Simia troglodytes Blumenbach, 1776
Troglodytes troglodytes (Blumenbach, 1776)
Troglodytes niger E. Geoffroy, 1812
Pan niger (E. Geoffroy, 1812) Anthropopithecus troglodytes (Sutton,
The common chimpanzee (Pan troglodytes), also known as the robust
chimpanzee, is a species of great ape. Colloquially, the common
chimpanzee is often called the chimpanzee (or "chimp"), though this
term can be used to refer to both species in the genus Pan: the common
chimpanzee and the closely related bonobo, formerly called the pygmy
chimpanzee. Evidence from fossils and
DNA sequencing shows both
species of chimpanzee are the sister taxon to the modern human
The common chimpanzee is covered in coarse black hair, but has a bare
face, fingers, toes, palms of the hands, and soles of the feet. It is
considered more robust than the bonobo, weighing between 40 and
65 kg (88 and 143 lb) and measuring about 63 to 94 cm
(25 to 37 in). Its gestation period is eight months. The infant
is weaned at about three years old, but usually maintains a close
relationship with its mother for several more years; it reaches
puberty at the age of eight to 10. Its lifespan in the wild is 36
years and its lifespan in captivity is about 50 years.
The common chimpanzee lives in groups which range in size from 15 to
150 members, although individuals travel and forage in much smaller
groups during the day. The species lives in a male-dominated, strict
hierarchy, which means disputes can generally be settled without the
need for violence. Nearly all chimpanzee populations have been
recorded using tools, modifying sticks, rocks, grass and leaves and
using them for acquiring honey, termites, ants, nuts and water. The
species has also been found creating sharpened sticks to spear Senegal
bushbabies out of small holes in trees.
The common chimpanzee is listed on the
IUCN Red List
IUCN Red List as an endangered
species. Between 170,000 and 300,000 individuals are estimated across
its range in the forests and savannahs of West and Central Africa. The
biggest threats to the common chimpanzee are habitat loss, poaching
2 Evolutionary history
5.1 Group structure
5.2 Mating and parenting
5.4 Tool use
6 Chimpanzees and humans
6.1 In culture
6.2 Field study
6.3 Genome Project
6.5 Link with human immunodeficiency virus type 1
6.6 Status and conservation
7 See also
9 External links
The common chimpanzee was named
Simia troglodytes by Johann Friedrich
Blumenbach in 1776;
Lorenz Oken moved it to the new genus Pan in 1816.
The species name troglodytes is a reference to the Troglodytae
(literally "cave-goers"), an African people described by Greco-Roman
geographers. Blumenbach first used it in his De generis humani
varietate nativa liber ("[Book] on the natural varieties of the human
genus") in 1776. This book was based on his dissertation
presented one year before (it had a date 16 September 1775 printed on
its title page) to the
University of Göttingen
University of Göttingen for internal use
only, thus the dissertation did not meet the conditions for
published work in the sense of zoological nomenclature.
The English name "chimpanzee" is first recorded in 1738. It is derived
Tshiluba language term kivili-chimpenze, with a meaning of
"mockman" or possibly just "ape". The colloquialism "chimp" was most
likely coined some time in the late 1870s.
view • discuss • edit
Earliest stone tools
Earliest exit from Africa
Earliest fire use
Earliest in Europe
Axis scale: million years
Also see: Life timeline and Nature timeline
Despite a large number of
Homo fossil finds, chimpanzee fossils (genus
Pan) were not described until 2005. Existing chimpanzee populations in
Central Africa do not overlap with the major human fossil
sites in East Africa. However, chimpanzee fossils have now been
reported from Kenya. This would indicate that both humans and members
of the Pan clade were present in the
East African Rift
East African Rift Valley during
the Middle Pleistocene.
