New Papilio paper

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adamcotton
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New Papilio paper

Post by adamcotton »

The Papilio paper was finally published on 30 March, and can be downloaded from
https://authors.elsevier.com/c/1gqTf3m3nNAEK5
before 18th May 2023, when it will be placed behind a paywall.

I have been busy preparing a summary for you all, and have pasted the text here:
(A fully formatted version can be sent on request)

Summary of changes to taxonomy of genus Papilio
This is a summary of the taxonomic changes made in Condamine et al. (2023), which was published online on 30 March in the journal Molecular Phylogenetics and Evolution.

Full citation:
Condamine, F.L., Allio, R., Reboud, E.L., Dupuis, J.R., Toussaint, E.F.A., Mazet, N., Hu, S-J., Lewis, D.S., Kunte, K., Cotton, A.M. & Sperling, F.A.H. 2023. A comprehensive phylogeny and revised taxonomy illuminate the origin and diversification of the global radiation of Papilio (Lepidoptera: Papilionidae), Molecular Phylogenetics and Evolution, doi: https://doi.org/10.1016/j.ympev.2023.107758

Genus-group changes
Genus Papilio Linnaeus, 1758 is considered to consist of 14 subgenera, some of which have been treated as separate genera. Treatment of the monophyletic Papilio as a single genus is recommended in order to convey as much information as possible about relationships between species to non-expert end-users of the names through the use of subgeneric names. This allows systematists to refer clearly to their refined taxonomic concepts without putting the burden of increased confusion from name changes onto the users in other fields of these genus and species names.

The subgenera recognised here are:
Heraclides Hübner, [1819] (Type Species: Papilio thoas Linnaeus, 1771)
Chilasa Moore, [1881] (Type Species: Papilio dissimilis Linnaeus, 1758)
Pterourus Scopoli, 1777 (Type Species: Papilio troilus Linnaeus, 1758)
Alexanoria Koçak & Kemal, 2002 (Type Species: Papilio alexanor Esper, [1800])
Euchenor Igarashi, 1979 (Type Species: Papilio euchenor Guérin-Méneville, [1830])
Druryia Aurivillius, 1881 (Type Species: Papilio antimachus Drury, [1782])
Eleppone Hancock, 1979 (Type Species: Papilio anactus Macleay, 1826)
Araminta Moore, 1886 (Type Species: Papilio demolion Cramer, 1776)
Sinoprinceps Hancock, 1983 (Type Species: Papilio xuthus Linnaeus, 1767)
Papilio Linnaeus, 1758 (Type Species: Papilio machaon Linnaeus, 1758)
Nireopapilio Cotton & Nakae, 2020 (Type species: Papilio nireus Linnaeus, 1758)
Achillides Hübner, [1819] (Type Species: Papilio paris Linnaeus, 1758)
Princeps Hübner, [1807] (Type Species: Papilio demodocus Esper, 1799)
Menelaides Hübner, [1819] (Type Species: Papilio polytes Linnaeus, 1758)

The two species sometimes placed in subgenus Agehana Matsumura, 1936 (Type Species: Papilio maraho Shiraki & Sonan, 1934) are confirmed as belonging to subgenus Pterourus as per Wu et al. (2015).
The recently discovered species, Papilio natewa Tennent, Chandra & Müller, 2018 from Fiji is confirmed as sister to Papilio anactus MacLeay, 1826 and placed in subgenus Eleppone.
The Papilio demolion group of species is found not to be closely related to the species of subgenus Menelaides and is placed in its own subgenus, Araminta.
Papilio zalmoxis Hewitson, [1864] is shown to belong to subgenus Nireopapilio rather than subgenus Druryia, where it was previously placed.
Subgenus Princeps is confirmed as comprising only the Papilio demoleus group of species.
Papilio tree.jpg
Papilio tree.jpg (727.43 KiB) Viewed 70649 times
Species-group changes
(Not all species in genus Papilio are listed here, subspecies are only listed for species with changes.)

Subgenus Pterourus Scopoli, 1777
Papilio victorinus Doubleday, 1844
(separate species to Papilio menatius (Hübner, [1819]))
ssp. morelius Rothschild & Jordan, 1906 (Type locality: Guerrero)
ssp. victorinus Doubleday, 1844 (Type locality: the west coast of America)
ssp. vulneratus Butler, 1872 (Type locality: Costa Rica)

Subgenus Euchenor Igarashi, 1979
Papilio euchenor Guérin-Méneville, 1830
Papilio depilis Rothschild, 1895
(separate species to Papilio euchenor Guérin-Méneville, [1830])
ssp. depilis Rothschild, 1895 (Type locality: New Britain)
ssp. neohannoveranus Rothschild, 1898 (Type locality: New Hanover, Bismarck Archipelago)
ssp. novohibernicus Rothschild, 1896 (Type locality: Neu Mecklenburg (= New Ireland)) [not ‘1898’]

