Editing Slow loris (section)

==Taxonomy and systematics==
Although many previous classifications recognized as few as a single all-inclusive species, there are now at least eight that are considered valid: 
{{Species table |no-note=y |genus=[[Nycticebus]] |authority-name=[[Étienne Geoffroy Saint-Hilaire|É Geoffroy]] |authority-year=1812 |species-count=eight|narrow-percent=75}}
{{Species table/row
|name=[[Bangka slow loris]] |binomial=N. bancanus
|image= |image-size=123px |image-alt=
|authority-name=[[Marcus Ward Lyon Jr.|Lyon]] |authority-year=1906 |authority-not-original=yes
|range=[[Borneo]] and [[Bangka Island]]s in southeastern Asia |range-image= |range-image-size=172px
|size=About {{convert|26|cm|in|0|abbr=on}} long, with no tail<ref name="KayanPaper"/>
|habitat=Forest<ref name="IUCNBangkaslowloris"/>
|hunting=Insects, gum, nectar, and fruit<ref name="PrimatesPerspective2833"/>
|iucn-status=CR |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCNBangkaslowloris"/>
}}
{{Species table/row
|name=[[Bengal slow loris]] |binomial=N. bengalensis
|image=File:Captive N. bengalensis from Laos with 6-week baby.JPG |image-size=123px |image-alt=Brown and white loris
|authority-name=[[Bernard Germain de Lacépède|Lacépède]] |authority-year=1800 |authority-not-original=yes
|range=Southeastern Asia |range-image=File:Bengal Slow Loris area.png |range-image-size=172px
|size={{convert|26|–|38|cm|in|0|abbr=on}} long, with [[vestigial]] tail<ref name="ADWBengalslowloris"/>
|habitat=Forest<ref name="IUCN_N._bengalensis"/>
|hunting=Resin and gum, as well as nectar, fruit, invertebrates, bark, and bird eggs<ref name="ADWBengalslowloris"/>
|iucn-status=EN |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCN_N._bengalensis"/>
}}
{{Species table/row
|name=[[Bornean slow loris]] |binomial=N. borneanus
|image=File:Nycticebus borneanus (Nycticebus coucang borneanus) - Museo Civico di Storia Naturale Giacomo Doria - Genoa, Italy - DSC02519.JPG |image-size=180px |image-alt=Gray loris
|authority-name=[[Marcus Ward Lyon Jr.|Lyon]] |authority-year=1906 |authority-not-original=yes
|range=Borneo |range-image= |range-image-size=172px
|size=About {{convert|26|cm|in|0|abbr=on}} long, with no tail<ref name="KayanPaper"/>
|habitat=Forest<ref name="IUCNBorneanslowloris"/>
|hunting=Insects, gum, nectar, and fruit<ref name="PrimatesIndonesia25"/>
|iucn-status=VU |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCNBorneanslowloris"/>
}}
{{Species table/row
|name=[[Javan slow loris]] |binomial=N. javanicus
|image=File:Coucang.jpg |image-size=180px |image-alt=Light brown loris
|authority-name=[[Étienne Geoffroy Saint-Hilaire|É Geoffroy]] |authority-year=1812
|range=[[Java]] in southeastern Asia |range-image=File:Javan Slow Loris area.png |range-image-size=180px
|size={{convert|28|–|31|cm|in|0|abbr=on}} long, with vestigial tail<ref name="PrimatesIndonesia2122"/>
|habitat=Forest<ref name="IUCN_N._javanicus"/>
|hunting=Nectar, gum, insects, fruit, lizards, and eggs<ref name="PrimatesIndonesia2122"/>
|iucn-status=CR |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCN_N._javanicus"/>
}}
{{Species table/row
|name=[[Kayan River slow loris]] |binomial=N. kayan
|image=File:Slow Loris.jpg |image-size=180px |image-alt=Brown loris
|authority-name=[[Rachel A. Munds|Munds]], [[K. Anne-Isola Nekaris|Nekaris]], [[Susan M. Ford (zoologist)|Ford]] |authority-year=2013
|range=Borneo |range-image= |range-image-size=172px
|size=About {{convert|27|cm|in|0|abbr=on}} long, with no tail<ref name="KayanPaper"/>
|habitat=Forest<ref name="IUCNKayanslowloris"/>
|hunting=Unknown<ref name="PrimatesIndonesia30"/>
|iucn-status=VU |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCNKayanslowloris"/>
}}
{{Species table/row
|name=[[Philippine slow loris]] |binomial=N. menagensis
|image=File:Nycticebus menagensis.png |image-size=120px |image-alt=Gray photo of a loris
|authority-name=[[Richard Lydekker|Lydekker]] |authority-year=1893
|range=Borneo and nearby islands |range-image=File:Range Nycticebus menagensis.png |range-image-size=172px
|size=About {{convert|27|cm|in|0|abbr=on}} long, with no tail<ref name="KayanPaper"/>
|habitat=Forest<ref name="IUCN_N._menagensis"/>
|hunting=Insects, nectar, gum, and fruit<ref name="SlowLorisCranium"/>
|iucn-status=VU |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCN_N._menagensis"/>
}}
{{Species table/row
|name=[[Sumatran slow loris]] |binomial=N. hilleri
|image= |image-size=180px |image-alt=
|authority-name=[[Witmer Stone|Stone]] and [[James A. G. Rehn|Rehn]] |authority-year=1902 |authority-not-original=yes
|range=[[Sumatra]] in southeastern Asia |range-image= |range-image-size=172px
|size={{convert|26|–|30|cm|in|0|abbr=on}} long, with no tail<ref name="SlowLoris2007"/>
|habitat=Forest<ref name="IUCNSumatranslowloris"/>
|hunting=Insects, nectar, gum, and fruit<ref name="IUCNSumatranslowloris"/>
|iucn-status=EN |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCNSumatranslowloris"/>
}}
{{Species table/row
|name=[[Sunda slow loris]] |binomial=N. coucang
|image=File:Nycticebus coucang 004.jpg |image-size=130px |image-alt=Brown loris
|authority-name=[[Pieter Boddaert|Boddaert]] |authority-year=1785 |authority-not-original=yes
|range=Southeastern Asia |range-image=File:Sunda Loris area.png |range-image-size=180px
|size={{convert|27|–|38|cm|in|0|abbr=on}} long, with no tail<ref name="ADWSundaslowloris"/>
|habitat=Forest<ref name="IUCN_N._coucang"/>
|hunting=Sap, gum, nectar, stems, and fruit, as well as arthropods and insects<ref name="ADWSundaslowloris"/>
|iucn-status=EN |population=Unknown
|direction={{decrease|Population declining}}<ref name="IUCN_N._coucang"/>
}}
{{Species table/end}}

