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25 October 2016
Vol. 23, Supplement 1
Molecular phylogeny of the NotodontidaeSubfamilies inferred from
28S rRNA sequences
(Lepidoptera, Noctuoidea, Notodontidae)
Hideki Kobayashi and Masaru Nonaka
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Molecular phylogeny of the Notodontidae: Subfamilies inferred
from 28S rRNA sequences (Lepidoptera, Noctuoidea, Notodontidae)
Hideki KOBAYASHI 1) and Masaru NONAKA 2) 3)1) Higashiyama
2-2-1-1003, Meguro-ku, Tokyo, 153-0043 Japan2) The University
Museum, The University of Tokyo, Hongo 7-3-1, Bunkyo-ku, Tokyo,
113-0033 Japan3) Department of Biological Sciences, Graduate School
of Science and Engineering, Tokyo Metropolitan University, 1-1
Minami-Osawa, Hachioji-shi, Tokyo, 192-0397, Japan
Abstract In order to reconstruct the subfamilies of
Notodontidae, we have analyzed the nucleotide sequences of a part
of the 28S ribosomal RNA gene. A total of 324 species in 141 genera
of the Notodontidae, excluding Afrotropic species, were analyzed
together with some out-group taxa. We have recognized ten
subfamilies in it, i.e. Platychasmatinae, Scranciinae, Pygaerinae,
Dudusinae, Thaumetopoeinae, Periergosinae, Heterocampinae,
Spataliinae, Phalerinae and Notodontinae. We accept two other
subfamilies not analyzed here, Dioptinae and Nystaleinae, which are
restricted to the Neotropic and Nearctic. Thus the family
Notodontidae has become to comprise twelve subfamilies, excluding
the Afrotropic region. In the course of the present study, we have
recognized some new taxa and revived genera according to their
molecular monophylies with corroborative morphology.
New taxa by Kobayashi: Periergosinae nom. nov.; Neodrymoniaini
tribe nov. (Notodontinae); Leptophalera gen. nov. (Phalerinae);
Metrioptera gen. nov. (Notodontinae); Griseodonta gen. nov.
(Notodontinae); Biula pinratanai sp. nov. (Spataliinae)
Revived genera:Heterocampinae: Hemifentonia Kiriakoff, 1967.
Spataliinae: Allata Walker, 1862; Pseudallata Kiriakoff, 1968;
Epiptilodontis Kiriakoff, 1964; Inouella Kiriakoff, 1967; Besaia
Walker, 1865, Odnarda Kiriakoff, 1962; Subniganda Kiriakoff, 1962;
Ogulina Kiriakoff, 1962; Curuzza Kiriakoff, 1962; Dypna Kiriakoff,
1962; Cutuza Kiriakoff, 1962; Torigea Matsumura, 1934; Mangea
Kishida, 2004; Torona Walker, 1865. Notodontinae: Eufentonia
Matsumura, 1922; Viridifentonia Nakamura, 1974; Miostauropus
Kiriakoff, 1964; Palaeostauropus Okagaki & Nakamura, 1953;
Chlorostauropus Kiriakoff, 1968; Benbowia Kiriakoff, 1967;
Takadonta Matsumura, 1920; Hexafrenum Matsumura, 1925; Oligaeschra
Kiriakoff, 1964; Hyperaeschrella Strand, 1916; Fusapteryx
Matsumura, 1920; Microphalera Butler, 1885.
Key words Notodontidae, molecular phylogeny, 28SrRNA, subfamily,
tribe, new genera, revived genera, new species
Introduction
The concept of the family and subfamily in zoological
nomenclature is a man-made product and it does not necessarily
reflect the nature system. It has been constructed for the
convenience of our use, that is, description of the classification,
ecological observation, applied entomology for agriculture and
medicine, etc., though the modern phylogenetic study well explains
the course of the evolution; thus the range of a family or
subfamily is desirable to be stabilized. In the family
Notodontidae, however, the subfamilies are far from settled. The
classification of the subfamilies of Notodontidae has been made
based on mainly morphology of the imagoes, larvae and pupae. The
cladistics concept aims firstly to identify the monophyletic
taxonomic group, clade, and next, the phylogenetic relations of the
clades. To use solely morphological evaluation is not enough for
determining the clade and phylogeny, and the molecular study is
essential in the phylogeny today.
Miller (1991) published an article on the cladistics and
classification of the Notodontidae with the conclusion that was
based on the most massive morphological criteria till that time and
for now. He also precisely described the history of the
classification of the Notodontidae and the transition of the
subfamilial framework which had been very complicated. He proposed
nine subfamilies,
Tinea 23 (Supplement 1), October, 2016
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2 Hideki KOBAYASHI and Masaru NONAKA
Platychasmatinae, Thaumetopoeinae, Notodontinae (Notodontini,
Dicranurini), Pygaerinae, Dudusinae (Dudusini, Scranciini),
Heterocampinae, Nystaleinae, Dioptinae, Phalerinae, and incertae
sedis (Lirimiris, Hemiceratini). However, he did not study enough
species which distribute mainly in south Asia. Nakamura (2007)
proposed a new idea of the subfamilies mainly based on the pupal
morphology of the Japanese Notodontidae. He recognized four
subfamilies, Pygaerinae, Notodontinae (Dicranurini, Notodontini,
Stauropini, Dudusini), Heterocampinae (Phalerini, Pydnini,
Heterocampini) and Platychasmatinae. Miller’s Dudusinae and
Phalerinae are downgraded to tribes and positioned in Notodontinae
and Heterocampinae respectively. It is a very clear-cut
classification; however it omitted a lot of Asian and American
genera. Schintlmeister (2013) stated, “18 subfamilies are currently
recognized within the Notodontidae”, and presented a list of the
subfamilies, which contained his newly recognizing subfamilies, but
without the explanations. And yet he stated also “It was beyond the
focus of this work to establish a new classification in the
Notodontidae”. As seen above, the history of the classification of
the subfamilies tells the difficulty of the higher classification
of the Notodontidae. We have analyzed the nuclear 28S rRNA gene and
mitochondrial CO1 gene sequences focusing on the higher
classification and consequently we propose here a new
classification on the subfamilies of the Notodontidae. Since we
could not sampled Afrotropic genera, we anticipate the study of the
higher classification of the Afrotoropic fauna in the future.
Materials and methods
MaterialsThe samples are mainly Asian notodontids, with some
European, North and South American and Australian notodontids. We
have got the DNA sequences of 456 individual samples of 324 species
in 141 genera; 124, 7, 2 and 8 genera from Eurasia, North America,
South America and Australia, respectively. As the out-group, some
species of Sphingidae and Noctuoidea were sampled.Genomic DNA
extractionLegs of dried specimens, newly taken or stocked for
several years, preserved in 99% alcohol were used. We extracted the
genomic DNA using the Prep GEM insect DNA extraction Kit, following
the manufacture’s instruction (ZyGEM NZ Ltd, Hamilton, New
Zealand).PCR amplificationThe primers were designed to amplify the
5’ end region of the 28S rRNA gene which showed a suitable
diversification for higher classification of the family
Notodontidae. The primers were 28SF1b (GAACTTTGAAGAGAGAGTT) and
28SR1 (CAGGCATAGTTCACCATCTT), and 622-670 bp DNA sequences were
used for phylogenetic analysis. Some specimens were also analyzed
for the mitochondrial CO1 gene. The primers were LCO1490
(GGTCAACAAATCATAAAGATATTGG) and HCO2198
(TAAACTTCAGGGTGACCAAAAAATCA).
We used the KOD FX Neo polymerase following the manufacturer’s
instruction (TOYOBO Life Science, Tokyo, Japan). The PCR was
conducted as follows. 94℃ for 2 min; 30 cycles of 98℃ for 10 sec,
50℃ ｆｏｒ 30 sec, 68℃ for 1 min.
DNA sequencing and Phylogenetic analysis
The amplified fragments were treated with Antarctic Phosphatase
(New England Biolabs, Massachusetts, USA) and exonuclease I (New
England Biolabs, Massachusetts, USA). DNA sequence analysis was
performed on a 3100/3130xl Genetic Analyzer (Applied Biosystems
Japan Ltd., Tokyo, Japan) with BigDye Terminater v3.3 Cycle
Sequencing Kit (Applied Biosystems Japan Ltd.). Nucleotide
sequences were aligned by ClustalX ver. 2.1 on Windows PC and MEGA6
(Tamura et al., 2013). The phylogenetic trees were constructed by
the neighbor-joining (NJ) method (Higgins et al., 1992; Saitou and
Nei, 1987; Thompson et al., 1994). The bootstrap method with 100 or
1000 replications was conducted for the evaluation of stability of
the clades. We also used MEGA6 (Tamura et al., 2013), aligning
through Muscle and ClutalW, and making phylogenetic trees by way of
maximum-likelihood (ML) method (Felsenstein, 1981), Tamura
3-parameter model, gamma
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Molecular phylogeny of the Notodontidae 3
distributed with invariant sites, and the neighbor-joining (NJ)
tree using the P-distance method (Nei and Kumar, 2000). Figs 2, 4
and 5.2 are the trees by Muscle and ML. The other trees are
constructed by NJ.
Results and Discussion
We have recognized ten subfamilies in the family Notodontidae.
Those are Platychasmatinae Nakamura, 1956, Scranciinae Miller,
1991, Pygaerinae Duponchel 1845, Dudusinae Matsumura, 1929,
Thaumetopoeinae Aurivillius 1889 (1861), Periergosinae Kobayashi,
nom. nov., Heterocampinae Neumoegen & Dyar, 1894, Spataliinae
Matsumura, 1929, Phalerinae Butler, 1886, and Notodontinae
Stephens, 1829. The former five subfamilies are the basal and
rather primitive lineages and the latter five clades are derived
groups. The clades, which we consider as the subfamilies, are
almost stable in every phylogenetic tree depicted in our analyses,
but the branching order of them is slightly different. We,
consequently, made a lot of phylogenetic trees by the various
analysis-programs with cutting some clades or changing out-group
clades to search for the stability of the order considering the
morphological classification. Through comparison of all the
depicted trees that we have conducted in this study, we have
concluded the phylogenetic tree configured by the ClustalX/NJ has
represented the most acceptable and clearest phylogeny of
Notodontidae from the perspective of morphological corroboration.