Subgenus Druryia Aurivillius, 1881
Papilio humbloti Oberthür, 1888
(separate species to Papilio dardanus Yeats, 1776)
Papilio meriones C. Felder & R. Felder, 1864
(separate species to Papilio dardanus Yeats, 1776)

Subgenus Sinoprinceps Hancock, 1983
Papilio benguetana Joicey & Talbot, 1923 (Type locality: Benguet District, N. Luzon)
(separate species to Papilio xuthus Linnaeus, 1767)

Subgenus Papilio Linnaeus, 1758
Papilio kahli F. & R. Chermock, 1937 (Type locality: Riding Mts., Manitoba)
(separate species to Papilio machaon Linnaeus, 1758)
Papilio bairdii Edwards, 1866
(separate species to Papilio machaon Linnaeus, 1758)
ssp. dodi McDunnough, 1939 (Type locality: Dorothy, Alberta)
ssp. oregonia Edwards, 1876 (Type locality: Columbia River)
ssp. bairdii Edwards, 1866 (Type locality: Fort Whipple, near Prescott, Arizona)

Subgenus Nireopapilio Cotton & Nakae, 2020
Papilio zalmoxis Hewitson, [1864]
(Moved from subgenus Druryia Aurivillius, 1881 and placed between epiphorbas and oribazus groups)

Subgenus Achillides Hübner, [1819]
ulysses group
Papilio ulysses Linnaeus, 1758
(split into 3 species, ssp. retained in P. ulysses are listed for clarity)
ssp. ampelius Rothschild, 1908 (Type locality: Buru)
ssp. ambelauensis (Schäffler & Schmidbauer 2004) (Type locality: Ambelau Is.)
ssp. reikoae Kariya, 1988 (Type locality: Manipa Is.)
ssp. penelope (Dufek, Schäffler & Schmidbauer, 2005) (Type locality: Kelang Isl.)
ssp. ulysses Linnaeus, 1758 (Type locality: Asia [= Ambon])
ssp. scylla (Dufek, Schäffler & Schmidbauer, 2005) (Type locality: Saparua Isl.)
ssp. waterstradti Goode, 2012 (Type locality: Moluques, Ile Gebi [= Gebe Island])
ssp. dirce Jordan, 1909 (Type locality: Misol)
ssp. physkon Fruhstorfer, 1909 (Type locality: Waigiu)
ssp. calypso (Schäffler & Schmidbauer 2004) (Type locality: Salawati)
ssp. denticulatus Joicey & Talbot, 1916 (Type locality: Biak, Schouten Is.)
ssp. autolycus C. Felder & R. Felder, 1865 (Type locality: Nova Guinea: Dorey)
ssp. oxyartes Fruhstorfer, 1909 (Type locality: Aru Island)
ssp. melanotica Jordan, 1909 (Type locality: Dampier) [Hagen, 1897 originally described melanotica as an aberration. The name is available as a subspecies from Jordan, 1909]
ssp. telemachus Montrouzier, 1856 (Type locality: Woodlark)
ssp. joesa Butler, 1869 (Type locality: Queensland)
Papilio orsippus Godman & Salvin, 1888
(separate species to Papilio ulysses Linnaeus, 1758)
ssp. gabrielis Rothschild, 1898 (Type locality: St. Gabriel, Admiralty Islands)
ssp. kallinikos Fruhstorfer, 1903 (Type locality: Neu-Mecklenburg = Neu-Irland)
ssp. ambiguus Rothschild, 1895 (Type locality: New Britain)
ssp. orsippus Godman & Salvin, 1888 (Type locality: Solomon Islands, Aola in Guadalcanar Island)
ssp. rothschildianus Fruhstorfer, 1909 (Type locality: Solomon Islands: Guizo) [Replacement Name for georgius Rothschild, 1908]
Papilio telegonus C. Felder & R. Felder, 1860
(separate species to Papilio ulysses Linnaeus, 1758)
ssp. dohertius Rothschild, 1898 (Type locality: Obi)
ssp. telegonus C. Felder & R. Felder, 1860 (Type locality: Batschian)
ssp. morotaicus Rothschild, 1908 (Type locality: Mira, Morty I. (= Morotai))
palinurus group
Papilio daedalus C. Felder & R. Felder, 1861
(separate species to Papilio palinurus Fabricius, 1787)
ssp. daedalus C. Felder & R. Felder, 1861 (Type locality: Luzon (Manila))
ssp. angustatus Staudinger, 1888 (Type locality: Palawan)
bianor group
Papilio hermeli Nuyda, 1992 (Type locality: Mt Halcon, 1800 m, Northern Mindoro)
(separate species to Papilio chikae Igarashi, 1965)
Papilio polyctor Boisduval, 1836 (Type locality: Cachemire)
(separate species to Papilio bianor Cramer, 1777)
Papilio ryukyuensis Fujioka, 1975
(confirmed separate species to Papilio dehaanii C. Felder & R. Felder, 1864)
ssp. ryukyuensis Fujioka, 1975 (Type locality: Okinawa Islands, Japan)
ssp. amamiensis (Fujioka, 1981) (Type locality: Amami Island, Japan) [‘author, year’ in ()]
Papilio dehaanii C. Felder & R. Felder, 1864
(confirmed separate species to Papilio bianor Cramer, 1777)
ssp. tokaraensis Fujioka, 1975 (Type locality: Nakanoshima, Tokara Islands, Japan)
ssp. hachijonis Matsumura, 1919 (Type locality: Hachijo Islands, Japan)
ssp. dehaanii C. Felder & R. Felder, 1864 (Type locality: Japonia)
maackii group
Papilio syfanius Oberthür, 1886
(confirmed separate species to Papilio maackii Ménétriés, 1858)

