View source for Fork-marked lemur

{{Short description|Genus of Madagascan primates}}
{{Automatic taxobox
| image = Phaner pallescens 1985.JPG
| image_caption = [[Pale fork-marked lemur]] (''P.&nbsp;pallescens'')
| image_alt = Lemur with black stripes over its eyes clings to a vertical tree branch.
| status = CITES_A1
| status_system = CITES
| status_ref = {{R|CITES}}
| taxon = Phaner
| authority = [[John Edward Gray|J. E. Gray]], 1870
| type_species = ''[[Masoala fork-marked lemur|Lemur furcifer]]''
| type_species_authority = [[Henri Marie Ducrotay de Blainville|Blainville]], 1839
| subdivision_ranks = Species
| subdivision = {{Plainlist| 
* ''[[Masoala fork-marked lemur|Phaner furcifer]]''
* ''[[Pale fork-marked lemur|Phaner pallescens]]''
* ''[[Pariente's fork-marked lemur|Phaner parienti]]''
* ''[[Amber Mountain fork-marked lemur|Phaner electromontis]]''}}
| range_map = Phaner range map.svg
| range_map_caption = Distribution of ''Phaner'':
'''{{color|#e41a1c|red}}''' = ''P.&nbsp;furcifer''{{R|IUCN_Phaner_furcifer}}<br/>
'''{{color|#4daf4a|green}}''' = ''P.&nbsp;pallescens''{{R|IUCN_Phaner_pallescens}}<br/>
'''{{color|#984ea3|purple}}''' = ''P.&nbsp;parienti''{{R|IUCN_Phaner_parienti}}<br/>
'''{{color|#ff7f00|orange}}''' = ''P.&nbsp;electromontis''{{R|IUCN_Phaner_electromontis}}
| range_map_alt = Map of Madagascar, off the southeast coast of Africa, with a range covering parts of the west, northwest, north, and northeast.
}}

'''Fork-marked lemurs''' or '''fork-crowned lemurs''' are [[Strepsirrhini|strepsirrhine]] primates; the four [[species]] comprise the [[genus]] '''''Phaner'''''. Like all [[lemur]]s, 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 [[Ring-tailed lemur|genus ''Lemur'']] in 1839, later moved between the genera ''[[Dwarf lemur|Cheirogaleus]]'' and ''[[Mouse lemur|Microcebus]]'', and given their own genus in 1870 by [[John Edward Gray]]. Only one species (''[[Masoala fork-marked lemur|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 {{convert|350|g|oz}} or more. They are the most [[Phylogenetics|phylogenetically]] distinct of the cheirogaleids, and considered a [[sister group]] to the rest of the family. Aside from their [[Anatomical terms of location#Dorsal and ventral|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, making repeated calls at the beginning and end of the night. Like the other members of their family, they are [[Nocturnality|nocturnal]], and sleep in tree holes and nests during the day. [[Monogamy in animals|Monogamous pairing]] is typical for fork-marked lemurs, and females are dominant. Females are thought to have only one offspring every two years or more.

These species live in a wide variety of habitats, ranging from [[Tropical and subtropical dry broadleaf forests|dry deciduous forests]] to [[rainforest]]s, and run [[Quadrupedalism|quadrupedally]] across branches. Their diet consists primarily of tree [[Natural gum|gum]] and other [[exudate]]s, though they may obtain some of their [[protein]] and [[nitrogen]] by hunting small [[arthropod]]s later at night. All four species are [[Endangered species|endangered]]. Their populations are in decline due to [[habitat destruction]]. Like all lemurs, they are protected against commercial trade under [[CITES#Appendix I|CITES Appendix I]].