Fig. 1.1 and Fig. 1.2 trees were constructed with Mimas christophi
(Staudinger, 1887) of Sphingidae used as the out-group; Fig. 1.3
and Fig. 1.4 trees were constructed with some quadrifid noctuoid
species added. Fig. 1.1 and Fig. 1.3 trees are made by ClustalX/NJ;
Fig. 1.2 and Fig. 1.4 are Muscle-aligned ML trees. The five basal
clades are stably reproduced in the all trees, but the five derived
groups exchange some of the member genera in different programs
trees, that is, the divergence among the five derived groups is
rather shallow, and showing a limited nucleotide sequence
divergence of the part of the 28S ribosomal RNA gene that we
analyzed. We take Fig. 1.1 as the well representing trees to
explain our conclusion, and we use Fig. 1.1 as a working
hypothesis. In Figs 1.5 and 6~10, finer phylogenetic trees are
given for each subfamily. Fig. 1.2, which was based on Muscle/ML,
has two different branching orders compared to Fig. 1.1. The first
is Dudusinae which are branching just before the Spataliinae. The
Dudusinae are a family of the basal lineage morphologically, and
Figs 1.1, 1.3 and 1.4 depict it. And the second, two derived clade,
Periergosinae and Heterocampinae change their position. In Fig.
1.3, the Platychasmatinae branches next to the Scranciinae and the
Dudusinae branch next to the Thaumetopoeinae, but they keep the
position of the basal lineage; two derived clades, Spataliinae and
Heterocampinae change their place. In Fig. 1.4, the
Platychasmatinae, though only one species was analyzed, sink in the
Thaumetopoeinae. However, this is not acceptable from the
morphological viewpoint. We will discuss about this in detail in
the Platychasmatinae section. In Figs 1.2 and 1.4, both conducted
by Mus/ML, Harpyia milhauser stands as an independent clade out of
Dicranurini. We think the genus Harpyia
Tab. 1. Subfamilies in the Notodontidae
PlatychasmatinaeScranciinaePygaerinaeDudusinaeThaumetopoeinaePeriergosinaeHeterocampinaeSpataliinae
Spataliini Ceirini
Notodontidae
Phalerinae Notodontinae Neodrymoniaini Stauropini Dicranurini
Notodontini Dioptinae Nystaleinae
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4 Hideki KOBAYASHI and Masaru NONAKA
has evolved rather fast, and its position out of the Dicranurini
in Fig. 1.2 and 1.4 is presumably due to the long branch attraction
(LBA) (Philippe and Laurent, 1998). And the clade containing
Destolmia, which is positioned in the Heterocampinae in Fig. 1.1
and Fig. 1.3 (-NJ) it is placed in the Stauropini of the
Notodontinae in Figs 1.2 and 1.4 (-ML). We will state about this
problem in the Stauropini section.
Notodontidae Stephens, 1829Notodontidae Stephens, 1829. Nom. Br.
Insects: 39. Type genus: Notodonta Ochsenheimer, 1810.
Nowadays the monophyly of Notodontidae in Noctuoidea is
indisputable. The morphological study, e.g. Miller (1991), and the
molecular, e.g. Zahili et al (2011), ascertain this. In our study,
the monophyly of the Notodontidae in the Noctuoidea, including the
most controversial clades, Platychasmatinae and Thaumetopoeinae, is
depicted in Fig. 1.3 and Fig. 1.4. Kitching & Rawlins (1998)
comprehensively compiled the morphological classification of the
Noctuoidea and stated about monophyly of the Notodontidae. They
accepted nine subfamilies. The diagnostic characters of the
Notodontidae in the Noctuoidea are the trifid cubital vein of
forewing (M2 arising midway between M1 and M3) and the male
genitalia with socii on microscopic observation. The metathoracic
tympanum has the obliquely ventrally faced membrane without nodular
sclerite, which forms the roof of the tympanal cavity. The
metascutum is swollen to form a metascutal bulla (The Dudusinae
lack the metascutal bulla). The first abdominal segment lacks the
counter-tympanal hood. The tibial spurs of legs are serrate. The
larvae have two MD setae on A1 (except the Dioptinae) and seta X on
the anal shield (The Dudusinae lack seta X). As regarding two MD on
metathorax, Miller (1991) reckoned it as a reliable synapomorphy
for Noctuoidea, except the Dioptinae that has one MD. Below we
explain the subfamilies that we have accepted.
Tab. 2. Transition of the Subfamily Classification
Miller, 1991 Nakamura, 2007 Kobayashi et Nonaka, 2016
ScranciinaePlatychasmatinae Platychasmatinae
PlatychasmatinaePygaerinae Pygaerinae PygaerinaeDudusinae Dudusinae
Dudusini Scranciini Thaumetopoeinae Thaumetopoeini Thaumetopoeinae
PeriergosinaeHeterocampinae Heterocampinae Heterocampinae
Heretocampini Heretocampini Spataliinae Spatalia Spataliini Pydnini
Ceirini Stauropini Phalerinae Phalerini PhalerinaeNotodontinae
Notodontinae Notodontinae Dudusini Neodrymoniaini Stauropini
Stauropini Dicranurini Dicranurini Dicranurini Notodontini
Notodontini NotodontiniDioptinae DioptinaeNystaleinae
Nystaleinae
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Molecular phylogeny of the Notodontidae 5
Scranciinae Gargetta, Hijracona, Lasioceros, Phycidopsis,
Pseudogargetta, Thacona (Schintlmeister & Witt, 2015 contained
seven African genera including Thacona to the Scranciinae).
Platychasmatinae Cyphanta, Paracyphanta, Platychasma,
Sphetta.Pygaerinae Clostera, Gluphisia, Gonoclostera,
Micromelalopha, Pygaera, Rhegmatophila.Dudusinae Dudusa,
Talsolepis.Thaumetopoeinae Adrallia, Aglaosoma, Anaphe,
Arctiomorpha, Axiocleta, Coenostegia,
Cynosarga, Epanaphe, Epicoma, Epicomana, Gazalina,
Helianthocampa, Hypsoides, Lomatosticha, Marane, Mesodrepta,
Nesanaphe, Ochrogaster, Oligoclona, Oxymetopa, Paradrallia,
Pseudohypsoides, Sthenadelpha, Tanystola, Teara, Tearina,
Thaumetopoea, Themerastis, Traumatocampa, Trichiocercus,
Zastonia.
Periergosinae Chadisra, Chalepa, Ellida, Franzdaniela,
Gangarides, Hupodonta, Periergos, Rachia, Teleclita.
Heterocampinae Betashachia, Disphragis, Fentonia, Hemifentonia,
Heterocampa, Hyparpax, Lochmaeus, Neopheosia, Oligocentria,
Schizura, Wilemanus.
Spataliinae Spataliini Allata, Ginshachia, Pseudallata, Rosama,
Spatalia. Ceirini Antiphalera, Besaia, Bireta, Biula, Brykia,
Changea, Curuzza, Cutuza, Dypna,
Epiptilodontis, Eushachia, Honveda, Hyperaeschra, Inouella,
Liccana, Mangea, Mimopydna, Norraca, Norracana, Odnarda, Ogulina,
Pterostoma, Pterotes, Ramesa, Saliocleta, Spatalina, Subniganda,
Togaritensha, Torigea, Torona.
Tribe unknown Ambadra, Baradesa, Cashara, Higena, Metaschalis,
Niganda, Ortholomia, Pydnella, Sumeria, Turnaca.
Phalerinae Leptophalera, Phalera, Teinophalera, Uropyia.
Notodontinae
Neodrymoniaini Disparia, Egonociades, Epistauropus, Eufentonia,
Libido, Megaceramis, Mesophalera, Metrioptera, Neodrymonia,
Pseudofentonia, Snellenita, Theinophalera, Viridifentonia.
Stauropini Benbowia, Cascera, Cerasana, Chlorostauropus,
Cnethodonta, Destolmia, Formofentonia, Gallaba, Hapigia, Hylaeora,
Miostauropus, Neola, Netria, Omichlis, Oxoia, Palaeostauropus,
Rodneya, Scythrophanes, Somera, Stauroplitis, Stauropus,
Syntypistis, Zaranga.
Dicranurini Cerura, Dicranura, Furcula, Harpyia, Kamalia,
Liparopsis, Neocerura, Neoharpyia, Pseudohoplitis, Shachia,
Teleclita.
Notodontini Acmeshachia, Allodonta, Allodontoides, Cleapa,
Drymonia, Epinotodonta, Epodonta, Eriodonta, Euhampsonia,
Fusadonta, Fusapteryx, Gangaridopsis, Griseodonta, Hagapteryx,
Hexafrenum, Himeropteryx, Hiradonta, Homocentridia,
Hyperaeschrella, Jurivalentinia, Leucodonta, Leucolopha,
Lophocosma, Lophontosia, Melagonia, Mimesiosomera, Metriaeschra,
Microphalera, Nadata, Nephodonta, Nerice, Norracoide, Notodonta,
Odontosia, Odontosiana, Odontosina, Oligaeschra, Paradrymonia,
Peridea, Periphalera, Phalerodonta, Pheosia, Pheosiopsis, Ptilodon,
Ptilodontosia, Ptilophora, Rachiades, Schintlmeistera, Semidonta,
Shaka, Takadonta, Togaritensha, Togepteryx.
