Macaca+fascicularis

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 * Long-tailed Macaque ** ** //Macaca fascicularis// (Raffles, 1821) **

Two adult females macaques and a juvenile cuddling together. Photo by Amanda Tan, used with permission. = 1. Introduction =

The long-tailed macaque, also known as the c rab-eating macaque, is a native species in Singapore. Like what its name suggest, the animal can be easily identified by its long tail. Commonly encountered in parks, reserves and occasionally residential estates, the long-tailed macaque is not unfamiliar to most of us. While many native species were lost due to urbanisation and habitat destruction, the highly adaptable long-tailed macaque continue to exist in our landscape. The sight of these wild animals jumping between trees, hanging from the trees with their long tail and grooming each other is a beautiful sight to behold. Not forgetting the little ones that love chasing and wrestling with one another, that makes our neighbourhood even livelier.

Don't stop here, read on to find out more about these lovely critters!

Etymology
The [|generic epithet] //Macaca// originates from the Portuguese word //macaco//, derived from //makaku//. //Makaku// is a Fiot (West African language) word, and //kaku// means monkey in Fiot. The [|specific epithet] //fascicularis// means a small band or stripe in Latin. The epithets were given by Sir Thomas Raffles, who did not explain the use of terms 1].

= 2. Description =

The head and body of an adult long-tailed macaque measure to 45cm, with tail up to 56cm 2]. It has a silky, golden brown coat which becomes pale grey to whitish below 2]. The face is brownish to pinkish, with distinct eyelids above the eyes, sometimes also observed faintly below the eyes [2]. You might also observe an irregular tuft of hairs behind the ears [3]. These features vary among individuals, and are often used by researchers to distinguish different troops, and even members of the same troop.

Identification features of the long-tailed macaque labelled. Photo by Joys Tan. Unlike adults, infants are born with black fur and pinkish facial skin with minimal hair, with the fur slowly changing to golden brown when they are around 2 to 3 months old [3]. Interestingly, the transition from black to brown coat begins with the limbs, then the body and finally, the crown [3].

An infant macaque with black fur and pinkish face. Photo by Amanda Tan, used with permission. = = =** 3. Biology **=

Habitat Preference
Being a forest edge species, long-tailed macaques inhabit the edges of diverse habitat types such as mangroves, swamp, riverine and secondary forests [4, 5, 6]. Their strong adaptability to disturbed habitats has allowed them to successfully thrive in the human modified landscape such as Singapore [7, 8].

Diet
Long-tailed macaques are primarily frugivorous, but can become an opportunistic omnivore when fruit availability is limited [9]. While their diet consists mainly of fruits, they also feed on flowers, leaves, stems, insects and human food.

An adult female macaque feeding on flowers. Photo by Joys Tan. When human food is available, it is often preferred over natural food as it is an excellent source of energy, easily digestible and accessible. In Singapore, at least 50% of our long-tailed macaques fed on human food sources, probably due to proximity to human settlements [10]. Therefore, open food sources in settlements can attract macaques to visit, and should be avoided.

Other than that, food can also be directly obtained from humans who deliberately feed them. It is however, important to note that, such actions are not desirable as it encourages macaques to approach humans for food, fuelling human-macaque conflict (Read more at Section 4). Furthermore, these unnatural foods cause health problems in them. Hence, please do NOT feed them.

(Left) An adult male macaque feeding on rice from leftover food boxes on the roadside. (Right) An adult male macaque feeding on peanuts, given by passer-bys. Photos by Joys Tan.

Activity Pattern
Being diurnal animals, long-tailed macaques are active from dawn to dusk. Here’s a typical day for macaques [10, 11, 12, 13] : 1) At dawn, they move out of their sleeping sites to look for food. 2) Around noon, macaques rest and sometimes take a little nap on branches of trees or under trees to avoid the heat. When it gets a little too hot, they like to cool themselves down in a nearby pool of water (And yes, they can swim too!).

Macaques cooling themselves in a pond at MacRitchie Reservoir Park. An awesome pool party, isn't it! Photo by Lai Chui Ting, used with permission. media type="youtube" key="beEAKVZsJe8" width="568" height="344" align="center"

Long-tailed macaques (Crab-eating macaques) are excellent swimmers! 3) They would then feed again during the second feeding peak in the late afternoon.  4) Approaching dusk, the macaques were also observed to engage in play, grooming and inactivity, before heading back to their sleeping sites. These behaviours can occur throughout the day, but more happen more frequently at the end of their day. An adult female macaque being groomed by two other females. Photo by Joys Tan. 5) When the Sun sets, most of them would be at or around their sleeping sites, slowing moving up to the tree and finding their positions. Some of them prefer to cuddle and sleep together, while some prefer to enjoy the space to themselves alone. Macaques have not just one but a few sleeping sites, and the leader of the troop gets to decide where the troop sleeps for the day!  A juvenile macaque snoozing on a tree branch, possible a sweet dream? Photo by Sabrina Jabbar, used with permission.