==Taxonomy==

Fork-marked lemurs were first documented in 1839 by [[Henri Marie Ducrotay de Blainville]] when he [[Species description|described]] the [[Masoala fork-marked lemur]] (''P.&nbsp;furcifer'') as ''Lemur furcifer''.{{LoM3 Sfn|pp=211–212}}{{MSW3 Sfn|Groves|id=12100027|heading=Phaner}} The [[holotype]] is thought to be MNHN&nbsp;1834-136, a female specimen taken from Madagascar by French naturalist [[:de:Jules Prosper Goudot|Jules Goudot]]. The source of this specimen is unknown, but thought to be [[Antongil Bay]].{{Sfn|Tattersall|1982|p=131}} In 1850, [[Isidore Geoffroy Saint-Hilaire]] moved the fork-marked lemurs to the genus ''Cheirogaleus'' ([[dwarf lemur]]s),{{Sfn|Szalay|Delson|1980|p=177}} but they were also commonly listed in the genus ''Microcebus'' ([[mouse lemur]]s).{{Sfn|Osman Hill|1953|p=347}} In 1870, [[John Edward Gray]] assigned fork-marked lemurs to their own genus, ''Phaner'',{{LoM3 Sfn|pp=211–212}} after initially including them and the mouse lemurs in the genus ''Lepilemur'' ([[sportive lemur]]s). Although French naturalist [[Alfred Grandidier]] accepted Gray's new genus (while also lumping the other cheirogaleids in ''Cheirogaleus'' and illustrating the [[Skull|cranial]] similarities between cheirogaleids and ''Lepilemur'') in 1897,{{Sfn|Masters|Silvestro|Génin|DelPero|2013|pp=214–215}} the genus ''Phaner'' was not widely accepted. In the early 1930s, [[Ernst Schwarz (zoologist)|Ernst Schwarz]], [[Guillaume Grandidier]], and others resurrected the name, citing characteristics that were intermediate between ''Cheirogaleus'' and ''Microcebus''.{{Sfn|Osman Hill|1953|p=325}}

Until the late 20th&nbsp;century, there was only one recognized species of fork-marked lemur,{{LoM3 Sfn|pp=211–212}} although size and coloration differences had been noted previously.{{Sfn|Tattersall|1982|p=131}} After comparing museum specimens, paleoanthropologist [[Ian Tattersall]] and physical anthropologist [[Colin Groves]] recognized three new subspecies in 1991: the [[Pale fork-marked lemur]] (''P.&nbsp;f.&nbsp;pallescens''), [[Pariente's fork-marked lemur]] (''P.&nbsp;f.&nbsp;parienti''), and the [[Amber Mountain fork-marked lemur]] (''P.&nbsp;f.&nbsp;electromontis'').{{LoM3 Sfn|pp=211–212}}{{Sfn|Groves|Tattersall|1991|p=39}} In 2001, Groves elevated all four subspecies to species status{{LoM3 Sfn|pp=211–212}}{{Sfn|Groves|2001|p=71}} 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,{{Sfn|Tattersall|2007|p=16}} the arrangement is generally accepted.{{LoM3 Sfn|pp=211–212}}

In December&nbsp;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 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.{{R|BBC_News_13Dec2010|CI_13Dec2010}}

===Etymology===
The etymology of the genus ''Phaner'' puzzled researchers for many years. Gray often created mysterious and unexplained [[Taxonomy (biology)|taxonomic names]]. In 1904, [[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 language|Greek]] φανερός (''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&nbsp;November&nbsp;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 lemur]]s) and ''Azema'' (for ''[[Brown mouse lemur|M.&nbsp;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.{{Sfn|Dunkel|Zijlstra|Groves|2012|pp=66–67}}

Fork-marked lemurs were called "fork-marked dwarf lemurs" by [[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.{{Sfn|Dunkel|Zijlstra|Groves|2012|pp=66–67}} These lemurs get their [[common name]] from the distinctive forked stripe on their head.{{LoM3 Sfn|pp=211–212}}