Other than the [[pygmy slow loris]] in sister genus ''[[Xanthonycticebus]]'',<ref name=Xanth/> the group's closest relatives are the [[slender loris]]es of southern India and Sri Lanka. Their next closest relatives are the African [[Lorisidae|lorisids]], the [[potto]]s, [[false potto]]s, and [[angwantibo]]s. They are less closely related to the remaining [[Lorisoidea|lorisoids]] (the various types of [[galago]]), and more distantly to the [[lemur]]s of [[Madagascar]]. Their evolutionary history is uncertain since their [[fossil]] record is patchy and [[molecular clock]] studies have given inconsistent results.

===Evolutionary history===
{{cladogram|align=left
|title=Closest living relatives of slow lorises{{Sfn|Perelman|Johnson|Roos|Seuánez|2011|loc=figs. 1, 2}}{{Sfn|Seiffert|Simons|Ryan|Attia|2005|loc=fig. 3}}<ref name=Xanth/>
|clades={{clade|style=font-size:85%; line-height:90%; width:40em;
|label1=[[Lemuriformes]]
|1={{clade
  |label1=[[Lorisoidea]]
  |1={{clade
     |label1=[[Lorisidae]]
     |1={{clade
        |label1=[[Perodicticinae]]
        |1={{clade
           |1=[[Angwantibo]]s (''Arctocebus'')
           |2=[[Potto]]s (''Perodicticus'')
           }}
        |label2= [[Lorisinae]]
        |2={{clade
           |1=[[Slender loris]]es (''Loris'')
           |2={{clade
              |1='''Slow lorises (''Nycticebus'')'''
              |2=[[Pygmy slow loris]] (''Xanthonycticebus'')
              }}
           }}
        }}
     |2=[[Galago]]s (Galagidae)
     }}
  |2=[[Lemur]]s (Lemuroidea)
  }}
}}
}}