Social Organization
A healthy long-tailed macaque troop consists of 15-35 individuals. Each troop consists of multi males and multi females 14, 15, 16]. Within the troop, they are organised based on social hierarchy or dominance ranks. High ranking individuals usually have greater access to food and protection from predation, as compared to lower ranking individuals. The highest ranking male and female are often known as alpha male and alpha female respectively. A long-tailed macaque troop in Upper Seletar Reservoir Park, Singapore. Photo by Joys Tan. The social ranks of macaques are inherited through the matrilineal lineage 14]. This means that females usually inherit the ranks from their mothers. If the mother is a high ranking individual within the troop, her daughters would be too. Among the siblings, their dominance depend on the birth order 17]. Interestingly, younger females are usually higher in rank than their elder sisters 17]. On the other hand, males usually leave the troop to join another troop and negotiate their rank within the new troop 10]. This is one reason why you sometimes see a lone macaque wandering around in the urban areas. What should you do when you see a lone macaque? Find out more here.

Mating System
Long-tailed macaques have a polygamy and polyandry mating system in the macaques 14]. This means that each male and female have more than one mating partners. Males usually mate within high ranking females to maintain their social ranks while females mate with any available males when fertile.

= 4. Relationship with Humans =

Human-Macaque Conflict
Increasing proximity of human settlements to the forest has led to a closer interface, as well as greater opportunities for interaction between humans and macaques. When interactions between humans and macaques negatively impact each other, human-macaque conflict happens 7] . Human-macaque conflict for various species of macaques has been reported for barbary macaques in Gibraltar, formosan rock macaques in Taiwan and long-tailed macaques in Hong Kong as well as Singapore 12, 18, 19] //.//

Macaques can negatively impact humans in the form of raiding crops, houses and refuse sites for food, when accessible 20, 12, 21] . When humans behave inappropriately towards macaques, macaques can react aggressively to defend themselves, which may lead to distress or harm for people. Such actions arises from the lack of understanding of humans towards macaques, or wild animals in general.

Humans also impact macaques negatively. In the shared space of humans and macaques, wild macaques can become used to the presence of humans. Feeding by humans causes them to associate humans with food. As macaques learn to approach humans, it may lead to aggression, contact and often, conflict 8, 13]. Gradually, they are recognised as nuisance, and often, measures such as trapping and culling are implemented. These measures directly decimate their population and destabilises the troop structure.

So, how should we behave around macaques? Continue reading to find out more!

Facial Expressions and Body Languages
Like any other primates, macaques use facial expressions and body languages to communicate their emotions to recipients. However, this is often misinterpreted by humans, which can result in unnecessary fear and resent towards the animals. Here’s a guide to help you understand them and know how to behave around them 22] !

When macaques are //relaxed//, they maintain an impassive face and avoid eye-contact. Sometimes, their furrowed brows (especially females) might make them look angry or unhappy, but usually not the case. When a macaque is comfortable around you, it may look at you and 'coo' gently, a way of asking to be groomed. However, please do not make contact. Others may stand not their hind legs to beg for food. Please do not feed them and find out why you should not here.

A relaxed female macaque with her infant. Photo by Joys Tan. When //uncomfortable// with your presence, macaques display several types of facial expressions and body language to prompt you to move away. They do not usually make contact with humans, especially aggressively, as it is a big risk for them to take on opponents bigger than them. Hence, attacks do not usually happen unless you have ignored all the other prompts.

Below are some signs that you can look out for in a distressed macaque. Please be nice and back off when you see such signs.

Facial expressions and body languages of macaques that may indicate that they are uncomfortable about your presence. The recipient should move away to give it more space. Photo and content by Amy Klegarth, used with permission.

Living with Monkeys
Information by Amy Klegarth and Sabrina Jabbar 22, 23] :

The DOs:
 * Proper waste disposal: keep the trash and food out of sight of monkeys.
 * Monkey proof waste disposal points.
 * Behave appropriately around them. Refer to above section on Facial Expressions and Body Languages.

