Jump to content

Fork-marked lemur

From Wikipedia, the free encyclopedia

Fork-marked lemur
Lemur with black stripes over its eyes clings to a vertical tree branch.
Pale fork-marked lemur(P. pallescens)
CITESAppendix I(CITES)[1]
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Strepsirrhini
Family: Cheirogaleidae
Genus: Phaner
J. E. Gray,1870
Type species
Lemur furcifer
Species
Map of Madagascar, off the southeast coast of Africa, with a range covering parts of the west, northwest, north, and northeast.
Distribution ofPhaner:

red=P. furcifer[2]
green=P. pallescens[3]
purple=P. parienti[4]
orange=P. electromontis[5]

Fork-marked lemursorfork-crowned lemursarestrepsirrhineprimates; the fourspeciescomprise thegenusPhaner.Like alllemurs,they are native toMadagascar,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 thegenusLemurin 1839, later moved between the generaCheirogaleusandMicrocebus,and given their own genus in 1870 byJohn 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 familyCheirogaleidae,weighing around 350 grams (12 oz) or more. They are the mostphylogeneticallydistinct of the cheirogaleids, and considered asister groupto the rest of the family. Aside from theirdorsalforked stripe, they have dark rings around their eyes, and large membranous ears. Males have ascent glandon their throat, but only use it duringsocial 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 arenocturnal,and sleep in tree holes and nests during the day.Monogamous pairingis 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 fromdry deciduous foreststorainforests,and runquadrupedallyacross branches. Their diet consists primarily of treegumand otherexudates,though they may obtain some of theirproteinandnitrogenby hunting smallarthropodslater at night. All four species areendangered.Their populations are in decline due tohabitat destruction.Like all lemurs, they are protected against commercial trade underCITES Appendix I.

Taxonomy[edit]

Fork-marked lemurs were first documented in 1839 byHenri Marie Ducrotay de Blainvillewhen hedescribedtheMasoala fork-marked lemur(P. furcifer) asLemur furcifer.[6][7]Theholotypeis thought to be MNHN 1834-136, a female specimen taken from Madagascar by French naturalistJules Goudot.The source of this specimen is unknown, but thought to beAntongil Bay.[8]In 1850,Isidore Geoffroy Saint-Hilairemoved the fork-marked lemurs to the genusCheirogaleus(dwarf lemurs),[9]but they were also commonly listed in the genusMicrocebus(mouse lemurs).[10]In 1870,John Edward Grayassigned fork-marked lemurs to their own genus,Phaner,[6]after initially including them and the mouse lemurs in the genusLepilemur(sportive lemurs). Although French naturalistAlfred Grandidieraccepted Gray's new genus (while also lumping the other cheirogaleids inCheirogaleusand illustrating thecranialsimilarities between cheirogaleids andLepilemur) in 1897,[11]the genusPhanerwas not widely accepted. In the early 1930s,Ernst Schwarz,Guillaume Grandidier,and others resurrected the name, citing characteristics that were intermediate betweenCheirogaleusandMicrocebus.[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, paleoanthropologistIan Tattersalland physical anthropologistColin Grovesrecognized three new subspecies in 1991: thePale fork-marked lemur(P. f. pallescens),Pariente's fork-marked lemur(P. f. parienti), and theAmber 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 MittermeierofConservation Internationaland conservation geneticistEdward E. Louis Jr.announced the possibility of a new species of fork-marked lemur in the protected area ofDarainain 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 afterFanamby,a key conservation organization working in that protected forest.[16][17]

Etymology[edit]

The etymology of the genusPhanerpuzzled researchers for many years. Gray often created mysterious and unexplainedtaxonomic names.In 1904,Theodore Sherman Palmerattempted to document the etymologies of all mammalian taxa, but could not definitively explain the origins of the generic namePhaner,noting only that it derived from theGreekφανερός (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 comedyThe Palace of TruthbyW. 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 generaMirza(giant mouse lemurs) andAzema(forM. rufus,now a synonym forMicrocebus) were both described in the same publication and equally Enigma tic, 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" byHenry Ogg Forbesin 1894 and "fork-crowned mouse lemur" by English missionary and naturalistJames Sibreein 1895. Literature searches by Dunkelet 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 theircommon namefrom the distinctive forked stripe on their head.[6]

Evolution[edit]

Competing phylogenies
Lepilemuridae

Lepilemur(sportive lemurs)

Cheirogaleidae

Phaner(fork-marked lemurs)

