Editing Red panda (section)

== Taxonomy ==
[[File:Anh48 0368 1.jpg|thumb|Watercolour painting of a red panda commissioned by [[Thomas Hardwicke]] {{circa|1820}}<ref name=Lowther2021>{{cite journal |author=Lowther, D. A. |year= 2021 |title= The first painting of the Red Panda (''Ailurus fulgens'') in Europe? Natural history and artistic patronage in early nineteenth-century India |journal= Archives of Natural History |volume= 48 |issue=2 |pages=368–376 |doi=10.3366/anh.2021.0728|s2cid= 244938631}}</ref>|alt=Watercolour of a red panda on branch with three separate depictions of the paws at the top]]
The red panda was [[Species description|described]] and named in 1825 by [[Frederic Cuvier]], who gave it its current [[scientific name]] ''Ailurus fulgens''. Cuvier's description was based on [[zoological specimen]]s, including skin, paws, jawbones and teeth "from the mountains north of India", as well as an account by [[Alfred Duvaucel]].<ref>{{cite book |author=Cuvier, F. |year=1825 |chapter=Panda |page=LII 1–3 |chapter-url=https://archive.org/details/HistoirenaturelVGeof/page/LII |editor1=Geoffroy Saint-Hilaire, E. |editor2=Cuvier, F. |title=Histoire naturelle des mammifères, avec des figures originales, coloriées, dessinées d'après des animaux vivans: publié sous l'autorité de l'administration du Muséum d'Histoire naturelle |volume=5 |publisher=A. Belin |location=Paris |language=fr}}</ref><ref>{{cite book |author=Cuvier, G. |year=1829 |title=Le règne animal distribué d'après son organisation |volume=1 |publisher=Chez Déterville|location= Paris |chapter=''Le Panda éclatant'' |page=138 |chapter-url=https://archive.org/stream/lergneanimaldi01regn#page/138/mode/2up |language=fr}}</ref> The red panda was described earlier by [[Thomas Hardwicke]] in 1821, but his paper was only published in 1827.<ref name="Introduction2022" /><ref>{{cite journal |author=Hardwicke, T. |year=1827 |title=Description of a new genus of the class Mammalia, from the Himalaya chain of hills between Nepal and the Snowy Mountains |journal=The Transactions of the Linnean Society of London |volume=XV |pages=161–165 |url=https://archive.org/stream/transactionsofli15182627linn#page/160/mode/2up}}</ref> In 1902, [[Oldfield Thomas]] described a skull of a male red panda specimen under the name ''Ailurus fulgens styani'' in honour of [[Frederick William Styan]] who had collected this specimen in [[Sichuan]].<ref name="thomas_1902"/>

===Subspecies and species===
The modern red panda is the [[monotypic taxon|only recognised species]] in the genus ''Ailurus''. It is traditionally divided into two [[subspecies]]: the Himalayan red panda (''A. f. fulgens'') and the Chinese red panda (''A. f. styani''). The Himalayan subspecies has a straighter profile, a lighter coloured forehead and [[ochre]]-tipped hairs on the lower back and rump. The Chinese subspecies has a more curved forehead and sloping snout, a darker coat with a less white face and more contrast between the tail rings.<ref name="taxonomy"/>

