Fork-marked lemur

Fork-marked lemurs or fork-crowned lemurs are strepsirrhine

primates; the four species comprise the genus Phaner. Like all lemurs, Fork-marked lemur
they are native to Madagascar, where they are found only in the west,
north, and east sides of the island. They are named for the two black
stripes which run up from the eyes, converge on the top of the head,
and run down the back as a single black stripe. They were originally
placed in the genus Lemur in 1839, later moved between the genera
Cheirogaleus and Microcebus, and finally given their own genus in
1870 by British zoologist John Edward Gray. Only one species (Phaner
furcifer) was recognized, until three subspecies described in 1991 were
promoted to species status in 2001. New species may yet be identified,
particularly in northeast Madagascar.

Fork-marked lemurs are among the least studied of all lemurs and are
some of the largest members of the family Cheirogaleidae, weighing
around 350 grams (0.77 lb) or more. They are the most
phylogenetically distinct of the cheirogaleids, and considered a sister
group to the rest of the family. Aside from their dorsal forked stripe,
they have dark rings around their eyes, and large membranous ears.
Males have a scent gland on their throat, but only use it during social
grooming, not for marking territory. Instead, they are very vocal, Pale fork-marked lemur (P. pallescens)
making repeated calls at the beginning and end of the night. Like the Conservation status
other members of their family, they are nocturnal, and sleep in tree
holes and nests during the day. Monogamous pairing is typical for fork- CITES Appendix I (CITES)[1]
marked lemurs, and females are dominant. Females are thought to have
Scientific classification
only one offspring every two years or more.
Kingdom: Animalia
These lemurs live in a wide variety of habitats, ranging from dry
deciduous forests to rainforests, and run quadrupedally across branches. Phylum: Chordata
Their diet consists primarily of tree gum and other exudates, though Class: Mammalia
they may obtain some of their protein and nitrogen by hunting small
arthropods later at night. Three of the four species are endangered and Order: Primates
the other is listed as vulnerable. Their populations are in decline due to Suborder: Strepsirrhini
habitat destruction. Like all lemurs, they are protected against
commercial trade under CITES Appendix I. Family: Cheirogaleidae
Genus: Phaner
Gray, 1870
Contents Type species
1 Taxonomy Lemur furcifer
1.1 Etymology Blainville , 1839
1.2 Evolution
2 Description Species
3 Distribution and habitat
4 Behavior Phaner furcifer
5 Ecology Phaner pallescens
6 Conservation Phaner parienti
7 References
7.1 Literature cited Phaner electromontis
8 External links
Taxonomy
Fork-marked lemurs were first
documented in 1839 by French
zoologist Henri Marie Ducrotay de
Blainville when he described the
Masoala fork-marked lemur
(P. furcifer) as Lemur furcifer.[6][7] The
holotype is thought to be MNHN 1834-
136, a female specimen taken from
Madagascar by French naturalist Justin
Goudot. The source of this specimen is
unknown, but thought to be Antongil
Bay.[8] In 1850, French zoologist
Isidore Geoffroy Saint-Hilaire moved
In 1897, Alfred Grandidier the fork-marked lemurs to the genus
demonstrated the similarities Cheirogaleus (dwarf lemurs),[9] but Distribution of Phaner:
between Lepilemur (middle they were also commonly listed in the
column, bottom two) and the genus Microcebus (mouse lemurs).[10] red = P. furcifer[2]
cheirogaleids, particularly
In 1870, British zoologist John Edward green = P. pallescens[3]
Phaner (middle, top).
Gray assigned fork-marked lemurs to purple = P. parienti[4]
orange = P. electromontis[5]
their own genus, Phaner,[6] after
initially including them and the mouse
lemurs in the genus Lepilemur (sportive lemurs). Although French naturalist Alfred Grandidier accepted Gray's
new genus (while also lumping the other cheirogaleids in Cheirogaleus and illustrating the cranial similarities
between cheirogaleids and Lepilemur) in 1897,[11] the genus Phaner was not widely accepted. In the early
1930s, zoologists Ernst Schwarz, Guillaume Grandidier, and others resurrected the name, citing characteristics
that were intermediate between Cheirogaleus and Microcebus.[12]

