] It originally bred
from the southern parts of eastern and central Canada
south to eastern Kansas, Oklahoma, Mississippi, and
Georgia in the United States, but the primary breeding
range was in southern Ontario and the Great Lakes states
south through states north of the Appalachian Mountains.[41] Though the western forests were ecologically
similar to those in the east, these were occupied by bandtailed pigeons, which may have kept out the passenger
pigeons through competitive exclusion.[12]
The passenger pigeon wintered from Arkansas, Tennessee, and North Carolina south to Texas, the Gulf
Coast, and northern Florida, though ocks occasionally
wintered as far north as southern Pennsylvania and Connecticut. It preferred to winter in large swamps, particularly those with alder trees; if swamps were not available,
forested areas, particularly with pine trees, were favored
roosting sites. There were also sightings of passenger pigeons outside of its normal range, including in several
Western states, Bermuda, Cuba, and Mexico, particularly
during severe winters.[41][42][43] It has been suggested that
some of these extralimital records may be considered as
such more on the basis of the paucity of observers in what
was then unsettled country than on the actual extent of
wandering passenger pigeons, and that the bird may have
appeared anywhere on the continent except for the far
west.[30] There were also records of stragglers in Scotland,
Ireland, and France, although these birds may have been
escaped captives, or the records simply incorrect.[20][41]
More than 130 passenger pigeon fossils have been found
scattered across 25 states and provinces of the United
States, including in the La Brea Tar Pits of California.
These records date as far back as 100,000 years ago in
the Pleistocene era, during which the pigeons range extended to several western states that were not a part of
its modern range. The abundance of the species in these
regions and during this time is unknown.[41][44][45]
4 ECOLOGY AND BEHAVIOR
Ecology and behavior
with inconceivable velocity, mounted perpendicularly so as to resemble a vast column, and,

when high, were seen wheeling and twisting
within their continued lines, which then resembled the coils of a gigantic serpent... Before
sunset I reached Louisville, distant from Hardensburgh fty-ve miles. The Pigeons were
still passing in undiminished numbers and continued to do so for three days in succession.[46]
These ocks were frequently described as being so dense
that they blackened the sky and as having no sign of subdivisions. The ocks ranged from only 1.0 m (3.3 ft)
above the ground in windy conditions to as high as 400 m
(1,300 ft). These migrating ocks were typically in narrow columns that twisted and undulated, and they were
reported as being in nearly every conceivable shape.[42]
A skilled yer, the passenger pigeon is estimated to have
averaged 100 km/h (62 mph) during migration. It ew
with quick, repeated aps that increased the birds velocity the closer the wings got to the body. It was equally
as adept and quick at ying through a forest as through
open space. A ock was also adept at following the lead
of the pigeon in front of it, and ocks swerved together
to avoid a predator. When landing, the pigeon apped its
wings repeatedly before raising them at the moment of
landing. The pigeon was awkward when on the ground,
and moved around with jerky, alert steps.[47]
Live male in Whitmans aviary, 1896/98
The passenger pigeon was nomadic, constantly migrating

in search of food, shelter, or nesting grounds.[20] In his
1831 Ornithological Biography, American naturalist and
artist John James Audubon described a migration he observed in 1813 as follows:
I dismounted, seated myself on an eminence, and began to mark with my pencil, making a dot for every ock that passed. In a short
time nding the task which I had undertaken
impracticable, as the birds poured in in countless multitudes, I rose and, counting the dots
then put down, found that 163 had been made
in twenty-one minutes. I traveled on, and still
met more the farther I proceeded. The air was
literally lled with Pigeons; the light of noonday was obscured as by an eclipse; the dung fell
in spots, not unlike melting akes of snow, and
the continued buzz of wings had a tendency to
lull my senses to repose... I cannot describe
to you the extreme beauty of their aerial evolutions, when a hawk chanced to press upon
the rear of the ock. At once, like a torrent,
and with a noise like thunder, they rushed into
a compact mass, pressing upon each other towards the center. In these almost solid masses,
they darted forward in undulating and angular
lines, descended and swept close over the earth
Illustration of migrating ocks, Frank Bond, 1920
The passenger pigeon was one of the most social of all

land birds.[48] Estimated to have numbered three to ve
billion at the height of its population, it may have been
the most numerous bird on Earth; researcher Arlie W.
Schorger believed that it accounted for between 25 and
40 percent of the total land bird population in the United
States.[49] The passenger pigeons historic population is
roughly the equivalent of the number of birds that overwinter in the United States every year in the early 21st
century.[50] One ock in 1866 in southern Ontario was
described as being 1.5 km (1 mi) wide and 500 km (300
mi) long, took 14 hours to pass, and held in excess of
3.5 billion birds.[51] Such a number would likely represent a large fraction of the entire population at the time,
or perhaps all of it.[12] Most estimations of numbers were
based on single migrating colonies, and it is unknown how
7
many of these existed at a given time. American writer
Christopher Cokinos has suggested that if the birds ew
single le, they would have stretched around the earth 22
times.[52] A 2014 genetic study (based on coalescent theory) suggested that the passenger pigeon population uctuated rapidly across the last million years, due to their
dependence on availability of mast (which itself uctuates). The study suggested the bird was not always abundant, mainly persisting at around 1/10,000 the amount of
the several billions estimated in the 1800s.[53][54] Some
early accounts also suggest that the appearance of ocks
in great numbers was an irregular occurrence.[32] These
large uctuations in population may have been the result
of a disrupted ecosystem.
Alert parent bird posing deantly towards the camera

Juvenile (left), male (center), female (right), Louis Agassiz
Fuertes, 1910
A communally roosting species, the passenger pigeon

chose roosting sites that could provide shelter and enough
food to sustain their large numbers for an indenite period. The time spent at one roosting site may have depended on the extent of human persecution, weather conditions, or other, unknown factors. Roosts ranged in size
and extent, from a few acres to 260 km2 (100 square
miles) or greater. Some roosting areas would be reused
for subsequent years, others would only be used once.[20]
The passenger pigeon roosted in such numbers that even
thick branches on a tree would break under the strain.
The birds frequently piled on top of each others backs
to roost. They rested in a slumped position that hid their
feet. They slept with their bills concealed by the feathers
in the middle of the breast while holding their tail at a 45degree angle.[47] Dung could accumulate under a roosting
site to a depth of over 0.3 m (1.0 ft).[38]
If the pigeon became alert, it would often stretch out its
head and neck in line with its body and tail, then nod its
head in a circular pattern. When aggravated by another
pigeon, it raised it wings threateningly, but passenger pigeons almost never actually fought. The pigeon bathed in
shallow water, and afterwards lay on each side in turn and
raised the opposite wing to dry it.[47] The passenger pigeon drank at least once a day, typically at dawn, by fully
inserting its bill into lakes, small ponds, and streams. Pigeons were seen perching on top of each other to access
water, and if necessary, the species could alight on open
water to drink.[38] One of the primary causes of natural

mortality was the weather, and every spring many individuals froze to death after migrating north too early. In
captivity, a passenger pigeon was capable of living at least
15 years; Martha, the last known living passenger pigeon,
was at least 17 and possibly as old as 29 when she died.
It is undocumented how long a wild pigeon lived.[55]
The bird is believed to have played a signicant ecological role in the composition of presettlement forests of
eastern North America. For instance, while the passenger pigeon was extant, forests were dominated by white
oaks. This species germinated in the fall, therefore producing acorns during the spring to be devoured and spread
by the pigeons. The absence of the passenger pigeons
seed dispersal may have led to the modern dominance of
red oaks. At roosting sites, few plants grew for years after the pigeons left. Also, the immense amount of dung
present at these sites increased both the frequency and
intensity of forest res.[50] With the large numbers in the
ocks, the excrement they produced was enough to destroy surface-level vegetation, and along with the breaking of treelimbs under their collective weight, the passenger pigeons could do signicant damage to forests. Due
to these inuences, some ecologists have considered the
passenger pigeon a keystone species.[53] The American
chestnut trees that provided much of the mast on which
the passenger pigeon fed was itself almost driven to extinction by an imported Asian fungus (chestnut blight)
around 1905. As many as thirty billion trees are thought
to have died as a result in the following decades, but this

did not aect the passenger pigeon, which was already
extinct in the wild at the time.[20]
4.1
Diet
Acorns in South Carolina, among the diet of this bird
Beeches and oaks produced the mast needed to support nesting and roosting ocks.[56] The passenger pigeon
changed its diet depending on the season. In the fall, winter, and spring, it mainly ate beechnuts, acorns, and chestnuts. During the summer, berries and softer fruits, such
as blueberries, grapes, cherries, mulberries, pokeberries,
and bunchberry, became the main objects of its consumption. It also ate worms, caterpillars, snails, and other invertebrates, particularly while breeding.[20][43] Additionally, the passenger pigeon took advantage of cultivated
grains, particularly buckwheat, when it found them. The
species was especially fond of salt, which it ingested either from brackish springs or salty soil.[56] Mast occurs
in large quantities in dierent places at dierent times,
and rarely in consecutive years, which is one of the reasons why the large ocks were constantly on the move. As
mast is produced during autumn, there would have to be
a large amount of it left by the summer, when the young
were reared. It is unknown how they located this uctuating food source, but their eyesight and ight powers
aided them in surveying large areas for places that could
provide food enough for a temporary stay.[12][20]
The passenger pigeon foraged in ocks of tens or hundreds of thousands of individuals that overturned leaves,
dirt, and snow with their bills in a frantic search for large
quantities of food. One observer described the motion
of such a ock in search of mast as having a rolling appearance, as birds in the back of the ock ew overhead
to the front of the ock, dropping leaves and grass in
ight.[20][43] The ocks had wide leading edges to better
scan the landscape for food sources.[56] When nuts on a
tree loosened from their caps, a pigeon would land on a
branch and, while apping vigorously to stay balanced,
grab the nut, pull it loose from its cap, and swallow it
whole. Collectively, a foraging ock was capable of removing nearly all fruits and nuts from their path. Birds
Internal organs of Martha, the last individual: cr. denotes the

crop, gz. the gizzard, 1915
in the back of the ock ew to the front in order to pick

over unsearched ground; however, birds never ventured
far from the ock and hurried back if they became isolated. It is believed that the pigeons used social cues in
order to identify abundant sources of food, and a ock
of pigeons that saw others feeding on the ground often
joined them.[43] During the day, the birds left the roosting forest to forage on more open land.[42] They regularly
ew 100 to 130 km (62 to 81 mi) away from their roost
daily in search of food, and some pigeons reportedly traveled as far as 160 km (99 mi), leaving the roosting area
early and returning at night.[20][50]
The passenger pigeon had a very elastic mouth and throat,
allowing for increased capacity, and a joint in the lower
bill enabled it to swallow acorns whole. The bird was also
able to store large quantities of food in its crop, which
was capable of expanding to about the size of an orange, causing the neck to bulge. This allowed a bird to
quickly grab any food it discovered in the highly competitive ock. The crop was described as being capable
of holding at least 17 acorns or 28 beechnuts, 11 grains
of corn, 100 maple seeds, plus other material; it was estimated that a passenger pigeon needed to eat about 61
cm3 (3.7 in3 ) of food a day in order to survive. If shot,
a pigeon with a crop full of nuts would fall to the ground
with a sound described as like the rattle of a bag of marbles. After feeding, the pigeons perched on branches and
digested the food stored in their crop overnight.[20][43][50]
The pigeon could eat and digest 0.100 kg (0.22 lb) of
4.2
Reproduction
acorns per day.[57] At the historic population of three billion passenger pigeons, this amounted to 210,000,000 L
(55,000,000 US gal) of food a day.[50] The pigeon was
also able to regurgitate food from its crop when more desirable food became available.[39]
4.2
Reproduction
9
lation of these nesting areas, but most accounts mention
colonies containing millions of birds. The largest nesting
area ever recorded was in central Wisconsin in 1871; it
was reported as covering 2,200 km2 (850 sq mi), with
the number of birds nesting there estimated to be around
136,000,000. As well as these cities, there were regular reports of much smaller ocks or even individual pairs
setting up a nesting site.[20][58] The birds do not seem to
have formed as vast breeding colonies at the periphery of
their range.[32]
Courtship took place at the nesting colony.[48] Unlike
other pigeons, courtship took place on a branch or perch.
The male, with a ourish of the wings, made a keck
call while near a female. The male then gripped tightly to
the branch and vigorously apped his wings up and down.
When the male was close to the female, he then pressed
against her on the perch with his head held high and pointing at her.[47] If receptive, the female pressed back against
the male.[48] When ready to mate, the pair preened each
other. This was followed by the birds billing, in which the
female inserted its bill into and clasped the males bill,
shook for a second, and separated quickly while standing next to each other. The male then scrambled onto
the females back and copulated, which was then followed
by soft clucking and occasionally more preening.[48] John
James Audubon described the courtship of the passenger
pigeon as follows:
Nesting captive bird, wary of the photographer
Other than nding roosting sites, the migrations of the