In 2020, results of a genetic analysis of red panda samples showed that the red panda populations in the Himalayas and China were separated about 250,000 years ago. The researchers suggested that the two subspecies should be treated as distinct species. Red pandas in southeastern Tibet and northern Myanmar were found to be part of ''styani'', while those of southern Tibet were of ''fulgens'' in the strict sense.<ref name=Hu>{{Cite journal |last1=Hu|first1=Y. |last2=Thapa|first2=A. |last3=Fan |first3=H. |last4=Ma |first4=T. |last5=Wu|first5=Q. |last6=Ma|first6=S. |last7=Zhang|first7=D. |last8=Wang|first8=B. |last9=Li|first9=M. |last10=Yan|first10=L. |last11=Wei|first11=F. |name-list-style=amp |title=Genomic evidence for two phylogenetic species and long-term population bottlenecks in red pandas |year=2020 |journal=Science Advances |volume=6 |issue=9 |page=eaax5751 |doi=10.1126/sciadv.aax5751 |pmid=32133395 |pmc=7043915 |bibcode=2020SciA....6.5751H}}</ref> [[DNA sequencing]] of 132 red panda faecal samples collected in [[Northeast India]] and China also showed two distinct clusters indicating that the [[Brahmaputra River|Siang River]] constitutes the boundary between the Himalayan and Chinese red pandas.<ref name="Joshi">{{cite journal |author1=Joshi, B. D. |name-list-style=amp |author2=Dalui, S. |author3=Singh, S. K. |author4=Mukherjee, T. |author5=Chandra, K. |author6=Sharma, L. K. |author7=Thakur, M. |year=2021 |title=Siang river in Arunachal Pradesh splits Red Panda into two phylogenetic species |journal=Mammalian Biology |volume=101 |issue=1 |pages=121–124 |doi=10.1007/s42991-020-00094-y |s2cid=231811193}}</ref> They probably diverged due to [[glaciation]] events on the southern [[Tibetan Plateau]] in the [[Pleistocene]].<ref>{{cite journal |author1=Dalui, S. |name-list-style=amp |author2=Singh, S. K. |author3= Joshi, B. D. |author4=Ghosh, A. |author5=Basu, S. |author6=Khatri, H. |author7=Sharma, L. K.|author8=Chandra, K. |author9=Thakur, M. |year=2021 |title=Geological and Pleistocene glaciations explain the demography and disjunct distribution of Red Panda (''A. fulgens'') in eastern Himalayas |journal=Scientific Reports |volume=11 |issue=1 |page=65 |doi=10.1038/s41598-020-80586-6 |pmid=33420314 |pmc=7794540 |doi-access=free}}</ref>

===Phylogeny===
The placement of the red panda on the [[evolutionary tree]] has been debated. In the early 20th century, various scientists placed it in the [[Family (biology)|family]] [[Procyonidae]] with [[raccoon]]s and their allies. At the time, most prominent biologists also considered the red panda to be related to the giant panda, which would eventually be found to be a [[bear]]. A 1982 study examined the similarities and differences in the skull between the red panda and the giant panda, other bears and procyonids, and placed the species in its own family [[Ailuridae]]. The author of the study considered the red panda to be more closely related to bears.<ref name="taxonomy">{{cite book |author=Groves, C. |year=2021 |chapter=The taxonomy and phylogeny of ''Ailurus'' |title=Red Panda: Biology and Conservation of the First Panda |editor=Glatston, A. R. |pages=95–117 |edition=Second |publisher=Academic Press |place=London |isbn=978-0-12-823753-3}}</ref> A 1995 [[mitochondrial DNA]] analysis revealed that the red panda has close affinities with procyonids.<ref>{{cite journal |author1=Pecon-Slattery, J. |name-list-style=amp |author2=O'Brien, S. J. |year=1995 |title=Molecular phylogeny of the red panda (''Ailurus fulgens'') |journal=The Journal of Heredity |volume=86 |issue=6 |pages=413–422 |doi=10.1093/oxfordjournals.jhered.a111615 |pmid=8568209}}</ref> Further genetic studies in 2005, 2018 and 2021 have placed the red panda within the [[clade]] [[Musteloidea]], which also includes Procyonidae, [[Mustelidae]] ([[weasel]]s and relatives) and [[Mephitidae]] ([[skunk]]s and relatives).<ref name="Flynn2005">{{Cite journal |author1=Flynn, J. J. |author2=Finarelli, J. A. |author3=Zehr, S. |author4=Hsu, J. |author5=Nedbal, M. A. |title=Molecular phylogeny of the Carnivora (Mammalia): Assessing the impact of increased sampling on resolving enigmatic relationships |doi=10.1080/10635150590923326 |journal=Systematic Biology |volume=54 |issue=2 |pages=317–337 |year=2005 |pmid=16012099 |doi-access=free |name-list-style=amp}}</ref><ref name="Law-2018">{{Cite journal |author1=Law, C. J. |author2=Slater, G. J. |author3=Mehta, R. S. |name-list-style=amp |date=2018 |title=Lineage Diversity and Size Disparity in Musteloidea: Testing Patterns of Adaptive Radiation Using Molecular and Fossil-Based Methods|journal=Systematic Biology |volume=67|issue=1 |pages=127–144 |doi=10.1093/sysbio/syx047|pmid=28472434 |doi-access=free}}</ref><ref name=Hassanin>{{cite journal|author1=Hassanin, A.|author2=Veron, G.|author3=Ropiquet, A.|author4=van Vuuren, B. J.|author5=Lécu, A.|author6=Goodman, S. M.|author7=Haider, J.|author8=Nguyen, T. T.|year=2021|title=Evolutionary history of Carnivora (Mammalia, Laurasiatheria) inferred from mitochondrial genomes|journal=PLOS ONE |volume=16|issue=2|page=e0240770|doi=10.1371/journal.pone.0240770 |pmid=33591975 |pmc=7886153 |bibcode=2021PLoSO..1640770H |doi-access=free}}</ref>