Nystaleinae DioptinaeIncertae sedis Lirimiris, Hemiceras,
Rosema.
Tab. 3. Recognized genera in the Notodontidae
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6 Hideki KOBAYASHI and Masaru NONAKA
①
②
③
④
⑤
⑥
⑦
⑧
⑨
⑩-1
⑩-2
⑩-3
⑩-4
Sphingidae Mimas christophi801Platychasma virgo798
ArimayamaLasioceras sola1138 Sulawesi 0Phycidopsis albovittata1185
OkGargetta eucharius090 Vietnam Micromelalopha adrian875
NanliGluphisia crenata347 Hokkaido Clostera mahatoma961 Yunnan
03Gonoclostera timoniorum1000 AkDudusa celebensis775 Sulawesi
Tarsolepis japonica307 Taiwan Aglaosoma variegata1212 ThauGazalina
crysoloppha657 YunnanThaumetopoia processinea1072 DEpicoma
contristis Thau 1226 VHupodonta corticalis396 AkashiRachia
striata984 Yunnan 0131 Gangarides wallacei790 SulawesGangarides
puerariaePeriergos dispar1047 Nanling 0Chadisra bipartita294 Taiwan
0Ellida viridimixta Miyagi 610Ellida arcuata244 Akashina
072Destolmia lineata1246 Vic 0305Formofentonia rotundata773
SulCascera bella153 Papua 0626 foLochmaeus manteo740 Louisiana
Hyperax aurora728 Lousiana 051Schizura bedia735 Louisiana
09Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria semirufescens726
Neopheosia fasciata300 Taiwan Hemifentonia mandschurica910
NWilemanus bidentatus934 SaitamFentonia excurvata1060
NanlingBetashachia angustipennis915 NDisphragis sabaria188 FGynia
0Heterocapma astalte741 LouisiaSchizula leptinoides714
QuebecHeterocampa umbrata727 LousianPterostomia pterostromia920
NaOrtholomia ocica1160 Sulawesi Baradessa lithosioides283
LaosRosama xmagnum515 Yunnan Allata laticostalis535 Yunnan Spatalia
doeriesi954 Nomugi 08Ginshachia hoebe1013 Shaanxi 0Changea
yanguifei543 Yunnan 10Eushachia acyptera964 Yunnan 0Saliocleta
odnarda1158 SulawesTorigea plumosa866 Nyugasa 071Besaia argentea502
Yunnan 0807Dypna furax889 Nanling 1108 foCutuza srtaminea419 Happo
28S1Pterostoma sinica1015 Shaanxi Spatalina umbrosa958 Yunnan
03Mimopydna pallida619 Miyagi 08Norracana sp252 Laos 0816
foraRamesa tosta260 Laos 0927 foraCuruzza brykyi1099 Nanling
101Besaia melanius514 Yunnan 0621Besaia godrica921 Nanling
0816Liccana argyrosticta1058 NanliMimopydna sikkima086 Vietnam
1Phalera buchephala1073 DeutschUropyia metriculodina319
TaiwaDisparia diluta571 Hiekawa 051Viridifentonia
plagioviridis08Libido bipunctata1165 Taiwan 0Pseudofentonia
argentifera911 Neodrymonia okanoi917 Nanling Eufentonia
nihonica1085 YakushMesophalera libra856 Nanling 0Neodrymonia
comes824 Nanling 0Epistauropus vinaceus787 SulawZaranga permagna245
Akashina 0Neola semiaurata1245 VicAust 0Hyleora capcina1248 Vic
0305 fVaneeckeia pallidifascia1098 NSyntypistis subgeneris333
TaiwBembowia sp285 Laos 0807 foralNeostauropus alternus771
SulawStauropus sikkimensis312 TaiwaMiostauropu mioides893
NanlingPalaeostauropus obliterata935 Harpyia milhauseri1080
DeutschPseudohoplitis vernalis278 LaoShachia circumscripta638
HokkaLiparopsis formosana835 NanlinFurcula bicuspis418 Happo
0320Cerura erminea1082 Deutsche 03Lophocosma sarantoja816
ShisraDrymonia japonica696 Todai 062Fusadonta basilinea376
AkashinTogepteryx dorsoalbida923 NanlPeridea aliena434 Happo 0307
fPeridea graeseri978 Yunnan 030Pheosiopsis luscinicola004
TaiRachiades lichenicolor002 TaiwEpodonta lineata796 Akahsina
0Himeropteryx miraculosa1119 NiNerice hoenei1022 Shaanxi
0307Nephodonta tsushimensis228 TsuAcmeshachia gigantea672
YunnanFusapteryx ladislai1114 Gifu 1Pheosiopsis plutenkoi1023
ShaaLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingHexafrenum maculifer876 NanlinEpinotodonta fumosa705 Gumma
0Takadonta takamukui1095 DaibosNadata gibbosa1116 Washington
Epinotodonta griseotincta443 YAllodonta plebeja1034 Shaanxi
Eriodonta amagisana822 Abo 071Periphalera melanius445
YunnanEuhampsonia splendida603 MiyagPhalerodonta manleyi1118
NikkoPtilodon hoegei Ashoro931 0802Notodonta torva928 Ashoro
0719Pheosia gnoma1079 Deutsch 1016Pheosia tremura1081 Deutsch
10Odontosia sieversii819 Abo 071Ptilophora rufula204 Taiwan
06Odontosina morosa542 Yunnan 10Microphalera grisea324 Taiwan
Hagapteryx kishidai144 Shaanxi
49
89
15
3656
95
15
94
7
2524
36
7
100
100100
31
72
24
31
10
50
72
82
10083
73
100
71
10099
78
69
57
48
90
59
60
3
7
2819
49
15
17
9
18
7
5
55
33
77
39
63
21
9
63
45
97
6
51
7
46
1
41
2
53
8078
100
21
22
31
43
3
3561
55
92
98
63
74
7
8
2
100
9454
71
31
8
24
22
36
40
17
76
47
7
9556
30
2
2
43
24
9
55
92
6943
19
27
60
32
0
0
47
73
91
82
56
31
16
3
0
Sphingidae Mimas christophi801Platychasma virgo798
ArimayamaLasioceras sola1138 Sulawesi 0Phycidopsis albovittata1185
OkGargetta eucharius090 Vietnam Micromelalopha adrian875
NanliGluphisia crenata347 Hokkaido Clostera mahatoma961 Yunnan
03Gonoclostera timoniorum1000 AkDudusa celebensis775 Sulawesi
Tarsolepis japonica307 Taiwan Aglaosoma variegata1212 ThauGazalina
crysoloppha657 YunnanThaumetopoia processinea1072 DEpicoma
contristis Thau 1226 VHupodonta corticalis396 AkashiRachia
striata984 Yunnan 0131 Gangarides wallacei790 SulawesGangarides
puerariaePeriergos dispar1047 Nanling 0Chadisra bipartita294 Taiwan
0Ellida viridimixta Miyagi 610Ellida arcuata244 Akashina
072Destolmia lineata1246 Vic 0305Formofentonia rotundata773
SulCascera bella153 Papua 0626 foLochmaeus manteo740 Louisiana
Hyperax aurora728 Lousiana 051Schizura bedia735 Louisiana
09Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria semirufescens726
Neopheosia fasciata300 Taiwan Hemifentonia mandschurica910
NWilemanus bidentatus934 SaitamFentonia excurvata1060
NanlingBetashachia angustipennis915 NDisphragis sabaria188 FGynia
0Heterocapma astalte741 LouisiaSchizula leptinoides714
QuebecHeterocampa umbrata727 LousianPterostomia pterostromia920
NaOrtholomia ocica1160 Sulawesi Baradessa lithosioides283
LaosRosama xmagnum515 Yunnan Allata laticostalis535 Yunnan Spatalia
doeriesi954 Nomugi 08Ginshachia hoebe1013 Shaanxi 0Changea
yanguifei543 Yunnan 10Eushachia acyptera964 Yunnan 0Saliocleta
odnarda1158 SulawesTorigea plumosa866 Nyugasa 071Besaia argentea502
Yunnan 0807Dypna furax889 Nanling 1108 foCutuza srtaminea419 Happo
28S1Pterostoma sinica1015 Shaanxi Spatalina umbrosa958 Yunnan
03Mimopydna pallida619 Miyagi 08Norracana sp252 Laos 0816
foraRamesa tosta260 Laos 0927 foraCuruzza brykyi1099 Nanling
101Besaia melanius514 Yunnan 0621Besaia godrica921 Nanling
0816Liccana argyrosticta1058 NanliMimopydna sikkima086 Vietnam
1Phalera buchephala1073 DeutschUropyia metriculodina319
TaiwaDisparia diluta571 Hiekawa 051Viridifentonia
plagioviridis08Libido bipunctata1165 Taiwan 0Pseudofentonia
argentifera911 Neodrymonia okanoi917 Nanling Eufentonia
nihonica1085 YakushMesophalera libra856 Nanling 0Neodrymonia
comes824 Nanling 0Epistauropus vinaceus787 SulawZaranga permagna245
Akashina 0Neola semiaurata1245 VicAust 0Hyleora capcina1248 Vic
0305 fVaneeckeia pallidifascia1098 NSyntypistis subgeneris333
TaiwBembowia sp285 Laos 0807 foralNeostauropus alternus771
SulawStauropus sikkimensis312 TaiwaMiostauropu mioides893
NanlingPalaeostauropus obliterata935 Harpyia milhauseri1080
DeutschPseudohoplitis vernalis278 LaoShachia circumscripta638
HokkaLiparopsis formosana835 NanlinFurcula bicuspis418 Happo
0320Cerura erminea1082 Deutsche 03Lophocosma sarantoja816
ShisraDrymonia japonica696 Todai 062Fusadonta basilinea376
AkashinTogepteryx dorsoalbida923 NanlPeridea aliena434 Happo 0307
fPeridea graeseri978 Yunnan 030Pheosiopsis luscinicola004
TaiRachiades lichenicolor002 TaiwEpodonta lineata796 Akahsina
0Himeropteryx miraculosa1119 NiNerice hoenei1022 Shaanxi
0307Nephodonta tsushimensis228 TsuAcmeshachia gigantea672
YunnanFusapteryx ladislai1114 Gifu 1Pheosiopsis plutenkoi1023
ShaaLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingHexafrenum maculifer876 NanlinEpinotodonta fumosa705 Gumma
0Takadonta takamukui1095 DaibosNadata gibbosa1116 Washington
Epinotodonta griseotincta443 YAllodonta plebeja1034 Shaanxi
Eriodonta amagisana822 Abo 071Periphalera melanius445
YunnanEuhampsonia splendida603 MiyagPhalerodonta manleyi1118
NikkoPtilodon hoegei Ashoro931 0802Notodonta torva928 Ashoro
0719Pheosia gnoma1079 Deutsch 1016Pheosia tremura1081 Deutsch
10Odontosia sieversii819 Abo 071Ptilophora rufula204 Taiwan
06Odontosina morosa542 Yunnan 10Microphalera grisea324 Taiwan
Hagapteryx kishidai144 Shaanxi
49
89
15
3656
95
15
94
7
2524
36
7
100
100100
31
72
24
31
10
50
72
82
10083
73
100
71
10099
78
69
57
48
90
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3
7
2819
49
15
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18
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5
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33
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7
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100
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3561
55
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71
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6943
19
27
60
32
0
0
47
73
91
82
56
31
16
3
0
Fig. 1.1. Phylogenetic tree. Notodontidae subfamilies (28SrRNA,
NJ, out-gproup Sphingidae). ①Platychasmatinae; ②Scranciinae;
③Pygaerinae; ④Dudusinae; ⑤Thaumetopoeinae; ⑥Periergosinae;
⑦Heterocampinae; ⑧Spataliinae; ⑨Phalerinae; ⑩Notodontinae.