passenger pigeon were connected with nding places appropriate for this communally breeding bird to nest and
raise its young. It is not certain how many times a year
the birds bred; once seems most likely, but some accounts
suggest more. The nesting period lasted around four to
Nest and egg in Whitmans aviary
six weeks. The ock arrived at a nesting ground around
March in southern latitudes, and some time later in more
northern areas.[20][49] The pigeon had no site delity, ofThither the countless myriads resort, and
ten choosing to nest in a dierent location each year.[55]
prepare to fulll one of the great laws of naThe formation of a nesting colony did not necessarily
ture. At this period the note of the Pigeon is
take place until several months after the pigeons arrived
a soft coo-coo-coo-coo much shorter than that
on their breeding grounds, typically during late March,
of the domestic species. The common notes
April, or May.[58]
resemble the monosyllables kee-kee-kee-kee,
The colonies, which were known as cities, were imthe rst being the loudest, the others gradumense, ranging from 49 ha (120 acres) to thousands of
ally diminishing In power. The male assumes
hectares in size, and were often long and narrow in shape
a pompous demeanor, and follows the female,
(L-shaped), with a few areas untouched for unknown reawhether on the ground or on the branches, with
sons. Due to the topography, they were rarely continuspread tail and drooping wings, which it rubs
ous. Since no accurate data were recorded, it is not posagainst the part over which it is moving. The
sible to give more than estimates on the size and popubody is elevated, the throat swells, the eyes
10

sparkle. He continues his notes, and now and
then rises on the wing, and ies a few yards
to approach the fugitive and timorous female.
Like the domestic Pigeon and other species,
they caress each other by billing, in which action, the bill of the one is introduced transversely into that of the other, and both parties
alternately disgorge the contents of their crop
by repeated eorts.[46]
been swept clean, due to all the twigs being collected at

the same time, yet this area would also have been covered in dung.[20][39][59] As both sexes took care of the
nest, the pairs were monogamous for the duration of the
nesting.[47]
After observing captive birds, Wallace Craig found that

this species did less charging and strutting than other pigeons (as it was awkward on the ground), and thought it
probable that no food was transferred during their brief
billing (unlike in other pigeons), and he therefore considered Audubons description partially based on analogy
with other pigeons as well as imagination.[40][47]
Live nestling or squab
Generally, the eggs were laid during the rst two weeks
of April across the pigeons range.[58] Each female laid its
egg immediately or almost immediately after the nest was
completed; sometimes the pigeon was forced to lay it on
the ground if the nest was not complete.[60] The normal
clutch size appears to have been a single egg, but there
is some uncertainty about this, as two have also been reported from the same nests.[20] Occasionally, a second female laid its egg in another females nest, resulting in two
eggs being present.[61] The egg was white and oval shaped
and averaged 40 mm (1.6 in) by 34 mm (1.3 in) in size.[59]
If the egg was lost, it was possible for the pigeon to lay a
replacement egg within a week.[60] A whole colony was
known to re-nest after a snowstorm forced them to abanPreserved egg, Musum de Toulouse
don their original colony.[55] The egg was incubated by
both parents for 12 to 14 days, with the male incubatNests were built immediately after pair formation and
and the female
took two to four days to construct; this process was highly ing it from midmorning to midafternoon
[20][60]
incubating
it
for
the
rest
of
the
time.
[58]
synchronized within a colony.
The female chose the
nesting site by sitting on it and icking its wings. The Upon hatching, the nestling (or squab) was blind and
male then carefully selected nesting materials, typically sparsely covered with yellow, hairlike down.[60] The
twigs, and handed them to the female over her back. The nestling developed quickly and within 14 days weighed
male then went in search of more nesting material while as much as its parents. During this brooding period both
the female constructed the nest beneath herself. Nests parents took care of the nestling, with the male attending
were built between 2.0 and 20.1 m (6.6 and 65.9 ft) above in the middle of the day and the female at other times.
the ground, though typically above 4.0 m (13.1 ft), and The nestlings were fed crop milk (a substance similar to
were made of 70 to 110 twigs woven together to create curd, produced in the crops of the parent birds) exclua loose, shallow bowl through which the egg could eas- sively for the rst days after hatching. Adult food was
ily be seen. This bowl was then typically lined with ner gradually introduced after three to six days. After 13 to
twigs. The nests were about 15 cm (5.9 in) wide, 6.1 cm 15 days, the parents fed the nestling for a last time and
(2.4 in) high, and 1.9 cm (0.75 in) deep. Though the nest then abandoned it, leaving the nesting area en masse. The
has been described as crude and imsy compared to those nestling begged in the nest for a day or two, before climbof many other birds, remains of nests could be found at ing from the nest and uttering to the ground, whereafter
sites where nesting had taken place several years prior. it moved around, avoided obstacles, and begged for food
Nearly every tree capable of supporting nests had them, from nearby adults. It was another three or four days beoften more than 50 per tree; one hemlock was recorded fore it edged.[20][61] The entire nesting cycle lasted about
as holding 317 nests. The nests were placed on strong 30 days.[39] It is unknown whether colonies re-nested afbranches close to the tree trunks. Some accounts state ter a successful nesting.[55] The passenger pigeon sexuthat ground under the nesting area looked as if it had ally matured during its rst year and bred the following
11
spring.[61]
4.3
Predators and parasites
Two parasites have been recorded on passenger pigeons.

One species of phtilopterid louse, Columbicola extinctus,
was originally thought to have lived on just passenger pigeons and to have become coextinct with them. This was
proven inaccurate in 1999 when C. extinctus was rediscovered living on band-tailed pigeons.[62][63] This, and
the fact that the related louse C. angustus is mainly found
on cuckoo-doves, further supports the relation between
these pigeons, as the phylogeny of lice broadly mirrors
that of their hosts.[15] Another louse, Campanulotes defectus, was thought to have been unique to the passenger pigeon, but is now believed to have been a case of
a contaminated specimen, as the species is considered to
be the still-extant Campanulotes avus of Australia.[63]
There is no record of a wild pigeon dying of either disease or parasites.[55]
5 Relationship with humans
Immature bird; the young were vulnerable to predators after leaving the nest
Nesting colonies attracted large numbers of predators, including American minks, American weasels, American
martens, and raccoons that preyed on eggs and nestlings,
birds of prey, such as owls, hawks, and eagles that preyed
on nestlings and adults, and wolves, foxes, bobcats, bears,
and mountain lions that preyed on injured adults and
fallen nestlings. Hawks of the genus Accipiter and falcons pursued and preyed upon pigeons in ight, which in
turn executed complex aerial maneuvers to avoid them;
Coopers hawk was known as the great pigeon hawk
due to its successes, and these hawks allegedly followed
migrating passenger pigeons.[48] While many predators
were drawn to the ocks, individual pigeons were largely
protected due to the sheer size of the ock, and overall little damage could be inicted on the ock by predation.[48]
Despite the number of predators, nesting colonies were so
large that they were estimated to have a 90% success rate
if not disturbed.[55] After being abandoned and leaving
the nest, the very fat juveniles were vulnerable to predators until they were able to y. The sheer number of juveniles on the ground meant that only a small percentage of
them were killed; predator satiation may therefore be one
of the reasons for the extremely social habits and communal breeding of the species.[20][28]
Billing pair by John James Audubon, from The Birds of America, 18271838
The passenger pigeon played a religious role for some

northern Native American tribes. The Wyandot people
(or Huron) believed that every twelve years during the
Feast of the Dead, the souls of the deceased changed into
passenger pigeons, which were then hunted and eaten.[64]
Before hunting the juvenile pigeons, the Seneca people
made an oering of wampum and brooches to the old
passenger pigeons, these were placed in a small kettle or
other receptacle by a smoky re.[64] The Ho-Chunk people considered the passenger pigeon to be the bird of the
12
5 RELATIONSHIP WITH HUMANS
chief, as they were served whenever the chieftain gave

a feast.[65] The Seneca people believed that a white pigeon was the chief of the passenger pigeon colony, and
that a Council of Birds had decided that the pigeons had
to give their bodies to the Seneca because they were the
only birds that nested in colonies. The Seneca developed
a pigeon dance as a way of showing their gratitude.[65]
French explorer Jacques Cartier was the rst European
to report on passenger pigeons, during his voyage in
1534.[66] The bird was subsequently observed and noted
by historical gures such as Samuel de Champlain and
Cotton Mather. Most early accounts dwell on the vast
number of pigeons, the resulting darkened skies, and the
enormous amount of hunted birds (50,000 birds were reportedly sold at a Boston market in 1771).[52] The early
colonists thought that large ights of pigeons would be
followed by ill fortune or sickness. When the pigeons
wintered outside of their normal range, some believed
that they would have a sickly summer and autumn.[67]
In the 18th and 19th centuries, various parts of the pigeon were thought to have medicinal properties. The
blood was supposed to be good for eye disorders, the powdered stomach lining was used to treat dysentery, and the
dung was used to treat a variety of ailments, including
headaches, stomach pains, and lethargy.[68] Though they
did not last as long as the feathers of a goose, the feathers of the passenger pigeon were frequently used for bedding. Pigeon feather beds were so popular that for a time
in Saint-Jrme, Quebec, every dowry included a bed and
Painting of a male, K. Hayashi, c. 1900
pillows made of pigeon feathers. In 1822, one family in
Chautauqua County, New York, killed 4,000 pigeons in
a day solely for this purpose.[69]
trations of the passenger pigeon were often drawn after
The passenger pigeon was featured in the writings of stued birds, and Charles R. Knight is the only serious
many signicant early naturalists, as well as accompa- artist known to have drawn the species from life. He did
nying illustrations. Mark Catesbys 1731 illustration, the so on at least two occasions; in 1903 he drew a bird posrst published depiction of this bird, is somewhat crude, sibly in one of the three aviaries with surviving birds, and
according to some later commentators. The original wa- some time before 1914, he drew Martha, the last individtercolor that the engraving is based on was bought by ual, in Cincinnati Zoo. The bird has been written about
[upper-alpha 1]
and ction) and
the British royal family in 1768, along with the rest of (including in poems, songs,
Catesbys watercolors. The naturalists Alexander Wil- illustrated by many notable writers and artists, and is deson and John James Audubon both witnessed large pi- picted in art to this day, for example in Walton Ford's
geon migrations rst hand, and published detailed ac- 2002 painting Falling Bough, and National Medal of
counts wherein both attempted to deduce the total num- Arts winner John A. Ruthven's 2014 mural in Cincinnati,
ber of birds involved. The most famous and often re- which commemorates the 100th anniversary of Marthas
[40][70][75][76]
The passenger pigeon mural on the
produced depiction of the passenger pigeon is Audubons death.
wall
of
a
six-story
building in Cincinnati which can be
illustration (handcolored aquatint) in his book The Birds
[46][52][70]
seen
in
the
2014
documentary
From Billions To None by
of America, published between 1827 and 1838.
[77][78][79][80][81]
David
Mrazek
and
Joel
Greenberg.
Audubons image has been praised for its artistic qualities, but criticized for its supposed scientic inaccuracies. As Wallace Craig and R. W. Shufeldt (among others) pointed out, the birds are shown perched and billing 5.1 Hunting
one above the other, whereas they would instead have
source of food
done this side by side, the male would be the one pass- The passenger pigeon was an important
[82]
for
the
people
of
North
America.
The
indigenous
peoing food to the female, and the males tail would not be
ples
ate
pigeons,
and
tribes
near
nesting
colonies
would
spread. Craig and Shufeldt instead cited illustrations by
the juAmerican artist Louis Agassiz Fuertes and Japanese artist sometimes move to live closer to them and eat
[83]
Many
veniles,
killing
them
at
night
with
long
poles.
K. Hayashi as more accurate depictions of the bird. IllusNative Americans were careful not to disturb the adult
5.1
Hunting
13
Depiction of a shooting in Louisiana, Smith Bennett, 1875
pigeons, and instead ate only the juveniles as they were