Subgenus Menelaides Hübner, [1819]
helenus group
Papilio hystaspes C. Felder & R. Felder, 1862 (Type locality: Luzon)
(confirmed separate species to Papilio helenus Linnaeus, 1758)
Papilio daksha Moore, 1889
(separate species to Papilio helenus Linnaeus, 1758)
ssp. daksha Moore, 1889 (Type locality: Nilgiris)
ssp. mooreanus Rothschild, 1895 (Type locality: Ceylon)
(probably a separate species, but placed here due to lack of material to sequence)
Papilio helenus Linnaeus, 1758
(restricted to mainland Asia, Taiwan & Japan)
ssp. helenus Linnaeus, 1758 (Type locality: Asia [= Canton, China])
ssp. fortunius Fruhstorfer, 1908 (Type locality: Lake Candidius)
ssp. nicconicolens Butler, 1881 (Type locality: Nikko, Central Japan)
Papilio palawanicus Staudinger, 1888
(separate species to Papilio helenus Linnaeus, 1758)
ssp. palawanicus Staudinger, 1888 (Type locality: Palawan)
ssp. boloboca Page & Treadaway, 1996 (Type locality: Boloboc, Sanga Sanga I.)
ssp. enganius Doherty, 1891 (Type locality: Engano)
ssp. sinabangana Goode & Burk, 2013 (Type locality: Sinabang, Simeulue Island, Indonesia)
Papilio mangarinus Rothschild, 1908
(separate species to Papilio helenus Linnaeus, 1758)
ssp. mangarinus Rothschild, 1908 (Type locality: South Flores)
ssp. tambora Rothschild, 1908 (Type locality: Sumbawa)
ssp. jindanus Rothschild, 1908 (Type locality: Sumba)
Papilio biseriatus Rothschild, 1895 (Type locality: Dili)
(separate species to Papilio helenus Linnaeus, 1758)
(Papilio sataspes C. Felder & R. Felder, 1865 is internal to this group and sister to Papilio palawanicus)
polytes group
Papilio protenor Cramer, 1775
(confirmed basal to the polytes group)
Papilio alphenor Cramer, 1776
(separate species to Papilio polytes Cramer, 1777; sister to P. ambrax Boisduval, 1832 & P. phestus Guérin-Méneville, 1830)
ssp. ledebouria Eschscholtz, 1821 (Type locality: Manilla)
ssp. steffi (Page & Treadaway, 2003) (Type locality: Cavan Cavan, Sibutu, 5m)
ssp. perversus Rothschild, 1895 (Type locality: Sangir Island; Talaut; Siao Island)
ssp. polycritos Fruhstorfer, 1902 (Type locality: Sula-Besi)
ssp. nicomachus Fruhstorfer, 1909 (Type locality: Buru)
ssp. alphenor Cramer, 1776 (Type locality: China [loc. err.])
ssp. nicanor C. Felder & R. Felder, 1864 (Type locality: Batjan, Halmaheira)
ssp. muratai Kishida, 1933 (Type locality: Koror Island)
Papilio ambrax Boisduval, 1832
(separate species to Papilio phestus Guérin-Méneville, 1830)
ssp. ambrax Boisduval, 1832 (Type locality: Nouvelle-Guinée)
ssp. artanus Rothschild, 1908 (Type locality: Suer, Mefor I., Geelvink Bay)
ssp. epirus Wallace, 1865 (Type locality: Aru Islands)
ssp. dunali Montrouzier, 1856 (Type locality: Woodlark)
ssp. egipius Miskin, 1876 (Type locality: Rockingham Bay, Queensland)
Papilio phestus Guérin-Méneville, 1830
(separate species to Papilio ambrax Boisduval, 1832)
ssp. reductus Rothschild, 1915 (Type locality: Manus, Admiralty Islands)
ssp. parkinsoni Honrath, 1886 (Type locality: Neu-Pommern)
ssp. phestus Guérin-Méneville, 1830 (Type locality: Nouvelle-Guinée [loc. err.])
ssp. minusculus Ribbe, 1898 (Type locality: Schortlands)
Papilio polytes Linnaeus, 1758
(restricted to mainland Asia, Taiwan & Japan, plus some islands)
ssp. thibetanus Oberthür, 1886 (Type locality: Chapa, Thibet)
ssp. polytes Linnaeus, 1758 (Type locality: Asia [= southern China])
ssp. pasikrates Fruhstorfer, 1908 (Type locality: Formosa, Takan)
ssp. polycles Fruhstorfer, 1903 (Type locality: Okinawa and Ishigaki)
ssp. romulus Cramer, 1775 (Type locality: Côtes de Coromandel & Ceylon)
ssp. rubidimacula Talbot, 1932 (Type locality: Tibet (? south-east), Yunnan: Teng-yueh-Ting)
ssp. latreilloides Yoshino, 2018 (Type locality: Weixi County, 2,500 m, North Yunnan, China)
ssp. stichioides Evans, 1932 (Type locality: Andamans)
ssp. nikobarus C. Felder, 1862 (Type locality: Nankauri, Nicobar Is.)
Papilio theseus Cramer, 1777 [incorrectly treated as Papilio javanus in the paper]
(separate species to Papilio polytes Cramer, 1777)
ssp. theseus Cramer, 1777 (Type locality: la côte Occidentale de Sumatra)
