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SPECIES
Systematics
Species description. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Taxonomy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Distribution and Evolution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4-6
Significance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Natural History
Habitat use and Home range. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Food habits. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
Social organizations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7-8
Reproduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8-9
Survival and Mortality. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9-11
Competition With Other Species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11
Pursuant to the Endangered Species Act of 1973, 16 U.S.C. 1531 et seq., the Conservation of
Endangered Primates Organization hereby formally petition the United States Secretary of
Interior and the United States Fish and Wildlife Service to list the Japanese macaque (Macaca
fuscata) as endangered throughout its range. We submit this petition as an interested party
under 5 U.S.C. 553(e) and 50 C.F.R. 424.14.
In accordance with the Endangered Species and the Administrative Procedures Act, the Center
also requests that critical habitat be designated concurrently with the final listing rule.
The Center of Old and New World Primates was incorporated in 1994 to contribute to the
scientific understanding and conservation of old and new world primates particularly in Africa
and Asia. The center has been vital in the efforts to protect Aye Aye (Daubentonia
madagascariensis), Ring tail lemur (Lemur catta), Brown Spider Monkey (Ateles hybridus), and
Tarsier (Tarsius).
1
Executive Summary
Japanese macaque (Macaca fuscata), also known as the Snow Monkey, are endemic to Japan
found on three of four islands: Honshu, Shikoku, and Kyushu. They are the only known primate
to travel as far north compared to humans and are keystone species responsible for seed
dispersals, pruning which in turn stimulates plant growth, and their feces provide resources for
unique species of dung beetles. The extinction of the Japanese macaque would be a tremendous
loss to Japans ecosystems, culture, and economy.
Habitats for the Japanese macaque range from deciduous trees, sub-tropical, and broad leaf
evergreen forest along with their source of food. However, with increase in lumbar plantation
and loss of broad leaf forest and increase monotone conifer forest is drastically effecting the
Japanese macaques population. Increase in habitat destruction had led the macaques to extend
their range even further looking for resources. Increase in range outside the norm has resulted
large numbers of death with numbers being around 5,000 to 20,000 a year for various reasons
such as population control and farmers protecting their crops. Another 15,000 macaques are
believed to be sold or traded in the pet trade. The Conservation of Endangered Primates
Organization and the Center of Old and New World Primates petition to list the Japanese
macaque under the U.S. Endangered Species Act.
The Japanese macaque crossed a dry land connection taking them from the Korean peninsula to
Kyusha-West Honshu area in Japan around 0.63-0.43 million years ago. The Japanese macaque
diverged from Eastern rhesus monkey, Macaca mulatta, 0.31-0.88 million years ago and it is
presumed allopatric distribution separated the Japanese macaque from other macaque relatives
during the Pleistocene. Two subspecies of the Japanese macaque are recognized, the M. F.
fuscata and M. F. yakui.
The Japanese macaque population has been dangerously low, due to World War II and extensive
hunting (independent of each other), and population rose again thanks to Japanese government
intervention in 1947 protecting the species from hunting. Presently, population of the macaque
are estimated around 50,000 and considered stable though the population is declining at a slow
rate. Decline in population is presumed to be largely on habitat destruction with broad leaf forest
being torn down for lumbar plantations. An exact population status and trends lack sufficient
data and more studies are needed.
The causes of decline are largely due to anthropogenic factors and threatened destruction with
disease following though other factors may be possible. Japanese macaques are suffering from
high habitat destruction with a loss of 70% of their broadleaf forest being replace with conifer
lumbar plantations. Loss of habitat has led to increase in conflict with humans especially
farmers. Japanese macaques have raided farms for resources and easy food source resulting in
5,000 deaths by farmers. Japanese macaques were hunted to near extinction before the Japanese
government stepped in, where hunting and killing the macaque was illegal. Estimates of 10,000
Japanese macaques were killed to control population growth.