{{Clade gallery
|main-caption = Phylogenetic relationship of the red panda based on analysis of
|header1 = Six genes from 76 Carnivora species<ref name="Flynn2005"/>
|header2 = 46 genes from 75 musteloid species<ref name="Law-2018"/>
|header3 = Mitogenomes from 220 mammal species<ref name=Hassanin/>
|align=left |width=370px |height=340px |width2=275px
|cladogram1 = {{clade |style=font-size:85%;line-height:85% |label1=[[Caniformia]]
|1={{clade
  |1 =[[Canidae]] [[File:Dogs, jackals, wolves, and foxes (Plate XI).jpg|60px|African golden wolf]]
  |label2=[[Arctoidea]]
  |2={{clade
    |1=[[Ursidae]] [[File:Recherches pour servir à l'histoire naturelle des mammifères (Pl. 50) (white background).jpg|50px|Giant panda]]
    |2={{clade
      |1=[[Pinnipedia]] [[File:Cambridge Natural History Mammalia Fig 230 white background.jpg|70px|Common seal]]
      |label2=[[Musteloidea]]
      |2={{clade
        |1=[[Ailuridae]] [[File:Ailurus fulgens - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|60px|Red panda]]          
        |2={{clade
          |1=[[Mephitidae]] [[File:Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen (Plate CXXI-) (white background).jpg|50px|Striped skunk]]
          |2={{clade
             |1=[[Procyonidae]] |image1=[[File:Wild animals of North America, intimate studies of big and little creatures of the mammal kingdom (Page 410) (white background).jpg|50px|Common raccoon]]
             |2=[[Mustelidae]] |image2=[[File:Fitch white background.png|60px|European polecat]]
}} }} }} }} }} }} }}
|cladogram2 = {{clade |style=font-size:85%;line-height:85%; |label1=[[Musteloidea]]
 |1={{clade
   |1=[[Mephitidae]] [[File:Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen (Plate CXXI-) (white background).jpg|50px|Striped skunk]]
   |2={{clade
    |1=[[Ailuridae]] [[File:Ailurus fulgens - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|60px|Red panda]]
    |2={{clade
      |1=[[Procyonidae]] |image1=[[File:Wild animals of North America, intimate studies of big and little creatures of the mammal kingdom (Page 410) (white background).jpg|50px|Common raccoon]]
      |2=[[Mustelidae]] |image2=[[File:Fitch white background.png|60px|European polecat]]
}} }} }} }}
|cladogram3 = {{clade | style=font-size:85%;line-height:85% |label1=[[Caniformia]]
 |1={{clade
   |1 =[[Canidae]] [[File:Dogs, jackals, wolves, and foxes (Plate XI).jpg|60px|African golden wolf]]
   |label2=[[Arctoidea]]
   |2={{clade
    |1=[[Ursidae]] [[File:Recherches pour servir à l'histoire naturelle des mammifères (Pl. 50) (white background).jpg|50px|Giant panda]]
    |2={{clade
      |1=[[Pinnipedia]] [[File:Cambridge Natural History Mammalia Fig 230 white background.jpg|70px|Common seal]]
      |label3=[[Musteloidea]]
      |3={{clade
        |1={{clade
          |1=[[Ailuridae]] |image1=[[File:Ailurus fulgens - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|60px|Red panda]]          
          |2=[[Mephitidae]] [[File:Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen (Plate CXXI-) (white background).jpg|50px|Striped skunk]]
    }}
 |2={{clade
   |1=[[Procyonidae]] |image1=[[File:Wild animals of North America, intimate studies of big and little creatures of the mammal kingdom (Page 410) (white background).jpg|50px|Common raccoon]]
   |2=[[Mustelidae]] |image2=[[File:Fitch white background.png|60px|European polecat]]
}} }} }} }} }} }} }}
{{clear}}