afraid that the adults might desert their nesting grounds; in
some tribes, disturbing the adult pigeons was considered
a crime.[84] Away from the nests, large nets were used to
capture adult pigeons, sometimes up to 800 at a time.[85]
Low-ying pigeons could be killed by throwing sticks or
stones. At one site in Oklahoma, the pigeons leaving their
roost every morning ew low enough that the Cherokee
could throw clubs into their midst, which caused the lead
pigeons to try to turn aside and in the process created
a blockade that resulted in a large mass of ying, easily
hit pigeons.[86] Among the game birds, passenger pigeons
were second only to the wild turkey (Meleagris gallopavo)
in terms of importance for the Native Americans living in
the southeastern United States. The birds fat was stored,
often in large quantities, and used as butter. Archaeological evidence supports the idea that Native Americans ate
the pigeons frequently prior to colonization.[87]
After European colonization, the passenger pigeon was
hunted more intensely and with more sophisticated methods than the more sustainable methods practiced by the
natives.[28] Yet it has also been suggested that the species
was rare prior to 1492, and that the subsequent increase
in their numbers may be due to the decrease in the Native
American population (who, as well as hunting the birds,
competed with them for mast) caused by European immigration, and the supplementary food (agricultural crops)
the immigrants provided.[88] It was of particular value
on the frontier, and some settlements counted on the
pigeon to support their population.[89][90] The avor of
the esh of passenger pigeons varied depending on how
they were prepared. In general, juveniles were thought
to taste the best, followed by birds fattened in captivity and birds caught in September and October. It was
common practice to fatten trapped pigeons before eating
them or storing their bodies for winter.[82] Dead pigeons
were commonly stored by salting or pickling the bodies;
other times, only the breasts of the pigeons were kept, in
which case they were typically smoked. In the early 19th
century, commercial hunters began netting and shooting
the birds to sell as food in city markets, and even as pig
1881 spread showing methods of trapping pigeons for shooting

contests
fodder. Once pigeon meat became popular, commercial

hunting started on a prodigious scale.[90][91]
Passenger pigeons were shot with such ease that many
did not consider them to be a game bird, as an amateur hunter could easily bring down six with one shotgun
blast; a particularly good shot with both barrels of a shotgun at a roost could kill 61 birds.[92][93] The birds were
frequently shot either in ight during migration or immediately after, when they commonly perched in dead,
exposed trees.[92] Hunters only had to shoot toward the
sky without aiming, and many pigeons would be brought
down.[28] The pigeons proved dicult to shoot head-on,
so hunters typically waited for the ocks to pass overhead
before shooting them. Trenches were sometimes dug and
lled with grain so that a hunter could shoot the pigeons
along this trench.[94] Hunters largely outnumbered trappers, and hunting passenger pigeons was a popular sport
for young boys.[95] In 1871, a single seller of ammunition provided three tons of powder and 16 tons (32,000
lb) of shot during a nesting. In the latter half of the
19th century, thousands of passenger pigeons were captured for use in the sports shooting industry. The pigeons
were used as living targets in shooting tournaments, such
as "trap-shooting", the controlled release of birds from
special traps. Competitions could also consist of people
standing regularly spaced while trying to shoot down as
many birds as possible in a passing ock.[28][96] The pigeon was considered so numerous that 30,000 birds had
to be killed to claim the prize in one competition.[39]
14
Pigeon net in Canada, by James Pattison Cockburn, 1829
There were a wide variety of other methods used to capture and kill passenger pigeons. Nets were propped up
to allow passenger pigeons entry, then closed by knocking loose the stick that supported the opening, trapping
twenty or more pigeons inside.[97] Tunnel nets were also
used to great eect, and one particularly large net was capable of catching 3,500 pigeons at a time.[98] These nets
were used by many farmers on their own property as well
as by professional trappers.[99] Food would be placed on
the ground near the nets to attract the pigeons. Decoy or
Stool pigeons (sometimes blinded by having their eyelids sewn together) were tied to a stool. When a ock of
pigeons passed by, a cord would be pulled that made the
stool pigeon utter to the ground, making it seem as if
it had found food, and the ock would be lured into the
trap.[28][100][101] Salt was also frequently used as bait, and
many trappers set up near salt springs.[102] At least one
trapper used alcohol-soaked grain as bait to intoxicate the
birds and make them easier to kill.[86] Another method
of capture was to hunt at a nesting colony, particularly
during the period of a few days after the adult pigeons
abandoned their nestlings, but before the nestlings could
y. Some hunters used sticks to poke the nestlings out
of the nest, while others shot the bottom of a nest with
a blunt arrow to dislodge the pigeon. Others cut down
a nesting tree in such a way that when it fell, it would
also hit a second nesting tree and dislodge the pigeons
within.[103] In one case, 6 km2 (1,500 acres) of large trees
were speedily cut down to get birds, and such methods
were common.[28] An extreme method, practiced only by
particularly unscrupulous hunters, was to set re to the
base of a tree nested with pigeons; the adults would ee
and the juveniles would fall to the ground.[104][105] Sulfur
was sometimes burned beneath the nesting tree to suocate the birds, which fell out of the tree in a weakened
state.[106]
By the mid-1800s, railroads had opened new opportunities for pigeon hunters. While previously it had proved
too dicult to ship masses of pigeons to eastern cities, the
access provided by the railroad permitted pigeon hunting
to become commercialized.[91] An extensive telegraph
system was introduced in the 1860s, which improved
Trapper Albert Cooper with blind decoy pigeons for luring wild
birds, c. 1870
communication across the United States, making it easier to spread information about the whereabouts of pigeon ocks.[96] After being opened up to the railroads,
the town of Plattsburg, New York is estimated to have
shipped 1.8 million pigeons to larger cities in 1851 alone
at a price of 31 to 56 cents a dozen. By the late 1800s,
the trade of passenger pigeons had become commercialized. Large commission houses employed trappers
(known as pigeoners) to follow the ocks of pigeons
year-round.[107] A single hunter is reported to have sent
three million birds to eastern cities during his career.[108]
In 1874, at least 600 people were employed as pigeon
trappers, a number which grew to 1,200 by 1881. Pigeons were caught in such numbers that by 1876, shipments of dead pigeons were unable to recoup the costs
of the barrels and ice needed to ship them.[109] The price
of a barrel full of pigeons dropped to below fty cents,
due to overstocked markets. Passenger pigeons were instead kept alive so their meat would be fresh when the
birds were killed, and sold once their market value had
increased again. Thousands of birds were kept in large
pens, though the bad conditions led many to die from lack
of food and water, and by fretting (gnawing) themselves;
many rotted away before they could be sold.[52]
5.2
Decline and conservation attempts
Hunting of passenger pigeons was documented and depicted in contemporaneous newspapers, wherein various
trapping methods and uses were featured. The most often
reproduced of these illustrations was captioned Winter
sports in northern Louisiana: shooting wild pigeons, and
published in 1875. Passenger pigeons were also seen as
agricultural pests, since entire crops could be destroyed
by feeding ocks. The bird was described as a perfect scourge by some farming communities, and hunters
were employed to wage warfare on the birds to save
grain, as shown in another newspaper illustration from
1867 captioned as Shooting wild pigeons in Iowa.[96]
When comparing these pests to the bison of the Great
Plains, it is possible to infer that the valuable resource
needed was not the species of animals but the agriculture
which was consumed by said animal. The crops that were
eaten were seen as marketable calories, proteins, and nutrients all grown for the wrong species.[110][111]
5.2
15
east, though there were still millions of birds in the 1850s.
The population must have been decreasing in numbers for
many years, though this went unnoticed due to the apparent vast number of birds, which clouded their decline.[52]
In 1856 Bndict Henry Rvoil may have been one of the
rst writers to voice concern about the fate of the passenger pigeon, after witnessing a hunt in 1847:
Everything leads to the belief that the pigeons, which cannot endure isolation and are
forced to ee or to change their way of living
according to the rate at which North America is
populated by the European inow, will simply
end by disappearing from this continent, and,
if the world does not end this before a century,
I will wager... that the amateur of ornithology
will nd no more wild pigeons, except those in
the Museums of Natural History.[52]
Decline and conservation attempts
Life drawing by Charles R. Knight, 1903
Male and female by Louis Agassiz Fuertes, frontispiece of

William Butts Mershon's 1907 The Passenger Pigeon
By the 1870s, the decrease in birds was noticeable, especially after the last large scale nestings and subsequent
slaughters of millions of birds in 1874 and 1878. By this
time, large nestings only took place in the north, around
the Great Lakes. The last large nesting was in Petoskey,
Michigan, in 1878 (following one in Pennsylvania a few
days earlier), where 50,000 birds were killed each day for
nearly ve months. The surviving adults attempted a second nesting at new sites, but were killed by professional
hunters before they had a chance to raise any young. Scattered nestings are reported into the 1880s, but the birds
were now wary, and commonly abandoned their nests if
persecuted.[12][34][52]
The notion that the species could be driven to extinction

was alien to the early colonists, both because the number
of birds did not appear to diminish, but also because the
concept of extinction itself was yet to be dened. The
bird seems to have been slowly pushed westwards since By the time of these last nestings, laws had already been
the arrival of Europeans, becoming scarce or absent in the enacted to protect the passenger pigeon, but these proved
16
ineective, as they were unclearly framed and hard to

enforce. H. B. Roney, who had witnessed the Petoskey
slaughter, led campaigns to protect the pigeon, but was
met with resistance, and accusations that he was exaggerating the severity of the situation. Few oenders were
prosecuted, mainly some poor trappers, but the large enterprises were not aected.[52] In 1857, a bill was brought
forth to the Ohio State Legislature seeking protection for
the passenger pigeon, yet a Select Committee of the Senate led a report stating that the bird did not need protection, being wonderfully prolic, and dismissing the
suggestion that the species could be destroyed.[112] Public protests against trap-shooting erupted in the 1870s, as
the birds were badly treated before and after such contests. Conservationists were ineective in stopping the
slaughter. A bill was passed in the Michigan legislature
making it illegal to net pigeons within 3 km (2 miles) of a
nesting area. In 1897, a bill was introduced in the Michigan legislature asking for a 10-year closed season on passenger pigeons. Similar legal measures were passed and
then disregarded in Pennsylvania. The gestures proved
futile, and by the mid-1890s, the passenger pigeon had almost completely disappeared, and was probably extinct as
a breeding bird in the wild.[96][108] Small ocks are known
to have existed at this point, since large numbers of birds
were still being sold at markets. Thereafter, only small
groups or individual birds were reported, many of which
were shot on sight.[52]
5.3
Last survivors
this is the most often cited last wild specimen, in 2014,

writer Joel Greenberg pointed out two later records, one
of which involves a male shot in 1902 in Indiana, that
was stued but later destroyed. The reliability of later accounts are in question. US President Theodore Roosevelt
claimed to have seen a bird in Michigan in 1907.[52] Ornithologist Alexander Wetmore claimed that he saw a pair
ying near Independence, Kansas, in April 1905.[114][115]
In 1910, the American Ornithologists Union oered a
reward of $3,000 for discovering a nest the equivalent
of $76,990.05 in 2015.[116][117]
Whitmans aviary with passenger pigeons and other species,