===Evolution===
{{cladogram|title=Competing phylogenies
|caption=Fork-marked lemurs are either a [[sister group]] within Cheirogaleidae (top—Weisrock ''et al.'' 2012){{Sfn|Weisrock|Smith|Chan|Biebouw|2012|p=1626}} or more closely related to sportive lemurs (bottom—Masters ''et al.'' 2013).{{Sfn|Masters|Silvestro|Génin|DelPero|2013|p=209}}
|cladogram={{Clade|style=font-size:85%; line-height:75%; width:34em;
|1={{Clade
   |label1=Lepilemuridae
   |1=''Lepilemur'' ([[sportive lemur]]s)
   |label2=[[Cheirogaleidae]]
   |2={{Clade
      |1='''''Phaner'' (fork-marked lemurs)'''
      |2={{Clade
         |1=''Cheirogaleus'' ([[dwarf lemur]]s)
         |2={{Clade
            |1=''Allocebus'' ([[hairy-eared dwarf lemur]])
            |2={{Clade
               |1=''Mirza'' ([[giant mouse lemur]]s)
               |2=''Microcebus'' ([[mouse lemur]]s)
               }}
            }}
         }}
      }}
   }}
}}
|cladogram2={{Clade|style=font-size:85%; line-height:75%; width:32em;
|1={{Clade
   |1={{Clade
      |1='''''Phaner''''' ('''fork-marked lemurs''')
      |2=''Lepilemur'' (sportive lemurs)
      }}
   |2={{Clade
      |1=''Cheirogaleus'' (dwarf lemurs)
      |2={{Clade
         |1=''Allocebus'' (hairy-eared dwarf lemur)
         |2={{Clade
            |1=''Mirza'' (giant mouse lemurs)
            |2=''Microcebus'' (mouse lemurs)
            }}
         }}
      }}
   }}
}}
}}
Within the family [[Cheirogaleidae]], fork-marked lemurs are the most [[Phylogenetics|phylogenetically]] distinct, although their placement remained uncertain until recently.{{Sfn|Chatterjee|Ho|Barnes|Groves|2009|p=5 of 19}} One uniting characteristic ([[synapomorphy]]) among all cheirogaleids, to the exclusion of other lemurs, is the branching of the [[Common carotid artery|carotid artery]] along with how it enters the skull{{Sfn|Ankel-Simons|2007|p=177}}—a trait which is shared by fork-marked lemurs.{{Sfn|Chatterjee|Ho|Barnes|Groves|2009|p=5 of 19}} Analyses based on [[Morphology (biology)|morphology]], [[immunology]], and [[Repeated sequence (DNA)|repetitive DNA]] have given contradictory placements of ''Phaner'', while studies in 2001 and 2008 either lacked data or yielded poor resolution of their placement.{{Sfn|Weisrock|Smith|Chan|Biebouw|2012|p=1626}}

A study in 2009 of seven [[Mitochondrial DNA|mitochondrial genes]] (mtDNA) and three [[Nuclear DNA|nuclear genes]] grouped fork-marked lemurs with sportive lemurs (family Lepilemuridae), offering a host of explanations, such as a possible hybridization ([[introgression]]) following the initial split between the families.{{Sfn|Chatterjee|Ho|Barnes|Groves|2009|p=5 of 19}} A study published in 2013 also grouped fork-marked lemurs with sportive lemurs{{Sfn|Masters|Silvestro|Génin|DelPero|2013|p=214}} when it used 43&nbsp;morphological traits and mtDNA.{{Sfn|Masters|Silvestro|Génin|DelPero|2013|pp=203–204}} If correct, this would make the family Cheirogaleidae [[Paraphyly|paraphyletic]].{{Sfn|Masters|Silvestro|Génin|DelPero|2013|p=214}} Broad agreement between two [[Evolution of lemurs|lemur phylogeny]] studies—one in 2004 using [[Retrotransposon#SINEs|SINE]] analysis and another in 2012 using [[Locus (genetics)|multilocus]] phylogenetic tests—gave 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.{{Sfn|Weisrock|Smith|Chan|Biebouw|2012|p=1626}} The split between ''Phaner'' and the rest of the cheirogaleids is thought to have occurred approximately 38&nbsp;[[myr|mya]] (million years ago), not long after the [[Adaptive radiation|radiation]] of most of the major lemur groups on Madagascar, roughly 43&nbsp;mya.{{Sfn|Roos|Schmitz|Zischler|2004|p=10653}}{{Sfn|Masters|Silvestro|Génin|DelPero|2013|p=213}}

==Description==
[[File:Katzenmaki (Chirogaleus furcifer).jpg|thumb|right|alt=Black-and-white drawing of two fork-marked lemurs walking quadrupedally through the trees.|''P.&nbsp;furcifer'', first described in 1839, was illustrated in ''[[Brehms Tierleben]]''.]]
Of the mostly small, [[Nocturnality|nocturnal]] lemurs in family Cheirogaleidae, the genus ''Phaner'' contains some of the largest species, along with ''Cheirogaleus''.{{LoM3 Sfn|pp=211–212}} Their body weight ranges between {{convert|350|and|500|g|lb|abbr=on}},{{Sfn|Tattersall|1982|p=132}} and their head-body length averages between {{convert|23.7|and|27.2|cm|in|abbr=on}}, with a tail length between {{convert|31.9|and|40.1|cm|in|abbr=on}}.{{LoM3 Sfn|pp=218, 226, & 228}}