Until the late 20th century, there was only one recognized species of fork-marked lemur,[6] although size and
coloration differences had been noted previously.[8] After comparing museum specimens, paleoanthropologist
Ian Tattersall and physical anthropologist Colin Groves recognized three new subspecies in 1991: the Pale fork-
marked lemur (P. f. pallescens), Pariente's fork-marked lemur (P. f. parienti), and the Amber Mountain fork-
marked lemur (P. f. electromontis).[6][13] In 2001, Groves elevated all four subspecies to species status[6][14]
based on noticeable color, size, and body proportion differences between the fragmented populations. Although
Tattersall disagreed with this promotion, citing inadequate information for the decision,[15] the arrangement is
generally accepted.[6]

In December 2010, Russell Mittermeier of Conservation International and conservation geneticist Edward E.
Louis, Jr. announced the possibility of a new species of fork-marked lemur in the protected area of Daraina in
northeast Madagascar. In October, a specimen was observed, captured, and released, although genetic tests have
yet to determine if it is in fact a new species. The specimen demonstrated a slightly different color pattern from
other fork-marked lemur species. If shown to be a new species, they plan to name it after Fanamby, a key
conservation organization working in that protected forest.[16][17]

Etymology

The etymology of the genus Phaner puzzled researchers for many years. Gray often created mysterious and
unexplained taxonomic names. In 1904, American zoologist Theodore Sherman Palmer attempted to document
the etymologies of all mammalian taxa, but could not definitively explain the origins of the generic name
Phaner, noting only that it derived from the Greek (phaneros) meaning "visible, evident". In 2012,
Alex Dunkel, Jelle Zijlstra, and Groves attempted to solve the mystery. Following some initial speculation, a
search of the general literature published around 1870 revealed the source: the British comedy The Palace of
Truth by W. S. Gilbert, which premiered in London on 19 November 1870, nearly one and a half weeks prior to
the date written on the preface of Gray's manuscript (also published in London). The comedy featured
characters bearing three names: King Phanor (sic), Mirza, and Azema. Since the genera Mirza (giant mouse
lemurs) and Azema (for M. rufus, now a synonym for Microcebus) were both described in the same publication
and equally enigmatic, the authors concluded that Gray had seen the comedy and then based the names of three
lemur genera on its characters.[18]

Fork-marked lemurs were called "fork-marked dwarf lemurs" by Scottish naturalist Henry Ogg Forbes in 1894
and "fork-crowned mouse lemur" by English missionary and naturalist James Sibree in 1895. Literature
searches by Dunkel et al. also uncovered other names, such as "fork-lined lemur" and "squirrel lemur", during
the early 1900s. By the 1970s, reference to dwarf and mouse lemurs had ended, and the "fork-crowned" prefix
became popular between 1960 and 2001. Since then, the "fork-marked" prefix has become more widely
used.[18] These lemurs get their common name from the distinctive forked stripe on their head.[6]

Evolution

Within the family Cheirogaleidae, Competing phylogenies

fork-marked lemurs are the most
phylogenetically distinct, although Lepilemuridae
Lepilemur (sportive lemurs )
their placement remained
uncertain until recently.[21] One Phaner (fork-marked lemurs)
uniting characteristic
(synapomorphy) among all
Cheirogaleus (dwarf lemurs )
cheirogaleids, to the exclusion of
other lemurs, is the branching of Cheirogaleidae
the carotid artery along with how Allocebus (hairy-eared dwarf lemur )
it enters the skull[22]a trait
which is shared by fork-marked Mirza (giant mouse lemurs )
lemurs.[21] Analyses based on
morphology, immunology, and Microcebus (mouse lemurs )
repetitive DNA have given
contradictory placements of
Phaner, while studies in 2001 and Phaner (fork-marked lemurs )
2008 either lacked data or yielded
poor resolution of their Lepilemur (sportive lemurs)
placement.[19]