DNA evidence suggests the bonobo and common chimpanzee species
separated from each other less than one million years ago (similar in
Homo sapiens and Neanderthals). The chimpanzee
line split from the last common ancestor of the human line around six
million years ago. Because no species other than
Homo sapiens has
survived from the human line of that branching, both chimpanzee
species are the closest living relatives of humans. The lineage of
humans and chimpanzees diverged from that of the gorilla about seven
million years ago. A 2003 study argues the common chimpanzee should be
included in the human branch as
Homo troglodytes, and notes experts
say many scientists are likely to resist the reclassification,
especially in the emotionally-charged and often disputed field of
Four subspecies of the common chimpanzee have been recognised,
with the possibility of a fifth:
Central chimpanzee or tschego, P. t. troglodytes, in Cameroon, the
Central African Republic, Equatorial Guinea, Gabon, the Republic of
the Congo, and the Democratic Republic of the Congo
Western chimpanzee, P. t. verus, in Guinea, Guinea-Bissau, Mali,
Senegal, Sierra Leone, Liberia, Ivory Coast, and Ghana
Cameroon chimpanzee, P. t. ellioti (also known as P. t.
Nigeria and Cameroon
Eastern chimpanzee, P. t. schweinfurthii, in the Central African
Republic, South Sudan, the Democratic Republic of the Congo, Uganda,
Rwanda, Burundi, Tanzania, and Zambia
Southeastern chimpanzee, P. t. marungensis, in Burundi, Rwanda,
Tanzania, and Uganda:
Colin Groves argues that this subspecies is the
result of enough variation between the northern and southern
populations of P. t. schweinfurthii.
Close-up of face, at Kibale National Park, Uganda
The skeleton of chimpanzee in La Plata Museum, Argentine
The adult male common chimpanzee weighs between 40 and 60 kg (88
and 132 lb), the female weighs 32 to 47 kg (71 to
104 lb). However, large wild males can weigh up to 70 kg
(150 lb) and males in captivity, such as Travis the Chimp, have
reached 91 kg (201 lb). Head-body length (from
the nose to the rump while on all fours) ranges from 63 to 94 cm
(25 to 37 in). Males can measure up to 1.6 m
(5 ft 3 in) tall while standing and females up to 1.3 m
(4 ft 3 in) tall. Their bodies are covered by coarse, black
hair, except for the face, fingers, toes, palms of the hands, and
soles of the feet. Both its thumbs and big toes are opposable,
allowing a precise grip. The common chimpanzee is both arboreal and
terrestrial, and spends its nights in the trees, while most daylight
hours are spent on the ground.
Close-up of hand at Augsburg Zoo
Close-up of foot at Augsburg Zoo
Its habitual gait is quadrupedal, using the soles of its feet and
resting on its knuckles, but it can walk upright for short distances.
The common chimpanzee is a 'knuckle walker', like the gorilla and the
bonobo, in contrast to the quadrupedal locomotion of the
orangutan, a 'palm walker' that uses the outside edge of its palms. It
is the anatomically closest relative of the human.
The common chimpanzee is a highly adaptable species. It lives in a
variety of habitats, including dry savanna, evergreen rainforest,
montane forest, swamp forest and dry woodland-savanna mosaic.
In Gombe, the chimpanzee lives in subalpine moorland, open woodland,
semideciduous forest, evergreen forest, and grassland with scattered
trees. At Bossou, the chimpanzee inhabits multistage secondary
deciduous forests, which have grown after shifting cultivation, as
well as primary forests and grasslands. At Taï, it can be found
in the last remaining tropical rain forest in Ivory Coast.
The chimpanzee has an advanced cognitive map of its home range and can
repeatedly find food. The chimpanzee makes a night nest in a tree
in a new location every night, with every chimpanzee in a separate
nest other than infants or juvenile chimpanzees, which sleep with
Leopard predation is apparently a significant cause
of mortality in chimpanzees at Taï and Lopé National Parks.
Chimps are generally hostile towards leopards and may mob the
predators and even kill their cubs. Lions may have preyed on the
chimpanzees at Mahale Mountains National Park, where at least four
chimpanzees could have fallen prey to them. Although no other
instances of lion predation on chimpanzees have been recorded, the
larger group sizes of savanna chimps may have developed as a response
to threats from these big cats. Isolated cases of cannibalism have
also been documented.