ssp. melanides Haan, 1840 (Type locality: Banjermasin)
ssp. rubeus (Page & Treadaway, 2011) (Type locality: Indonesia, Karimata Island)
ssp. javanus C. Felder, 1862 (Type locality: Java)
ssp. vigellius Fruhstorfer, 1909 (Type locality: Bawean)
ssp. messius Fruhstorfer, 1909 (Type locality: Lombok)
ssp. sotira Jordan, 1909 (Type locality: Sumbawa)
ssp. timorensis C. Felder & R. Felder, 1864 (Type locality: Timor)
ssp. alcindor Oberthür, 1879 (Type locality: Celebes)
ssp. kurokawai Nakae, 2011 (Type locality: Batuatas Island) [not ‘2013’]
ssp. tucanus Jordan, 1909 (Type locality: Binongka and Kalidupa, Toekan Bessi Islands)
memnon group
Papilio diophantus Grose-Smith, 1883 (Type locality: Sumatra)
(confirmed as basal to the memnon group)
Papilio janaka Moore, 1857 (Type locality: Darjeeling)
(separate species to Papilio bootes Westwood, 1842)
Papilio deiphobus Linnaeus, 1758
ssp. rumanzovia Eschscholtz, 1821 (Type locality: Manilla)
(ssp. status of rumanzovia within Papilio deiphobus confirmed)
Papilio agenor Linnaeus, 1758
(separate species to Papilio memnon Linnaeus, 1758)
ssp. parinda (Moore, 1881) (Type locality: Ceylon)
(ssp. of Papilio agenor Linnaeus, 1758)
ssp. polymnestor Cramer, 1775 (Type locality: Côte de Coromandel)
(ssp. of Papilio agenor Linnaeus, 1758)
ssp. thunbergii Siebold, 1824 (Type locality: Dezima, near Nangasaki)
ssp. iriomotensis Fujioka, 2012 (Type locality: Komi~Yubu, Iriomote Is., Okinawa-ken, SW Japan)
ssp. heronus Fruhstorfer, 1902 (Type locality: Formosa)
ssp. agenor Linnaeus, 1758 (Type locality: Asia [= Canton, China])
ssp. nicobarensis Hachitani, 1986 (Type locality: Great Nicobar Is., Nicobar Iss.)
Papilio memnon Linnaeus, 1758
(restricted to SE Asian islands)
ssp. mayo Atkinson, [1874] (Type locality: Andaman Islands)
(ssp. of Papilio memnon Linnaeus, 1758)
ssp. anceus Cramer, 1779 (Type locality: Côte Occidentale de Sumatra)
ssp. caeruleus van Eecke, 1914 (Type locality: Udjung Laukè, Mata-n-Amurèn und Sinabang, Simalur)
ssp. babimemnon Hachitani, 1993 (Type locality: Babi Is.)
ssp. oceani Doherty, 1891 (Type locality: Engano)
ssp. christianae Manguin, 1996 (Type locality: Belitung)
ssp. singkepensis Manguin, 1997 (Type locality: Singkep)
ssp. suisei Hachitani, 1985 (Type locality: Jemaja Is., Anambas)
ssp. bunguranomemnon Hachitani, 1989 (Type locality: Ranai, Bunguran I.)
ssp. tambelanomemnon Hachitani, 1987 (Type locality: Tambelan Is.)
ssp. memnon Linnaeus, 1758 (Type locality: Asia [= Java])
ssp. lowii Druce, 1873 (Type locality: Borneo [loc. err.])
(confirmed ssp. of Papilio memnon Linnaeus, 1758)
ssp. eos Hachitani, 1991 (Type locality: Kangean Id.) [elevation of Infrasubspecific name]
ssp. perlucidus Fruhstorfer, 1903 (Type locality: Sapit, Lombok)
ssp. clathratus Rothschild, 1896 (Type locality: Sambawa)
ssp. kalaomemnon Hachitani, 1987 (Type locality: Kalao Island, Indonesia)
ssp. merapu Doherty, 1891 (Type locality: Koloki, Pada Dalung, Sumba)
ssp. subclathratus Fruhstorfer, 1903 (Type locality: Flores)
ssp. extremus Tsukada & Nishiyama, 1980 (Type locality: Alor)
nephelus group
Papilio castor Westwood, 1842
ssp. mahadeva Moore, [1879] (Type locality: Moolai to Moolat, Upper Tenasserim)
(confirmed ssp. of Papilio castor Westwood, 1842)
Papilio chaon Westwood, 1844
(separate species to Papilio nephelus Boisduval, 1836)
ssp. chaon Westwood, 1844 (Type locality: Assam)
ssp. rileyi Fruhstorfer, 1913 (Type locality: Chungking, Szetchuan, Westchina)
ssp. chaonulus Fruhstorfer, 1902 (Type locality: Hainan, Five Finger Mts.)
ssp. annulus Pendlebury, 1936 (Type locality: Peninsula Siam: Nakon Sri Tamarat. Khao Ram, Khao Luang; Trang. Ban Chong)
Papilio nephelus Boisduval, 1836
(restricted to Malay Peninsula and Sundaic islands)
ssp. sunatus Corbet, 1940 (Type locality: Côte Malaye) [Replacement Name for saturnus Guérin-Méneville, 1840]
ssp. albolineatus Forbes, 1885 (Type locality: Borneo)
ssp. uranus Weymer, 1885 (Type locality: Nias)
ssp. tellonus Fruhstorfer, 1906 (Type locality: Hafenort Poeloe-Tello, Batu-Inseln)