A study in 2009 of seven Cheirogaleus (dwarf lemurs)

mitochondrial genes (mtDNA) and
three nuclear genes grouped fork- Allocebus (hairy-eared dwarf lemur)
marked lemurs with sportive
lemurs (family Lepilemuridae), Mirza (giant mouse lemurs)
offering a host of explanations,
such as a possible hybridization
Microcebus (mouse lemurs)
(introgression) following the
initial split between the
Fork-marked lemurs are either a sister group within Cheirogaleidae (topWeisrock
families.[21] A study published in et al. 2012)[19] or more closely related to sportive lemurs (bottomMasters et al.
2013 also grouped fork-marked 2013).[20]
lemurs with sportive lemurs [23]

when it used 43 morphological traits and mtDNA.[24] If correct, this would make the family Cheirogaleidae
paraphyletic.[23] However, broad agreement between two lemur phylogeny studiesone in 2004 using SINE
analysis and another in 2012 using multilocus phylogenetic testsgave strong support for a sister group
relationship between fork-marked lemurs and the rest of the cheirogaleids and a more distant relationship with

sportive lemurs.[19] The split between Phaner and the rest of the cheirogaleids is thought to have occurred
sportive lemurs.[19] The split between Phaner and the rest of the cheirogaleids is thought to have occurred
approximately 38 mya (million years ago), not long after the radiation of most of the major lemur groups on
Madagascar, roughly 43 mya.[25][26]

Description
Of the mostly small, nocturnal lemurs in family Cheirogaleidae, the
genus Phaner contains some of the largest species, along with
Cheirogaleus.[6] Their body weight ranges between 350 and 500 g (0.77
and 1.10 lb),[27] and their head-body length averages between 23.7 and
27.2 cm (9.3 and 10.7 in), with a tail length between 31.9 and 40.1 cm
(12.6 and 15.8 in).[28]

Fork-marked lemurs' dorsal (back) fur is either light brown or light

grayish-brown, while their ventral (underside) fur can be yellow, cream,
white, or pale brown.[6][8] A black stripe extends from the tail, along the
dorsal midline to the head, where it forks at the top of the head in a
distinguishing Y-shape leading to the dark rings around both eyes, and
sometimes extends down the snout. The dorsal stripe varies in width
and darkness.[6][29] The base of the tail is the same color as the dorsal
fur[27] and is usually tipped in black;[6][27] the tail is bushy.[27][30] The P. furcifer, first described in 1839, was
[30] illustrated in Brehms Tierleben.
lemurs' ears are relatively large and membranous. Males have a
scent gland on the middle of their throat,[27] which is approximately
20 mm (0.79 in) wide and pink in color. Females have a narrow, bare
patch of white skin in the same location, but theirs does not appear to produce secretions.[31]

These lemurs have relatively long hindlegs. For gripping tree trunks and large
branches, they have large hands and feet with extended pads on the digits, as
well as claw-like nails.[30][32] They have a long tongue which assists obtaining
the gum and nectar,[30][32] as well as a long caecum, which helps digest
gums.[6][30] Their procumbent (forward-facing) lemuriform toothcomb (formed
by the lower incisors and canines) is long[32] and more compressed, with
significantly reduced interdental spaces to minimize the accumulation of gum
between the teeth.[33]