Slow lorises (genus ''Nycticebus'') are [[Strepsirrhini|strepsirrhine]] [[primate]]s and are related to other living [[Lorisoidea|lorisoids]], such as the [[pygmy slow loris]] (''Xanthonycticebus''), [[slender loris]]es (''Loris''), [[potto]]s (''Perodicticus''), [[false potto]]s (''Pseudopotto''), [[angwantibo]]s (''Arctocebus''), and [[galago]]s (family Galagidae), and to the [[lemur]]s of [[Madagascar]].{{Sfn|Phillips|Walker|2002|p=91}}<ref name=Xanth/> They are most closely related to the pygmy slow loris, followed by the slender lorises of [[South Asia]], the angwantibos, pottos and false pottos of [[Central Africa|Central]] and [[West Africa]].{{Sfn|Perelman|Johnson|Roos|Seuánez|2011|loc=figs. 1, 2}}{{Sfn|Seiffert|Simons|Ryan|Attia|2005|loc=fig. 3}}<ref name=Xanth/> Lorisoids are thought to have evolved in Africa, where most living species occur;{{Sfn|Seiffert|Simons|Attia|2003|p=421}}{{Sfn|Seiffert|Simons|Ryan|Attia|2005|p=11400}} later, one group may have migrated to Asia and evolved into the slender and slow lorises of today.{{Sfn|Phillips|Walker|2002|pp=93–94}}

Lorises first appear in the Asian fossil record in the [[Miocene]], with records in [[Thailand]] around 18 million years ago (mya){{Sfn|Mein|Ginsburg|1997|pp=783, 805–806}} and in [[Pakistan]] 16&nbsp;mya.{{Sfn|Flynn|Morgan|2005|p=100}} The Thai record is based on a single tooth that most closely resembles living slow lorises and that is tentatively classified as a species of ''Nycticebus''. The species is named [[? Nycticebus linglom|?&nbsp;''Nycticebus linglom'']], using [[open nomenclature]] (the preceding "?" indicates the tentative nature of the assignment).{{Sfn|Mein|Ginsburg|1997|pp=805–806}}

Several lorises are found in the [[Sivalik Hills|Siwalik]] deposits of Pakistan, including ''[[Nycticeboides]]'' and ''[[Microloris]]''; they date from 16 to 8 mya Most are small, but an unnamed form dating to 15–16&nbsp;mya is comparable in size to the largest living slow lorises.{{Sfn|Flynn|Morgan|2005|pp=100–107}} Molecular clock analysis suggests that slow lorises may have started [[speciation|evolving into distinct species]] about 10&nbsp;mya.{{Sfn|Perelman|Johnson|Roos|Seuánez|2011|loc=table 1, fig. 2}} They are thought to have reached the islands of [[Sundaland]] when the [[Sunda Shelf]] was exposed at times of low sea level, creating a [[land bridge]] between the mainland and islands off the coast of Southeast Asia.{{Sfn|Groves|1971|p=52}}

====Discovery and taxonomy====
{{Quote box
| quote = ... it had the face of a bear, the hands of a monkey and moved like a sloth ...
| source = American zoologist [[Dean Conant Worcester]], describing the Bornean slow loris in 1891.{{Sfn|Worcester|Bourns|1905|pp=683–684}}
| align = right
}}
The earliest known mention of a slow loris in scientific literature is from 1770, when Dutchman Arnout Vosmaer (1720–1799) described a specimen of what we know today as ''N.&nbsp;bengalensis''<!--not coucang: Osman Hill writes of the "Bengalese race of slow loris", our N. bengalensis--> that he had received two years earlier. The French naturalist [[Georges-Louis Leclerc, Comte de Buffon]], later questioned Vosmaer's decision to affiliate the animal with sloths, arguing that it was more closely aligned with the lorises of [[Sri Lanka|Ceylon]] (now Sri Lanka) and Bengal.{{Sfn|Osman Hill|1953b|p=45}}  The word "loris" was first used in 1765 by Buffon as a close equivalent to a Dutch name, ''loeris''.  This [[etymology]] was later supported by the [[physician]] [[William Baird (physician)|William Baird]] in the 1820s, who noted that the Dutch word ''loeris'' signified "a clown".{{Sfn|Osman Hill|1953b|pp=44–45}}

In 1785, the Dutch physician and naturalist [[Pieter Boddaert]] was the first to officially [[species description|describe]] a species of slow loris using the name ''Tardigradus coucang''.{{Sfn|Boddaert|1785|p=67}}{{Sfn|Elliot|1913|p=21}}{{Sfn|Osman Hill|1953b|p=46}} This species was based on the "tailless maucauco" described by [[Thomas Pennant]] in 1781, which is thought to have been based on a Sunda slow loris, and on Vosmaer's description of a Bengal slow loris.{{Sfn|Pocock|1939|p=171}} Consequently, there has been some disagreement over the identity of ''Tardigradus coucang''; currently the name is given to the Sunda slow loris.{{Sfn|Thomas|1922}}{{Sfn|Groves|1971|p=49}} The next slow loris species to be described was ''Lori bengalensis'' (currently ''Nycticebus bengalensis''), named by [[Bernard Germain de Lacépède]] in 1800.{{Sfn|Lacépède|1800|p=68}}{{Sfn|Husson|Holthuis|1953|p=213}}