ssp. siporanus Hagen, 1898 (Type locality: nördlichen Mentawej-Insel Pora oder Si-Pora)
ssp. nephelus Boisduval, 1836 (Type locality: Célèbes [loc. err. = Java])
aegeus group
(species group contents changed)
Papilio amynthor Boisduval, 1859
Papilio schmeltzi Herrich-Schäffer, 1869
Papilio tydeus C. Felder & R. Felder, 1860
Papilio godeffroyi Semper, 1866
Papilio oritas Godman & Salvin, 1879
(separate species to Papilio aegeus Donovan, 1805)
ssp. websteri Grose-Smith, 1894 (Type locality: New Britain)
ssp. oritas Godman & Salvin, 1879 (Type locality: New Ireland)
ssp. byronensis Talbot, 1932 (Type locality: New Hannover)
Papilio bridgei Mathew, 1886
Papilio erskinei Mathew, 1886
Papilio weymeri Niepelt, 1914
Papilio gambrisius Cramer, 1777
Papilio aegeus Donovan, 1805
ssp. komos Fruhstorfer, 1904 (Type locality: Dammer)
(ssp. of Papilio aegeus Donovan, 1805)
ssp. inopinatus Butler, 1883 (Type locality: Maroe Island, Timor Laut)
(ssp. of Papilio aegeus Donovan, 1805, not a separate species)
canopus group
(separate to fuscus group)
Papilio canopus Westwood, 1842
(separate species to Papilio fuscus Goeze, 1779)
ssp. buisi Kalis, 1941 (Type locality: Tjandikoesoema, Bali)
ssp. umbrosus Rothschild, 1894 (Type locality: Sambawa)
ssp. sumbanus Rothschild, 1896 (Type locality: Patadala, Sumba)
ssp. modestia Tsukada & Nishiyama, 1980 (Type locality: Reo N.W. Flores Is.)
ssp. alorensis Rothschild, 1894 (Type locality: Alor)
ssp. vollenhovii C. Felder & R. Felder, 1865 (Type locality: Archipelagus Malayicus?)
ssp. hypsiclides Rothschild, 1894 (Type locality: Wetter I.)
ssp. kallon Fruhstorfer, 1904 (Type locality: Roma)
ssp. croton Fruhstorfer, 1904 (Type locality: Dammer Island)
ssp. canopinus Rothschild, 1895 (Type locality: Moa Island)
ssp. babberensis Fruhstorfer, 1903 (Type locality: Babber Island)
ssp. tenimberensis Rothschild, 1896 (Type locality: Timorlaut)
ssp. canopus Westwood, 1842 (Type locality: Melville Island)
Papilio hypsicles Hewitson, 1868
(separate species to Papilio fuscus Goeze, 1779)
ssp. nomus Gabriel, 1936 (Type locality: Loh Is., Torres Isl. N. of New Hebrides)
ssp. hypsicles Hewitson, 1868 (Type locality: New Hebrides)
ssp. burgessi Samson, 1982 (Type locality: Futuna, Vanuatu [New Hebrides])
Papilio cilix Godman & Salvin, 1879
(separate species to Papilio fuscus Goeze, 1779)
ssp. lamponius Fruhstorfer, 1904 (Type locality: New Pomerania)
ssp. cilix Godman & Salvin, 1879 (Type locality: New Ireland)
Papilio capaneus Westwood, 1843
(separate species to Papilio fuscus Goeze, 1779)
ssp. rotalita (Swinhoe, 1893) (Type locality: Ké Island [loc. err. = Aru])
ssp. beccarii Oberthür, 1880 (Type locality: Doreï)
ssp. indicatus Butler, 1876 (Type locality: Port Moresby, New Guinea)
ssp. capaneus Westwood, 1843 (Type locality: Australia)
ssp. hasterti Ribbe, 1907 (Type locality: Bougainville. Strandgegend bei Gieta)
ssp. relmae Tennent, 1999 (Type locality: road from Noro to Munda, New Georgia)
ssp. gyrei Tennent, 1999 (Type locality: north-east of Auki, Malaita)
ssp. xenophilus Mathew, 1886 (Type locality: Ugi, Solomon Islands)
fuscus group
Papilio prexaspes C. Felder & R. Felder, 1865
Papilio hipponous C. Felder & R. Felder, 1862
Papilio fuscus Goeze, 1779
(restricted to islands west of New Guinea)
ssp. motomurai Hanafusa, 1989 (Type locality: Kangean Is., Indonesia)
ssp. porrothenus Jordan, 1909 (Type locality: Kalao and Dyampea (= Djampea))
ssp. pertinax Wallace, 1865 (Type locality: Macassar)
ssp. lunifer Rothschild, 1894 (Type locality: Sangir Is. (Sangi, Sanghi, Sanghir))
ssp. talyabona Talbot, 1932 (Type locality: Sula Islands)
ssp. metagenes Fruhstorfer, 1904 (Type locality: Binongko, Kaledupa Dez.)
ssp. fuscus Goeze, 1779 (Type locality: not stated)
ssp. ombiranus Rothschild, 1898 (Type locality: Obi)
ssp. lapathus Fruhstorfer, 1904 (Type locality: Batjan)
ssp. offakus Fruhstorfer, 1904 (Type locality: Waigiu)
ssp. thomsonii Butler, 1884 (Type locality: Ké Dulan)
Papilio albinus Wallace, 1865
Papilio woodfordi Godman & Salvin, 1888
Papilio heringi Niepelt, 1924