1896/98
Most captive passenger pigeons were kept for exploitative purposes, but some were housed in zoos and aviaries.
See also: Martha (pigeon)
Audubon alone claimed to have brought 350 birds to EngThe last recorded nest and egg in the wild were col- land in 1830, distributing them among various noblemen,
and the species is also known to have been kept at London
Zoo. Being common birds, these attracted little interest, until the species became rare in the 1890s. By the
turn of the 20th century, the last known captive passenger pigeons were divided in three groups; one in Milwaukee, one in Chicago, and one in Cincinnati. There are
claims of a few further individuals having been kept in
various places, but these accounts are not considered reliable today. The Milwaukee group was kept by David
Buttons, the last conrmed wild passenger pigeon, Cincinnati
Whittaker, who began his collection in 1888, and posZoo
sessed fteen birds some years later, all descended from
[12][118]
lected in 1895 near Minneapolis. The last wild individ- a single pair.
ual in Louisiana was discovered among a ock of mourn- The Chicago group was kept by Professor Charles Otis
ing doves in 1896, and subsequently shot. Many late Whitman, whose collection began with passenger pigeons
sightings are thought to be false or due to confusion with bought from Whittaker beginning in 1896. He had an
mourning doves.[52] The last fully authenticated record of interest in studying pigeons, and kept his passenger pia wild passenger pigeon was near Sargents, Pike County, geons with other pigeon species. Whitman brought his
Ohio, on March 22 or 24, 1900, when a female bird was pigeons with him from Chicago to Massachusetts by railkilled by a boy named Press Clay Southworth with a BB car each summer. By 1897, Whitman had bought all of
gun.[34][113] The boy had not recognized the bird as a pas- Whittakers birds, and upon reaching a maximum of 19
senger pigeon, but his parents identied it, and sent it to a individuals, he gave seven back to Whittaker in 1898.
taxidermist. The specimen, nicknamed buttons due to Around this time, a series of photographs were taken of
the buttons used instead of glass eyes, was donated to the these birds; 24 of the photos survive to this day. Some
Ohio Historical Society by the family in 1915. Though of these images have been reproduced in various media,
5.4
Extinction causes
copies of which are now kept at the Wisconsin Historical Society. It is unclear exactly where, when, and by
whom these photos were taken, but some appear to have
been taken in Chicago in 1896, others in Massachusetts
in 1898, the latter by a J. G. Hubbard. By 1902, Whitman
owned sixteen birds. Many eggs were laid by his pigeons,
but few hatched, and many hatchlings died. A newspaper
inquiry was published that requested fresh blood to the
ock which had now ceased breeding. By 1907, he was
down to two female passenger pigeons that died that winter, and was left with two infertile male hybrids, whose
subsequent fate is unknown. By this time, only four (all
males) of the birds Whitman had returned to Whittaker
were alive, and these died between November 1908 and
February 1909.[118][119]
Martha, the last passenger pigeon, alive in 1912
The Cincinnati Zoo, one of the oldest zoos in the US, kept
passenger pigeons from its beginning in 1875. The zoo
kept more than twenty individuals, in a ten-by-twelvefoot cage.[118] Passenger pigeons do not appear to have
been kept at the zoo due to their rarity, but to enable
guests to have a closer look at a native species.[120] Recognizing the decline of the wild populations, Whitman and
the Cincinnati Zoo consistently strove to breed the surviving birds, including attempts at making a rock dove foster
passenger pigeon eggs.[121] In 1902, Whitman gave a female passenger pigeon to the zoo; this was possibly the
individual later known as Martha, which would become
the last living member of the species. Other sources argue that Martha was hatched at the Cincinnati Zoo, had
lived there for 25 years, and was the descendant of three
17
pairs of passenger pigeons purchased by the zoo in 1877.
It is thought this individual was named Martha because
her last cage mate was named George, thereby honoring
George Washington and his wife Martha, though it has
also been claimed she was named after the mother of a
zookeepers friends.[118][122]
In 1909, Martha and her two male companions at the
Cincinnati Zoo became the only known surviving passenger pigeons. One of these males died around April
that year, followed by George, the remaining male, on
July 10, 1910.[120] It is unknown whether the remains of
George were preserved. Martha soon became a celebrity
due to her status as an endling, and oers of a $1,000
reward for nding a mate for her brought even more visitors to see her. During her last four years in solitude
(her cage was 5.4 by 6 m (18 by 20 ft)), Martha became
steadily slower and more immobile; visitors would throw
sand at her to make her move, and her cage was roped o
in response.[118][123] Martha died of old age on September 1, 1914, and was found lifeless on the oor of her
cage.[37][124] It was claimed that she died at 1 p.m., but
other sources suggest she died some hours later.[118] Depending on the source, Martha was between 17 and 29
years old at the time of her death, although 29 is the generally accepted gure.[125] At the time, it was suggested
that Martha might have died from an apoplectic stroke,
as she had suered one a few weeks before dying.[126]
Her body was frozen into a block of ice and sent to
the Smithsonian Institution in Washington, where it was
skinned, dissected, photographed, and mounted.[37][104]
As she was molting when she died, she proved dicult
to stu, and previously shed feathers were added to the
skin. Martha was on display for many years, but after a
period in the museum vaults, she was put back on display
at the Smithsonians National Museum of Natural History
in 2015.[118] A memorial statue of Martha stands on the
grounds of the Cincinnati Zoo, in front of the Passenger Pigeon Memorial Hut, formerly the aviary wherein
Martha lived, now a National Historic Landmark. Incidentally, the last specimen of the extinct Carolina parakeet, named Incus, died in Marthas cage in 1918; the
stued remains of that bird are exhibited in the Memorial Hut.[34][127]
5.4 Extinction causes

The main reasons for the extinction of the passenger pigeon were the massive scale of hunting, the rapid loss
of habitat, and the extremely social lifestyle of the bird,
which made it highly vulnerable to the former factors.
Deforestation was driven by the need to free land for agriculture and expanding towns, but also due to the demand
for lumber and fuel. About 728,000 km2 (180 million
acres) were cleared for farming between 1850 and 1910.
Though there are still large woodland areas in eastern
North America, which support a variety of wildlife, it
was not enough to support the vast number of passen-
18

1800s, a combination which would have led to the rapid
extinction of the species. A similar scenario may also explain the rapid extinction of the Rocky Mountain locust
(Melanoplus spretus) during the same period.[53] It has
also been suggested that after the population was thinned
out, it would be harder for few or solitary birds to locate
suitable feeding areas.[12] In addition to the birds killed
or driven away by hunting during breeding seasons, many
nestlings were also orphaned before being able to fend for
themselves. Other, less convincing contributing factors
have been suggested at times, including mass drownings,
Newcastle disease, and migrations to areas outside their
original range.[1][28]
The extinction of the passenger pigeon aroused public interest in the conservation movement, and resulted in new
laws and practices which prevented many other species
from becoming extinct.[34] The rapid decline of the passenger pigeon has inuenced later assessment methods
of the extinction risk of endangered animal populations.
The International Union for the Conservation of Nature
(IUCN) has used the passenger pigeon as an example in
cases where a species was declared at risk for extinction
even though population numbers are high.[111]
Martha at the Smithsonian Museum, 2015
5.5 Re-creation of the species
ger pigeons needed to sustain the population. In contrast,

very small populations of nearly extinct birds, such as the
kakapo (Strigops habroptilus) and the takah (Porphyrio
hochstetteri), have been enough to keep those species alive
to the present. The combined eects of intense hunting
and deforestation has been referred to as a "Blitzkrieg"
against the passenger pigeon, and it has been labeled one
of the greatest and most senseless human-induced extinctions in history.[28][53][96] As the ocks dwindled in
size, the passenger pigeon population decreased below
the threshold necessary to propagate the species.[128]
Taxidermied male and female, Laval University Library
Pigeons being shot to save crops in Iowa, 1867
The 2014 study that demonstrated natural uctuations in

population numbers prior to human arrival also showed
that the species routinely recovered from lows in the population, and suggested that one of these lows may have coincided with the intensied exploitation by humans in the
Today, more than 1,532 passenger pigeon skins (along

with 16 skeletons) are in existence, spread across many
institutions all over the world.[32][33] It has been suggested that the passenger pigeon should be revived when
available technology allows it (a concept which has been
termed "de-extinction"), using genetic material from such
specimens. In 2003, the Pyrenean ibex (Capra pyrenaica
pyrenaica, a subspecies of the Spanish ibex) was the rst
extinct animal to be cloned back to life; the clone lived for
only seven minutes before dying of lung defects.[129][130]
A hindrance to cloning the passenger pigeon is the fact
that the DNA of museum specimens has been contaminated and fragmented, due to exposure to heat and oxygen. American geneticist George M. Church has proposed that the passenger pigeon genome can be reconstructed by piecing together DNA fragments from dier-
6.2
References
19
ent specimens. The next step would be to splice these

genes into the stem cells of rock pigeons (or band-tailed
pigeons), which would then be transformed into egg and
sperm cells, and placed into the eggs of rock pigeons, resulting in rock pigeons bearing passenger pigeon sperm
and eggs. The ospring of these would have passenger
pigeon traits, and would be further bred to favor unique
features of the extinct species.[129][131][132]
(1244):
445.
Bibcode:1918Sci....48..445O.
doi:10.1126/science.48.1244.445.
[6] Swainson, W. J. (1827). Mr. Swainson on several new
groups in Ornithology.. The Zoological Journal. 3: 362.
[7] Bangs, O. (1906). The names of the passenger pigeon
and the mourning dove. Proceedings of the Biological Society of Washington. 19: 4344.
The general idea of re-creating extinct species has been [8] Schenk, E. T.; McMasters, J. H. (1956). Procedure in
Taxonomy (Third ed.). Stanford, California: Stanford
criticized, since the large funds needed could be spent on
University Press. p. 89. ISBN 978-0-8047-3867-5.
conserving currently threatened species and habitats, and
because conservation eorts might be viewed as less ur[9] Blockstein 2002, p. 4
gent. In the case of the passenger pigeon, since it was
very social, it is unlikely that enough birds could be cre- [10] Miller, W. J. (January 16, 1969). The Biology and Natural History of the Mourning Dove. Should Doves be
ated for revival to be successful, and it is unclear whether
Hunted in Iowa?. Ames, IA: Ames Audubon Society. Rethere is enough appropriate habitat left for its reintroductrieved April 23, 2013.
tion. Furthermore, the parent pigeons that would raise
the cloned passenger pigeons would belong to a dierent [11] Brewer, T. M. (1840). Wilsons American Ornithology:
species, with a dierent way of rearing young.[129][131]
with Notes by Jardine; to which is Added a Synopsis of
6
6.1
American Birds, Including those Described by Bonaparte,

Audubon, Nuttall, and Richardson. Boston: Otis, Broaders, and Company. p. 717.
Bibliography
[12] Hume, J. P.; Walters, M. (2012). Extinct Birds. London:

T & AD Poyser. pp. 144146. ISBN 978-1-4081-57251.
Notes
[1] John Herald, a bluegrass singer, wrote a song dedicated to the extinction of the species and Martha, the
species endling, that he titled Martha (Last of the Passenger Pigeons)".[71][72] In connection with the centenary
of Marthas death, the song was cited as evidence of her
iconic stature a symbol of the wanton slaughter of
these pigeons and the human-caused extinction of the
species.[73][74]
6.2
References
[1] BirdLife International (2012). "Ectopistes migratorius".