In 1812, [[Étienne Geoffroy Saint-Hilaire]] named the genus ''Nycticebus'',{{Sfn|Geoffroy Saint-Hilaire|1812|p=163}} naming it for its [[Nocturnality|nocturnal]] behavior.  The name derives from the {{langx|grc|νυκτός|(nyktos)}}, [[genitive case|genitive]] form of {{lang|grc|νύξ}} ({{transliteration|grc|nyx}}, "night"), and {{lang|grc|κῆβος}} ({{transliteration|grc|kêbos}}, "monkey").<ref>{{LSJ|nu/c|νύξ|ref}}</ref><ref>{{LSJ|kh{{=}}bos|κῆβος|shortref}}</ref>{{Sfn|Palmer|1904|p=465}} Geoffroy also named ''Nycticebus javanicus'' in this work.{{Sfn|Geoffroy Saint-Hilaire|1812|p=164}} Later 19th-century authors also called the slow lorises ''Nycticebus'', but most used the species name ''tardigradus'' (given by Linnaeus in 1758 in the [[10th edition of Systema Naturae|10th edition of ''Systema Naturæ'']]) for slow lorises, until [[mammalogy|mammalogists]] [[Witmer Stone]] and [[James A. G. Rehn]] clarified in 1902 that Linnaeus's name actually referred to a slender loris.{{Sfn|Stone|Rehn|1902|pp=136–139}}

Several more species were named around 1900, including ''Nycticebus menagensis'' (originally ''Lemur menagensis'') by [[Richard Lydekker]] in 1893{{Sfn|Lydekker|1893|pp=24–25}} and ''Nycticebus pygmaeus'' by [[J. Lewis Bonhote|John James Lewis Bonhote]] in 1907.{{Sfn|Nekaris|Jaffe|2007|pp=187–196}} However, in 1939 [[Reginald Innes Pocock]] consolidated all slow lorises into a single [[species]], ''N.&nbsp;coucang'',{{Sfn|Pocock|1939|p=165}} and in his influential 1953 book ''Primates: Comparative Anatomy and Taxonomy'', primatologist [[William Charles Osman Hill]] also followed this course.{{Sfn|Osman Hill|1953a|pp=156–163}} In 1971 [[Colin Groves]] recognized the [[pygmy slow loris]] (''N.&nbsp;pygmaeus'') as a separate species,{{Sfn|Groves|1971|p=45}} and divided ''N.&nbsp;coucang'' into four [[subspecies]],{{Sfn|Groves|1971|pp=48–49}} while in 2001 Groves opined there were three species (''N.&nbsp;coucang'', ''N.&nbsp;pygmaeus'', and ''N.&nbsp;bengalensis''), and that ''N.&nbsp;coucang'' had three subspecies (''Nycticebus coucang coucang'', ''N.&nbsp;c.&nbsp;menagensis'', and ''N.&nbsp;c.&nbsp;javanicus'').{{Sfn|Groves|2001|p=99}}

[[File:Slow Loris.jpg|thumb|The Kayan River slow loris<ref name="KayanSlowLoris">{{cite web|last=Nekaris|first=Anna|title=Experts gather to tackle slow loris trade|url=http://www.nocturama.org/tag/nycticebus-kayan/|website=Prof Anna Nekaris' Little Fireface Project|publisher=nocturama.org|access-date=30 April 2013|date=23 January 2013|quote=Anna Nekaris, ... who described the new Kayan slow loris, presented the results of her research highlighting the differences between the species.|archive-date=1 February 2014|archive-url=https://web.archive.org/web/20140201181531/http://www.nocturama.org/tag/nycticebus-kayan/|url-status=live}}</ref><ref name=ITIS-kayan>{{ITIS |id=944112|taxon=''Nycticebus kayan''|access-date=28 January 2016}}</ref> (''[[Nycticebus kayan|N.&nbsp;kayan]]'') was distinguished from ''[[Nycticebus menagensis|N. menagensis]]'' in 2012.]]
In 2006, the Bornean slow loris was elevated to the species level (as ''Nycticebus menagensis'') based on [[molecular phylogenetics|molecular analysis]] of [[Nucleic acid sequence|DNA sequences]] of the [[D-loop]] and the [[cytochrome b|cytochrome ''b'']] gene.{{Sfn|Chen|Pan|Groves|Wang|2006|p=1198}}  In 2008, Groves and Ibnu Maryanto confirmed the promotion of the fifth species, the Javan slow loris, to species status, a move that had been suggested in previous studies from 2000.  They based their decision on an analysis of cranial [[morphology (biology)|morphology]] and characteristics of [[fur|pelage]].{{Sfn|Groves|Maryanto|2008|p=120}}  Species differentiation was based largely on differences in morphology, such as size, fur color, and head markings.{{Sfn|Chen|Pan|Groves|Wang|2006|p=1188}}