Cheirogaleus(dwarf lemurs)

Phaner(fork-marked lemurs)

Lepilemur(sportive lemurs)

Cheirogaleus(dwarf lemurs)

Allocebus(hairy-eared dwarf lemur)

Mirza(giant mouse lemurs)

Microcebus(mouse lemurs)

Fork-marked lemurs are either asister groupwithin Cheirogaleidae (top—Weisrocket al.2012)[19]or more closely related to sportive lemurs (bottom—Masterset al.2013).[20]

Within the familyCheirogaleidae,fork-marked lemurs are the mostphylogeneticallydistinct, although their placement remained uncertain until recently.[21]One uniting characteristic (synapomorphy) among all cheirogaleids, to the exclusion of other lemurs, is the branching of thecarotid arteryalong with how it enters the skull[22]—a trait which is shared by fork-marked lemurs.[21]Analyses based onmorphology,immunology,andrepetitive DNAhave given contradictory placements ofPhaner,while studies in 2001 and 2008 either lacked data or yielded poor resolution of their placement.[19]

A study in 2009 of sevenmitochondrial genes(mtDNA) and threenuclear genesgrouped 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.[21]A study published in 2013 also grouped fork-marked lemurs with sportive lemurs[23]when it used 43 morphological traits and mtDNA.[24]If correct, this would make the family Cheirogaleidaeparaphyletic.[23]Broad agreement between twolemur phylogenystudies—one in 2004 usingSINEanalysis and another in 2012 usingmultilocusphylogenetic tests—gave strong support for asister grouprelationship between fork-marked lemurs and the rest of the cheirogaleids and a more distant relationship with sportive lemurs.[19]The split betweenPhanerand the rest of the cheirogaleids is thought to have occurred approximately 38mya(million years ago), not long after theradiationof most of the major lemur groups on Madagascar, roughly 43 mya.[25][26]

Description[edit]

Black-and-white drawing of two fork-marked lemurs walking quadrupedally through the trees.
P. furcifer,first described in 1839, was illustrated inBrehms Tierleben.

Of the mostly small,nocturnallemurs in family Cheirogaleidae, the genusPhanercontains some of the largest species, along withCheirogaleus.[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 theirventral(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 lemurs' ears are relatively large and membranous.[30]Males have ascent glandon the middle of theirthroat,[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 thegumandnectar,[30][32]as well as a longcaecum,which helps digest gums.[6][30]Their procumbent (forward-facing)lemuriformtoothcomb(formed by the lowerincisorsandcanines) is long[32]and more compressed, with significantly reduced interdental spaces to minimize the accumulation of gum between the teeth.[33]

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 itsmaxilla(upper jaw), which are parallel and do not converge towards the front of the mouth.[14]The fork-marked lemurdental formulais2.1.3.32.1.3.3 × 2 = 36;on each side of the mouth, top and bottom, there are two incisors, one canine, threepremolars,and threemolars—a total of 36 teeth.[34]Their upper first incisor (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 upperanteriorpremolars (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, pointedcuspthat 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 largeparaconeand a smallerprotocone.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 (M1–2) have a developedhypocone,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 (M2and M2) having reduced functionality compared to those of mouse lemurs.[9]

Males have relatively smalltestescompared to other lemurs, and their canine teeth are the same size as those seen in females. During thedry season,females can weigh more than males. Both patterns ofsexual dimorphismare consistent with the theory ofsexual selectionformonogamous speciesand female dominance respectively.[36]Females have two pairs ofnipples.[36]

Distribution and habitat[edit]

Fork-marked lemurs are found in the west, north, and east of Madagascar, but their distribution is discontinuous.[6][30]Their habitat ranges fromdry deciduous forestson the western coast of the island torainforestin the east.[32]They are also commonly found insecondary 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 southernspiny forestsin the dry southern part of the island, and only recently have been reported from the southeastern rainforest atAndohahela 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 theMasoala Peninsulain the northeast of the island,[38]while the Amber Mountain fork-marked lemur is located in the far north of the island, particularly atAmber Mountain National Park.[39]Pariente's fork-marked lemur is found in theSambirano regionin the northwest,[40]and the pale fork-marked lemur is in the west of the island.[41]

Behavior[edit]

View of a male fork-marked lemur in a tree from underneath, showing the scent gland on the throat.
Males, such as thisP. pallescens,have ascent glandon their throat, which they only use duringsocial grooming.