-
Molecular phylogeny of the Notodontidae 7
①
②
③
④
⑤
⑥
⑦
⑧
⑨LBA
⑩-1
⑩-2
⑩-3
⑩-4
Sphingidae Mimas christophi801Platychasma virgo798
ArimayamaPhycidopsis albovittata1185 OkGargetta eucharius090
VietnamLasioceras sola1138 Sulawesi 0Micromelalopha adrian875
NanliGluphisia crenata347 HokkaidoGonoclostera timoniorum1000
AkClostera mahatoma961 Yunnan 03Aglaosoma variegata1212
ThauGazalina crysoloppha657 YunnanEpicoma contristis Thau 1226
VThaumetopoia processinea1072 DLochmaeus manteo740
LouisianaSchizura bedia735 Louisiana 09Hyperax aurora728 Lousiana
051Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria
semirufescens726Wilemanus bidentatus934 SaitamFentonia
excurvata1060 NanlingNeopheosia fasciata300 TaiwanHemifentonia
mandschurica910 NBetashachia angustipennis915 NDisphragis
sabaria188 FGynia 0Heterocapma astalte741 LouisiaHeterocampa
umbrata727 LousianSchizula leptinoides714 QuebecPeriergos
dispar1047 Nanling 0Gangarides wallacei790 SulawesGangarides
puerariaeHupodonta corticalis396 AkashiRachia striata984 Yunnan
0131Chadisra bipartita294 Taiwan 0Ellida arcuata244 Akashina
072Ellida viridimixta610 Miyagi 0Tarsolepis japonica307
TaiwanDudusa celebensis775 SulawesiGinshachia hoebe1013 Shaanxi
0Spatalia doeriesi954 Nomugi 08Allata laticostalis535 YunnanRosama
xmagnum515 Yunnan Pterostomia pterostromia920 NaBaradessa
lithosioides283 LaosOrtholomia ocica1160 SulawesiPterostoma
sinica1015 ShaanxiChangea yanguifei543 Yunnan 10Eushachia
acyptera964 Yunnan 0Saliocleta odnarda1158 SulawesTorigea
plumosa866 Nyugasa 071Besaia argentea502 Yunnan 0807Dypna furax889
Nanling 1108 foCutuza srtaminea419 Happo 28S1Spatalina umbrosa958
Yunnan 03Mimopydna pallida619 Miyagi 08Besaia godrica921 Nanling
0816Liccana argyrosticta1058 NanliMimopydna sikkima086 Vietnam
1Norracana sp252 Laos 0816 foraRamesa tosta260 Laos 0927 foraBesaia
melanius514 Yunnan 0621Curuzza brykyi1099 Nanling 101Uropyia
metriculodina319 TaiwaPhalera buchephala1073 DeutschHarpyia
milhauseri1080 DeutschPseudohoplitis vernalis278 LaoShachia
circumscripta638 HokkaLiparopsis formosana835 NanlinCerura
erminea1082 Deutsche 03Furcula bicuspis418 Happo 0320Zaranga
permagna245 Akashina 0Neola semiaurata1245 VicAust 0Hyleora
capcina1248 Vic 0305 fDestolmia lineata1246 Vic 0305Formofentonia
rotundata773 SulCascera bella153 Papua 0626 foSyntypistis
subgeneris333 TaiwVaneeckeia pallidifascia1098 NBembowia sp285 Laos
0807 foralNeostauropus alternus771 SulawPalaeostauropus
obliterata935Miostauropu mioides893 NanlingStauropus sikkimensis312
TaiwaDisparia diluta571 Hiekawa 051Libido bipunctata1165 Taiwan
0Pseudofentonia argentifera911Viridifentonia
plagioviridis08Neodrymonia okanoi917 NanlingEufentonia nihonica1085
YakushMesophalera libra856 Nanling 0Neodrymonia comes824 Nanling
0Epistauropus vinaceus787 SulawDrymonia japonica696 Todai
062Lophocosma sarantoja816 ShisraPheosiopsis luscinicola004
TaiEpodonta lineata796 Akahsina 0Rachiades lichenicolor002
TaiwLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingEpinotodonta fumosa705 Gumma 0Hexafrenum maculifer876
NanlinAllodonta plebeja1034 ShaanxiEpinotodonta griseotincta443
YNadata gibbosa1116 WashingtonTakadonta takamukui1095
DaibosPheosiopsis plutenkoi1023 ShaaFusapteryx ladislai1114 Gifu
1Phalerodonta manleyi1118 NikkoEuhampsonia splendida603
MiyagPeriphalera melanius445 YunnanEriodonta amagisana822 Abo
071Togepteryx dorsoalbida923 NanlPeridea graeseri978 Yunnan
030Peridea aliena434 Happo 0307 fAcmeshachia gigantea672
YunnanFusadonta basilinea376 AkashinNephodonta tsushimensis228
TsuHimeropteryx miraculosa1119 NiNerice hoenei1022 Shaanxi
0307Ptilodon hoegei Ashoro931 0802Notodonta torva928 Ashoro
0719Pheosia gnoma1079 Deutsch Pheosia tremura1081 Deutsch
Odontosina morosa542 Yunnan Ptilophora rufula204 Taiwan 06Odontosia
sieversii819 Abo 071Hagapteryx kishidai144 ShaanxiMicrophalera
grisea324 Taiwan
100
100
6057
80
94
98
100
100
60
100
100
93
59
48
57
66
51
100
87
96
79
32
90
64
70
54
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40
34
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75
64
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15
4276
92
52
98
23
95
49
60
59
79
37
39
24
92
61
58
43
2518
48
8
8
31
49
36
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63
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9
35
5
52
22
3
57
10
28
41
6
23
14
11
1
3
0
1
0
1
0
Sphingidae Mimas christophi801Platychasma virgo798
ArimayamaPhycidopsis albovittata1185 OkGargetta eucharius090
VietnamLasioceras sola1138 Sulawesi 0Micromelalopha adrian875
NanliGluphisia crenata347 HokkaidoGonoclostera timoniorum1000
AkClostera mahatoma961 Yunnan 03Aglaosoma variegata1212
ThauGazalina crysoloppha657 YunnanEpicoma contristis Thau 1226
VThaumetopoia processinea1072 DLochmaeus manteo740
LouisianaSchizura bedia735 Louisiana 09Hyperax aurora728 Lousiana
051Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria
semirufescens726Wilemanus bidentatus934 SaitamFentonia
excurvata1060 NanlingNeopheosia fasciata300 TaiwanHemifentonia
mandschurica910 NBetashachia angustipennis915 NDisphragis
sabaria188 FGynia 0Heterocapma astalte741 LouisiaHeterocampa
umbrata727 LousianSchizula leptinoides714 QuebecPeriergos
dispar1047 Nanling 0Gangarides wallacei790 SulawesGangarides
puerariaeHupodonta corticalis396 AkashiRachia striata984 Yunnan
0131Chadisra bipartita294 Taiwan 0Ellida arcuata244 Akashina
072Ellida viridimixta610 Miyagi 0Tarsolepis japonica307
TaiwanDudusa celebensis775 SulawesiGinshachia hoebe1013 Shaanxi
0Spatalia doeriesi954 Nomugi 08Allata laticostalis535 YunnanRosama
xmagnum515 Yunnan Pterostomia pterostromia920 NaBaradessa
lithosioides283 LaosOrtholomia ocica1160 SulawesiPterostoma
sinica1015 ShaanxiChangea yanguifei543 Yunnan 10Eushachia
acyptera964 Yunnan 0Saliocleta odnarda1158 SulawesTorigea
plumosa866 Nyugasa 071Besaia argentea502 Yunnan 0807Dypna furax889
Nanling 1108 foCutuza srtaminea419 Happo 28S1Spatalina umbrosa958
Yunnan 03Mimopydna pallida619 Miyagi 08Besaia godrica921 Nanling
0816Liccana argyrosticta1058 NanliMimopydna sikkima086 Vietnam
1Norracana sp252 Laos 0816 foraRamesa tosta260 Laos 0927 foraBesaia
melanius514 Yunnan 0621Curuzza brykyi1099 Nanling 101Uropyia
metriculodina319 TaiwaPhalera buchephala1073 DeutschHarpyia
milhauseri1080 DeutschPseudohoplitis vernalis278 LaoShachia
circumscripta638 HokkaLiparopsis formosana835 NanlinCerura
erminea1082 Deutsche 03Furcula bicuspis418 Happo 0320Zaranga
permagna245 Akashina 0Neola semiaurata1245 VicAust 0Hyleora
capcina1248 Vic 0305 fDestolmia lineata1246 Vic 0305Formofentonia
rotundata773 SulCascera bella153 Papua 0626 foSyntypistis
subgeneris333 TaiwVaneeckeia pallidifascia1098 NBembowia sp285 Laos
0807 foralNeostauropus alternus771 SulawPalaeostauropus
obliterata935Miostauropu mioides893 NanlingStauropus sikkimensis312
TaiwaDisparia diluta571 Hiekawa 051Libido bipunctata1165 Taiwan
0Pseudofentonia argentifera911Viridifentonia
plagioviridis08Neodrymonia okanoi917 NanlingEufentonia nihonica1085
YakushMesophalera libra856 Nanling 0Neodrymonia comes824 Nanling
0Epistauropus vinaceus787 SulawDrymonia japonica696 Todai
062Lophocosma sarantoja816 ShisraPheosiopsis luscinicola004
TaiEpodonta lineata796 Akahsina 0Rachiades lichenicolor002
TaiwLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingEpinotodonta fumosa705 Gumma 0Hexafrenum maculifer876
NanlinAllodonta plebeja1034 ShaanxiEpinotodonta griseotincta443