Infectious and non-infectious disease has effected the population of Japanese macaques and
those in captivity as well; some serious diseases are affecting captive Japanese macaques. In
2
2008, Tetanus outbreak killed 16 (25%) Japanese macaques due to a contaminated environment
and Measles virus killed 53 within days on contracting the virus presumably from visitors.
Japanese macaque encephalomyelitis (JME) is a new threat to the macaque first seen in 1986
with death being the final symptom.
3
Systematics
Species Description
The Japanese macaque is found on three out of the four islands of Japan. Distinct characteristics
are their pelage that ranges from yellow-brown, brown, and white with shades of gray with a
pink-red face and posterior, and a stump tail (Gron 2007; Flannery 2007). Newborns have grey
to dark fur with pale pink faces that will resemble adult pelage when they are four to five months
old (DPIPWE 2011). Males and females have cheek pouches to carry food as they forage the
forest ground (Robinson Library 2017). Sexual dimorphism is present, but small, between males
and female with males weighing more, 24.91 lb., and having longer body length, 22.4 inches,
versus females, 18.52 lb., and 20.58 inches (Gron 2007). Body fat in males and females differed
with females have higher median fat mass (9%) versus males (7%) where macaques in colder
regions weighed more (Hamada et al. 2003). Tail length in males measured 3.64 inches and 3.11
inches in females (Gron 2007). Life span varies between sex with average being around 15 to 20
years (Fedigan & Zohar 1997) with males living to 28 years and females to 32 years with 25
being considered old age in females (Gron 2007; Fedigan & Asquith 1991).
Taxonomy
The taxonomic validity of the Macaca fuscata, Japanese macaque, isnt questioned. Belongs in
the Macaca mulatta group distinctive by short-tail with pink-red face (Academic Press 2005).
Two subspecies of the Japanese macaque are recognized (ITIS 2017): M. F. fuscata and M. F.
yakui.
During the Pleistocene, it is presumed the Japanese macaque diverged from the eastern rhesus
macaque (Macaca mulatta) between 0.31-0.88 million years ago (Marmi et al. 2004). Fooden &
Aimi (2006) discovered evidence that the divergence between M. mulata and M. fuscata and
traveled to Japan during glacial intervals around 0.63-0.43 million years ago through a dry land
connection between the Korean peninsula and Kyusha-West Honshu area. Mitochondrial DNA
(mtDNA) from both species were collected to test and analyze for speciation and sub speciation
resulting in high genetic similarities with two fixed (Marmi et al. 2004). Due to lack of
correlation between genetic and geographic distances line up with rapid dispersion of species
macaques after they arrived in Japan (Marmi et al. 2004). According to Richard et al. (1989), the
Japanese macaque, Long-tail macaque, Bonnet macaque, and Toque macaque are the results of
allopatric distribution.
Figure 1. Japanese macaque geographic distribution in Japan (Nakagawa & Nakamichi 2010).
4
Figure 2. Geographic range in 2008 of Japanese macaques (Watanabe & Tokita 2008).
Figure 3. Range of Japanese macaques on Shimokita Peninsula over 80 years (Primate isolated
population 2010)
5
Significance
Japanese macaques are native only in Japan, with the exception of Texas where a troop was
transported for study (Gron 2007) so distribution is restricted. The popular name of Japanese
macaque is the snow monkey and well known to be the monkeys seen in hot springs. The
macaques are keystone species responsible for seed dispersals, skilled pruning which stimulates
plant growth, and their feces provide resources for unique dung beetles that are also crucial
decomposers and seed dispersal (Enari & Sakamaki-Enari 2013).
They are used and preferred as laboratory animals in Japan in the fields of neuroscience,
virology, and reproduction due their high intelligence, social behavior, and gentle nature (Isa et
al. 2009).
Japan has several monkey parks where the Japanese macaque can be observed and have allowed
the macaques to become comfortable with humans (Bengsch 2016). Japanese macaques edge of
their habitat remains close to human dwellings since food is easier to acquire (Bengsch 2016).