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 inKirindy Forest.[42]Like the other cheirogaleids, these lemurs are nocturnal, sleeping in tree hollows (typically in largebaobab 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 runningquadrupedallyacross 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 10 m (33 ft), but they are typically seen running along branches at a height of 3 to 4 m (9.8 to 13.1 ft).[37]While running, they can leap 4 to 5 m (13 to 16 ft) horizontally between tree branches without losing height or as much as 10 m (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]Theireye shinecreates a unique pattern among lemurs because they tend to bob their heads up and down and from side to side.[6]

Illustration showing the profile of 9 lemur species from both Cheirogaleidae or Lepilemuridae, demonstrating the similarities in skull shape
In 1897,Alfred Grandidierdemonstrated the similarities betweenLepilemur(middle column, bottom two) and the cheirogaleids, particularlyPhaner(middle, top).

These lemurs areterritorial,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 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 aHoncall (contact call between male-female pairs),KiandKiucalls (more excited contact calls that identify the caller), and aKeacall (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 andforagingtogether as monogamous pairs, althoughpolygamyand 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]Duringsocial 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 thering-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. Femalesproduce milkfor 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 theirdispersalat maturity. Females have not been observed giving birth in consecutive years.[36]

Ecology[edit]

These lemurs have a specialized diet of tree gums andsap.[6][54]Their diet consists mainly of gum from trees in the genusTerminalia(known locally as "Talinala" ),[6][42][55]which are oftenparasitizedby beetlelarvaethat 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 fromCommiphoraspecies andColvillea racemosa,budexudatesfromZanthoxylum tsihanimposa,sap from baobab trees (Adansoniaspecies),[32]nectar fromCrateva greveanaflowers, the sugary excretions from bugs (familyMachaerotidae) which feed on trees of the genusRhopalocarpus,[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 toestivateor accumulate fat reserves for the dry season.[45]

Madagascar harrier-hawk sits perched over a tree hole.
TheMadagascar harrier-hawkmay prey on fork-marked lemurs by extracting them from their sleeping holes.

To meet theirproteinrequirements and obtainnitrogen,these lemurs also hunt smallarthropods.In captivity,P. furciferstrongly favoredpreying mantisesandmothsof the familySphingidaewhile ignoringgrasshoppers,larva of the moth genusCoeloptera,and smallreptiles.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 asgalagos.[56]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 aresympatricwith fork-marked lemurs. In western Madagascar,interspecific competitionis reduced by restricting activity to specific levels of thecanopy,such as using only the highest sleeping sites at least 8 m (26 ft) above the ground. Competition with other cheirogaleids, such as thegray mouse lemur(Microcebus murinus) andCoquerel's giant mouse lemur(Mirza coquereli), is most intense forTerminaliagum during the dry season, but fork-marked lemurs always drive the other lemur species off.[42]Studies ofP. pallescensat 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]

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

Conservation[edit]

In 2020, theInternational Union for Conservation of Nature(IUCN) listed all four species asendangered.[57][58][59][60]Before this assessment, it was assumed that their population was in decline due tohabitat destructionfor the creation of pasture and agriculture. Measures of theirpopulation densityvary 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 theAfrican 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 underCITES Appendix Ias 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[61]to 25 years.[62]

References[edit]