IUCN Red List of Threatened Species. IUCN. 2012:
e.T22690733A39226308. doi:10.2305/IUCN.UK.20121.RLTS.T22690733A39226308.en. Retrieved 27 August
2016.
[2] Catesby, M. (1729). Natural History of Carolina, Florida
and the Bahama Islands. 1. London: W. Innys and R.
Manby. p. 23.
[3] Aldrich, J. W. (1993). Classication and Distribution.
In Baskett, T.S., Sayre, M.W., Tomlinson, R.E., Mirarchi, R.E. Ecology and management of the Mourning Dove.
Harrisburg, PA: Stackpole Books. p. 48. ISBN 978-08117-1940-7.
[4] Hemming, F. (1952). Proposed use of the plenary powers to secure that the name Columba migratoria Linnaeus,
1766, shall be the oldest available name for the Passenger
Pigeon, the type species of the genus Ectopistes Swainson,
1827. Bulletin of Zoological Nomenclature. 9: 8084.
doi:10.5962/bhl.part.10238.
[5] Oberholser, H. C. (1918).
name of the Passenger Pigeon.
The scientic
Science.
48
[13] Shapiro, B.; Sibthorpe, D.; Rambaut, A.; Austin, J.;

Wragg, G. M.; Bininda-Emonds, O. R. P.; Lee, P. L. M.;
Cooper, A. (2002). Flight of the Dodo (PDF). Science.
295 (5560): 1683. doi:10.1126/science.295.5560.1683.
PMID 11872833.
[14] Fulton, T. L.; Wagner, S. M.; Fisher, C.; Shapiro, B.
(2012). Nuclear DNA from the Extinct Passenger Pigeon (Ectopistes migratorius) Conrms a Single Origin of
New World Pigeons. Annals of Anatomy. Elsevier. 194
(1): 5257. doi:10.1016/j.aanat.2011.02.017. PMID
21482085.
[15] Johnson, K. P.; Clayton, D. H.; Dumbacher, J. P.;
Fleischer, R. C. (2010).
The ight of the Passenger Pigeon: phylogenetics and biogeographic history of an extinct species. Molecular Phylogenetics
and Evolution. Academic Press. 57 (1): 4558.
doi:10.1016/j.ympev.2010.05.010. PMID 20478386.
[16] Fulton, T. L.; Wagner, S. M.; Shapiro, B. (2012). Case
Study: Recovery of Ancient Nuclear DNA from Toe Pads
of the Extinct Passenger Pigeon. Ancient DNA. Methods
in Molecular Biology. 840. p. 29. doi:10.1007/978-161779-516-9_4. ISBN 978-1-61779-515-2.
[17] Hung, C. M.; Lin, R. C.; Chu, J. H.; Yeh, C. F.; Yao, C. J.;
Li, S. H. (2013). The De Novo Assembly of Mitochondrial Genomes of the Extinct Passenger Pigeon (Ectopistes
migratorius) with Next Generation Sequencing. Plos
One. 8 (2): e56301. doi:10.1371/journal.pone.0056301.
PMC 3577829 . PMID 23437111.
[18] Deane, R. (1908). The Passenger Pigeon (Ectopistes migratorius) in Connement. The Auk. 25 (2): 181183.
doi:10.2307/4070695. JSTOR 4070695.
20
[19] Atkinson, G. E. (1907). Mershon, W. B, ed. The Passenger Pigeon. New York: The Outing Publishing Co. p.
188.
[20] Fuller 2014, pp. 3047
[21] Harper, D. (2012). Passenger (n.)". Online Etymology
Dictionary. Retrieved April 23, 2013.
6 BIBLIOGRAPHY
[40] Craig, W. (1911). The expressions of emotion in

the pigeons. III. The Passenger Pigeon (Ectopistes migratorius Linn.)". The Auk. 4. 28 (4): 408427.
doi:10.2307/4071160. JSTOR 4071160.
[41] Blockstein 2002, p. 3
[22] Schorger 1955, p. 251
[23] Schorger 1955, pp. 252253
[44] Howard, H. (1937). A Pleistocene Record of the Passenger Pigeon in California. Condor. 39 (1): 1214.
doi:10.2307/1363481. JSTOR 1363481.
[24] Tatum, J.; Rementer, J.; the Culture Preservation Committee (2010). Extinct Birds the Lenape Knew. Culture
and History of the Delaware Tribe. Delaware Tribe of Indians. Archived from the original on October 6, 2012.
Retrieved April 23, 2013.
[25] Omiimii. Ojibwe Peoples Dictionary. Department of
American Indian Studies, University of Minnesota. Retrieved March 2, 2012.
[26] Costa, D. J. (2005). Wolfart, H. C., ed. The St. Jrme
Dictionary of Miami-Illinois (pdf). Papers of the 36th
Algonquian Conference. Winnipeg: University of Manitoba. pp. 107133. Retrieved April 23, 2013.
[27] Schorger 1955, p. 255
[28] Fuller 2014, pp. 7288
[29] Fuller 2014, pp. 150161
[31] Gibbs, D.; Barnes, E.; Cox, J. (2001). Pigeons and Doves:
A Guide to the Pigeons and Doves of the World. Sussex:
Pica Press. pp. 318319. ISBN 1-873403-60-7.
[32] Greenway, J. C. (1967). Extinct and Vanishing Birds of
the World. New York: American Committee for International Wild Life Protection 13. pp. 304311. ISBN
978-0-486-21869-4.
[33] Hume, J. P.; van Grouw, H. (2014). Colour aberrations
in extinct and endangered birds. Bulletin of the British
Ornithologists Club. 134: 168193.
[34] Department of Vertebrate Zoology, National Museum of
Natural History (March 2001). The Passenger Pigeon.
Encyclopedia Smithsonian. Smithsonian Institution. Retrieved April 22, 2013.
[35] Fuller 2014, pp. 162168
[36] Shufeldt, R. W. (1914). Osteology of the Passenger Pigeon (Ectopistes migratorius)". The Auk. 31 (3): 358362.
doi:10.2307/4071953. JSTOR 4071953.
[37] Shufeldt, R. W. (1915). Anatomical and other notes on
the Passenger Pigeon (Ectopistes migratorius) lately living
in the Cincinnati Zoological Gardens. The Auk. 32: 29
41. doi:10.2307/4071611. JSTOR 4071611.
[39] Fuller, E. (2001). Extinct Birds (Revised ed.). Ithaca,
New York: Comstock Publishing Associates. pp. 9697.
ISBN 0-8014-3954-X.
[45] Chandler, R. M. (1982). A Second Pleistocene Passenger Pigeon from California (PDF). Condor. Cooper Ornithological Society. 84 (2): 242. doi:10.2307/1367681.
JSTOR 1367681.
[46] Audubon, J. J. (1835). Ornithological biography, or, an
account of the Habits of the Birds of the United States of
America. Edinburgh: A. Black. pp. 319327.
[49] Schorger 1955, p. 205
[50] Ellsworth, J. W.; McComb, B. C. (2003). Potential Eects of Passenger Pigeon Flocks on the Structure and Composition of Presettlement Forests of Eastern North America. Conservation Biology. WileyBlackwell. 17 (6): 15481558. doi:10.1111/j.15231739.2003.00230.x.
[51] Sullivan, J.; Sutton, B.; Cronon, W. (April 2004). The
Passenger Pigeon: Once There Were Billions. Hunting
for Frogs on Elston, and Other Tales from Field & Street.
Chicago, Illinois: University of Chicago Press: 210113.
ISBN 978-0-226-77993-5. Retrieved February 29, 2012.
[52] Fuller 2014, pp. 5069
[53] Hung, C. M.; Shaner, P. J. L.; Zink, R. M.;
Liu, W. C.; Chu, T. C.; Huang, W. S.; Li,
S. H. (2014). Drastic population uctuations explain the rapid extinction of the passenger pigeon.
Proceedings of the National Academy of Sciences. 111
(29): 1063610641. Bibcode:2014PNAS..11110636H.
doi:10.1073/pnas.1401526111.
[54] Williams, S. C. P. (2014). Humans not solely to blame
for passenger pigeon extinction. Science & AAAS.
American Association for the Advancement of Science.
Retrieved June 17, 2014.
[57] Neumann, T. W. (1985). Human-wildlife competition
and the passenger pigeon: Population growth from system destabilization. Human Ecology. 13 (4): 389410.
doi:10.1007/BF01531152.
6.2
References

[62] Clayton, D. H.; Price, R. D. (1999). Taxonomy of
New World Columbicola (Phthiraptera: Philopteridae)
from the Columbiformes (Aves), with Descriptions of
Five New Species (PDF). Annals of the Entomological
Society of America. Entomological Society of America.
92 (5): 675685. doi:10.1093/aesa/92.5.675.
21
[79] Passenger Pigeon Symposium: From Billions to None.

Wisconsin Academy of Arts and Sciences. November 1,
2014. Retrieved March 3, 2016.
[80] Theres an Important Lesson in From Billions to None, a
New Documentary About the Extinct Passenger Pigeon.
Chicago Reader. Retrieved March 3, 2016.
[81] Mrazek, D. From Billions to None Preview (Video).
Retrieved March 3, 2016.
[82] Schorger 1955, p. 129
[63] Price, R. D.; Clayton, D. H.; Adams, R. J. (2000).

Pigeon Lice Down Under: Taxonomy of Australian
Campanulotes (Phthiraptera: Philopteridae), with a description of C. durdeni n.sp (PDF). Journal of Parasitology. American Society of Parasitologists. 86 (5): 94850.
doi:10.2307/3284803.
[83] Schorger 1955, pp. 133134
[64] Schorger 1955, p. 135
[87] Schorger 1955, p. 134
[65] Schorger 1955, p. 136
[88] Mann 2005, pp. 315318.
[66] Edey, M. (December 22, 1961). Once there were billions, now there are none. Life. 51 (25): 169176. ISSN
0024-3019. Retrieved June 16, 2015.
[89] Schorger 1955, p. 130
[67] Schorger 1955, p. 12
[91] Schorger 1955, p. 144
[68] Schorger 1955, pp. 132133
[92] Schorger 1955, p. 186
[69] Schorger 1955, p. 132
[93] Schorger 1955, p. 193
[70] Fuller 2014, pp. 124147
[94] Schorger 1955, p. 192
[71] Herald, J. Lyrics to Martha (Last of the Passenger Pigeons)"". Johnherald.com. Retrieved April 28, 2013.
[95] Schorger 1955, p. 198
[72] Gebhart, Parrish (October 17, 2010). Martha: Last of

the Passenger Pigeons (video). Retrieved December 12,
2013 via YouTube.
[73] Harvey, C.; Newbern, E. (August 29, 2014). 13 Memories of Martha, the Last Passenger Pigeon. Audubon
Magazine. Retrieved March 3, 2016.
[84] Schorger 1955, p. 137

[85] Schorger 1955, p. 139
[86] Schorger 1955, p. 168
[90] Schorger 1955, p. 131
[96] Hume, J. P. (2015). Large-scale live capture of Passenger Pigeons Ectopistes migratorius for sporting purposes:
overlooked illustrated documentation. Bulletin of the
British Ornithologists Club. 2. 135: 174184.
[97] Schorger 1955, p. 169
[98] Schorger 1955, p. 172
[99] Schorger 1955, p. 170