Japanese macaques are traditionally used in Japanese art and are famously seen in the 17th
century carving over the door of Tsh-g shrine where the macaques are used to depict
Confuciuss code of conduct (Bengsch 2016). They are also the monkeys seen in the see no
evil, hear no evil, speak no evil (IMA nd). In folklore, the Japanese macaques are described as
being greedy, evil, trickster, cunning, witty, humorous, and lovable where their roles in stories
varying from scared mediators to messengers for deities and humans (Brazil 2017).
6
Natural History
Food Habits
The Japanese Macaque are omnivores. Their diet includes a wide variety plants and animals that
change with the availability and seasons (Hardman 2011). Typically, from June to November
they consume fruits and seeds (20%), from April to May and December through March they
consume flowers and nectar, and in the winter, they consume fibrous leafs (their main diet), and
opportunistically fungi (Hardman 2011). Hill (1997) found similar results with 35.0% of
foraging on leaves and shoots, 30.2% on fleshy fruit, 13.2% on seeds, and 5.5% on flowers.
Invertebrates and other animal matter accounted for 10.3% of foraging and fungi for 4.6%.
Sawada et al. (2013) found that 67 possible fungal species in 31 genera were eaten supporting
that fungi makes a small portion of their diet (2.2% annually). Fungi that was eaten without
being examined i.e. sniffed, nibbled, handled carefully, were less likely to be poisoned, although
there isnt a correlation that examining behavior allowed the macaques to distinguish poisonous
from nonpoisonous fungi (Sawada et al. 2013). They have also been found to eat insects from
grooming each other (Monkeyworld), bird eggs, crabs, with most foraging done on ground floor
(Blue Planet Biome). 213 plant species were identified that are consumed along with soil and
fish (anecdotal) when primary food source wasnt available (Encyclopedia of Life). Increased
availability led to them feeding more on fruits and seeds and on mature leaves if fruits and seeds
were low; when temperatures dropped, they resorted to herbs and staying in sunny areas to
conserve energy (Hanya 2004). There is anecdotal data found on Koshima Island in 1979
according to Watanabe (1989) were an adult macaque was eating raw fish and passing the habit
to the females in his troop over a period of six years. Watanabe (1989) concluded that the
consumption of raw fish was perhaps due to nutrition conditions in their habitat.
Social Organization
Japanese macaques live in troops along with class system hierarchies where dominance is passed
on through the maternal line and heavily centered on (Robinson Library 2017; Hays 2013).
Yamagiwa (2010) added that troop formation is based on dominance rank and leadership system.
Troop size varies in size from 10 to 160 individuals with average being around 40 and depends
largely on amount of resources available such as food (Robinson Library 2017). Females remain
7
in the troop through their lives whereas males go from troop to troop to mate in their lives
starting at 4-5 years of age (Robinson Library 2017). When a male leaves his natal troop, he may
join an all-male troop, a new troop, or spend a period in solitary (Yamagiwa & Hill 1998).
Hierarchy plays a large role in the Japanese macaques, which includes a dominant alpha male
and female followed by lower ranked males and females throughout the troop with the alpha
male siring the most offspring (Robinson Library 2017; Soltis et al. 2001). According to
Yamagiwa (2010) females gain dominant ranks vie two ways: one is a daughter is higher among
subordinates but below her mother and second maturing daughters younger daughters are higher
in rank versus their older sisters. The class hierarchy is flexible allowing movement up and down
in ranks and because the ranks arent strict this allows the troop to share tight spaces and huddle
for warmth without clashing (Hays 2013). Territory ranges of troops averaged at 1.4 square miles
and could be smaller in areas with larger food resources (Robinson Library 2017). Troop size
varied according to the habitat they inhabited with mean group size of 34.9 in cool-temperature
forest, 74.8 in warm-temperature forest, and 27.1 in warm/subtropical forest (Yamagiwa & Hill
1998).