Some interesting repeated speciation patterns were noted in Asian Papilio. Firstly, as in other genera, South Indian taxa are often separate species to those in the rest of mainland Asia. There is repeated speciation between mainland Asian taxa and Sundaic islands (Sumatra, Borneo, Java and other islands) in both subgenus Achillides (the paris group) and Menelaides (e.g. the helenus, polytes, nephelus and memnon groups). Notably, speciation also occurred in taxa from several species groups between New Guinea and the Bismarck Archipelago (New Britain, New Ireland etc).
Chuck
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Re: New Papilio paper

Post by Chuck »

Astonishing.

Uh oh, I do have my "fuscus" name labeled. Fortunately, I never name labeled any of my zelicaon/ machaon/ whatevers.

Looks like the guys who "have every species of Papilio" but didn't care about subspecies better get busy.
eurytides
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Re: New Papilio paper

Post by eurytides »

Regarding Papilio kahli, Julian Dupuis has done a lot of molecular work on the machaon/polyxenes group here in North America. It is of hybrid origin and Dupuis et al places it under Papilio machaon kahli.

https://journals.plos.org/plosone/artic ... =printable

I have not had a chance to look at the new paper in detail. Kahli specimens are not easy to obtain and I do wonder which analysis was more robust.
eurytides
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Re: New Papilio paper

Post by eurytides »

Nevermind. I just downloaded the paper and realize that both Julian Dupuis and his former PhD supervisor Felix Sperling are authors on this paper. I'm not completely surprised to be honest. Dr. Sperling is a giant in the field.
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Re: New Papilio paper

Post by daveuk »

My head hurts.
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Re: New Papilio paper

Post by adamcotton »

eurytides wrote: Sat Apr 01, 2023 3:33 am Dr. Sperling is a giant in the field.
Yes, I have been assisting Felix for very many years. He was a pioneer in Papilio genetics, and many of the samples used in a large number of analyses which were combined in this paper came from me. As well as Felix's own machaon group studies we were both heavily involved in the Heraclides, Achillides and Menelaides studies among others.

Adam.
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Re: New Papilio paper

Post by adamcotton »

I should clarify that I am not directly involved in the DNA analysis or building the trees and other analyses of the sequences. As well as providing samples I verify the identity of specimens and work on morphological taxonomy and nomenclature. Geneticists are not always so familiar with the actual taxa and how to distinguish them from similar species.