[74] McLendon, R. (September 1, 2011). Ode to Martha, the
last passenger pigeon. Mother Nature Network. Narrative [100] Schorger 1955, pp. 177179
Content Group. Retrieved March 3, 2016. One of eastern North Americas most iconic animals vanished forever [101] Paxson, H. D. (1917). The last of the Wild Pigeon in
Bucks County. Collection of papers read before the Bucks
on Sept. 1, 1914. Now, 97 years later, the passenger piCounty Historical Society. 4: 367382.
geon has become an icon for something else: manmade
extinction.
[102] Schorger 1955, p. 173
[75] Shufeldt, R. W. (1921). Published gures and plates of
the extinct passenger pigeon. Scientic Monthly (5 ed.). [103] Schorger 1955, p. 141
12: 458481. Bibcode:1921SciMo..12..458S.
[104] Yeoman, B. (2014). Why the Passenger Pigeon Went
Extinct. Audubon Magazine. Retrieved August 26, 2014.
[76] Milner, R. (2012). Charles R. Knight: The Artist Who Saw
Through Time. New York: Abrams Books. p. 138. ISBN [105] Schorger 1955, p. 142
978-0-8109-8479-0.
[106] Schorger 1955, p. 167
[77] Ruthven, J. Martha, the Last Passenger Pigeon (Video).
Retrieved March 3, 2016 via YouTube.
[107] Schorger 1955, p. 145
[78] Kogan, R. (September 10, 2014). "'Billions brings pi- [108] Ehrlich, P. R.; Dobkin, D. S.; Wheye, D. (1988). The
geons extinction tale to screen From Billions to None.
Passenger Pigeon. Stanford University. Retrieved March
Chicago Tribune. Retrieved March 3, 2016.
3, 2012.
22
[109] Schorger 1955, p. 146
EXTERNAL LINKS
[129] Lewis, T. (2013). How to bring extinct animals back to

life. NBC News. Retrieved 25 August 2013.
[110] Whaples, R. (2015). A Feathered River Across the Sky:

The Passenger Pigeons Flight to Extinction. The Inde- [130] ""Bringing Back the Passenger Pigeon Meeting convened
pendent Review. 19 (3): 4436.
at Harvard Medical School in Boston. The Long Now
Foundation. Retrieved April 10, 2015.
[111] Jackson, J. A.; Jackson, B. (2007). Extinction: the Passenger Pigeon, last hopes, letting go. The Wilson Journal [131] Zimmer, C. (2013). Bringing them back to life. Naof Ornithology. 119 (4): 767772. doi:10.1676/1559tional Geographic. Retrieved October 29, 2015.
4491(2007)119[767:etpplh]2.0.co;2. JSTOR 20456089.
[132] Landers, J. (2013). Scientists look to revive the long[112] Hornaday, W. T. (1913). Our Vanishing Wild Life. Its Exextinct passenger pigeon. The Washington Post. Retermination and Preservation. New York: Charles Scribtrieved November 6, 2014.
ners Sons. Retrieved February 29, 2012. at Project
Gutenberg.
[113] Reeve, S. (March 2001). Going Down in History. Geographical. Campion Interactive Publishing. 73 (3): 60
64. ISSN 0016-741X.
[114] Wetmore, A. (October 1936). Game Birds of Prairie,
Forest and Tundra. National Geographic. p. 495.
[115] McKinley, D. (1960). A History of the Passenger Pigeon in Missouri. Auk. 77 (4): 399420.
doi:10.2307/4082414. JSTOR 4082414.
[116] Stukel, E. D. (JanuaryFebruary 2005). Passenger Pigeon. South Dakota Game Fish & Parks. Retrieved August 7, 2016.
[117] Reward for Wild Pigeons. Ornithologists Oer $3,000
for the Discovery of Their Nests. The New York Times.
Boston, Massachusetts. April 4, 1910. Retrieved February 29, 2012.
[118] Fuller 2014, pp. 92121
[119] Rothschild, W. (1907).
Extinct Birds.
Hutchinson & Co. pp. 167170.
London:
6.3 Cited texts

Blockstein, D. E. (2002). Passenger Pigeon Ectopistes migratorius". In Poole, A.; Gill, F. The Birds
of North America. Philadelphia: The Birds of North
America, Inc., Cornell Lab of Ornithology. p. 611.
Retrieved March 3, 2016. (subscription required
(help)).
Fuller, E. (2014). The Passenger Pigeon. Princeton
and Oxford: Princeton University Press. ISBN 9780-691-16295-9.
Mann, C. C. (2005). The Articial Wilderness.
1491: New Revelations of the Americas Before
Columbus. New York: Alfred A. Knopf. pp. 315
318. ISBN 1-4000-4006-X.
Schorger, A. W. (1955). The Passenger Pigeon:
Its Natural History and Extinction. Madison, WI:
University of Wisconsin Press. ISBN 1-930665-962.
[120] Schorger 1955, p. 28

[121] D'Elia, J. (2010). Evolution of Avian Conservation
Breeding with Insights for Addressing the Current Extinction Crisis. Journal of Fish and Wildlife Management. Fish and Wildlife Service. 1 (2): 189210.
doi:10.3996/062010-JFWM-017.
7 External links
Project Passenger Pigeon: Lessons from the Past for
a Sustainable Future
[122] Schorger 1955, p. 27
The Demise of the Passenger Pigeon (as broadcast

on National Public Radio's Day to Day)
[123] Shell, H. R. (May 2004). The Face of Extinction.

Natural History. American Museum of Natural History.
113 (4): 72. ISSN 0028-0712.
360 Degree View of Martha, the Last Passenger Pigeon (Smithsonian Institution)
[124] Schorger 1955, p. 29

[125] Schorger 1955, p. 30
[126] Last Passenger Pigeon Dies. El Paso Morning Times. El
Paso, Texas. September 14, 1914. p. 6.
[127] Martha Passenger Pigeon Memorial Hut. Cincinnati,
Ohio: Roadside America. Retrieved February 29, 2012.
[128] Halliday, T. (1980). The extinction of the passenger pigeon Ectopistes migratorius and its relevance to contemporary conservation. Biological Conservation. 17 (2): 157
162. doi:10.1016/0006-3207(80)90046-4.
23
Text and image sources, contributors, and licenses
8.1
Text
Passenger pigeon Source: https://en.wikipedia.org/wiki/Passenger_pigeon?oldid=737181321 Contributors: Vicki Rosenzweig, Pinkunicorn, Eclecticology, William Avery, Zippy, Olivier, Frecklefoot, JohnOwens, Dante Alighieri, Shyamal, Tannin, Jimfbleak, Stevenj,
JohnCastle, Furrykef, Alight, Omegatron, Wetman, Catskul, Dale Arnett, Pigsonthewing, Kristof vt, Smallweed, Cholling, Mervyn, Hadal,
UtherSRG, Wereon, Kent Wang, JerryFriedman, Pengo, Alan Liefting, Ancheta Wis, Andy, Abigail-II, Wwoods, Everyking, Michael
Devore, Bobblewik, Andycjp, Alexf, Gdr, Yath, Sonjaaa, Williamb, Beland, MistToys, Kaldari, Mukerjee, Phil Sandifer, Howardjp,
Klemen Kocjancic, Esperant, Gazpacho, DanielCD, Rich Farmbrough, Guanabot, Fungus Guy, Vsmith, Xezbeth, Martpol, Bender235,
Evice, Mr. Billion, Dralwik, Circeus, Apostrophe, Phils, Anthony Appleyard, Hackwrench, Ashley Pomeroy, Kurieeto, Esrob, Stillnotelf,
Bart133, Aranae, SidP, Saga City, ReyBrujo, Shoey, Mikeo, Fred Condo, Richard Arthur Norton (1958- ), Firsfron, Woohookitty, MagicBez, Etacar11, Jpers36, Kosher Fan, Bennetto, Grika, John Hill, Isnow, Paxsimius, Graham87, Rjwilmsi, Alaney2k, Bruce1ee, Cjoev,
Leon Robbins, SchuminWeb, Eubot, Hottentot, Bubbleboys, RexNL, Preslethe, Hatch68, DVdm, Patientone, Gdrbot, Bgwhite, YurikBot,
Wavelength, Hairy Dude, NTBot~enwiki, Huw Powell, Phantomsteve, Igo4U, CambridgeBayWeather, Royalbroil, NawlinWiki, Dysmorodrepanis~enwiki, Spike Wilbury, Phoenix79, Lexicon, Apokryltaros, Brian Crawford, Melly42, Zwobot, Sforandy, Kortoso, DeadEyeArrow, Bota47, Asarelah, Doncram, Elkman, Strolls, Wknight94, StuRat, Wsiegmund, Npeters22, Vicarious, RenamedUser jaskldjslak904,
Sneftel, Mmcannis, Bibliomaniac15, True Pagan Warrior, SmackBot, Fireworks, CJLippert, Hux, Jereykopp, Pmaas, ScaldingHotSoup,
Jrockley, Delldot, Kintetsubualo, Ian Rose, Peter Isotalo, Gilliam, Ohnoitsjamie, Hmains, Teemu08, Hibernian, George Church, Ryecatcher773, Colonies Chris, Brinerustle, Scwlong, Tsca.bot, Abyssal, Nixeagle, Snowmanradio, Ritchie333, Bigturtle, Savidan, Kevlar67,
Bigheadjer, Devoblue, Salamurai, Thistheman, Kukini, Thor Dockweiler, Esrever, Xdamr, Swatjester, BrownHairedGirl, Agradman, John,
Butko, Mgiganteus1, IronGargoyle, Bensecs, BillFlis, Beetstra, Muadd, Mr Stephen, Dicklyon, Waggers, RomeoVoid, Peyre, Atakdoug,
Iridescent, Clarityend, Cotinis, DavidOaks, Ewulp, Civil Engineer III, Courcelles, Boucher4, Dixonsej, BruceGrubb, Ogmedina, J Milburn, Tpetross, CmdrObot, Iuio, Schweiwikist, AshLin, Neelix, Beastie Bot, AndrewHowse, Jac16888, Cydebot, Clayoquot, Gogo Dodo,
Khatru2, Casliber, Thijs!bot, Epbr123, Gaijin42, Felix Portier~enwiki, Missvain, Escarbot, The Obento Musubi, Manushand, Leuqarte,
Ironiridis, JAnDbot, AOB, Sarasays, Matthew Fennell, Michig, Maias, Tpholland, Magioladitis, WolfmanSF, Dekimasu, Doug Coldwell,
Bigdan201, Pvmoutside, Wormcast, LorenzoB, Urco, Woolla, Atulsnischal, Ulkomaalainen, R'n'B, Kylie Larson, Dinkytown, Pharaoh of
the Wizards, DrKay, Paul1953h, Rufous-crowned Sparrow, Johnbod, Worthadonkey, Ferahgo the Assassin, Jfer1958, Num1dgen, Dorftrottel, Scewing, Gregarious Oddfellow, VolkovBot, ABF, Je G., Bovineboy2008, MatsT, Philip Trueman, Fran Rogers, Oshwah, GDonato, Jdcrutch, Sandhillcrane, Broadbot, Sidewinder1, Vector Potential, Rontrigger, SieBot, Brenont, Calliopejen1, MeegsC, FunkMonk,
Aillema, Cupzonia, Ametrine, Mimihitam, Baseball Bugs, Gajeam, Lightmouse, Polbot, Amelia9mm, AngelLeliel, Manway, IdreamofJeanie, Mygerardromance, Faulknerfan, Pinkadelica, Grandmasterkush, ClueBot, Deanlaw, Cygnis insignis, SeaValeYen, Sting au, Mild
Bill Hiccup, Polybandrous, Mayy may y, Niceguyedc, Deadmanwalking28, Stylteralmaldo, Excirial, Gnome de plume, Jusdafax, Star
Mississippi, Nicols10~enwiki, 7&6=thirteen, Whatabout this, JasonAQuest, Dndnerd, PCHS-NJROTC, Berean Hunter, Wannabe99,
Rossen4, Tdslk, DumZiBoT, , Hotdogcolors, XLinkBot, Wikiuser100, Rogerthepigeon, Greasywheel, Cmr08, MystBot, Good
Olfactory, Addbot, DOI bot, GD 6041, Leszek Jaczuk, Cst17, Lost on Belmont, ThreeOfCups, Sussmanbern, Christopher140691,
Erutuon, Tide rolls, Lightbot, First Light, Ettrig, Drpickem, Luckas-bot, Yobot, Fraggle81, The Earwig, Richigi, AnomieBOT, AaRH,
Krasss, Piano non troppo, Materialscientist, Citation bot, Ruby2010, Quebec99, LilHelpa, Xqbot, Gigemag76, Tad Lincoln, Tomwsulcer,
GrouchoBot, Peeball, Energybender, Zupcic, FrescoBot, Simuliid, Archaeodontosaurus, Dger, MonickerToo, Craig Pemberton, JMilty,
Cjtsmith, Fredisfree, AstaBOTh15, Pinethicket, Metricmike, Edderso, Jonesey95, Simple Catherine Crerar, , Merlion444, White Shadows, Dutchmonkey9000, Gerda Arendt, Kgrad, Lotje, Dahendrix, Randomfactoids, Clarkcj12, YoMOTH3r, Warrah, Reaper Eternal,
YrHelperInfonut, Diannaa, Innotata, RjwilmsiBot, Galloping Moses, Wembwandt, Steve03Mills, Mukogodo, EmausBot, WikitanvirBot,
EricthePinko, Immunize, Luer, GoingBatty, Winner 42, Wikipelli, Ryguy611, Wertoy36, Thecheesykid, ZroBot, Tomgazer, Leaest
of Futures, Jimmieh, Jackson Montana, SpencerCollins, Rcsprinter123, Brandmeister, L Kensington, Totaldrama24, Donner60, Fanyavizuri, Cregagh, HandsomeFella, Gamecallmaster, Metamorphosed Fossil, ClueBot NG, Cwmhiraeth, BOMBINI, BarrelProof, Cestertonio,
Audreykkb, Helpful Pixie Bot, BG19bot, Maryphillips1952, MusikAnimal, Fabaceae, Davidiad, Mark Arsten, FiveColourMap, CitationCleanerBot, Harizotoh9, Ryday, Klilidiplomus, BattyBot, Millennium bug, EinarLugnet, Estaab, MrPurple20, Khazar2, Esszet, EagerToddler39, Dexbot, BigJolly9, Abcdefhijklmnopqrstufwxyz, Jamesx12345, Richard Bruce Bradford, Corinne, Joshua G Smyth, Passengerpigeon, Brh131, Ghent12, TFA Protector Bot, Hansmuller, Finnusertop, Sam Sailor, Colbey84, Inababy, Alanrhobson, Coreyemotela,
Strangejames, Scottyboyfa, Monkbot, Prof. Mc, Anderswarr, Burklemore1, Tim Hough, Ryanhacker, FACBot, Crossark, Terrapericolosa, Elmidae, LL221W, Hehehehehehehhehe, Continentaleurope, Cephas, My Chemistry romantic, Vincenttino, 15extinction15E, 13C
ext2015, Gulumeemee, Asdfsfasf and Anonymous: 397
8.2
Images
File:Acorns_small_to_large_cropped.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/5/58/Acorns_small_to_
large_cropped.jpg License: CC BY 2.0 Contributors: This le was derived from Acorns small to large.jpg: <a href='//commons.
wikimedia.org/wiki/File:Acorns_small_to_large.jpg' class='image'><img alt='Acorns small to large.jpg' src='https://upload.
wikimedia.org/wikipedia/commons/thumb/d/d8/Acorns_small_to_large.jpg/50px-Acorns_small_to_large.jpg' width='50' height='33'
srcset='https://upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Acorns_small_to_large.jpg/75px-Acorns_small_to_large.jpg
1.5x, https://upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Acorns_small_to_large.jpg/100px-Acorns_small_to_large.jpg 2x'
data-le-width='2048' data-le-height='1365' /></a>
Original artist: Acorns_small_to_large.jpg: David Hill
File:Alabama_bird_day_book_(1915)_(14751712302).jpg Source: https://upload.wikimedia.org/wikipedia/commons/3/3c/Alabama_
bird_day_book_%281915%29_%2814751712302%29.jpg License: No restrictions Contributors: https://www.flickr.com/photos/
internetarchivebookimages/14751712302/ Original artist: Internet Archive Book Images
File:Bird-lore_(1913)_(14568824740).jpg Source:
https://upload.wikimedia.org/wikipedia/commons/b/b1/Bird-lore_%281913%
29_%2814568824740%29.jpg License: No restrictions Contributors: https://www.flickr.com/photos/internetarchivebookimages/
14568824740/ Original artist: J. G. Hubbard Internet Archive Book Images
File:Bird_lore_(1913)_(14562367319).jpg Source:
https://upload.wikimedia.org/wikipedia/commons/b/ba/Bird_lore_%281913%
29_%2814562367319%29.jpg License: Public domain Contributors: https://www.flickr.com/photos/internetarchivebookimages/
24
8 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES
14562367319/ Original artist: Internet Archive Book Images