Reproduction
Soltis et al. (2001) found that Japanese macaques live in large, coed social groups with
reproductive strategies for both males and females occurring simultaneously. Mating season
ranges from September to April and Japanese macaques only give birth once a year, though
twins are possible, in April through May with offspring born in equal proportions, but by
adulthood the ratio is 2:5 females to every male (Gron 2007; Hays 2013; Fedigan & Zohar 1997;
DPIPWE 2011). Gestation period is averaged 176.3 days (Rigaill et al. 2015). According to
Garcia et al. (2009), seasonal breeding has a cost on the energetic demands of mating, and that
higher condition (i.e. fatter) females could afford the demands of lactation and reproduced more
rapidly. In a 35-year study, Fedigan and Asquith (1991) found the mean fecundity was 0.36
with a mean survivorship of offspring at 0.86, however fecundity declined after the age 20 and
25 in females, with 25 being considered old age. Sperm production in males and cycling in
females begin around 4.5-5 years of age (Fedigan & Zohar 1997).
Social ranks play a large role in reproduction of Japanese macaques. Mating is done by
consortship with pair mating, traveling together, and resting for average of 1.6 days during
mating season with higher ranking males consortship lasting longer versus lower ranking males
(Gron 2007). High ranking male dominated and monopolized the females in one season most of
the offspring being sired by high ranking males (Soltis et al. 2001). To avoid competition with
high ranking males and prevent them from disrupting consortship, low ranking males would
mate with females when the high rank males were not present (Soltis et al. 2001; Gron 2007).
Reproduction relies heavily on females since they decide whether they will mate (Gron 2007).
Soltis et al. (2001) found that female preference affects conception when females mate with
certain males, but not with others (Huffman 1992). Rank of males did not seem to influence
female preference of a mate and appeared to prefer middle ranking males (Soltis et al. 2001;
Huffman 1992). Incentive for females to choose higher ranking males would be mate-guarding
and prevention of mating with lower ranking males (Gron 2007). High-ranking males mate with
most females within troop (Soltis et al. 2001).
8
Females advertise fertility status through sexual signals and behaviors such as mounting, holding
behavior, and auditory signals example being estrus calls to indicate follicular phase (Rigaill et
al. 2015). Mounting/mating position is popularly seen where the male mounts the female with
male leg grasping the females leg (Gron 2007). Estrus and copulation calls were found to
influence female and male mating choices according to Rigaill et al. (2015). Estrus calls was
associated with reproductive status with significant increase during the first two months of
pregnancy (Rigaill et al. 2015). Females become redder during the estrus phase with the face
becoming less red during pregnancy and decreasing from the first to second month of pregnancy
(Rigaill et al. 2015). Rigaill et al. (2015) found that males and females that didnt mate had low
participation in sexual and social behaviors, especially females during early pregnancy indicating
that males are able to distinguish cycling and pregnant females.
Mortality factors are numerous for Japanese macaque (see Table 1). Due to large habitat range
the Japanese macaque has little predators with exception of feral dogs, grey wolves (Canis lupus)
that became extinct in the 20th century, and humans (AZ Animals 2008; Robinson Library 2017).
The biggest cause of death and largest threats amongst the macaque presently is hunting, killing
to reduce numbers, and habitat destruction by humans (AZ Animals 2008; Fedigan & Zohar
1997). There are seven categories for cause of death in Japanese macaques being: predator
related, infectious disease, non-infectious disease, neonatal mortality, social stress, human
related, and unknown disappearances (see Table 1.) (Fedigan & Zohar 1997). Mortality rates
differ between males and females with males at a significant higher risk of adolescent death
beginning around 4.5-5 years of age when they reach sexual maturity (see Figure 4) (Fedigan &
Zohar 1997). Males that have reached sexual maturity will migrate to mate with females of other
troops or just immigrate increasing the risk of contracting infectious disease and human-related
death (Table 1.) (Fedigan & Zohar 1997). As the males migrate, either alone or in pairs, they are
traveling in less familiar territories rendering them vulnerable to predators and unknown
elements, whereas females tend to remain in their known range allowing longer life spans
(Fedigan & Zohar 1997).