The genus is distinguished from other cheirogaleids by the toothrows on its

maxilla (upper jaw), which are parallel and do not converge towards the front
of the mouth.[14] The fork-marked lemur dental formula is 2.1.3.3
2.1.3.3
2 = 36; on
Fork-marked lemurs are
distinguished by the dorsal each side of the mouth, top and bottom, there are two incisors, one canine, three
black stripe that forks on the premolars, and three molarsa total of 36 teeth.[34] Their upper first incisor
crown of their head. (I1) is long and curved towards the middle of the mouth (unique among
lemurs),[14][34] while the second upper incisor (I2) is small with a gap
(diastema) between the two.[34] The upper canines are large, with their tips
curved. [14][34] Their upper anterior premolars (P2) are caniniform (canine-shaped)[30][32] and more pronounced
than in any other living lemur.[35] The next upper premolar (P3) is very small,[14] with a single, pointed cusp
that contacts the lingual cingulum (a crest or ridge on the tongue side), which circles the base of the tooth. The
two cusps on the last upper premolar (P4) are a large paracone and a smaller protocone. Like other
cheirogaleids, their first lower premolar (P2) is caniniform and large, while the cingulids (ridges) on the three
lower premolars are more developed compared to most other cheirogaleids. The first two upper molars (M12)
have a developed hypocone, and the buccal cingulum (a crest or ridge on the cheek side) is well developed on
all three upper molars.[35] The molars are relatively small compared to other cheirogaleids, with the second
upper and lower molars (M2 and M2) having reduced functionality compared to those of mouse lemurs.[9]
Males have relatively small testes compared other lemurs, and their canine teeth are the same size as those seen
in females. During the dry season, females can weigh more than males. Both patterns of sexual dimorphism are
consistent with the theory of sexual selection for monogamous species and female dominance respectively.[36]
Females have two pairs of nipples.[36]

Distribution and habitat

Fork-marked lemurs are found in the west, north, and east of Madagascar, but their distribution is
discontinuous.[6][30] Their habitat ranges from dry deciduous forests on the western coast of the island to
rainforest in the east.[32] They are also commonly found in secondary forest, but not in areas lacking
continuous forest cover.[37] They are most common in the west of the island.[30] Fork-marked lemurs are not
found in the southern spiny forests in the dry southern part of the island, and only recently have been reported
from the southeastern rainforest at Andohahela National Park, though this has not been confirmed. A team led
by E. E. Louis Jr. has suggested that undescribed varieties may also exist elsewhere on the island.[6]

The Masoala fork-marked lemur is found on the Masoala peninsula in the northeast of the island,[38] while the
Amber Mountain fork-marked lemur is located in the far north of the island, particularly at Amber Mountain
National Park.[39] Pariente's fork-marked lemur is found in the Sambirano region in the northwest,[40] and the
pale fork-marked lemur is in the west of the island.[41]

Behavior
Fork-marked lemurs are among the least studied of all lemurs, and little is
known about them.[6] Only the pale fork-marked lemur (P. pallescens) has been
studied relatively well, primarily by Pierre Charles-Dominique, Jean-Jacques
Petter, and Georges Pariente during two expeditions in the 1970s and a more
extensive 1998 study in Kirindy Forest.[42] Like the other cheirogaleids, these
lemurs are nocturnal, sleeping in tree hollows (typically in large baobab trees)
or abandoned nests built by giant mouse lemurs (Mirza coquereli) during the
day.[37][30][43] Some of the abandoned nests they sleep in are leaf-lined, and
fresh leaves are often added when young are born. As many as 30 sleeping sites
may be used over the course of a year, each for a variable length of time.[42]

At night, fork-marked lemurs visit the feeding sites within their range by
running quadrupedally across branches[30] at high speed over long
distances,[44] leaping from tree to tree without pausing.[6] They have been seen
on the ground (typically during chases following fights)[37][44] and as high as Males, such as this
10 m (33 ft), but they are typically seen running along branches at a height of 3 P. pallescens, have a scent
to 4 m (9.8 to 13.1 ft).[37] While running, they can leap 4 to 5 m (13 to 16 ft) gland of their throat, which
horizontally between tree branches without losing height or as much as 10 m they only use during social
grooming.
(33 ft) while falling a short distance.[45]