YNadata gibbosa1116 WashingtonTakadonta takamukui1095
DaibosPheosiopsis plutenkoi1023 ShaaFusapteryx ladislai1114 Gifu
1Phalerodonta manleyi1118 NikkoEuhampsonia splendida603
MiyagPeriphalera melanius445 YunnanEriodonta amagisana822 Abo
071Togepteryx dorsoalbida923 NanlPeridea graeseri978 Yunnan
030Peridea aliena434 Happo 0307 fAcmeshachia gigantea672
YunnanFusadonta basilinea376 AkashinNephodonta tsushimensis228
TsuHimeropteryx miraculosa1119 NiNerice hoenei1022 Shaanxi
0307Ptilodon hoegei Ashoro931 0802Notodonta torva928 Ashoro
0719Pheosia gnoma1079 Deutsch Pheosia tremura1081 Deutsch
Odontosina morosa542 Yunnan Ptilophora rufula204 Taiwan 06Odontosia
sieversii819 Abo 071Hagapteryx kishidai144 ShaanxiMicrophalera
grisea324 Taiwan
100
100
6057
80
94
98
100
100
60
100
100
93
59
48
57
66
51
100
87
96
79
32
90
64
70
54
98
40
34
65
75
64
62
15
4276
92
52
98
23
95
49
60
59
79
37
39
24
92
61
58
43
2518
48
8
8
31
49
36
25
1
14
6
21
6
9
46
12
16
25
17
38
22
51
3
53
46
73
66
86
63
8
79
58
7
23
26
21
18
56
24
64
60
6
58
87
63
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38
12
48
85
9
35
5
52
22
3
57
10
28
41
6
23
14
11
1
3
0
1
0
1
0
Fig. 1.2. Phylogenetic tree. Notodontidae subfamilies (28SrRNA,
ML, out-gproup Sphingidae). ①Platychasmatinae; ②Scranciinae;
③Pygaerinae; ④Dudusinae; ⑤Thaumetopoeinae; ⑥Periergosinae;
⑦Heterocampinae; ⑧Spataliinae; ⑨Phalerinae; ⑩Notodontinae.
-
8 Hideki KOBAYASHI and Masaru NONAKA
①
②
③
④
⑤
⑥
⑦
⑧
⑨
⑩-1
⑩-2
⑩-3
⑩-4
Sphingidae Mimas christophi801Noctuidae Amphipyra
schrenckiiNoctuidae Thysanoplusia intermEutelia gegeri Akashina
1003EuLymantriidae Lymantria mathuraLymantriidae Lymantria
fumida9Arctiidae Rhyparioides nebulosArctuiidae Pericaria
matronulaErebidae Sypnoides picta939 NoErebidae Catocala patala949
NoAganaidae Eupleuca membranariaAganaidae Asota sp1177
SulawesLasioceras sola1138 Sulawesi 0Phycidopsis albovittata1185
OkGargetta hampsoni sulawaesianaGargetta nagaensis090
VietnamPlatychasma virgo798 ArimayamaMicromelalopha adrian875
NanliClostera mahatoma961 Yunnan 03Gluphisia crenata347
HokkaidoGonoclostera timoniorum1000 AkAglaosoma variegata1212
ThauGazalina crysoloppha657 YunnanThaumetopoia processinea1072
DEpicoma contristis Thau 1226 VDudusa celebensis775
SulawesiTarsolepis japonica307 TaiwanTarsolepis fulgurifera506
YunnHupodonta corticalis396 AkashiRachia striata984 Yunnan
0131Gangarides wallacei790 SulawesChadisra bipartita294 Taiwan
0Ellida arcuata244 Akashina 072Periergos dispar1047 Nanling
0Ortholomia ocica1160 SulawesiBaradessa lithosioides283 LaosRosama
xmagnum515 Yunnan Allata laticostalis535 YunnanSpatalia doeriesi954
Nomugi 08Ginshachia hoebe1013 Shaanxi 0Pterostomia pterostromia920
NaPterostoma sinica1015 ShaanxiSaliocleta odnarda1158
SulawesSaliocleta acyptera964 YunnanBesaia sp286 Laos 0807
foraligTorigea plumosa866 Nyugasa 071Besaia argentea502 Yunnan
0807Dypna furax889 Nanling 1108 foCutuza srtaminea419 Happo
28S1Changea yanguifei543 Yunnan 10Spatalina umbrosa958 Yunnan
03Ramesa tosta260 Laos 0927 foraNorracana sp252 Laos 0816
foraMimopydna pallida619 Miyagi 08Curuzza brykyi1099 Nanling
101Besaia melanius514 Yunnan 0621Besaia godrica921 Nanling
0816Liccana argyrosticta1058 NanliMimopydna sikkima086 Vietnam
1Destolmia spAK293 1246 Vic 030Formofentonia rotundata773
SulCascera bella153 Papua 0626 foHeterocapma manteo740
LouisianHyperax aurora728 Lousiana 051Schizura bedia735 Louisiana
09Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria
semirufescens726Wilemanus bidentatus934 SaitamHemifentonia
mandschurica910 NNeopheosia fasciata300 TaiwanFentonia
excurvata1060 NanlingBetashachia angustipennis915 NDisphragis
sabaria188 FGynia 0Heterocapma astalte741 LouisiaSchizula
leptinoides714 QuebecHeterocampa umbrata727 LousianPhalera
buchephala1073 DeutschUropyia metriculodina319 TaiwaNeola
semiaurata1245 VicAust 0Hyleora capcina1248 Vic 0305 fVaneeckeia
pallidifascia1098 NSyntypistis subgeneris333 TaiwBembowia sp285
Laos 0807 foralNeostauropus alternus771 SulawStauropus
sikkimensis312 TaiwaMiostauropu mioides893 NanlingPalaeostauropus
obliterata935Disparia diluta571 Hiekawa 051Viridifentonia
plagioviridis08Libido bipunctata1165 Taiwan 0Pseudofentonia
argentifera911Neodrymonia okanoi917 NanlingEufentonia nihonica1085
YakushMesophalera libra856 Nanling 0Neodrymonia comes824 Nanling
0Epistauropus vinaceus787 SulawHarpyia milhauseri1080
DeutschPseudohoplitis vernalis278 LaoShachia circumscripta638
HokkaLiparopsis formosana835 NanlinFurcula bicuspis418 Happo
0320Cerura erminea1082 Deutsche 03Lophocosma sarantoja816
ShisraDrymonia japonica696 Todai 062Eriodonta amagisana822 Abo
071Periphalera melanius445 YunnanEuhampsonia splendida603
MiyagPhalerodonta manleyi1118 NikkoFusadonta basilinea376
AkashinTogepteryx dorsoalbida923 NanlPeridea aliena434 Happo 0307
fPeridea graeseri978 Yunnan 030Acmeshachia gigantea672
YunnanNephodonta tsushimensis228 TsuNerice hoenei1022 Shaanxi
0307Himeropteryx miraculosa1119 NiPheosiopsis luscinicola004
TaiRachiades lichenicolor002 TaiwEpodonta lineata796 Akahsina
0Fusapteryx ladislai1114 Gifu 1Pheosiopsis plutenkoi1023
ShaaLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingHexafrenum maculifer876 NanlinEpinotodonta fumosa705 Gumma
0Takadonta takamukui1095 DaibosNadata gibbosa1116
WashingtonEpinotodonta griseotincta443 YAllodonta plebeja1034
ShaanxiPtilodon hoegei Ashoro931 0802Notodonta albicosta414 Happo
0Pheosia gnoma1079 Deutsch 1016Pheosia tremura1081 Deutsch
10Odontosina morosa542 Yunnan 10Odontosia sieversii819 Abo
071Ptilophora rufula204 Taiwan 06Microphalera grisea324
TaiwanHagapteryx kishidai144 Shaanxi
99
83
91
76
26
99
89
100
70
70
100
55
70
46
29 64
65
98
2156
29
52
13
26
57
14
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100
24
21
15
10
33
18
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15
17
11
8
6
4
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27
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6
18
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68
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100
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0
37
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60
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4
034
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15
26
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0 100
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36
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18
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9
22
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5
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42
76
96
83
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47
26
11
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0
6
46
88
73
19
8
20
50
38
Sphingidae Mimas christophi801Noctuidae Amphipyra
schrenckiiNoctuidae Thysanoplusia intermEutelia gegeri Akashina
1003EuLymantriidae Lymantria mathuraLymantriidae Lymantria
fumida9Arctiidae Rhyparioides nebulosArctuiidae Pericaria
matronulaErebidae Sypnoides picta939 NoErebidae Catocala patala949
NoAganaidae Eupleuca membranariaAganaidae Asota sp1177
SulawesLasioceras sola1138 Sulawesi 0Phycidopsis albovittata1185
OkGargetta hampsoni sulawaesianaGargetta nagaensis090
VietnamPlatychasma virgo798 ArimayamaMicromelalopha adrian875
NanliClostera mahatoma961 Yunnan 03Gluphisia crenata347
HokkaidoGonoclostera timoniorum1000 AkAglaosoma variegata1212