===Fossil record===
[[File:Simocyon batalleri head.jpg|right|thumb|Reconstructed skull and head of ''[[Simocyon]]'', a large carnivorous early relative of the modern red panda|alt=Drawing of a skull (above) and head (below) of an extinct animal]] 
The family Ailuridae appears to have evolved in Europe in either the [[Late Oligocene]] or [[Early Miocene]], about {{mya|25|18}}. The earliest member ''[[Amphictis]]'' is known from its {{cvt|10|cm|in|frac=2}} skull and may have been around the same size as the modern species. Its [[dentition]] consists of sharp premolars and [[carnassial]]s (P4 and m1) and [[Molar (tooth)|molar]]s adapted for grinding (M1, M2 and m2), suggesting that it had a generalised [[carnivorous]] diet. Its placement within Ailuridae is based on the grooves on the side of its [[Canine tooth|canine teeth]]. Other early or [[Basal (phylogenetics)|basal]] ailurids include ''[[Alopecocyon]]'' and ''[[Simocyon]]'', whose fossils have been found throughout Eurasia and North America dating from the [[Middle Miocene]], the latter of which survived into the [[Early Pliocene]].  Both have similar teeth to ''Amphictis'' and thus had a similar diet.<ref name="Salesa">{{cite book |author1=Salesa, M. J. |author2=Peigné, S. |author3=Antón, M. |author4=Morales, J. |name-list-style=amp |year=2021 |chapter=The taxonomy and phylogeny of ''Ailurus'' |title=Red Panda: Biology and Conservation of the First Panda |editor=Glatston, A. R. |pages=15–29 |edition=Second |publisher=Academic Press |place=London |isbn=978-0-12-823753-3}}</ref> The [[cougar|puma]]-sized ''Simocyon'' was likely a tree-climber and shared a "false thumb"—an extended wrist bone—with the modern species, suggesting the appendage was an adaptation to [[arboreal locomotion]] and not to feed on [[bamboo]].<ref name="Salesa"/><ref name="PNAS-Salesa-2006">{{cite journal |last1=Salesa |first1=M. J. |last2=Mauricio |first2=A. |last3=Peigné |first3=S. |last4=Morales |first4=J. |name-list-style=amp |title=Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas |journal=PNAS |year=2006 |volume=103 |pmid=16387860 |issue=2 |pmc=1326154 |pages=379–382 |doi=10.1073/pnas.0504899102 |bibcode=2006PNAS..103..379S |doi-access=free}}</ref>

Later and more [[Apomorphy and synapomorphy|advanced]] ailurids are classified in the subfamily Ailurinae and are known as the "true" red pandas. These animals were smaller and more adapted for an [[omnivorous]] or [[herbivorous]] diet. The earliest known true panda is ''[[Magerictis]]'' from the Middle Miocene of Spain and known only from one tooth, a lower second molar. The tooth shows both ancestral and new characteristics having a relatively low and simple [[Crown (tooth)|crown]] but also a lengthened crushing surface with developed [[Cusp (anatomy)|tooth cusps]] like later species.<ref name="Wallace">{{cite book |author1=Wallace, S. C. |author2=Lyon, L. |name-list-style=amp |year=2021 |chapter=Systemic revision of the Ailurinae (Mammalia: Carnivora: Ailuridae): with a new species from North America |title=Red Panda: Biology and Conservation of the First Panda |editor=Glatston, A. R. |pages=31–52|edition=Second |publisher=Academic Press |place=London |isbn=978-0-12-823753-3}}</ref> Later ailurines include ''[[Pristinailurus bristoli]]'' which lived in eastern North America from the late Miocene to the Early Pliocene<ref name="Wallace"/><ref name="Wallace2004">{{cite journal |last1=Wallace |first1=S. C. |last2=Wang |first2=X. |name-list-style=amp |date=2004 |title=Two new carnivores from an unusual late Tertiary forest biota in eastern North America |journal=Nature |issue=7008 |pages=556–559 |volume=431 |doi=10.1038/nature02819 |pmid=15457257 |bibcode=2004Natur.431..556W |s2cid=4432191|url=http://doc.rero.ch/record/13466/files/PAL_E141.pdf }}</ref> and species of the genus ''[[Parailurus]]'' which first appear in Early Pliocene Europe, spreading across Eurasia into North America.<ref name="Wallace"/><ref>{{cite journal |author=Tedford, R. H. |author2=Gustafson, E. P. |name-list-style=amp |title=First North American record of the extinct panda ''Parailurus'' |journal=Nature |issue=5595 |pages=621–623 |volume=265 |doi=10.1038/265621a0 |year=1977 |bibcode=1977Natur.265..621T |s2cid=4214900}}</ref> These animals are classified as a [[sister taxon]] to the lineage of the modern red panda. In contrast to the herbivorous modern species, these ancient pandas were likely omnivores, with highly cusped molars and sharp [[premolar]]s.<ref name="Wallace"/><ref name="Wallace2004" /><ref>{{cite journal |last1=Sotnikova |first1=M. V. |year=2008|title=A new species of lesser panda ''Parailurus'' (Mammalia, Carnivora) from the Pliocene of Transbaikalia (Russia) and some aspects of ailurine phylogeny|journal=Paleontological Journal |volume=42 |issue=1|pages=90–99|doi=10.1007/S11492-008-1015-X|bibcode=2008PalJ...42...90S |s2cid=82000411}}</ref>