The DON'Ts: If you need help with monkey issues, contact the [|ACRES] 24-hour Wildlife Rescue Hotline on 9783-7782.
 * Do not scream - macaques will be frightened and upset.
 * Do not stare - macaques will take stare as a form of aggression.
 * Smile with your lips, not teeth - when you show your teeth, it means that you are either angry or afraid, hence making the macaques upset.
 * No wild gesturing - it's seen as aggression by the macaques.
 * Do not carry plastic bags - macaques have learnt to associate plastic bags with food, and hence they will go for you.
 * No tug-of-war with monkeys - wait for the macaques to lose interest in the item before retrieving it. If you attempt to snatch it from them, the big males may step in to help them.

Lone Macaques
A lone macaque in an urban area of Singapore. Photo by Sabrina Jabbar, used with permission. Information by Sabrina Jabbar 23] :

__ Why do I see lone macaques? __ As mention under Social Organization, lone macaques are usually males who leave the troop in search of a new troop. This can also be due to troop segregation and mass culling.

__ What should I do when I see a lone macaque? __ 1. Please leave it alone. 2. Ensure proper waste disposal and no food is left around. This will reduce distractions and help the macaque to move on faster. 3. Inform [|ACRES] so that they can monitor the situation.

__ How long will it take before they leave? __ It varies with the individual as well as availability of negative distractions such as food in the area. In certain cases where macaques have high human dependency, they will be less motivated to move on, thus taking a longer time. When the negative distractions are kept away from the macaques, they will eventually leave.

=** 5. Distribution **=

Global Distribution
The long-tailed macaque is a widely distributed non-human primate species in Southeast Asia, estimated number at 3 million 24, 25]. Its geographic range extends south from southeastern Bangladesh to the Indochinese and Malay Peninsular, Singapore, Philippines and to Timor 7, 26].

Global distribution of long-tailed macaques in Southeast Asia. Photo by [|IUCN].

Distribution in Singapore
In 1990s to early 2000s, there are less than 1,000 long-tailed macaques in Singapore 27, 28, 29]. In 2008, the number is estimated at 1,218 to 1,454 individuals, with a core population (ca.1,027 individuals) residing within Bukit Timah and Central Catchment Nature Reserves 8].

Despite this, there is no clear evidence of a large increase in population size nor density 8]. In fact, the density of long-tailed macaques within the Bukit Timah and Central Catchment Nature Reserves was reported to be lower than of more naturalistic population, that have lesser contact with humans 8]. As long-tailed macaques prefer to stay on the edge of forests or habitats, they are often conspicuous in our landscapes, which can be misleading to people that there is a dense population 8]. =** 6. Threats **=

Global Threats
Although accorded the conservation status of Least Concern in the IUCN Red List <span style="font-family: Arial,Helvetica,sans-serif; font-size: 12px; vertical-align: super;">26], long-tailed macaques face several threats which includes: 1) habitat loss and degradation 2) human-macaque conflict and 3) trapping and trade for pharmaceutical testing, research and development. It was suggested that the conservation status was devalued due to human-macaque conflict in Southeast Asia <span style="font-family: Arial,Helvetica,sans-serif; font-size: 12px; vertical-align: super;">30] .  Habitat loss and degradation is a widespread problem faced by many species worldwide. Often, this also means human settlements become in greater proximity to the forest. This creates more opportunities for macaques, who are edge-dwelling species, to come into contact with humans more frequently. This contributes to rising human-macaque conflict when macaques raid crops or houses in search of food and when humans alter behaviour of macaques through feeding <span style="font-family: Arial,Helvetica,sans-serif; font-size: 12px; vertical-align: super;">7, 26]. The largest threat from pharmaceutical trade comes from the Indochinese Region, especially Cambodia, where wild macaques are harvested and transported to China and USA for captive breeding (based on 2003-2004 figures) 26]. Exploitation of long-tailed macaque in international trade. Photo by [|British Union for the Abolition of Vivisection (BUAV)].

Threats in Singapore
The long-tailed macaques in Singapore face similar threats, with human-macaque conflict being a particularly pronounced pressure. This includes culling and roadkill.

__Culling__  Complaints on nuisance macaques from the public often lead to arbitrary culling of large numbers of macaques by the Agri-Food and Veterinary Authority (AVA). In 2013, 1860 feedbacks on macaques were received by AVA, and 570 monkeys were culled, representing one-third of the population. As of October 2014, about 150 monkeys were culled 27]. <span style="display: inline !important; font-size: 16px;">Although c ulling is adopted as the main solution to mitigate human-macaque conflict in Singapore, it is not scientifically proven to be successful or sustainable. // Is there a need to trap and kill monkeys in Singapore? // // Does culling mitigates human-macaque conflict in Singapore? // // How does AVA carry out its trapping operations? //  This informative short [|video] (3-mins) explains further. media type="custom" key="26909694"