A mother chimpanzee with young offspring eating Ficus fruit in Kibale
National Park, Uganda
The chimpanzee is an omnivorous frugivore. It prefers fruit above all
other food items and even seeks out and eats them when they are not
abundant. It also eats leaves and leaf buds, seeds, blossoms, stems,
pith, bark and resin. Insects and meat make up a small proportion of
their diet, estimated as 2%. While the common chimpanzee is
mostly herbivorous, it does eat honey, soil, insects, birds and their
eggs, and small to medium-sized mammals, including other
primates. The western red colobus ranks at the top of
preferred mammal prey. Other mammalian prey include red-tailed
monkeys, yellow baboons, blue duikers, bushbucks, and common
Despite the fact that common chimpanzees are known to hunt, and to
collect insects and other invertebrates, such food actually makes up a
tiny portion of their diet, from as little as 2% yearly to as much as
65 grams of animal flesh per day for each adult chimpanzee in peak
hunting seasons. This also varies from troop to troop and year to
year. However, in all cases, the majority of their diet consists of
fruits, leaves, roots, and other plant matter. Female
chimpanzees appear to consume much less animal flesh than males,
according to several studies. Goodall documented many occasions
Gombe Stream National Park
Gombe Stream National Park of chimpanzees and western red
colobus monkeys ignoring each other within close proximity.
It is suspected that human observers can influence chimpanzee
behavior. It is suggested that drones, camera traps and remote
microphones should be used rather than human observers.
Common chimpanzees live in communities that typically range from 20 to
more than 150 members, but spend most of their time traveling in
small, temporary groups consisting of a few individuals, "which may
consist of any combination of age and sex classes." Both males and
females sometimes travel alone. The common chimpanzee lives in a
fission-fusion society and may be found in groups of these types:
all-male, adult females and offspring, both sexes, or one female and
her offspring. Chimpanzees have complex social relationships and spend
a large amount of time grooming each other.
At the core of social structures are males, which roam around, protect
group members, and search for food. Males remain in their natal
communities, while females generally emigrate at adolescence. As such,
males in a community are more likely to be related to one another than
females are to each other. Among males is generally a dominance
hierarchy, and males are dominant over females. However, this
unusual fission-fusion social structure, "in which portions of the
parent group may on a regular basis separate from and then rejoin the
rest," is highly variable in terms of which particular individual
chimpanzees congregate at a given time. This is mainly due to
chimpanzees having a high level of individual autonomy within their
fission-fusion social groups. Also, communities have large ranges that
overlap with those of other groups.
Chimpanzees grooming one another
As a result, individual chimpanzees often forage for food alone, or in
smaller groups (as opposed to the much larger "parent" group, which
encompasses all the chimpanzees which regularly come into contact and
congregate into parties in a particular area). As stated, these
smaller groups also emerge in a variety of types, for a variety of
purposes. For example, an all-male troop may be organized to hunt for
meat, while a group consisting of lactating females serves to act as a
"nursery group" for the young. An individual may encounter certain
individuals quite frequently, but have run-ins with others almost
never or only in large-scale gatherings. Due to the varying frequency
at which chimpanzees associate, the structure of their societies is
Male chimpanzees exist in a linear dominance hierarchy. Top-ranking
males tend to be aggressive even during dominance stability. This
is likely due to the chimp’s fission-fusion society, with male
chimps leaving groups and returning after extended periods of time.
With this, a dominant male is unsure if any "political maneuvering"
has occurred and must re-establish his dominance. Thus, a large amount
of aggression occurs 5–15 minutes after a reunion. During
aggressive encounters, displays are preferred over attacks.
Male chimpanzees in Mahale National Park, Tanzania
Males maintain and improve their social ranks by forming coalitions,
which have been characterized as "exploitative" and are based on an
individual’s influence in agonistic interactions. Being in a
coalition allows males to dominate a third individual when they could
not by themselves, as politically apt chimps can exert power over
aggressive interactions regardless of their rank. Coalitions can also
give an individual male the confidence to challenge a dominant male.
The more allies a male has, the better his chance of becoming
dominant. However, most changes in hierarchical rank are caused by
Chimpanzee alliances can be very fickle and
one member may turn on another if it serves him.
Low-ranking males commonly switch sides in disputes between more
dominant individuals. Low-ranking males benefit from an unstable
hierarchy and have increased sexual opportunities. In
addition, conflicts between dominant males cause them to focus on each
other rather than the lower-ranking males. Social hierarchies among
adult females tend to be weaker. Nevertheless, the status of an adult
female may be important for her offspring. Females in Taï have
also been recorded to form alliances.
Social grooming appears to
be important in the formation and maintenance of coalitions. It is
more common among adult males than adult females.