Fork-marked lemurs' [[Anatomical terms of location#Dorsal and ventral|dorsal]] (back) fur is either light brown or light grayish-brown, while their [[Anatomical terms of location#Dorsal and ventral|ventral]] (underside) fur can be yellow, cream, white, or pale brown.{{LoM3 Sfn|pp=211–212}}{{Sfn|Tattersall|1982|p=131}} 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.{{LoM3 Sfn|pp=211–212}}{{Sfn|Tattersall|1982|pp=131–132}} The base of the tail is the same color as the dorsal fur{{Sfn|Tattersall|1982|p=132}} and is usually tipped in black;{{LoM3 Sfn|pp=211–212}}{{Sfn|Tattersall|1982|p=132}} the tail is bushy.{{Sfn|Tattersall|1982|p=132}}{{Sfn|Fleagle|2013|p=63}} The lemurs' ears are relatively large and membranous.{{Sfn|Fleagle|2013|p=63}} Males have a [[scent gland]] on the middle of their [[Gular skin|throat]],{{Sfn|Tattersall|1982|p=132}} which is approximately {{convert|20|mm|in|abbr=on}} 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.{{Sfn|Charles-Dominique|Petter|1980|p=84}}

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.{{Sfn|Fleagle|2013|p=63}}{{Sfn|Charles-Dominique|Petter|1980|p=78}} They have a long tongue which assists obtaining the [[Natural gum|gum]] and [[nectar]],{{Sfn|Fleagle|2013|p=63}}{{Sfn|Charles-Dominique|Petter|1980|p=78}} as well as a long [[caecum]], which helps digest gums.{{LoM3 Sfn|pp=211–212}}{{Sfn|Fleagle|2013|p=63}} Their procumbent (forward-facing) [[Lemuriformes|lemuriform]] [[toothcomb]] (formed by the lower [[incisor]]s and [[Canine tooth|canines]]) is long{{Sfn|Charles-Dominique|Petter|1980|p=78}} and more compressed, with significantly reduced interdental spaces to minimize the accumulation of gum between the teeth.{{Sfn|Szalay|Seligsohn|1977|pp=77–78, & 80}}

[[File:Phaner furcifer 1868.jpg|thumb|upright|left|alt=Illustration of a fork-marked lemur positioned horizontally on a branch.|Fork-marked lemurs are distinguished by the dorsal black stripe that forks on the crown of their head.]]

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.{{Sfn|Groves|2001|p=71}} The fork-marked lemur [[Dentition#Dental formula|dental formula]] is {{DentalFormula|upper=2.1.3.3|lower=2.1.3.3|total=36}}; on each side of the mouth, top and bottom, there are two incisors, one canine, three [[premolar]]s, and three [[Molar (tooth)|molar]]s—a total of 36&nbsp;teeth.{{Sfn|Swindler|2002|p=73}} Their upper first incisor (I<sup>1</sup>) is long and curved towards the middle of the mouth (unique among lemurs),{{Sfn|Groves|2001|p=71}}{{Sfn|Swindler|2002|p=73}} while the second upper incisor (I<sup>2</sup>) is small with a gap ([[diastema]]) between the two.{{Sfn|Swindler|2002|p=73}} The upper canines are large, with their tips curved.{{Sfn|Groves|2001|p=71}}{{Sfn|Swindler|2002|p=73}} Their upper [[Anatomical terms of location#anterior and posterior|anterior]] premolars (P<sup>2</sup>) are caniniform (canine-shaped){{Sfn|Fleagle|2013|p=63}}{{Sfn|Charles-Dominique|Petter|1980|p=78}} and more pronounced than in any other living lemur.{{Sfn|Swindler|2002|pp=75–76}} The next upper premolar (P<sup>3</sup>) is very small,{{Sfn|Groves|2001|p=71}} with a single, pointed [[Cusp (anatomy)|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 (P<sup>4</sup>) are a large [[Molar (tooth)#Morphology|paracone]] and a smaller [[Molar (tooth)#Morphology|protocone]]. Like other cheirogaleids, their first lower premolar (P<sub>2</sub>) 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 (M<sup>1–2</sup>) have a developed [[Molar (tooth)#Morphology|hypocone]], and the buccal cingulum (a crest or ridge on the cheek side) is well developed on all three upper molars.{{Sfn|Swindler|2002|pp=75–76}} The molars are relatively small compared to other cheirogaleids, with the second upper and lower molars (M<sup>2</sup> and M<sub>2</sub>) having reduced functionality compared to those of mouse lemurs.{{Sfn|Szalay|Delson|1980|p=177}}