__Roadkills__ The victims of roadkill - a female macaque and her baby. Photo by Sabrina Jabbar, story reblogged by [|Monday Morgue]. Long-tailed macaques often fall victim to roadkills. They move from one point to another through the canopy of trees. However in Singapore, as trees are trimmed to reduce tree falls and accidents, it gets much harder for them to do so. Instead, they attempt to cross the roads. Unfortunately, vehicles driving at high speed along roads and expressways often fail to see them or stop in time to avoid them. Hence, a note to all drivers: do drive slowly and carefully, and look out for any wild animals crossing the road, especially near the nature reserves. //"Crossing a road may be simple for you, but for them, it is a battle they have to face just to make it out alive"// (Sabrina Jabbar, 2014)

=** 7. Taxonomy **=

Taxonavigation
Ranks are discarded as they do not contain meaningful taxonomic information.

The following shows the classification of //M. fascicularis// 32]. Each taxon displayed is a subset of the taxon above it. For example, 'Primates' is group under 'Mammalia'. Animalia Chordata Mammalia Primates Cercopithecidae // Macaca // // M. fascicularis //

Type Specimen
The type specimen of //M. f. fascicularis// was not preserved. However, the type specimen was described and documented by Raffles in 1821. It originated from Sumatra, restricted to ‘neighbourhood of Bencoolen [=Bengkulu]” <span style="font-family: Arial,Helvetica,sans-serif; vertical-align: super;">33].

An extract of the original description of //M. fascicularis// (Raffles,1821). Under under a Creative Commons Attribution-Noncommercial License. The holotype was likely an adult female. Five specimens were sent by Raffles from Sumatra or Java to two museums in London, three from Sumatra to the museum of Zoological Society of London and, two from either Sumatra or Java to the museum of East India Company. When both museums were disbanded, specimens were transferred to the British Museum of Natural History, London. However, the holotypes were not preserved.

Diagnosis
The long-tailed macaque is most easily identified by its long tail, that is much longer than its head to body length 25]. When using relative tail length as a index value to compare tail length across body sizes, it ranged from 1.163 +/- 0.1555 in adult females to 1.176+/- 0.1337 in adult males 25].

Although there is only one species of macaque in Singapore, we can find the its sister species in other parts of Southeast Asia. These are the close relatives of the long-tailed macaques. They belong to the same genus // Macaca //, but are different species. It is generally accepted that long-tailed macaques have 19 sister species within the group of //Macaca// 31, 33, 34, 35, 36, 37, 38]. They are //M. sylvanus//, //M. silenus, M. nemestrina, M. tonkeana, M. maura, M. ochreata, M. brunnescens, M. hecki, M. nigrescens, M. nigra, M. mulatta, M. cyclopis, M. fuscata, M. fascicularis, M. arctoides, M. radiate, M. sinica, M. assamensis// and //M. thibetana.//

A notable species is the [|Japanese macaque] (//Macaca fuscata//). Often known as the Snow monkeys, they are found residing within areas of Japan where snow fall is common. Unlike the long-tailed macaques, their coat colour can vary from grey to yellow brown or brown. Additionally, they are easily differentiated from the long-tailed macaques by their short tail.

Baby Japanese snow monkeys enjoying a dip in the spring! Look at the thick fur coat that helps them to stay warm. Photo by [|Oscar Tarneberg], used with permission.

Another example is the [|Barbary macaque] (//Macaca sylvanus//), the only non-human primate species found in Europe. Its coat varies from grey to yellow to brown. The absence of tail in the Barbary macaque differentiates it from the long-tailed macaque. This is also the reason why it is sometimes known as Barbary ape or Rock ape, although it is not an ape.

A juvenile barbary macaque carried by an adult. Note the absence of tail. Photo by Amy Klegarth, permission pending.

Subspecies
Although being debated, it is generally accepted that there are 10 subspecies of long-tailed macaques 33, 34]. They are distinguished by three main external characters of post-infant stage: 1) colour of the crown, thigh and coat at the back of the body, 2) lengths of head, body and tail and, 3) lateral facial crest (side whiskers) pattern 33].

Do you know the differences between the subspecies of //M. fascicularis//? Here's a brief summary adapted from Fooden (1995) and Brandon-Jones //et al, (2//004). 33, 34].

Due to the lack of images from reliable resources, only the Burmese long-tailed macaque (//M. f. aurea//) is shown below. The image is used with permission, from Amanda Tan, who studies the Burmese long-tailed macaque in Thailand. Some of these populations are known to use stone tools.