Chimpanzees have been described as highly territorial and are known to
kill other chimps, although Margaret Power wrote in her 1991 book
The Egalitarians that the field studies from which the aggressive data
came, Gombe and Mahale, use artificial feeding systems that increased
aggression in the chimpanzee populations studied, so might not reflect
innate characteristics of the species as a whole. In the years
following her artificial feeding conditions at Gombe, Jane Goodall
described groups of male chimps patrolling the borders of their
territory, brutally attacking chimps which had split off from the
Gombe group. A study published in 2010 found that the chimpanzees wage
wars over land, not mates. Patrol parties from smaller groups are
more likely to avoid contact with their neighbors. Patrol parties from
large groups even take over a smaller group's territory, gaining
access to more resources, food, and females.
Mating and parenting
Common chimpanzee infant and mother
Chimpanzees mate throughout the year, although the number of females
in oestrus varies seasonally in a group. Female chimps are
more likely to come into oestrus when food is readily available.
Oestrous females exhibit sexual swellings. Chimps tend to be
promiscuous, and during estrus females mate with several males in her
community, and males have large testicles for sperm competition.
However, other forms of mating also exist. A community's dominant
males sometimes restrict reproductive access to females. A male and
female can form consortship and mate outside their community. In
addition, females sometimes leave their communities and mate with
males from neighboring communities.
These alternative mating strategies give females more mating
opportunities without losing the support of the males in their
community. Infanticide has been recorded in chimp communities in
Gombe, Mahale, and Kibale National Parks. Male chimps practice
infanticide on unrelated young to shorten the interbirth intervals in
the females. Also, accounts of infanticide by females have been
reported; cases of female infanticide may be related to the dominance
hierarchy in females or simply isolated pathological
Care for the young is provided mostly by their mothers. The survival
and emotional health of the young is dependent on maternal care.
Mothers provide their young with food, warmth, and protection, and
teach them certain skills. In addition, a chimp’s future rank may be
dependent on its mother’s status. For their first 30 days,
infants cling to their mother's bellies. Newborn chimps are helpless;
their grasping reflex is not strong enough to support them for more
than a few seconds. Infants are unable to support their own weight for
their first two months and need their mothers' support.
When they reach five to six months, infants ride on their mothers’
backs. They remain in continual contact for the rest of their first
year. When they reach two years of age, they are able to move and sit
independently. By three years, infants move farther away from
their mothers. By four to six years, chimps are weaned and infancy
The juvenile period for chimps lasts from their sixth to ninth years.
Juveniles remain close to their mothers, but they also have more
interactions with other members of their community. Adolescent females
move between groups and are supported by their mothers in agonistic
encounters. Adolescent males spend time with adult males in social
activities like hunting and boundary patrolling.
Chimpanzee alarm hoo
Chimpanzee alarm call, ogg/
Chimpanzee soft hoo
Chimpanzee soft hoo, ogg/
Chimpanzee "waa bark"
Chimpanzee "waa bark", ogg/
Problems playing these files? See media help.
Chimpanzees use a variety of facial expressions, postures and sounds
to communicate with each other. Chimps have expressive faces which
are important in close-up communications. When frightened, a "full
closed grin" causes nearby individuals to be fearful, as well. Other
facial expressions include the "lip flip", "pout", "sneer", and
"compressed-lips face". When submitting to a conspecific, a chimp
crunches, bobs, and extends a hand. When in an aggressive mode, a
chimp swaggers bipedally, hunched over and arms waving, in an attempt
to exaggerate its size. Chimps beat their hands and feet against
the trunks of large trees, an act known as "drumming".
Vocalizations are also important in chimp communication. The most
common and important call in adults is the "pant-hoot". These calls
are made when individuals are excited. Pant-hoots are made of four
parts, starting with soft "hoos" that get louder and louder and climax
into screams and sometimes barks; the former die down to soft "hoos"
again as the call ends. Submissive individuals will make
"pant-grunts" towards their superiors. Chimps use distance
calls to draw attention to danger, food sources, or other community
members. "Barks" may be made as "short barks" when hunting and
"tonal barks" when sighting large snakes.
Further information: Tool use by animals
Chimps using tools
Nearly all chimpanzee populations have been recorded using tools. They
modify sticks, rocks, grass, and leaves and use them when foraging for
honey, termites, ants, nuts, and water. Despite the lack of
complexity, forethought and skill are seen in making these tools and
should be considered such. While it has been known since Jane
Goodall's 1960s discovery that modern chimpanzees use tools, research
published in 2007 indicates chimpanzee stone tool use dates to at
least 4,300 years ago.