Nakamichi et al (1997) found that limb malformations had low mortality rates versus other
primates with only 28.2% mortality in malformed Japanese macaques (see Table 2), indicating
that not only was clinging not necessary for survival, but that maternal care was good enough to
ensure the malformed newborns reached adulthood. Furuya (1966) found in his study that limb
malformations were limited to hands and feet and occurred more in males than females. It
remains unknown as to what causes the malformations (Iwamoto 1967).
9
Table 1. Causes of death for Japanese macaques (Fedigan & Zohar 1997).
Figure 4. Survival curve for male and female Japanese macaques (Fedigan & Zohar 1997).
Table 2. Limp malformation mortality in Japanese macaques of normal and malformed newborns
(Nakamichi et al. 1997).
10
Competition with Other Species
The Japanese macaque doesnt compete with other species.
Historically, habitats of Japanese macaque have undergone bottleneck effect limiting where the
macaques have been found since the population doesnt inhabit mountainous areas, which covers
70% of Japan (Nakagawa & Nakamichi 2010). Population of macaques have grown in some
areas and decreased in others such as the coast (Nakagawa & Nakamichi 2010).
In the early 1900s, the Japanese macaque was hunted to near extinction but bounced back after
World War II and has steadily increased since 1978 (Radhakrishna et al. 2012). According to
Watanabe & Tokita (2008) the population trend is stable with no fragmentation for Japanese
macaque seen in Figure 2, however their range is increasing due to lumbar plantations. Rise of
lumbar plantation had led to a 70% loss of broadleaf forest since 1960, which directly affected
the population of macaques leading them to expand their range even further, but leading to bits
of fragmentation and increased conflict with humans (Enari & Sakamaki-Enari 2013; Robinson
Library 2017).
Listing Factors
Section 4(a)(1) of the Endangered Species Act (16 U.S.C. 1531 et seq.) and regulations (50 CFR
part 424) promulgated to implement the listing provisions of the Act set forth general listing
criteria. If a species existence is imperiled by one or more of the following five factors, it must
be listed as threatened or endangered. We hereby petition to list the Japanese macaques
11
(Macaca fuscata) as threatened in its entire range due to habitat destruction, disease and
predation, and scientific purposes.
In 2015, it was estimated 15,000 Japanese macaques were privately owned (PETA 2017). When
sold as pets, the Japanese macaque are separated from birth (hours to days) from parent and sold
as infants in diapers to buyers and to control behavior may have their teeth cut/removed and be
castrated (PETA 2017).
Disease. Threats of infectious and non-infectious disease in Japanese macaques are very serious
and are frequent in the population (Fedigan & Zohar 1997). Non-infectious diseases affecting the
Japanese macaque are cancer, heart disease, and heat stroke to name a few (Fedigan & Zohar
1997). Table 1 shows infectious disease that affects macaques resulting in death sometimes are
pneumonia, flue, valley ever, screwworm, etc. (Fedigan & Zohar 1997). Outbreak of the measles
virus had infected a group of 53 Japanese macaques where 12 have died within 2-23 days of
contracting the virus with symptoms of anorexia, diarrhea, and dermatitis (Choi et al. 1999).
Transmission of the measles virus is believed to have come from human visitors though mode of
12
transmission couldnt be determined (Choi et al. 1999). An emerging spontaneous threat to
Japanese macaques particularly in captivity is JME, Japanese macaque encephalomyelitis, which
first appeared in 1986 (Axthelm et al. 2011). JME appears around the four years of age with no
preference of gender and is characterized by paralysis in at least one limbs, ataxia, or motor
paresis resulting in death by euthanization (Axthelm et al. 2011). Tetanus was responsible for
mass death of captive Japanese macaques in 2008 resulting in 16 deaths (25%) (Nakano et al.
2012). Tetanus outbreak originated from severe contamination of environmental C. tetani
(Nakano et al. 2012).
Figure 5. Japanese macaque deaths due Tetanus during 2006-2009 in Kantou area of Japan
(Nakano et al. 2012).
13
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