Males have relatively small [[Testicle|testes]] compared to 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 [[Monogamy in animals|monogamous species]] and female dominance respectively.{{Sfn|Schülke|2003|p=1320}} Females have two pairs of [[nipple]]s.{{Sfn|Schülke|2003|p=1320}}

==Distribution and habitat==
Fork-marked lemurs are found in the west, north, and east of Madagascar, but their distribution is discontinuous.{{LoM3 Sfn|pp=211–212}}{{Sfn|Fleagle|2013|p=63}} Their habitat ranges from [[Tropical and subtropical dry broadleaf forests|dry deciduous forests]] on the western coast of the island to [[rainforest]] in the east.{{Sfn|Charles-Dominique|Petter|1980|p=78}} They are also commonly found in [[secondary forest]], but not in areas lacking continuous forest cover.{{Sfn|Harcourt|1990|pp=67–69}} They are most common in the west of the island.{{Sfn|Fleagle|2013|p=63}} Fork-marked lemurs are not found in the southern [[Madagascar spiny forests|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.&nbsp;E.&nbsp;Louis Jr. has suggested that undescribed varieties may also exist elsewhere on the island.{{LoM3 Sfn|pp=211–212}}

The Masoala fork-marked lemur is found on the [[Masoala Peninsula]] in the northeast of the island,{{LoM3 Sfn|pp=216–217}} while the Amber Mountain fork-marked lemur is located in the far north of the island, particularly at [[Amber Mountain National Park]].{{LoM3 Sfn|pp=228–229}} Pariente's fork-marked lemur is found in the [[Sambirano region]] in the northwest,{{LoM3 Sfn|pp=226–227}} and the pale fork-marked lemur is in the west of the island.{{LoM3 Sfn|pp=218–223}}

==Behavior==
[[File:Phaner pallescens 2008.jpg|thumb|upright|right|alt=View of a male fork-marked lemur in a tree from underneath, showing the scent gland on the throat.|Males, such as this ''P.&nbsp;pallescens'', have a [[scent gland]] on their throat, which they only use during [[social grooming]].]]
Fork-marked lemurs are among the least studied of all lemurs, and little is known about them.{{LoM3 Sfn|pp=211–212}} Only the pale fork-marked lemur (''P.&nbsp;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]].{{Sfn|Schülke|2003|p=1318}} Like the other cheirogaleids, these lemurs are nocturnal, sleeping in tree hollows (typically in large [[Adansonia|baobab trees]]) or abandoned nests built by giant mouse lemurs (''Mirza coquereli'') during the day.{{Sfn|Harcourt|1990|pp=67–69}}{{Sfn|Fleagle|2013|p=63}}{{Sfn|Charles-Dominique|Petter|1980|p=81}} 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&nbsp;sleeping sites may be used over the course of a year, each for a variable length of time.{{Sfn|Schülke|2003|p=1318}}

At night, fork-marked lemurs visit the feeding sites within their range by running [[Quadrupedalism|quadrupedally]] across branches{{Sfn|Fleagle|2013|p=63}} at high speed over long distances,{{Sfn|Schülke|2003|p=1319}} leaping from tree to tree without pausing.{{LoM3 Sfn|pp=211–212}} They have been seen on the ground (typically during chases following fights){{Sfn|Harcourt|1990|pp=67–69}}{{Sfn|Schülke|2003|p=1319}} and as high as {{convert|10|m|ft|abbr=on}}, but they are typically seen running along branches at a height of {{convert|3|to|4|m|ft|abbr=on}}.{{Sfn|Harcourt|1990|pp=67–69}} While running, they can leap {{convert|4|to|5|m|ft|abbr=on}} horizontally between tree branches without losing height or as much as {{convert|10|m|ft|abbr=on}} while falling a short distance.{{Sfn|Nowak|1999|p=70}}

Fork-marked lemurs are sensitive to light intensity,{{Sfn|Schülke|2003|p=1319}} and emerge at twilight, calling numerous times and answering their neighbors' calls before going off to forage.{{Sfn|Charles-Dominique|Petter|1980|p=78}}{{Sfn|Schülke|2003|p=1319}} 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.{{Sfn|Schülke|2003|p=1319}} Their [[Tapetum lucidum|eye shine]] creates a unique pattern among lemurs because they tend to bob their heads up and down and from side to side.{{LoM3 Sfn|pp=211–212}}

[[File:Histoire naturelle des mammifères, Tome VI, Atlas III, Plate 259.jpg|left|thumb|upright|alt=Illustration showing the profile of 9 lemur species from both Cheirogaleidae or Lepilemuridae, demonstrating the similarities in skull shape|In 1897, [[Alfred Grandidier]] demonstrated the similarities between ''Lepilemur'' (middle column, bottom two) and the cheirogaleids, particularly ''Phaner'' (middle, top).]]