A common chimpanzee from the
Kasakela chimpanzee community was the
first nonhuman animal reported making a tool, by modifying a twig to
use as an instrument for extracting termites from their
mound. At Taï, chimps simply use their hands to extract
termites. When foraging for honey, chimps use modified short
sticks to scoop the honey out of the hive, that is, if the bees are
stingless. For hives of the dangerous African honeybees, chimps use
longer and thinner sticks to extract the honey. Chimps also fish
for ants using the same tactic.
Ant dipping is difficult and some chimps never master it. West African
chimps crack open hard nuts with stones or branches. Some
forethought in this activity is apparent, as these items are not found
together or near a source of nuts. Nut cracking is also difficult and
must be learned. Chimps also use leaves as sponges or spoons to
A recent study revealed the use of such advanced tools as spears,
which West African chimpanzees in
Senegal sharpen with their teeth,
being used to spear
Senegal bushbabies out of small holes in
trees. An eastern chimpanzee has been observed using a modified
branch as a tool to capture a squirrel.
Eastern chimpanzee snatches a dead bushbuck antelope from a
baboon, Gombe Stream National Park
When hunting small monkeys such as the red colobus, the chimpanzee
hunts where the forest canopy is interrupted or irregular. This
allows it to easily corner the monkeys when chasing them in the
appropriate direction. Chimps may also hunt as a coordinated team, so
that they can corner their prey even in a continuous canopy.
During an arboreal hunt, each chimp in the hunting groups has a role.
"Drivers" serve to keep the prey running in a certain direction and
follow them without attempting to make a catch. "Blockers" are
stationed at the bottom of the trees and climb up to block prey that
take off in a different direction. "Chasers" move quickly and try to
make a catch. Finally, "ambushers" hide and rush out when a monkey
nears. While both adults and infants are taken, adult male
black-and-white colobus monkeys will attack the hunting chimps. In
Gombe, the chimpanzee also fears adult female colobus monkeys, and
prefers to snatch infants from their mother's bellies without harming
the mothers. Male chimps hunt more than females. When caught and
killed, the meal is distributed to all hunting party members and even
Chimpanzees and humans
Chimpanzee mask from Liberia
Chimpanzees are rarely represented in African culture, as the natives
regard them as too similar to humans and thus "too close for comfort."
They were even thought to have kidnapped and raped women. The Gio
Liberia and the
Hemba people of the Congo have created
blocky and crude masks of the animals. The mask may have a smile which
suggests drunken anger, insanity or horror. They wear these masks when
teaching young people how not to act; performing rituals where they
act wildly and uncivilized. They may also act out these rituals during
funerals, representing the "awful reality of death."
In Western popular culture, chimpanzees have been commonly stereotyped
as childlike companions, sidekicks or clowns. They are especially
suited for the latter role on account of their prominent facial
features, long limbs and fast movements, which humans often find
amusing. Accordingly, entertainment acts featuring chimpanzees dressed
up as humans have been traditional staples of circuses and stage
shows. Westerners have also been disturbed by the chimps'
resemblance to humans as well as their "frank sexuality".
Jane Goodall undertook the first long-term field study of the common
chimpanzee, begun in
Gombe Stream National Park
Gombe Stream National Park in 1960.
Other long-term study sites begun in 1960 include A. Kortlandt in the
Republic of the Congo
Republic of the Congo and
Junichiro Itani in Mahale
Mountains National Park in Tanzania. Current understanding of the
species' typical behaviours and social organization are formed largely
from Goodall's ongoing 50-year Gombe research study.
Chimpanzee genome project
NCBI genome ID
Number of chromosomes
Relationships among apes. The numbers in this diagram are branch
lengths, a measure of evolutionary distinctness. Based on protein
electrophoresis data of Goldman et al.
Human and common chimpanzee
DNA are very similar. After the completion
Human Genome Project, a
Chimpanzee Genome Project was
initiated. In December 2003, a preliminary analysis of 7600 genes
shared between the two genomes confirmed that certain genes, such as
the forkhead-box P2 transcription factor which is involved in speech
development, have undergone rapid evolution in the human lineage. A
draft version of the chimpanzee genome was published on September 1,
2005, in an article produced by the
Chimpanzee Sequencing and Analysis
DNA sequence differences between humans and chimpanzees is about
35 million single-nucleotide changes, five million insertion/deletion
events, and various chromosomal rearrangements. Typical human and
chimp protein homologs differ in only an average of two amino acids.