  1. ^"Checklist of CITES Species".CITES.UNEP-WCMC.Retrieved18 March2015.
  2. ^Andriaholinirina, N.; et al. (2012)."Phaner furcifer".IUCN Red List of Threatened Species.2012.Retrieved12 December2014.
  3. ^Andriaholinirina, N.; et al. (2012)."Phaner pallescens".IUCN Red List of Threatened Species.2012.Retrieved12 December2014.
  4. ^Andriaholinirina, N.; et al. (2012)."Phaner parienti".IUCN Red List of Threatened Species.2012.Retrieved12 December2014.
  5. ^Andriaholinirina, N.; et al. (2012)."Phaner electromontis".IUCN Red List of Threatened Species.2012.Retrieved12 December2014.
  6. ^abcdefghijklmnopqrstuvwMittermeier et al. 2010,pp. 211–212.
  7. ^Groves 2005,Phaner.
  8. ^abcTattersall 1982,p. 131.
  9. ^abSzalay & Delson 1980,p. 177.
  10. ^Osman Hill 1953,p. 347.
  11. ^Masters et al. 2013,pp. 214–215.
  12. ^Osman Hill 1953,p. 325.
  13. ^Groves & Tattersall 1991,p. 39.
  14. ^abcdeGroves 2001,p. 71.
  15. ^Tattersall 2007,p. 16.
  16. ^"New species of lemur discovered in Madagascar".BBC News.13 December 2010.Archivedfrom the original on 15 December 2014.Retrieved24 December2014.
  17. ^"New lemur: big feet, long tongue and the size of squirrel"(Press release). Conservation International. 13 December 2010.Archivedfrom the original on 13 December 2014.Retrieved24 December2014.
  18. ^abDunkel, Zijlstra & Groves 2012,pp. 66–67.
  19. ^abcWeisrock et al. 2012,p. 1626.
  20. ^Masters et al. 2013,p. 209.
  21. ^abcChatterjee et al. 2009,p. 5 of 19.
  22. ^Ankel-Simons 2007,p. 177.
  23. ^abMasters et al. 2013,p. 214.
  24. ^Masters et al. 2013,pp. 203–204.
  25. ^Roos, Schmitz & Zischler 2004,p. 10653.
  26. ^Masters et al. 2013,p. 213.
  27. ^abcdeTattersall 1982,p. 132.
  28. ^Mittermeier et al. 2010,pp. 218, 226, & 228.
  29. ^Tattersall 1982,pp. 131–132.
  30. ^abcdefghijFleagle 2013,p. 63.
  31. ^Charles-Dominique & Petter 1980,p. 84.
  32. ^abcdefghijCharles-Dominique & Petter 1980,p. 78.
  33. ^Szalay & Seligsohn 1977,pp. 77–78, & 80.
  34. ^abcdSwindler 2002,p. 73.
  35. ^abSwindler 2002,pp. 75–76.
  36. ^abcdefghijkSchülke 2003,p. 1320.
  37. ^abcdefghiHarcourt 1990,pp. 67–69.
  38. ^Mittermeier et al. 2010,pp. 216–217.
  39. ^Mittermeier et al. 2010,pp. 228–229.
  40. ^Mittermeier et al. 2010,pp. 226–227.
  41. ^Mittermeier et al. 2010,pp. 218–223.
  42. ^abcdSchülke 2003,p. 1318.
  43. ^abcdCharles-Dominique & Petter 1980,p. 81.
  44. ^abcdefghijklmnopqSchülke 2003,p. 1319.
  45. ^abNowak 1999,p. 70.
  46. ^abCharles-Dominique & Petter 1980,p. 88.
  47. ^Charles-Dominique & Petter 1980,pp. 88–90.
  48. ^Charles-Dominique & Petter 1980,pp. 84–85.
  49. ^Charles-Dominique & Petter 1980,pp. 81–83.
  50. ^Charles-Dominique & Petter 1980,pp. 86 & 93.
  51. ^Charles-Dominique & Petter 1980,pp. 83–84, 86.
  52. ^Charles-Dominique & Petter 1980,pp. 85–86.
  53. ^Klopfer & Boskoff 1979,pp. 136–137.
  54. ^Charles-Dominique & Petter 1980,p. 93.
  55. ^Charles-Dominique & Petter 1980,p. 77.
  56. ^Charles-Dominique & Petter 1980,p. 80.
  57. ^Louis, E. E.; Randriamampoinana, R.; Bailey, C. A.; Sefczek, T. M. (2020)."Phaner furcifer".The IUCN Red List of Threatened Species.Retrieved17 January2021.
  58. ^Borgerson, C. (2020)."Phaner pallescens".The IUCN Red List of Threatened Species.Retrieved17 January2021.
  59. ^Louis, E. E.; Mittermeier, R. A.; Randrianambinina, B.; Randriatahina, G.; Rasoloharijaona, S.; Wilmet, L.; Ratsimbazafy, J.; Volampeno, S.; Zaonarivelo, J. (2020)."Phaner parienti".The IUCN Red List of Threatened Species.Retrieved17 January2021.
  60. ^Sgarlata, G. M.; Le Pors, B.; Salmona, J.; Hending, D.; Chikhi, L.; Cotton, S. (2020)."Phaner electromontis".The IUCN Red List of Threatened Species.Retrieved17 January2021.
  61. ^Nowak 1999,p. 71.
  62. ^Weigl 2005,p. 44.

Literature cited[edit]

External links[edit]


Retrieved from "https://en.wikipedia.org/w/index.php?title=Fork-marked_lemur&oldid=1231471595"