ThauGazalina crysoloppha657 YunnanThaumetopoia processinea1072
DEpicoma contristis Thau 1226 VDudusa celebensis775
SulawesiTarsolepis japonica307 TaiwanTarsolepis fulgurifera506
YunnHupodonta corticalis396 AkashiRachia striata984 Yunnan
0131Gangarides wallacei790 SulawesChadisra bipartita294 Taiwan
0Ellida arcuata244 Akashina 072Periergos dispar1047 Nanling
0Ortholomia ocica1160 SulawesiBaradessa lithosioides283 LaosRosama
xmagnum515 Yunnan Allata laticostalis535 YunnanSpatalia doeriesi954
Nomugi 08Ginshachia hoebe1013 Shaanxi 0Pterostomia pterostromia920
NaPterostoma sinica1015 ShaanxiSaliocleta odnarda1158
SulawesSaliocleta acyptera964 YunnanBesaia sp286 Laos 0807
foraligTorigea plumosa866 Nyugasa 071Besaia argentea502 Yunnan
0807Dypna furax889 Nanling 1108 foCutuza srtaminea419 Happo
28S1Changea yanguifei543 Yunnan 10Spatalina umbrosa958 Yunnan
03Ramesa tosta260 Laos 0927 foraNorracana sp252 Laos 0816
foraMimopydna pallida619 Miyagi 08Curuzza brykyi1099 Nanling
101Besaia melanius514 Yunnan 0621Besaia godrica921 Nanling
0816Liccana argyrosticta1058 NanliMimopydna sikkima086 Vietnam
1Destolmia spAK293 1246 Vic 030Formofentonia rotundata773
SulCascera bella153 Papua 0626 foHeterocapma manteo740
LouisianHyperax aurora728 Lousiana 051Schizura bedia735 Louisiana
09Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria
semirufescens726Wilemanus bidentatus934 SaitamHemifentonia
mandschurica910 NNeopheosia fasciata300 TaiwanFentonia
excurvata1060 NanlingBetashachia angustipennis915 NDisphragis
sabaria188 FGynia 0Heterocapma astalte741 LouisiaSchizula
leptinoides714 QuebecHeterocampa umbrata727 LousianPhalera
buchephala1073 DeutschUropyia metriculodina319 TaiwaNeola
semiaurata1245 VicAust 0Hyleora capcina1248 Vic 0305 fVaneeckeia
pallidifascia1098 NSyntypistis subgeneris333 TaiwBembowia sp285
Laos 0807 foralNeostauropus alternus771 SulawStauropus
sikkimensis312 TaiwaMiostauropu mioides893 NanlingPalaeostauropus
obliterata935Disparia diluta571 Hiekawa 051Viridifentonia
plagioviridis08Libido bipunctata1165 Taiwan 0Pseudofentonia
argentifera911Neodrymonia okanoi917 NanlingEufentonia nihonica1085
YakushMesophalera libra856 Nanling 0Neodrymonia comes824 Nanling
0Epistauropus vinaceus787 SulawHarpyia milhauseri1080
DeutschPseudohoplitis vernalis278 LaoShachia circumscripta638
HokkaLiparopsis formosana835 NanlinFurcula bicuspis418 Happo
0320Cerura erminea1082 Deutsche 03Lophocosma sarantoja816
ShisraDrymonia japonica696 Todai 062Eriodonta amagisana822 Abo
071Periphalera melanius445 YunnanEuhampsonia splendida603
MiyagPhalerodonta manleyi1118 NikkoFusadonta basilinea376
AkashinTogepteryx dorsoalbida923 NanlPeridea aliena434 Happo 0307
fPeridea graeseri978 Yunnan 030Acmeshachia gigantea672
YunnanNephodonta tsushimensis228 TsuNerice hoenei1022 Shaanxi
0307Himeropteryx miraculosa1119 NiPheosiopsis luscinicola004
TaiRachiades lichenicolor002 TaiwEpodonta lineata796 Akahsina
0Fusapteryx ladislai1114 Gifu 1Pheosiopsis plutenkoi1023
ShaaLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingHexafrenum maculifer876 NanlinEpinotodonta fumosa705 Gumma
0Takadonta takamukui1095 DaibosNadata gibbosa1116
WashingtonEpinotodonta griseotincta443 YAllodonta plebeja1034
ShaanxiPtilodon hoegei Ashoro931 0802Notodonta albicosta414 Happo
0Pheosia gnoma1079 Deutsch 1016Pheosia tremura1081 Deutsch
10Odontosina morosa542 Yunnan 10Odontosia sieversii819 Abo
071Ptilophora rufula204 Taiwan 06Microphalera grisea324
TaiwanHagapteryx kishidai144 Shaanxi
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Fig. 1.3. Phylogenetic tree. Notodontidae subfamilies (28SrRNA,
NJ, out-gproup Noctuoidea). ①Platychasmatinae; ②Scranciinae;
③Pygaerinae; ④Dudusinae; ⑤Thaumetopoeinae; ⑥Periergosinae;
⑦Heterocampinae; ⑧Spataliinae; ⑨Phalerinae; ⑩Notodontinae.
-
Molecular phylogeny of the Notodontidae 9
①
②
③
④
⑤
⑥
⑦
⑧
⑨
⑩-1
⑩-2
⑩-3
⑩-4
Sphingidae Mimas christophi801Noctuidae Amphipyra
schrenckiiNoctuidae Thysanoplusia intermEutelia gegeri Akashina
1003EuLymantriidae Lymantria fumida9Lymantriidae Lymantria
mathuraAganaidae Asota sp1177 SulawesAganaidae Eupleuca
membranariaErebidae Catocala patala949 NoErebidae Sypnoides
picta939 NoArctuiidae Pericaria matronulaArctiidae Rhyparioides
nebulosPhycidopsis albovittata1185 OkLasioceras sola1138 Sulawesi
0Gargetta nagaensis090 VietnamGargetta hampsoni
sulawaesianaMicromelalopha adrian875 NanliGluphisia crenata347
HokkaidoGonoclostera timoniorum1000 AkClostera mahatoma961 Yunnan
03Dudusa celebensis775 SulawesiTarsolepis japonica307
TaiwanTarsolepis fulgurifera506 YunnPeriergos dispar1047 Nanling
0Gangarides wallacei790 SulawesChadisra bipartita294 Taiwan 0Ellida
arcuata244 Akashina 072Rachia striata984 Yunnan 0131Hupodonta
corticalis396 AkashiAglaosoma variegata1212 ThauGazalina
crysoloppha657 YunnanPlatychasma virgo798 ArimayamaEpicoma
contristis Thau 1226 VThaumetopoia processinea1072 DHeterocapma
manteo740 LouisianSchizura bedia735 Louisiana 09Hyperax aurora728
Lousiana 051Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria
semirufescens726Wilemanus bidentatus934 SaitamNeopheosia
fasciata300 TaiwanHemifentonia mandschurica910 NFentonia
excurvata1060 NanlingBetashachia angustipennis915 NDisphragis
sabaria188 FGynia 0Heterocapma astalte741 LouisiaHeterocampa
umbrata727 LousianSchizula leptinoides714 QuebecGinshachia
hoebe1013 Shaanxi 0Spatalia doeriesi954 Nomugi 08Allata
laticostalis535 YunnanRosama xmagnum515 Yunnan Ortholomia ocica1160
SulawesiPterostomia pterostromia920 NaPterostoma sinica1015
ShaanxiBaradessa lithosioides283 LaosNorracana sp252 Laos 0816
foraMimopydna pallida619 Miyagi 08Besaia godrica921 Nanling
0816Mimopydna sikkima086 Vietnam Changea yanguifei543 Yunnan
10Besaia melanius514 Yunnan 0621Curuzza brykyi1099 Nanling
101Liccana argyrosticta1058 NanliSpatalina umbrosa958 Yunnan
03Ramesa tosta260 Laos 0927 foraSaliocleta odnarda1158
SulawesSaliocleta acyptera964 YunnanBesaia sp286 Laos 0807
foraligTorigea plumosa866 Nyugasa 071Besaia argentea502 Yunnan
0807Dypna furax889 Nanling 1108 foCutuza srtaminea419 Happo
28S1Uropyia metriculodina319 TaiwaPhalera buchephala1073
DeutschHarpyia milhauseri1080 DeutschPseudohoplitis vernalis278
LaoShachia circumscripta638 HokkaLiparopsis formosana835
NanlinCerura erminea1082 Deutsche 03Furcula bicuspis418 Happo
0320Disparia diluta571 Hiekawa 051Libido bipunctata1165 Taiwan
0Viridifentonia plagioviridis08Pseudofentonia
argentifera911Neodrymonia okanoi917 NanlingEufentonia nihonica1085
YakushEpistauropus vinaceus787 SulawMesophalera libra856 Nanling
0Neodrymonia comes824 Nanling Cascera bella153 Papua 0626
foDestolmia spAK293 1246 Vic 030Formofentonia rotundata773
SulHyleora capcina1248 Vic 0305 fNeola semiaurata1245 VicAust
0Syntypistis subgeneris333 TaiwVaneeckeia pallidifascia1098
NBembowia sp285 Laos 0807 foralNeostauropus alternus771
SulawPalaeostauropus obliterata935Miostauropu mioides893
NanlingStauropus sikkimensis312 TaiwaDrymonia japonica696 Todai
062Lophocosma sarantoja816 ShisraPheosiopsis luscinicola004
TaiEpodonta lineata796 Akahsina 0Rachiades lichenicolor002
TaiwLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingEpinotodonta fumosa705 Gumma Hexafrenum maculifer876
NanlinAllodonta plebeja1034 ShaanxiEpinotodonta griseotincta443
YNadata gibbosa1116 WashingtonTakadonta takamukui1095
DaibosFusapteryx ladislai1114 Gifu 1Pheosiopsis plutenkoi1023
ShaaPtilodon hoegei Ashoro931 0802Notodonta albicosta414 Happo