These lemurs are [[Territory (animal)|territorial]], with territory size dependent upon food availability,{{Sfn|Charles-Dominique|Petter|1980|p=88}} though territories typically cover {{convert|3|to|10|ha|acre}}. Because of their fast movement, individuals can easily defend their territories by traversing it within 5&nbsp;minutes.{{Sfn|Schülke|2003|p=1319}} Territory overlap is minimal between males, and the same pattern is seen in females, though males and females may overlap their territories.{{Sfn|Charles-Dominique|Petter|1980|p=88}} In areas where territory overlap occurs ("meeting areas"), several neighbors may gather and vocalize together without aggression.{{Sfn|Harcourt|1990|pp=67–69}}{{Sfn|Schülke|2003|p=1320}}{{Sfn|Charles-Dominique|Petter|1980|pp=88–90}} Multiple family groups may gather in these meeting areas, and females will often socialize with the other females and young.{{Sfn|Schülke|2003|p=1320}} Unlike other lemurs, fork-marked lemurs do not scent-mark, and instead use vocalizations during territorial confrontations.{{Sfn|Charles-Dominique|Petter|1980|pp=84–85}} They are considered very vocal animals, and have a complex range of calls.{{LoM3 Sfn|pp=211–212}} On average, males make approximately 30&nbsp;loud calls per hour,{{Sfn|Harcourt|1990|pp=67–69}}{{Sfn|Charles-Dominique|Petter|1980|p=81}} and are most vocal at dusk and dawn. Their high-pitched, whistling calls help researchers identify them in the field.{{LoM3 Sfn|pp=211–212}} 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.{{Sfn|Charles-Dominique|Petter|1980|pp=81–83}}

Males and females have been seen sleeping and [[Forage|foraging]] together as monogamous pairs, although [[Animal sexual behaviour#Polygamy|polygamy]] and solitary behavior has also been observed.{{Sfn|Charles-Dominique|Petter|1980|pp=86 & 93}} 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.{{Sfn|Schülke|2003|p=1319}} Nest sharing among pairs occurs one out of every three days.{{Sfn|Schülke|2003|p=1319}} During [[social grooming]] (allogrooming), the male allomarks females using a scent gland on the throat,{{Sfn|Charles-Dominique|Petter|1980|pp=83–84, 86}} and grooming sessions can last several minutes.{{Sfn|Schülke|2003|p=1319}} While feeding, females appear to be dominant, gaining first access to food.{{Sfn|Charles-Dominique|Petter|1980|pp=85–86}} Females are also dominant over non-resident males, indicating true female dominance, comparable to that seen in the [[ring-tailed lemur]] (''Lemur catta'').{{Sfn|Schülke|2003|p=1320}}

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.{{Sfn|Schülke|2003|p=1320}} Only one infant is born per season,{{Sfn|Harcourt|1990|pp=67–69}}{{Sfn|Schülke|2003|p=1320}} despite females having two pairs of nipples.{{Sfn|Schülke|2003|p=1320}} Infants are initially parked in unguarded tree holes while the mother forages.{{Sfn|Schülke|2003|p=1320}}{{Sfn|Klopfer|Boskoff|1979|pp=136–137}} 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 [[Lactation|produce milk]] for two years following the birth of the young.{{Sfn|Schülke|2003|p=1320}} The offspring may remain under the care of their parents for three years or more,{{Sfn|Schülke|2003|p=1319}} and there is no information about their [[Biological dispersal|dispersal]] at maturity. Females have not been observed giving birth in consecutive years.{{Sfn|Schülke|2003|p=1320}}