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Re: New Papilio paper

Post by Chuck »

Please explain how this works:

Papilio orsippus Godman & Salvin, 1888
...
ssp. orsippus Godman & Salvin, 1888 (Type locality: Solomon Islands, Aola in Guadalcanar Island)
ssp. rothschildianus Fruhstorfer, 1909 (Type locality: Solomon Islands: Guizo) [Replacement Name for georgius Rothschild, 1908]


So orsippus was a subspecific name, but now it's broken into a new species; I'm guessing since (ssp) orsippus was the earliest TL that it then became, by rules, the species name? Could they have named the newly recognized species something other than orsippus?

ssp georgius predates rothchildianus, so how does it lose out?
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Re: New Papilio paper

Post by joachim »

Thanks for posting. Now my head also hurtss.. Will keep it and check al the Papilio I have, maybe something is wrong ...


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Re: New Papilio paper

Post by adamcotton »

Chuck,

Hans Fruhstorfer (Entomologische Zeitschrift, 22(41): 169) wrote:
Pap. ulysses rothschildianus nom. nov.
für P. ull. georgius Rothsch. präokkupiert durch P.
eurypylus georgius Frühst, von der Insel Obi (Stuttgart
Ent. Zeitschr. 9. Novbr. 1907 p. 182.) — Patria: Salomonen,
Guizo.

This means the name Papilio ulysses georgius Rothschild, 1908 was correctly declared to be a junior homonym of Papilio eurypylus georgius Fruhstorfer, 1907, and as such is a permanently unavailable name (and cannot be used). Fruhstorfer replaced the taxon name georgius with rothschildianus as a result of this homonymy.

What has now happened in the Condamine et al. (2023) paper is that the taxa
gabrielis Rothschild, 1898, kallinikos Fruhstorfer, 1903, ambiguus Rothschild, 1895, orsippus Godman & Salvin, 1888 and rothschildianus Fruhstorfer, 1909
were all found to belong to a single separate species to Papilio ulysses Linnaeus, 1758. The oldest available name among the taxa belonging to the newly separated species becomes the valid species name, in this case Papilio orsippus Godman & Salvin, 1888. Any other name cannot be used for the species, and particularly a totally new name cannot be proposed unless there is absolutely no available name for the species. This can occasionally happen, but not in this case, or in any of the species separated in this paper.

I hope this explains the naming issues for Papilio orsippus.

Adam.
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Re: New Papilio paper

Post by joachim »

Thansk Adam!!!
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Re: New Papilio paper

Post by adamcotton »

Gerardo Lamas has kindly pointed out some nomenclatural errors in the paper, most of which I really should have spotted, especially that the name Papilio theseus Cramer, 1777 has priority over Papilio javanus C. Felder, 1862. This is a serious error, caused by my inattention to the dates of publication, when theseus was placed in the same species as javanus after the DNA analysis. No specimens of theseus from Sumatra were actually included in the original DNA analysis of Menelaides by Krushnamegh Kunte and I had to decide whether to include theseus in the island species or in Papilio polytes from mainland Asia without any DNA evidence. I should have noticed that actually theseus is the oldest name, and becomes the valid species name.

I edited the list above and marked the errors in RED with explanations.

Also note the following errors in Table 1 of the paper:

Papilio (Chilasa) paradoxa (Zincken, 1831) [‘author, year’ in ()]
Papilio (Druryia) filaprae Suffert, 1908 [not ‘Süffert’]
Papilio (Druryia) sjostedti Aurivillius, 1908 Ignore this change, see below
Papilio (Heraclides) andraemon (Hübner, [1823]) [‘author, year’ in ()]
Papilio (Papilio) joanae Heitzman, 1973 [not ‘1974’]

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Re: New Papilio paper

Post by Cabintom »

adamcotton wrote: Sun Apr 02, 2023 5:26 pm Papilio (Druryia) sjostedti Aurivillius, 1908 [not ‘sjoestedti’]
Why is this the case? (This is a question about why the code might render names original spelled with "ö" as "o" and not "oe".)
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Re: New Papilio paper

Post by adamcotton »

Under ICZN Code article 32.5.2.1 "ö" becomes "oe" only for names formed from German words. Bror Yngve Sjöstedt was Swedish, so his name becomes 'sjostedti' under the Code.

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Re: New Papilio paper

Post by teinopalpus »

Wow, huge changes. And some are rather surprising. Some expected are not here, f.e. I red many times that Chilasa clytia from Lesser Sundas ( echidna, takizawai ) are maybe something different. Area of Chilasa clytia is rather interesting.
Also probably Graphiums will follow as area of distribution of some species contain exactly that areas where Papilios were "separating" each another. Some species were already processed - G.euryphylus, G.sarpedon, but some - f.e. Graphium codrus probably waiting ....

I am feeling it starting to be "too much" ... New book about Mexican Papilionidae "created" also many new species and subspecies. If any new study about Africa will be published ( Thierry ? ) , I expected same earthquake.

At least we have also one REAL new species, and it is gem - Papilio natewa !!

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Re: New Papilio paper

Post by Chuck »

Thanks for all the ICZN insights Adam. I've read ICZN (quickly) but have not memorized it; I'm still playing catch-up on genomics since my last introduction to the topic 40 years ago. As for the errors- well, no publication goes unscathed, no matter how many eyes review it.