0Pheosia gnoma1079 Deutsch Pheosia tremura1081 Deutsch 10Odontosina
morosa542 Yunnan 10Ptilophora rufula204 Taiwan 06Odontosia
sieversii819 Abo 071Hagapteryx kishidai144 ShaanxiMicrophalera
grisea324 TaiwanPhalerodonta manleyi1118 NikkoEuhampsonia
splendida603 Periphalera melanius445 YunnanEriodonta amagisana822
Abo 071Peridea aliena434 Happo 0307 fPeridea graeseri978 Yunnan
030Togepteryx dorsoalbida923 NanlAcmeshachia gigantea672
YunnanFusadonta basilinea376 AkashinNephodonta tsushimensis228
TsuHimeropteryx miraculosa1119 NiNerice hoenei1022 Shaanxi 0307
100
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Sphingidae Mimas christophi801Noctuidae Amphipyra
schrenckiiNoctuidae Thysanoplusia intermEutelia gegeri Akashina
1003EuLymantriidae Lymantria fumida9Lymantriidae Lymantria
mathuraAganaidae Asota sp1177 SulawesAganaidae Eupleuca
membranariaErebidae Catocala patala949 NoErebidae Sypnoides
picta939 NoArctuiidae Pericaria matronulaArctiidae Rhyparioides
nebulosPhycidopsis albovittata1185 OkLasioceras sola1138 Sulawesi
0Gargetta nagaensis090 VietnamGargetta hampsoni
sulawaesianaMicromelalopha adrian875 NanliGluphisia crenata347
HokkaidoGonoclostera timoniorum1000 AkClostera mahatoma961 Yunnan
03Dudusa celebensis775 SulawesiTarsolepis japonica307
TaiwanTarsolepis fulgurifera506 YunnPeriergos dispar1047 Nanling
0Gangarides wallacei790 SulawesChadisra bipartita294 Taiwan 0Ellida
arcuata244 Akashina 072Rachia striata984 Yunnan 0131Hupodonta
corticalis396 AkashiAglaosoma variegata1212 ThauGazalina
crysoloppha657 YunnanPlatychasma virgo798 ArimayamaEpicoma
contristis Thau 1226 VThaumetopoia processinea1072 DHeterocapma
manteo740 LouisianSchizura bedia735 Louisiana 09Hyperax aurora728
Lousiana 051Oligocentria lignicolor736 LouSchizura unicornis734
LousianaTanassa semirufescens731 LoisiOligocentria
semirufescens726Wilemanus bidentatus934 SaitamNeopheosia
fasciata300 TaiwanHemifentonia mandschurica910 NFentonia
excurvata1060 NanlingBetashachia angustipennis915 NDisphragis
sabaria188 FGynia 0Heterocapma astalte741 LouisiaHeterocampa
umbrata727 LousianSchizula leptinoides714 QuebecGinshachia
hoebe1013 Shaanxi 0Spatalia doeriesi954 Nomugi 08Allata
laticostalis535 YunnanRosama xmagnum515 Yunnan Ortholomia ocica1160
SulawesiPterostomia pterostromia920 NaPterostoma sinica1015
ShaanxiBaradessa lithosioides283 LaosNorracana sp252 Laos 0816
foraMimopydna pallida619 Miyagi 08Besaia godrica921 Nanling
0816Mimopydna sikkima086 Vietnam Changea yanguifei543 Yunnan
10Besaia melanius514 Yunnan 0621Curuzza brykyi1099 Nanling
101Liccana argyrosticta1058 NanliSpatalina umbrosa958 Yunnan
03Ramesa tosta260 Laos 0927 foraSaliocleta odnarda1158
SulawesSaliocleta acyptera964 YunnanBesaia sp286 Laos 0807
foraligTorigea plumosa866 Nyugasa 071Besaia argentea502 Yunnan
0807Dypna furax889 Nanling 1108 foCutuza srtaminea419 Happo
28S1Uropyia metriculodina319 TaiwaPhalera buchephala1073
DeutschHarpyia milhauseri1080 DeutschPseudohoplitis vernalis278
LaoShachia circumscripta638 HokkaLiparopsis formosana835
NanlinCerura erminea1082 Deutsche 03Furcula bicuspis418 Happo
0320Disparia diluta571 Hiekawa 051Libido bipunctata1165 Taiwan
0Viridifentonia plagioviridis08Pseudofentonia
argentifera911Neodrymonia okanoi917 NanlingEufentonia nihonica1085
YakushEpistauropus vinaceus787 SulawMesophalera libra856 Nanling
0Neodrymonia comes824 Nanling Cascera bella153 Papua 0626
foDestolmia spAK293 1246 Vic 030Formofentonia rotundata773
SulHyleora capcina1248 Vic 0305 fNeola semiaurata1245 VicAust
0Syntypistis subgeneris333 TaiwVaneeckeia pallidifascia1098
NBembowia sp285 Laos 0807 foralNeostauropus alternus771
SulawPalaeostauropus obliterata935Miostauropu mioides893
NanlingStauropus sikkimensis312 TaiwaDrymonia japonica696 Todai
062Lophocosma sarantoja816 ShisraPheosiopsis luscinicola004
TaiEpodonta lineata796 Akahsina 0Rachiades lichenicolor002
TaiwLeucodonta bicolaria821 Abo 02Homocentridia picta870
NanlingEpinotodonta fumosa705 Gumma Hexafrenum maculifer876
NanlinAllodonta plebeja1034 ShaanxiEpinotodonta griseotincta443
YNadata gibbosa1116 WashingtonTakadonta takamukui1095
DaibosFusapteryx ladislai1114 Gifu 1Pheosiopsis plutenkoi1023
ShaaPtilodon hoegei Ashoro931 0802Notodonta albicosta414 Happo
0Pheosia gnoma1079 Deutsch Pheosia tremura1081 Deutsch 10Odontosina
morosa542 Yunnan 10Ptilophora rufula204 Taiwan 06Odontosia
sieversii819 Abo 071Hagapteryx kishidai144 ShaanxiMicrophalera
grisea324 TaiwanPhalerodonta manleyi1118 NikkoEuhampsonia
splendida603 Periphalera melanius445 YunnanEriodonta amagisana822
Abo 071Peridea aliena434 Happo 0307 fPeridea graeseri978 Yunnan
030Togepteryx dorsoalbida923 NanlAcmeshachia gigantea672
YunnanFusadonta basilinea376 AkashinNephodonta tsushimensis228
TsuHimeropteryx miraculosa1119 NiNerice hoenei1022 Shaanxi 0307
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Fig. 1.4. Phylogenetic tree. Notodontidae subfamilies (28SrRNA,
ML, out-gproup Noctuoidea). ①Platychasmatinae; ②Scranciinae;
③Pygaerinae; ④Dudusinae; ⑤Thaumetopoeinae; ⑥Periergosinae;
⑦Heterocampinae; ⑧Spataliinae; ⑨Phalerinae; ⑩Notodontinae.
LBA
LBA
-
10 Hideki KOBAYASHI and Masaru NONAKA
1. Platychasmatinae Nakamura, 1956Platychasmatinae Nakamura,
1956 [as Platycasmatinae]. Tinea 3: 143. Type genus: Platychasma
Butler, 1881.Platychasmatinae: Miller, 1911: 193; Kitching &
Rawlins, 1999: 363; Nakamura, 2007: 327; Schintlmeister, 2008:
409; Kobayashi, 2011: 138; Schintlmeister, 2013: 17.
The branching order of the Platychasmatinae is slightly
different in some phylogenetic tree that we conducted. In the tree
Fig. 1.1 and Fig. 1.2, the Platychasmatinae are branching first as
the sister clade with the rest of the Notodontidae. In Fig. 1.3
(with adding quadrifid noctuoids), the Platychasmatinae are
branching next to the Scranciinae. In Fig. 1.4, the
Platychasmatinae are in the subfamily Thaumetopoeinae, and this is
not adequate from the aspect of morphology. In Figs 1.51 and 1.52,
basal lineage tree, the Platychasmatinae are positioned next to the
Scranciinae. It is possible that the location in the Fig. 1.1 and
Fig. 1.2 is due to LBA, however all the phylogenetic trees
represent that it is certain that the Platychasmatinae are one of
the primitive lineage in the Notodontinae. The specific characters
of the Platychasmatinae are the forewing of the quadrifid cubital
vein (M2 arising close to or with M3), and the tibial spur apices
being smooth, both the exceptional characters in the Notodontidae
and affiliating the Noctuidae, however the male genitalia have
socii. Considering the genitalia, we think the Platychasmatinae are
positioned next to the Scranciinae. Miller (1911) studied the
morphology of Platychasma virgo Butler, 1881 precisely. He observed
the tympanum: absence of nodular sclerite, concaved epimeron, and
presence of metascutal bulla. These are synapomorphies for
Notodontidae+Doidae. The larvae have two MD setae on A1. This form
is for Notodontidae. He also stated the caterpillar showed
primitive character states, such as fully developed anal prolegs
and long secondary setae. He located the phylogenetic position of
the Platychasmatinae as the evolved clade next to the Pygaerinae
and Thaumetopoeinae, and as the sister clade of the rest of the
Notodontidae according to the result of the Hennig86 analysis with
using 174 characters. The branching order is not fixed for now and
it needs further molecular research with more samples and fitted
DNA sequence parts.