==Ecology==
These lemurs have a specialized diet of tree gums and [[Plant sap|sap]].{{LoM3 Sfn|pp=211–212}}{{Sfn|Charles-Dominique|Petter|1980|p=93}} Their diet consists mainly of gum from trees in the genus ''[[Terminalia (plant)|Terminalia]]'' (known locally as "Talinala"),{{LoM3 Sfn|pp=211–212}}{{Sfn|Schülke|2003|p=1318}}{{Sfn|Charles-Dominique|Petter|1980|p=77}} which are often [[Parasitism|parasitized]] by beetle [[larva]]e 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.{{LoM3 Sfn|pp=211–212}}{{Sfn|Charles-Dominique|Petter|1980|p=78}} They have also been documented eating gums from ''[[Commiphora]]'' species and ''[[Colvillea racemosa]]'', bud [[exudate]]s from ''[[Zanthoxylum tsihanimposa]]'', sap from baobab trees (''Adansonia'' species),{{Sfn|Charles-Dominique|Petter|1980|p=78}} nectar from ''[[Crateva greveana]]'' flowers, the sugary excretions from bugs (family [[Machaerotidae]]) which feed on trees of the genus ''[[Rhopalocarpus]]'',{{LoM3 Sfn|pp=211–212}}{{Sfn|Charles-Dominique|Petter|1980|p=78}}{{Sfn|Schülke|2003|p=1319}} and very small amounts of fruit.{{Sfn|Schülke|2003|p=1319}} Although fork-marked lemurs have widely varied forest habitat, gum and other plant exudates of other species are likely to dominate their diet.{{Sfn|Charles-Dominique|Petter|1980|p=78}} They are not known to [[Aestivation|estivate]] or accumulate fat reserves for the dry season.{{Sfn|Nowak|1999|p=70}}

[[File:Madagascar harrier-hawk 3.jpg|thumb|upright|left|alt=Madagascar harrier-hawk sits perched over a tree hole.|The [[Madagascar harrier-hawk]] may prey on fork-marked lemurs by extracting them from their sleeping holes.]]
To meet their [[protein]] requirements and obtain [[nitrogen]], these lemurs also hunt small [[arthropod]]s. In captivity, ''P.&nbsp;furcifer'' strongly favored [[Mantis|preying mantises]] and [[moth]]s of the family [[Sphingidae]] while ignoring [[grasshopper]]s, larva of the moth genus ''[[Coeloptera]]'', and small [[reptile]]s. 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 [[galago]]s.{{Sfn|Charles-Dominique|Petter|1980|p=80}} 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.{{Sfn|Schülke|2003|p=1319}}

Other nocturnal lemurs are [[Sympatry|sympatric]] with fork-marked lemurs. In western Madagascar, [[interspecific competition]] is reduced by restricting activity to specific levels of the [[Canopy (biology)|canopy]], such as using only the highest sleeping sites at least {{convert|8|m|ft|abbr=on}} 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.{{Sfn|Schülke|2003|p=1318}} Studies of ''P.&nbsp;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.{{Sfn|Schülke|2003|p=1319}}

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]]'').{{Sfn|Charles-Dominique|Petter|1980|p=81}} In one case, a family of fork-marked lemurs exhibited [[Mobbing (animal behavior)|mobbing behavior]] when they encountered a Malagasy tree boa.{{Sfn|Schülke|2003|p=1319}} [[Diurnality|Diurnal]] [[Bird of prey|raptors]], such as the [[Madagascar buzzard]] (''Buteo brachypterus'') and [[Madagascar cuckoo-hawk]] (''Aviceda madagascariensis'') hunt these lemurs at dusk,{{Sfn|Charles-Dominique|Petter|1980|p=81}}{{Sfn|Schülke|2003|p=1319}} and the hunting behavior of the [[Madagascar harrier-hawk]] (''Polyboroides radiatus'') suggests it might extract them from their sleeping holes. The [[Fossa (animal)|fossa]] (''Cryptoprocta ferox'') has also been seen attacking fork-marked lemurs, and remains have been found in their scat.{{Sfn|Schülke|2003|p=1319}}

==Conservation==
In 2020, the [[International Union for Conservation of Nature]] (IUCN) listed all four species as [[Endangered species|endangered]].<ref>{{cite web |last1=Louis |first1=E. E. |last2=Randriamampoinana |first2=R. |last3=Bailey |first3=C. A. |last4=Sefczek |first4=T. M. |title=Phaner furcifer |url=https://www.iucnredlist.org/species/16872/115570701 |publisher=The IUCN Red List of Threatened Species |access-date=17 January 2021 |date=2020}}</ref><ref>{{cite web |last1=Borgerson |first1=C. |title=Phaner pallescens |url=https://www.iucnredlist.org/species/16874/115571047 |publisher=The IUCN Red List of Threatened Species |access-date=17 January 2021 |date=2020}}</ref><ref>{{cite web |last1=Louis |first1=E. E. |last2=Mittermeier |first2=R. A. |last3=Randrianambinina |first3=B. |last4=Randriatahina |first4=G. |last5=Rasoloharijaona |first5=S. |last6=Wilmet |first6=L. |last7=Ratsimbazafy |first7=J. |last8=Volampeno |first8=S. |last9=Zaonarivelo |first9=J. |title=Phaner parienti |url=https://www.iucnredlist.org/species/16873/115570887 |publisher=The IUCN Red List of Threatened Species |access-date=17 January 2021 |date=2020}}</ref><ref>{{cite web |last1=Sgarlata |first1=G. M. |last2=Le Pors |first2=B. |last3=Salmona |first3=J. |last4=Hending |first4=D. |last5=Chikhi |first5=L. |last6=Cotton |first6=S. |title=Phaner electromontis |url=https://www.iucnredlist.org/species/16875/115571235 |publisher=The IUCN Red List of Threatened Species |access-date=17 January 2021 |date=2020}}</ref> 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&nbsp;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.{{Sfn|Harcourt|1990|pp=67–69}}