File:Bird_lore_(1913)_(14562557107).jpg Source:
https://upload.wikimedia.org/wikipedia/commons/0/03/Bird_lore_%281913%
29_%2814562557107%29.jpg License: Public domain Contributors: https://www.flickr.com/photos/internetarchivebookimages/
14562557107/ also at Wisconsin Historical society page on passenger pigeon photographs and photo page Original artist: J. G. Hubbard,
Internet Archive Book Images
File:Birdonnest.jpg Source: https://upload.wikimedia.org/wikipedia/commons/1/1f/Birdonnest.jpg License: Public domain Contributors:
https://archive.org/stream/birdlore151913nati/birdlore151913nati#page/101/mode/1up and http://www.wisconsinhistory.
org/whi/fullRecord.asp?id=53463&qstring=http%3A%2F%2Fwww.wisconsinhistory.org%2Fwhi%2Fresults.asp%3Fkeyword1%
3DPassenger%2520Pigeon%2520Specimen%2520Photographs%2C%25201896%26search_field1%3Dcollection_name%26search_
type%3Dadvanced%26sort_by%3Ddate%26boolean_type1%3Dand%26boolean_type2%3Dand Original artist: J. G. Hubbard
File:Commons-logo.svg Source: https://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License: CC-BY-SA-3.0 Contributors: ? Original artist: ?
File:Decoy_Passenger_Pigeon.jpg Source:
https://upload.wikimedia.org/wikipedia/commons/7/7d/Decoy_Passenger_Pigeon.jpg
License: Public domain Contributors: Paxson, Henry D. (1917). "The last of the Wild Pigeon in Bucks County". Collection of papers
read before the Bucks county historical society 4: 367-382. Original artist: Unknown<a href='//www.wikidata.org/wiki/Q4233718'
title='wikidata:Q4233718'><img
alt='wikidata:Q4233718'
src='https://upload.wikimedia.org/wikipedia/commons/thumb/f/ff/
Wikidata-logo.svg/20px-Wikidata-logo.svg.png' width='20' height='11' srcset='https://upload.wikimedia.org/wikipedia/commons/
thumb/f/ff/Wikidata-logo.svg/30px-Wikidata-logo.svg.png
1.5x,
https://upload.wikimedia.org/wikipedia/commons/thumb/f/ff/
Wikidata-logo.svg/40px-Wikidata-logo.svg.png 2x' data-le-width='1050' data-le-height='590' /></a>
File:Ectopistes_migratoriusAAP042CA.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/9/97/Ectopistes_
migratoriusAAP042CA.jpg License: Public domain Contributors: Birds of New York (New York State Museum. Memoir 12), Albany: University of the State of New York. Plates by Fuertes later reproduced in Birds of America (1917?) by Thomas Gilbert Pearson
(1873-1943) et al. Original artist: Louis Agassiz Fuertes
File:Ectopistes_migratoriusMCN2P28CA.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/d/d0/Ectopistes_
migratoriusMCN2P28CA.jpg License: Public domain Contributors: Orthogenetic Evolution in the Pigeons Original artist: K. Hayashi
File:Ectopistes_migratorius_(passenger_pigeon).jpg Source: https://upload.wikimedia.org/wikipedia/commons/0/00/Ectopistes_
migratorius_%28passenger_pigeon%29.jpg License: CC BY 2.0 Contributors: Ectopistes migratorius (passenger pigeon) 5 Original artist:
James St. John
File:Ectopistes_migratorius_ULaval_2.jpg Source: https://upload.wikimedia.org/wikipedia/commons/f/f5/Ectopistes_migratorius_
ULaval_2.jpg License: CC BY-SA 3.0 Contributors: Own work Original artist: Cephas
File:ExtinctDodoBird.jpeg Source: https://upload.wikimedia.org/wikipedia/commons/e/e0/ExtinctDodoBird.jpeg License: Public domain Contributors: ? Original artist: (anonymous artwork ; facsimile of 1626 painting by Roelant Savery)
File:Flock_of_passenger_pigeons.jpg Source: https://upload.wikimedia.org/wikipedia/commons/a/a8/Flock_of_passenger_pigeons.
jpg License: Public domain Contributors: http://www.jstor.org/stable/4073624?seq=1#page_scan_tab_contents Original artist: 1921
File:Free-to-read_lock_75.svg Source: https://upload.wikimedia.org/wikipedia/commons/8/80/Free-to-read_lock_75.svg License: CC0
Contributors:
Adapted
from
<a
href='//en.wikipedia.org/wiki/File:Open_Access_logo_PLoS_white_green.svg'
class='image'
title='Open_Access_logo_PLoS_white_green.svg'><img
alt='Open_Access_logo_PLoS_white_green.svg'
src='//upload.wikimedia.
org/wikipedia/commons/thumb/9/90/Open_Access_logo_PLoS_white_green.svg/9px-Open_Access_logo_PLoS_white_green.svg.png'
width='9' height='14' srcset='//upload.wikimedia.org/wikipedia/commons/thumb/9/90/Open_Access_logo_PLoS_white_green.svg/
14px-Open_Access_logo_PLoS_white_green.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/90/Open_Access_
logo_PLoS_white_green.svg/18px-Open_Access_logo_PLoS_white_green.svg.png 2x' data-le-width='640' data-le-height='1000'
/></a>
Original artist:
This version:Trappist_the_monk (talk) (Uploads)
File:Last_known_Passenger_Pigeon.jpg Source: https://upload.wikimedia.org/wikipedia/commons/8/8c/Last_known_Passenger_
Pigeon_-_Stierch.jpg License: CC BY 2.0 Contributors: Flickr: Last known Passenger Pigeon Original artist: Sarah Stierch
File:Map-Ectopistes-migratorius.png Source: https://upload.wikimedia.org/wikipedia/commons/8/87/Map-Ectopistes-migratorius.
png License: CC BY-SA 1.0 Contributors: Own work, based on map compiled by researcher Arlie W. Schorger in the early 1950s (published
in Schorger, A. W. (1955). The Passenger Pigeon: Its Natural History and Extinction. Madison, WI: University of Wisconsin Press. ISBN
1-930665-96-2.) Original artist: Valrie Chansigaud
File:Martha,_the_last_Passenger_Pigeon._Natural_History_Museum,_June,_2015._Digital_photo,_cropped_and_brightened.
jpg Source:
https://upload.wikimedia.org/wikipedia/commons/e/e7/Martha%2C_the_last_Passenger_Pigeon._Natural_History_
Museum%2C_June%2C_2015._Digital_photo%2C_cropped_and_brightened.jpg License: CC BY-SA 4.0 Contributors: Own work
Original artist: Ph0705
File:Martha_last_passenger_pigeon_1914.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/c/cc/Martha_last_
passenger_pigeon_1914.jpg License: Public domain Contributors: http://archive.org/stream/publishedfigures00shuf#page/466/mode/2up
Original artist: Enno Meyer
File:Mershon{}s_The_Passenger_Pigeon_(Audubon_plate,_crop).jpg Source: https://upload.wikimedia.org/wikipedia/commons/4/
4b/Mershon%27s_The_Passenger_Pigeon_%28Audubon_plate%2C_crop%29.jpg License: Public domain Contributors: https://archive.
org/stream/birdsofamericafr05audu#page/n39/mode/2up Original artist: John James Audubon
File:Mershon{}s_The_Passenger_Pigeon_(frontispiece,_crop).jpg Source: https://upload.wikimedia.org/wikipedia/commons/d/d5/
Mershon%27s_The_Passenger_Pigeon_%28frontispiece%2C_crop%29.jpg License: Public domain Contributors: Obtained from online
scan at passengerpigeon00mers, crop with some border of scanned paper, for relative colour balance. Slightly rotated. Original artist: Louis
Agassiz Fuertes
8.2
Images
25
File:Naturalis_Biodiversity_Center_-_RMNH.AVES.110088_-_Ectopistes_migratorius_(Linnaeus,_1766)_-_Passenger_
Pigeon_-_specimen_-_video.webm Source: https://upload.wikimedia.org/wikipedia/commons/9/9d/Naturalis_Biodiversity_Center_
-_RMNH.AVES.110088_-_Ectopistes_migratorius_%28Linnaeus%2C_1766%29_-_Passenger_Pigeon_-_specimen_-_video.webm
License: CC BY-SA 3.0 Contributors: Naturalis Original artist: Linnaeus, 1766
Pigeon_-_specimen_-_video.webm Source: https://upload.wikimedia.org/wikipedia/commons/6/6a/Naturalis_Biodiversity_Center_
Pigeon_-_specimen_-_video.webm Source: https://upload.wikimedia.org/wikipedia/commons/5/55/Naturalis_Biodiversity_Center_
File:Nestandegg.jpg Source:
https://upload.wikimedia.org/wikipedia/commons/f/f0/Nestandegg.jpg License:
Public domain Contributors:
https://archive.org/stream/birdlore151913nati/birdlore151913nati#page/96/mode/1up Original artist:
Unknown<a
href='//www.wikidata.org/wiki/Q4233718'
title='wikidata:Q4233718'><img
alt='wikidata:Q4233718'
src='https:
//upload.wikimedia.org/wikipedia/commons/thumb/f/ff/Wikidata-logo.svg/20px-Wikidata-logo.svg.png'
width='20'
height='11'
srcset='https://upload.wikimedia.org/wikipedia/commons/thumb/f/ff/Wikidata-logo.svg/30px-Wikidata-logo.svg.png
1.5x,
https://upload.wikimedia.org/wikipedia/commons/thumb/f/ff/Wikidata-logo.svg/40px-Wikidata-logo.svg.png 2x' data-le-width='1050'
data-le-height='590' /></a>
File:Passenger_Pigeon_Net_Cockburn_1829.jpg Source: https://upload.wikimedia.org/wikipedia/commons/7/76/Passenger_Pigeon_
Net_Cockburn_1829.jpg License: Public domain Contributors: This image is available from Library and Archives Canada under the reproduction reference number C-012539 and under the MIKAN ID number 2836130
Original artist: James Pattison Cockburn
File:Passenger_Pigeon_in_flight.jpg Source: https://upload.wikimedia.org/wikipedia/commons/c/c9/Passenger_Pigeon_in_flight.jpg
License: CC BY 2.0 Contributors: Extinct Passenger Pigeon (Ectopistes migratorius) Original artist: Jim, the Photographer from Springeld
PA, United States of America
File:Passenger_Pigeon_organs.jpg Source: https://upload.wikimedia.org/wikipedia/commons/f/fe/Passenger_Pigeon_organs.jpg License: Public domain Contributors: Schufeldt, R. W. (1915). Anatomical and other notes on the Passenger Pigeon (Ectopistes migratorius)
lately living in the Cincinnati Zoological Gardens. Auk 32(1): 29-41. https://archive.org/stream/jstor-4071611/4071611#page/n11/mode/
2up Original artist: R.W. Schufeldt
File:Passenger_Pigeon_skeleton.jpg Source: https://upload.wikimedia.org/wikipedia/commons/6/6c/Passenger_Pigeon_skeleton.jpg
License: Public domain Contributors: http://archive.org/stream/jstor-4071953/4071953#page/n1/mode/2up Original artist: Shufeldt, R.
W.
File:Passenger_pigeon.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/c/cb/Passenger_pigeon.jpg
License:
https://archive.org/stream/birdlore151913nati/birdlore151913nati#page/91/mode/1up
and
Public
domain
Contributors:
http://www.wisconsinhistory.org/whi/fullRecord.asp?id=53450&qstring=http%3A%2F%2Fwww.wisconsinhistory.org%2Fwhi%
2Fresults.asp%3Fkeyword1%3DPassenger%2520Pigeon%2520Specimen%2520Photographs%2C%25201896%26search_field1%
3Dcollection_name%26search_type%3Dadvanced%26sort_by%3Ddate%26boolean_type1%3Dand%26boolean_type2%3Dand
Original artist: J. G. Hubbard
File:Passenger_pigeon_aviary.jpg Source: https://upload.wikimedia.org/wikipedia/commons/4/44/Passenger_pigeon_aviary.jpg License: Public domain Contributors: http://www.wisconsinhistory.org/whi/fullRecord.asp?id=53454 Original artist: Hubbard, J. G.
File:Passenger_pigeon_capture.jpg Source: https://upload.wikimedia.org/wikipedia/commons/b/ba/Passenger_pigeon_capture.jpg License: Public domain Contributors: http://boc-online.org/passenger-pigeons-large-scale-live-capture/ Original artist: Frank Leslies Illustrated News (vol. LII, no. 1344, pp. 299300)
File:Passenger_pigeon_notes.jpg Source: https://upload.wikimedia.org/wikipedia/commons/d/d2/Passenger_pigeon_notes.jpg License:
Public domain Contributors: https://archive.org/stream/jstor-4071160/4071160#page/n9/mode/2up Original artist: Wallace Craig
File:Passenger_pigeon_shoot.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/8/89/Passenger_
pigeon_shoot.jpg
License:
Public
domain
Contributors:
https://www.washingtonpost.com/national/health-science/
scientists-look-to-revive-the-long-extinct-passenger-pigeon/2013/07/08/3d1323d4-b9a1-11e2-aa9e-a02b765ff0ea_story.html Original
artist: Smith Bennett[2]
File:Passenger_pigeon_shooting_in_Iowa.jpg Source: https://upload.wikimedia.org/wikipedia/commons/6/66/Passenger_pigeon_
shooting_in_Iowa.jpg License: Public domain Contributors: http://boc-online.org/passenger-pigeons-large-scale-live-capture/ Original
artist: Frank Leslies Illustrated News (vol. XXV, no. 625, p. 8)
File:Patagioenas_fasciata_-San_Luis_Obispo,_California,_USA-8_(1).jpg Source:
https://upload.wikimedia.org/wikipedia/
commons/b/ba/Patagioenas_fasciata_-San_Luis_Obispo%2C_California%2C_USA-8_%281%29.jpg License: CC BY 2.0 Contributors:
Band-tailed pigeon, (Patagioenas fasciata) Original artist: aroid from San Luis Obispo, CA, USA
File:Pigeon-of-passage.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/e2/Pigeon-of-passage.jpg License: Public domain Contributors: The Natural History of Carolina, Florida and Bahamas V. 1 (1754) Original artist: Mark Catesby, George Edwards
File:Pigeon_migrateur_MHNT.jpg Source: https://upload.wikimedia.org/wikipedia/commons/5/5e/Pigeon_migrateur_MHNT.jpg License: CC BY-SA 3.0 Contributors: Own work Original artist: Didier Descouens
File:Wikispecies-logo.svg Source: https://upload.wikimedia.org/wikipedia/commons/d/df/Wikispecies-logo.svg License: CC BY-SA
3.0 Contributors: Image:Wikispecies-logo.jpg Original artist: (of code) cs:User:-xfi File:Young_passenger_pigeon.jpg Source: https://upload.wikimedia.org/wikipedia/commons/d/d3/Young_passenger_pigeon.jpg License: Public domain Contributors: https://archive.org/stream/birdlore151913nati/birdlore151913nati#page/97/mode/1up (original photo
at [1]) Original artist: J. G. Hubbard
26
8 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES
File:Zenaida_macroura_-California-8-2c.jpg Source: https://upload.wikimedia.org/wikipedia/commons/7/75/Zenaida_macroura_