The earliest fossil record of the modern genus ''Ailurus'' dates no earlier than the [[Pleistocene]] and appears to have been limited to Asia. The modern red panda's lineage became adapted for a specialised bamboo diet, having molar-like premolars and more elevated cusps.<ref name="Wallace"/> The false thumb would secondarily gain a function in feeding.<ref name="Salesa"/><ref name="PNAS-Salesa-2006"/>

===Genomics===
Analysis of 53 red panda samples from Sichuan and Yunnan showed a high level of [[genetic diversity]].<ref>{{cite journal |author1=Su, B. |author2=Fu, Y. |author3=Wang, Y. |author4=Jin, L. |author5=Chakraborty, R. |name-list-style=amp |year=2001 |title=Genetic diversity and population history of the Red Panda (''Ailurus fulgens'') as inferred from mitochondrial DNA sequence variations |journal=Molecular Biology and Evolution |volume=18 |issue=6 |pages=1070–1076 |doi=10.1093/oxfordjournals.molbev.a003878 |pmid=11371595 |doi-access=free}}</ref> The full [[genome]] of the red panda was sequenced in 2017. Researchers have compared it to the genome of the giant panda to learn the genetics of [[convergent evolution]], as both species have false thumbs and are adapted for a specialised bamboo diet despite having the digestive system of a carnivore. Both pandas show modifications to certain limb development genes ([[DYNC2H1]] and [[PCNT]]), which may play roles in the development of the thumbs.<ref>{{cite journal |last1=Hu|first1=Y. |name-list-style=amp |author2=Wu, Q. |author3=Ma, S. |author4=Ma, T. |author5=Shan, L. |author6=Wang, X. |author7=Nie, Y. |author8=Ning, Z. |author9=Yan, L. |author10=Xiu, Y. |author11=Wei, F. |year=2017|title=Comparative genomics reveals convergent evolution between the bamboo-eating giant and red pandas |journal=Proceedings of the National Academy of Sciences |volume=114|issue=5|pages=1081–1086 |doi=10.1073/pnas.1613870114 |pmid=28096377 |pmc=5293045 |bibcode=2017PNAS..114.1081H |doi-access=free}}</ref> In switching from a carnivorous to a herbivorous diet, both species have reactivated taste receptor genes used for detecting [[Bitter (taste)|bitterness]], though the specific genes are different.<ref>{{cite journal|last1=Shan|first1=L. |last2=Wu|first2=Q. |last3=Wange|first3=L. |last4=Zhang|first4=L. |last5=Wei|first5=F. |year=2017|title=Lineage-specific evolution of bitter taste receptor genes in the giant and red pandas implies dietary adaptation |journal=Integrative Zoology |volume=13 |issue=2 |pages=152–159 |doi=10.1111/1749-4877.12291 |pmid=29168616 |pmc=5873442 |name-list-style=amp}}</ref>