Fork-marked lemurs are sensitive to light intensity,[44] and emerge at twilight,

calling numerous times and answering their neighbors' calls before going off to forage.[32][44] Just before dawn,
they also communicate again on their way to their sleep site. Cold temperatures can also cause individuals to
retire to their sleeping site as early as two hours before dawn.[44] Their eye shine creates a unique pattern
among lemurs because they tend to bob their heads up and down and from side to side.[6]

These lemurs are territorial, with territory size dependent upon food availability,[46] though territories typically
cover 3 to 10 hectares (7.4 to 24.7 acres). Because of their fast movement, individuals can easily defend their
territories by traversing it within 5 minutes.[44] Territory overlap is minimal between males, and the same
pattern is seen in females, though males and females may overlap their territories.[46] In areas where territory
overlap occurs ("meeting areas"), several neighbors may gather and vocalize together without
aggression.[37][36][47] Multiple family groups may gather in these meeting areas, and females will often
aggression.[37][36][47] Multiple family groups may gather in these meeting areas, and females will often
socialize with the other females and young.[36] Unlike other lemurs, fork-marked lemurs do not scent-mark,
and instead use vocalizations during territorial confrontations.[48] They are considered very vocal animals, and
have a complex range of calls.[6] On average, males make approximately 30 loud calls per hour,[37][43] and are
most vocal at dusk and dawn. Their high-pitched, whistling calls help researchers identify them in the field.[6]
As well as their stress call and fighting call, they emit a Hon call (contact call between male-female pairs), Ki
and Kiu calls (more excited contact calls that identify the caller), and a Kea call (a loud call shared between
males in adjacent territories). Females also make a "bleating" call when they have infants.[49]

Males and females have been seen sleeping and foraging together as monogamous pairs, although polygamy
and solitary behavior has also been observed.[50] At Kirindy Forest, pairs were observed staying together for
multiple seasons, though they were only seen foraging alone, with most interactions resulting from conflicts
over feeding sites.[44] Nest sharing among pairs occurs one out of every three days.[44] During social grooming
(allogrooming), the male allomarks females using a scent gland on the throat,[51] and grooming sessions can
last several minutes.[44] While feeding, females appear to be dominant, gaining first access to food.[52] Females
are also dominant over non-resident males, indicating true female dominance, comparable to that seen in the
ring-tailed lemur (Lemur catta).[36]

Mating has been observed to take place at the end of the dry season, in early November, and births were
inferred between late February and early March.[36] Only one infant is born per season,[37][36] despite females
having two pairs of nipples.[36] Infants are initially parked in unguarded tree holes while the mother
forages.[36][53] Older infants have not been observed clinging to the mother, and as they get older, they are
parked in vegetation until they can move independently. Females produce milk for two years following the
birth of the young.[36] The offspring may remain under the care of their parents for three years or more,[44] and
there is no information about their dispersal at maturity. Females have not been observed giving birth in
consecutive years.[36]

Ecology
These lemurs have a specialized diet of tree gums and sap.[6][54] Their diet consists mainly of gum from trees in
the genus Terminalia (known locally as "Talinala"),[6][42][55] which are often parasitized by beetle larvae that
burrow beneath the bark. Fork-marked lemurs either consume the gum as it seeps from cracks in the bark of
parasitized trees or gouge open the bark with their toothcomb to scoop it up directly with their long tongue.
Between March and May, gums compose the majority of the diet.[6][32] They have also been documented eating
gums from Commiphora species and Colvillea racemosa, bud exudates from Zanthoxylum tsihanimposa, sap
from baobab trees (Adansonia species),[32] nectar from Crateva greveana flowers, the sugary excretions from
bugs (family Machaerotidae) which feed on trees of the genus Rhopalocarpus,[6][32][44] and very small amounts
of fruit.[44] Although fork-marked lemurs have widely varied forest habitat, gum and other plant exudates of
other species are likely to dominate their diet.[32] They are not known to estivate or accumulate fat reserves for
the dry season.[45]