-California-8-2c.jpg License: CC BY-SA 2.0 Contributors: originally posted to Flickr as Dove by Almaden Lake and uploaded to commons
as Zenaida_macroura_-California-8.jpg Original artist:
derivative work:
Snowmanradio
Snowmanradio'>talk</a>)
8.3
(<a
Content license
Creative Commons Attribution-Share Alike 3.0
href='//commons.wikimedia.org/wiki/User_talk:Snowmanradio'
title='User
talk:

Passenger Pigeon

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Passenger Pigeon

Uploaded by

Copyright:

Available Formats

Passenger pigeon

Earliest published illustration of the species (a male), Mark

Swedish naturalist Carl Linnaeus coined the binomial

Passenger pigeons were hunted by Native Americans, but

In 1827 William John Swainson moved the passenger

The physically similar mourning dove is not closely related.

The passenger pigeon was a member of the pigeon and

The genus name, Ectopistes, translates as moving about

Turnaround video of an adult male specimen at Naturalis Biodiversity Center

Turnaround video of a juvenile female specimen at Naturalis

The juvenile passenger pigeon was similar in plumage to

Skeleton of a male bird, 1914

The passenger pigeon was physically adapted for speed,

Musical notes documenting male vocalizations, compiled by

Distribution and habitat

The passenger pigeon was found across most of North

they were possibly not restricted to one type, as long as

4 ECOLOGY AND BEHAVIOR

Ecology and behavior

with inconceivable velocity, mounted perpendicularly so as to resemble a vast column, and,

Live male in Whitmans aviary, 1896/98

The passenger pigeon was nomadic, constantly migrating

Illustration of migrating ocks, Frank Bond, 1920

The passenger pigeon was one of the most social of all

Alert parent bird posing deantly towards the camera

A communally roosting species, the passenger pigeon

water to drink.[38] One of the primary causes of natural

4 ECOLOGY AND BEHAVIOR

to have died as a result in the following decades, but this

Acorns in South Carolina, among the diet of this bird

Internal organs of Martha, the last individual: cr. denotes the

in the back of the ock ew to the front in order to pick

Nesting captive bird, wary of the photographer

Other than nding roosting sites, the migrations of the

4 ECOLOGY AND BEHAVIOR

been swept clean, due to all the twigs being collected at

After observing captive birds, Wallace Craig found that

Predators and parasites

Two parasites have been recorded on passenger pigeons.

5 Relationship with humans

The passenger pigeon played a religious role for some

5 RELATIONSHIP WITH HUMANS

chief, as they were served whenever the chieftain gave

Depiction of a shooting in Louisiana, Smith Bennett, 1875

pigeons, and instead ate only the juveniles as they were

1881 spread showing methods of trapping pigeons for shooting

fodder. Once pigeon meat became popular, commercial

5 RELATIONSHIP WITH HUMANS

Pigeon net in Canada, by James Pattison Cockburn, 1829

Decline and conservation attempts

Decline and conservation attempts

Life drawing by Charles R. Knight, 1903

Male and female by Louis Agassiz Fuertes, frontispiece of

The notion that the species could be driven to extinction

5 RELATIONSHIP WITH HUMANS

ineective, as they were unclearly framed and hard to

this is the most often cited last wild specimen, in 2014,

Whitmans aviary with passenger pigeons and other species,

Martha, the last passenger pigeon, alive in 1912

5.4 Extinction causes

5 RELATIONSHIP WITH HUMANS

Martha at the Smithsonian Museum, 2015

5.5 Re-creation of the species

ger pigeons needed to sustain the population. In contrast,