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|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,{{Sfn|Harcourt|1990|pp=67–69}} and their captive lifespan can range from 12{{Sfn|Nowak|1999|p=71}} to 25 years.{{Sfn|Weigl|2005|p=44}}

==References==
{{Reflist|colwidth=30em|refs=

<ref name="BBC_News_13Dec2010">{{cite news |url=http://news.bbc.co.uk/earth/hi/earth_news/newsid_9267000/9267996.stm |title=New species of lemur discovered in Madagascar |work=BBC News |date=13 December 2010 |access-date=24 December 2014 |archive-url=https://web.archive.org/web/20141215023158/http://news.bbc.co.uk/earth/hi/earth_news/newsid_9267000/9267996.stm |archive-date=15 December 2014 |url-status=live }}</ref>

<ref name="CI_13Dec2010">{{cite press release |title=New lemur: big feet, long tongue and the size of squirrel |url=http://www.conservation.org/newsroom/pressreleases/Pages/New-Lemur-BBC-Decade-of-discovery.aspx |publisher=Conservation International |date=13 December 2010 |access-date=24 December 2014 |archive-url=https://web.archive.org/web/20141213065529/http://www.conservation.org/NewsRoom/pressreleases/Pages/New-Lemur-BBC-Decade-of-discovery.aspx |archive-date=13 December 2014 |url-status=live }}</ref>

<ref name="IUCN_Phaner_parienti">{{cite iucn | url = https://www.iucnredlist.org/details/16873/0 | author = Andriaholinirina, N. | display-authors = etal | year = 2012 | title = ''Phaner parienti'' | access-date = 12 December 2014}}</ref>

<ref name="IUCN_Phaner_electromontis">{{cite iucn | url = https://www.iucnredlist.org/details/16875/0 | author = Andriaholinirina, N.  | display-authors = etal | year = 2012 | title = ''Phaner electromontis'' | access-date = 12 December 2014}}</ref>

<ref name="IUCN_Phaner_pallescens">{{cite iucn | url = https://www.iucnredlist.org/details/16874/0 | author = Andriaholinirina, N. | display-authors = etal | year = 2012 | title = ''Phaner pallescens'' | access-date = 12 December 2014}}</ref>

<ref name="IUCN_Phaner_furcifer">{{cite iucn | url = https://www.iucnredlist.org/details/16872/0 | author = Andriaholinirina, N. | display-authors = etal | year = 2012 | title = ''Phaner furcifer'' | access-date = 12 December 2014}}</ref>

<ref name="CITES">{{cite web | url = http://checklist.cites.org/#/en/search/scientific_name=Phaner | title = Checklist of CITES Species | website = CITES | publisher = UNEP-WCMC | access-date = 18 March 2015}}</ref>
}}

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{{Refend}}

==External links==
* [https://www.biodiversitylibrary.org/item/38172#page/148/mode/1up Original description of ''Phaner'' by J.E. Gray, 1870] – Biodiversity Heritage Library
* [http://hdl.handle.net/2027/ucm.5308588778?urlappend=%3Bseq=407 Original description of ''L.&nbsp;furcifer'' by Blainville, 1839] – HathiTrust Digital Library {{in lang|fr}}
* [http://gallica.bnf.fr/ark:/12148/bpt6k65389581/f71.image.r=.langEN Plate with skull used by Blainville, 1839] – Bibliothèque nationale de France (Gallica) {{in lang|fr}}

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[[Category:Lemurs]]
[[Category:Taxa named by John Edward Gray]]

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