About 30% of all human proteins are identical in sequence to the
corresponding chimp protein. Duplications of small parts of
chromosomes have been the major source of differences between human
and chimp genetic material; about 2.7% of the corresponding modern
genomes represent differences, produced by gene duplications or
deletions, during the roughly four to six million years since humans
and chimps diverged from their common evolutionary ancestor. Results
from human and chimp genome analyses, currently being conducted by
geneticists including David Reich, should help in understanding the
genetic basis of some human diseases.
Common chimpanzees have been known to attack humans. In
Uganda, several attacks on children have happened, some of them fatal.
Some of these attacks may be due to the chimpanzees being intoxicated
(from alcohol obtained from rural brewing operations) and mistaking
human children for the western red colobus, one of their favorite
Human interactions with chimpanzees may be especially
dangerous if the chimpanzees perceive humans as potential rivals.
At least six cases of chimpanzees snatching and eating human babies
A chimpanzee's great strength and sharp teeth mean that attacks, even
on adult humans, can cause severe injuries. This was evident after the
attack and near death of former
NASCAR driver St. James Davis, who was
mauled by two chimps before they were killed. Another example
of chimpanzees being aggressive toward humans occurred in 2009 in
Stamford, Connecticut, when a 200-pound (91 kg), 13-year-old pet
chimp named Travis attacked his owner's friend, who lost her hands,
eyelids, nose, and part of her maxilla from the attack.
Link with human immunodeficiency virus type 1
This Cameroonian chimpanzee was brought to a rescue centre after its
mother was killed by poachers.
Two types of human immunodeficiency virus (HIV) infect humans: HIV-1
and HIV-2. HIV-1 is the more virulent and easily transmitted, and is
the source of the majority of HIV infections throughout the world;
HIV-2 is largely confined to west Africa. Both types originated in
west and central Africa, jumping from primates to humans. HIV-1 has
evolved from a simian immunodeficiency virus (SIVcpz) found in the
common chimpanzee subspecies, P. t. troglodytes, native to southern
Cameroon. Kinshasa, in the Democratic Republic of Congo, has
the greatest genetic diversity of HIV-1 so far discovered, suggesting
the virus has been there longer than anywhere else. HIV-2 crossed
species from a different strain of SIV, found in the sooty mangabey
monkeys in Guinea-Bissau.
Status and conservation
Chimpanzee are a legally protected species in most of their range and
can be found both in and outside national parks. Between 172,700
and 299,700 individuals are thought to be living in the wild, a
decrease from about a million chimpanzees in the early 1900s.
The biggest threats to the common chimpanzee are habitat destruction,
poaching, and disease.
Chimpanzee habitats have been limited by
deforestation in both West and Central Africa. Road building has
caused habitat degradation and fragmentation of chimpanzee populations
and may allow poachers more access to areas that had not been
seriously affected by humans. While deforestation rates are low in
western Central Africa, selective logging may be done outside national
Chimpanzees are a common target for poachers. In Ivory Coast,
chimpanzees make up 1–3% of bushmeat sold in urban markets. They
are also taken in pet trades despite it being illegal in many
countries where they live. Chimpanzees are also hunted for
medicinal purposes in some areas. Capturing chimpanzees for
scientific research is still allowed in some countries, such as
Guinea. People sometimes kill chimpanzees that threaten their
crops. Chimps may also be unintentionally maimed or killed by
snares meant for other animals.
Infectious diseases are a main cause of death for chimpanzees. They
succumb to many diseases that afflict humans, because the two species
are so similar. As human populations grow, so does the risk of
disease transmission between humans and chimpanzees.
On 12 June 2015, the
U.S. Fish and Wildlife Service
U.S. Fish and Wildlife Service announced it will
classify all chimpanzees, both wild and captive, as endangered under
Species Act. Before this ruling, only wild chimpanzees
were listed as endangered, while captive chimpanzees were listed as
threatened under the act. The final rule was published in the
Federal Register of 16 June 2015 (80 FR 34499), and came into effect
90 days after publication on September 14, 2015.
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