A comparison between the lateral crest pattern (side whiskers) of the long-tailed macaque and Burmese long-tailed macaque. (Left) A side view of the facial features of the Common long-tailed macaque (M. f. fascicularis). Photo by Joys Tan. (Right) A side view of the facial features of the Burmese long-tailed macaque (M. f. aurea). Photo by Amanda Tan, used with permission.

Phylogeny
Until now, the phylogeny of genus //Macaca// remains debated <span style="font-family: Arial,Helvetica,sans-serif; line-height: 23px; vertical-align: super;">33, 34, 40, 41, 42, 43]. Majority of the studies based on morphological and molecular genetic analyses suggested four species groups within genus //Macaca//. However, the general consensus has only been reached for three groups, which are the //silenus//, //sinica// and //fascicularis// group. Why? Firstly, the number of sister species and species group under the genus //Macaca//, especially for Asian macaques, are not consistent. Secondly, the phylogenetic position among the Asian species group is still being debated. Lastly, the suggested position of //M. arctoides// is largely variable across studies 43]. __ Morphological Analysis __ Morphological analysis based on male genitalia morphology suggested the following classification 35] : **1) //silenus-sylvanus// group**: //M. sylvanus, M. silenus, M. nemestrina, M. tonkeana, M. maura, M. ochreata, M. brunnescens, M. hecki, M. nigra, M. nigrescens//; **2)** //**fascicularis**// **group**: //M. fascicularis, M. mulatta, M. fuscata, M. cyclopis//; **3) //sinica// group**: //M. sinica, M. radiata, M. assamensis, M. thibetana// and **4) //arctoides// group**: //M. arctoides//.

However, examination of dentition and cranium by Delson (1980) reclassified //M. arctoides// under sinica group and removed //M. sylvanus// from //silenus-sylvanus// group to form a sister taxon of the other three groups 43]. This conclusion is supported by molecular data 43].

__ Molecular Analysis __ Wide range of characters were used for molecular analysis across studies, including Y-chromosomal DNA 42], mitochondrial DNA 39, 42] , autosomal loci 42] , nuclear and genetic markers 43, 44]. However, results were not conclusive.

Examples of phylogenetic tree obtained from molecular markers are shown below 42]. Y-chrosomal DNA Tree shows paraphyly among //M. fascicularis// lineages while mitochondrial DNA Tree shows monophyly among //M. fascicularis// lineages.

Phylogeny of the genus Macaca illustrated on a likelihood tree derived from molecular analysis of Y-chrosomal DNA (TPSY and SRY loci) and mitochondrial DNA (12S/tRNA-VAL/16S), on the left and right respectively. Y-chrosomal DNA Tree shows paraphyly among //M. fascicularis// lineages while mitochondrial DNA Tree shows monophyly among //M. fascicularis// lineages. Figures adapted from Tosi //et al//., 2003, used with permission.

__ Singapore __ Long-tailed macaques in Singapore form a monophyletic group with two sub-populations in Bukit Timah Nature Reserve and Central Catchment Nature Reserve respectively 46]. This was established using neighbour-joining trees and parsimony trees conducted on mitochondrial DNA fragments (Cytb, 12s rRNA, COI, COII and COIII) 45].

Neighbour-joining trees depicted for the respective five mitochondrial DNA fragments, **A:** ctyb; **B:** 12s rRNA; **C:** COI; **D:** COII and **E:** COIII. Bootstrap values indicated at the nodes. Values vary from 98-100% (ctyb, COI, COII, COIII) and 68% (12s rRNA), which indicates a relatively strong support for monophyletic grouping. Photo by Schillaci //et al//., 2011, permission pending. Maximum parsimony trees depicted for the respective five mitochondrial DNA fragments, **A:** ctyb; **B:** 12s rRNA; **C:** COI; **D:** COII and **E:** COIII. Bootstrap values indicated at the nodes. Values vary from 89-99% (ctyb, COI, COII, COIII) and 40% (12s rRNA), which indicates a relatively strong support for monophyletic grouping. Photo by Schillaci //et al//., 2011, permission pending. = 8. Useful Links =

__ Species-related __ Long-tailed Macaque on [|Primate Info Net] Long-tailed Macaque on [|WildSingapore] Long-tailed Macaque on [|Ecology Asia] Burmese long-tailed Macaque on Project Sea Monkey [|Website] and [|Facebook Page]

__ Conflict Management __ Why you should not feed monkeys on [|WildSingapore] How macaques and humans can live together in Singapore on [|WildSingapore]

__ Activities __ Monthly monkey walks by [|NParks] and [|JGIS]

= 9. References =

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<span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">[2 ] Tan, R, 2009. Long-tailed Macaque, //Wild Singapore//. Retrieved from []. Accessed on 6 November 2014.

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