To meet their protein requirements and obtain nitrogen, these lemurs also hunt small arthropods. In captivity,
P. furcifer strongly favored preying mantises and moths of the family Sphingidae while ignoring grasshoppers,
larva of the moth genus Coeloptera, and small reptiles. Hunting usually occurs later at night, following gum
collection, and typically happens in the canopy or on tree trunks. Insects are captured by rapidly grasping them
with the hands, a stereotypic behavior seen in other members of their family, as well as galagos.[56] However,
the exudates of several tree species they are known to feed on are high in protein, so some fork-marked lemurs
may meet their protein requirements without preying on insects.[44]

Other nocturnal lemurs are sympatric with fork-marked lemurs. In western Madagascar, interspecific
competition is reduced by restricting activity to specific levels of the canopy, such as using only the highest
sleeping sites at least 8 m (26 ft) above the ground. Competition with other cheirogaleids, such as the gray
mouse lemur (Microcebus murinus) and Coquerel's giant mouse lemur (Mirza coquereli), is most intense for
 Terminalia gum during the dry season, but fork-marked lemurs always drive the
other lemur species off.[42] Studies of P. pallescens at Kirindy Forest found up
to a 20% drop in body mass during the dry season despite no changes in
exudate production, indicating flowers and insects have a significant impact on
the species' health.[44]

The Madagascar harrier-hawk
may prey on fork-marked
lemurs by extracting them from
their sleeping holes.
Fork-marked lemurs are thought to be preyed upon by large owls, such as the
Madagascar owl (Asio madagascariensis), and snakes like the Malagasy tree
boa (Sanzinia madagascariensis).[43] In one case, a family of fork-marked
lemurs exhibited mobbing behavior when they encountered a Malagasy tree
boa.[44] Diurnal raptors, such as the Madagascar buzzard (Buteo brachypterus)
and Madagascar cuckoo-hawk (Aviceda madagascariensis) hunt these lemurs at
dusk,[43][44] and the hunting behavior of the Madagascar harrier-hawk
(Polyboroides radiatus) suggests it might extract them from their sleeping
holes. The fossa (Cryptoprocta ferox) has also been seen attacking fork-marked
lemurs, and remains have been found in their scat.[44]

Conservation
In 2012, the International Union for Conservation of Nature (IUCN) listed P. parienti, P. electromontis, and
P. pallescens as endangered,[4][5][3] and P. furcifer as vulnerable.[2] Before this assessment, it was assumed that
their population was in decline due to habitat destruction for the creation of pasture and agriculture. Measures
of their population density vary widely, from 50 to 550 individuals per square kilometer (250 acres), but these
numbers are thought to reflect only small, gum-rich areas, and therefore only small, clustered populations with
an overall low population density.[37]

As with all lemurs, fork-marked lemurs were first protected in 1969 when they were listed as "Class A" of the
African Convention on the Conservation of Nature and Natural Resources. This prohibited hunting and capture
without authorization, which would only be given for scientific purposes or the national interest. They were
also protected under CITES Appendix I as of 1973. This strictly regulates their trade and forbids commercial
trade. Although enforcement is patchy, they are also protected under Malagasy law. Fork-marked lemurs are
rarely kept in captivity,[37] and their captive lifespan can range from 12[57] to 25 years.[58]

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54. Charles-Dominique & Petter 1980, p. 93.
55. Charles-Dominique & Petter 1980, p. 77.
56. Charles-Dominique & Petter 1980, p. 80.
57. Nowak 1999, p. 71.
58. Weigl 2005, p. 44.
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External links
Original description of Phaner by J.E. Gray, 1870 Biodiversity Heritage Library
Original description of L. furcifer by Blainville, 1839 HathiTrust Digital Library (in French)
Plate with skull used by Blainville, 1839 Bibliothque nationale de France (Gallica) (in French)

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