It's amazing what genomic studies are revealing.

Look how far Papilio pilumnus is from Papilio glaucus, who'd have thought?

On that line of thought, that being the genetic distance of two apparently similar appearing species: to my mind, this suggests either (1) convergent evolution, or (2) an ancestral species that looked like, or at least had significant genetic code for, a Tiger Swallowtail (as we know it.) So which is it?

In this case, I'd argue that it was an ancestral influence, but the North American Tigers are Clade 2, whereas the similar looking Papilio alexanor is Clade 3. So what does that tell us?

When we ignore outliers, Papilio clearly demonstrate a few basic designs: (1) light arced bands, (2) shiny blue/ green. I can see that it's not a great stretch (morphologically) between the banding of thoas, the yellow spots of polyxenes, and the stripes of glaucus. But I've not seen any suggestion of what a ancient ancestral Papilio might have looked like.
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Re: New Papilio paper

Post by adamcotton »

teinopalpus wrote: Mon Apr 03, 2023 11:40 am f.e. I red many times that Chilasa clytia from Lesser Sundas ( echidna, takizawai ) are maybe something different.
Nakae (2021 - Gekkan-Mushi, (602): 35-39) already separated Papilio echidna from P. clytia.
He wrote:
5. Papilio (Chilasa) echidna Boisduval, 1836, stat. rev.
Papilio dissimilis Variete Echidna Boisduval, 1836, Spec.
gén. Lépid.
1: 378. Type locality: Timor.
 Boisduval described echidna as a variety of Papilio
dissimilis
, which is a synonym of P. clytia Linnaeus, 1758,
but Rothschild (1895) treated it as a closely related but
separate species. Jordan (1909) united echidna and clytia
within the same species, while noting that the genitalia of
echidna differ from other subspecies of Papilio clytia. The
hindwings of echidna completely lack yellow or orange
marginal spots on both sides, these markings are conspicuous
on the underside and are at least present on the anal angle
of the upperside of all other subspecies of clytia. Instead all
marginal markings of echidna are the same dull white color
as the other markings on the wings. There is a significant
gap in distribution between mainland Asian clytia, which is
absent from Sumatra, Borneo, Java and the western Lesser
Sunda Islands, and echidna which is found from Flores to
Timor and Moa, much further east than the nearest population
of clytia in the Malay Peninsula and Singapore. The range of
echidna is also very far from the Philippine subspecies of P.
clytia
. For these reasons I am elevating Papilio echidna to
species status, and include subspecies takizawai from Flores
in this species.

He also separated Byasa mukoyamai Nakae, 2015 from B. hedistus (Jordan, 1928); Byasa ravana (Moore, [1858]) from B. dasarada (Moore, [1858]); Papilio arjuna Horsfield, 1828 and P. tamilana Moore, 1881 from P. paris Linnaeus, 1758 in the same paper.

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Re: New Papilio paper

Post by adamcotton »

Chuck wrote: Mon Apr 03, 2023 12:10 pm On that line of thought, that being the genetic distance of two apparently similar appearing species: to my mind, this suggests either (1) convergent evolution, or (2) an ancestral species that looked like, or at least had significant genetic code for, a Tiger Swallowtail (as we know it.) So which is it?

In this case, I'd argue that it was an ancestral influence, but the North American Tigers are Clade 2, whereas the similar looking Papilio alexanor is Clade 3. So what does that tell us?
Probably 'tiger stripes' are an old pattern across Papilionidae, as similar patterns occur in many genera.

Adam.
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Re: New Papilio paper

Post by Chuck »

FWIW, the recent changes left me questioning what institutions will do. This taxonomic change impacts some recent donations I made, along with an upcoming donation.

According to Cornell (reflecting on how they do things, it may vary):

1. Pinned specimens w/o ID labels go to "protem" (= purgatory = storage) until which time a student can ID and label it. So it's important to have ID labels on specimens. Note that Cornell has 80 YO papered specimens from at least one significant collection that still sit papered, so a set and pinned specimen is imperative if you want it to go into the reference collection.

2. Use new names on to-be-donated specimens. If there's an existing ID label, leave it there, and the new one goes under it.

3. The Header labels, which are those afixed to the unit pinning trays, will be updated as time allows and sections of the collection are re-collated.


Personally, for my reference collection, I won't change name labels until I get to the point that I need to handle these Papilio for a project.
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Re: New Papilio paper

Post by adamcotton »

Cabintom wrote: Mon Apr 03, 2023 5:59 am
adamcotton wrote: Sun Apr 02, 2023 5:26 pm Papilio (Druryia) sjostedti Aurivillius, 1908 [not ‘sjoestedti’]
Why is this the case? (This is a question about why the code might render names original spelled with "ö" as "o" and not "oe".)
Please see viewtopic.php?t=958 as it turns out that actually the name Papilio sjoestedti is correct under a prevailing usage clause in the ICZN Code.

Adam.
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