To help clarify species and subspecies boundaries, and to establish whether morphology-based classifications were consistent with evolutionary relationships, the [[Phylogenetics|phylogenetic]] relationships within the genus ''Nycticebus'' were investigated by Chen and colleagues using DNA sequences derived from the [[mitochondrial DNA|mitochondrial]] markers D-loop and cytochrome ''b''.{{Sfn|Chen|Pan|Groves|Wang|2006}} Previous molecular analyses using [[karyotype]]s,{{Sfn|Chen|Zhang|Shi|Liu|1993|pp=47–53}} [[restriction enzyme]]s,{{Sfn|Zhang|Chen|Shi|1993|pp=167–175}} and DNA sequences{{Sfn|Wang|Su|Lan|Liu|1996|pp=89–93}} were focused on understanding the relationships between a few species, not the phylogeny of the entire genus.{{Sfn|Chen|Pan|Groves|Wang|2006|p=1188}}  The analyses published in 2006 by Chen and colleagues' proved inconclusive, although one test suggested that ''N.&nbsp;coucang'' and ''N.&nbsp;bengalensis'' apparently share a closer evolutionary relationship with each other than with members of their own species, possibly due to [[Introgression|introgressive hybridization]] since the tested individuals of these two taxa originated from a region of [[sympatry]] in southern Thailand.{{Sfn|Chen|Pan|Groves|Wang|2006}}  This hypothesis was corroborated by a 2007 study that compared the variations in mitochondrial DNA sequences between ''N.&nbsp;bengalensis'' and ''N.&nbsp;coucang'', and suggested that there has been [[gene flow]] between the two species.{{Sfn|Pan|Chen|Groves|Wang|2007|pp=791–799}}

In 2012, two [[Synonym (taxonomy)|taxonomic synonyms]] (formerly recognized as subspecies) of ''N. menagensis''—''[[Nycticebus bancanus|N. bancanus]]'' and ''[[Nycticebus borneanus|N. borneanus]]''—were elevated to species status, and a new species—''[[Nycticebus kayan|N. kayan]]''—was also distinguished from the same.  Rachel Munds, Anna Nekaris and Susan Ford based these taxonomic revisions on distinguishable facial markings.{{Sfn|Munds|Nekaris|Ford|2013|p=46}}<ref name=2012BBC/>  With that, the ''N.&nbsp;menagensis'' [[species complex]] that had been collectively known as the Bornean slow loris became four species: the [[Philippine slow loris]] (''N. menagensis''),<ref name=ITIS-menagensis>{{ITIS |id=944109|taxon=''Nycticebus menagensis''|access-date=28 January 2016}}</ref> the [[Bornean slow loris]] (''N. borneanus''),<ref name=ITIS-borneanus>{{ITIS |id=944111|taxon=''Nycticebus borneanus''|access-date=28 January 2016}}</ref> the [[Bangka slow loris]] (''N. bancanus''),<ref name=ITIS-bancanus>{{ITIS |id=944110|taxon=''Nycticebus bancanus''|access-date=28 January 2016}}</ref> and the [[Kayan River slow loris]] (''N. kayan'').<ref name=ITIS-kayan/>

Nekaris and Nijman (2022) combined morphological, behavioural, karyotypical and genetic data and suggested that the pygmy slow loris is best placed in its own genus, ''Xanthonycticebus.''<ref name=Xanth>{{Cite journal |last1=Nekaris |first1=K. Anne-Isola |last2=Nijman |first2=Vincent |date=2022-03-23 |title=A new genus name for pygmy lorises, Xanthonycticebus gen. nov. (Mammalia, primates) |url=https://zse.pensoft.net/article/81942/ |journal=Zoosystematics and Evolution |language=en |volume=98 |issue=1 |pages=87–92 |doi=10.3897/zse.98.81942 |s2cid=247649999 |issn=1860-0743 |doi-access=free |access-date=26 March 2022 |archive-date=18 June 2022 |archive-url=https://web.archive.org/web/20220618095920/https://zse.pensoft.net/article/81942/ |url-status=live }}</ref>
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