Recognized genera of the Platychasmatinae
Platychasma Butler, 1881; Paracyphanta Sugi, 1994; Cyphanta
Walker, 1865; Sphetta Walker, 1865.
2. Scranciinae Miller, 1991 Scaranciini Miller, 1991. Bull. Am.
Mus. nat. Hist. 204: 184. Type genus: Scrancia Holland,
1893.Scranciinae: Schintlmeister, 2008: 47; Schintlmeister, 2013:
17; Schintlmeister, 2015: 14.
This subfamily is the second branch of the Fig. 1.1 tree, but as
stated in the Platychasmatinae section, this subfamily might be the
most primitive subfamily. Fig. 1.5, the cladogram of the basal
lineage group, gives the bootstrap value 91 to the Scranciinae
clade and it is the first branching clade. Miller (1991) settled
this clade as a tribe in the subfamily Dudusinae; however our study
showed this clade as a constantly independent clade apart from
Dudusinae clade. Miller added Scrancia and Gargetta to the
subfamily Dudusinae based on his newly identified numerous
synapomorphies, and gave them the tribe name Scranciini. Some of
the synapomorphies in his Scranciini are the protruding frons and
long legs in adults, the male genitalia of lacking socii and narrow
valvae, the larval reduced A3 prolegs. These can be used as the
synapomorphies of the Scranciinae. And counting the conformation of
the male genitalia, this subfamily is the nearest to the Noctuidae,
as Fig. 1.5 depicts. Miller considered the Dudusinae have two
tribes, Dudusini and Scranciini, based on the two synapomorphies,
i.e. presence of well-developed ocelli in adults and absence of
seta X on the anal shield. We think those apomorphies had occurred
two times in the primitive lineage; in the Scranciinae and in the
Dudusinae.
Recognized genera of the Scranciinae
Phycidopsis Hampson, 1893; Lasioceros Bethune-Baker, 1904;
Gargetta Walker, 1865; Pseudogargetta Bethune-Baker, 1904 ； Thacona
Walker, 1865; Hijracona Holloway, 2011; Schintlmeister & Witt,
2015 contained seven African genera to the Scranciinae including
Thacona.Scrancia Holland, 1893 is treated as a junior synonym of
Thacona Walker, 1864 (Schintlmeister &
-
Molecular phylogeny of the Notodontidae 11
Witt, 2015: 20). Porcica Walker, 1866 is a junior synonym of
Thacona Walker, 1864 (Holloway, 2011: 349)
3. Pygaerinae Duponchel 1845Pygaeridae Duponchel 1845. Cat.
méth. Lépid. Eur.: 95. Type genus: Pygaera Ochsenheimer,
1810.Melalophae Hübner, 1822. Verz. bekannter Schmett.: 162.
Unavailable.Gluphisiinae Packard, 1895. Mem. natn. Acad. Sci. 7:
88. Type genus: Gluphisia Boisduval, 1828.Melalophidae Grote, 1895.
Mitt. Roemermus. Hildesh. 1 (15): [3]. Type genus Melalopha Hübner,
1822.Pygaerinae: Miller, 1991: 176; Kitching & Rawlins, 1999:
363 ； Lafontaine & Schmidt, 2010: 10; Schintlmeister,
2008: 375; Kobayashi & Chen, 2011: 164; Kobayashi, 2011:
116; Schintlmeister, 2013: 17; Schintlmeister, 2015: 19.
The Pygaerinae are the third branch of the five basal clades.
This subfamily is well-known for its hairy larvae that had been
considered as the key character. Miller (1911) stated “Hairy larvae
occur throughout the Notodontidae, and a group formed solely on
that basis has no chance of being monophylylic.” He counted the
pleats of sacculus on the male genitalia. The four genera of our
Pygaerinae have the characteristic sacculus. Rhegmatophila, which
has the same sacculus, belongs to
①
②
③
④
⑤
⑥
Sphingidae Mimas christophi801Amphipyra schrenckii Nomugi
94Lymantria fumida Nomugi 942LymAsota egens Sulawesi
1147AganaCatocala patala Nomugi 949ErebPericaria matronula Nomugi
936Phycidopsis albovittata1185 OkLasioceras sola1138 Sulawesi
0Gargetta eucharius090 Vietnam Gargetta hampsoni788
sulawaesiGargetta divisa1136 Sulawesi 0Platychasma virgo798
ArimayamaGonoclostera timoniorum1000 AkGluphisia crenata470 Yunnan
07Gluphisia crenata347 Hokkaido Micromelalopha troglodita862
AMicromelalopha dorsimacula540 Micromelalopha sp277 Laos
0807Micromelalopha adrian875 NanliClostera anachoreta655
HokkaidClostera mahatoma469 Yunnan 08Clostera cultula1068 Deutsch
0Clostera sp1142 Sulawesi 0129 Clostera anastamosis691 AkiyosDudusa
celebensis775 Sulawesi Dudusa synopula552 Taiwan 1108Dudusa
sphingiformis544 TaiwanDudusa nobilis905 Nanling 0802Tarsolepis
punicea770 SulawesiTarsolepis japonica545 Taiwan Tarsolepis
taiwana008 Taiwan 1Tarsolepis brunnea857 Nanling Tarsolepis
fulgurifera506 YunnTarsolepis fulgurifera301 TaiwThaumetopoea
processionea DeutGazalina crysolopha233 Yunnan Epicoma contristis
Thau 1216 TAglaosoma variegata1212 ThauChadisra bipartita294 Taiwan
0Ellida arcuata244 Akashina 072Periergos dispar1047 Nanling
0Hupodonta lignea653 Hokkaido 0Rachia striata984 Yunnan 0131
Gangarides wallacei790 Sulawes
9565
82
50
10098
55
91
55
10079
28
52
81
48100
100
99
99
63
100
67
96
91
93
68
100
44
97
70
100
100
37
37
92
22
41
47
15
17
47
Fig. 1.51. Phylogenetic tree. Basal lineage (28SrRNA, NJ).
①Platychasmatinae; ②Scranciinae; ③Pygaerinae; ④Dudusinae;
⑤Thaumetopoeinae; ⑥Periergosinae;.
-
12 Hideki KOBAYASHI and Masaru NONAKA
the Pygaerinae. Gluphisia is a disputed genus for a long time.
Packard (1895) erected the Gluphisiinae for Gluphisia and Forbes
(1939, 1948) considered Gluphisia in the Gluphisiini belonging to
the Notodontinae. Miller placed Gluphisia in his Dicranurini in the
Notodontinae, and Nakamura (2007) also treated it as Miller. Our
molecular analysis has revealed Gluphisia belongs to the
Pygaerinae. The bootstrap value of the Pygaerinae which contains
Gluphisia is 97 and 95 in Figs 1.51 and 1.52. Schintlmeister (2003)
included Spatalia Hübner, 1819, Allata Walker, 1862, Metaschalis
Hampson, 1893, Ginshachia Matsumura, 1929, Pterotes Berg, 1901,
Rosama Walker, 1855 in his Pygaerinae; however their genitalia have
no such apomorphy as the Pygaerinae. Our phylogenetic tree shows
those genera belong to the Spataliinae. Schintlmeister & Witt
(2015) included African genera Peratodonta Aurivillius, 1904,
Janthinisca Kiriakoff, 1959 and Scalmicauda Holland, 1893 in the
subfamily Pygaerinae. Their male genitalia have not the pleats of
sacculus, but we have not conducted molecular examination of those
genera.
Recognized genera of the Pygaerinae
Micromelalopha Nagano, 1916; Clostera Samouelle, 1819; Gluphisia
Boisduval, 1828; Gonoclostera Butler, 1877; Pygaera Ochsenheimer,
1810; Rhegmatophila Standfuss, 1888.
①
②
③
④
⑤
⑥
Sphingidae Mimas christophi801Amphipyra schrenckii Nomugi
94Lymantria fumida Nomugi 942LymAsota egens Sulawesi
1147AganaCatocala patala Nomugi 949ErebPericaria matronula Nomugi
936Phycidopsis albovittata1185 OkLasioceras sola1138 Sulawesi
0Gargetta eucharius090 VietnamGargetta divisa1136 Sulawesi
0Gargetta hampsoni788 sulawaesiPlatychasma virgo798
ArimayamaGluphisia crenata347 HokkaidoGluphisia crenata470 Yunnan
07Gonoclostera timoniorum1000 AkMicromelalopha adrian875
NanliMicromelalopha sp277 Laos 0807Micromelalopha
dorsimacula540Micromelalopha troglodita862 AClostera anastamosis691
AkiyosClostera sp1142 Sulawesi 0129Clostera anachoreta655
HokkaidClostera cultula1068 Deutsch 0Clostera mahatoma469 Yunnan
08Dudusa celebensis775 SulawesiDudusa nobilis905 Nanling 0802Dudusa
sphingiformis544 TaiwanDudusa synopula552 Taiwan 1108Tarsolepis
japonica545 TaiwanTarsolepis punicea770 SulawesiTarsolepis
taiwana008 Taiwan 1Tarsolepis brunnea857 NanlingTarsolepis
fulgurifera506 YunnTarsolepis fulgurifera301 TaiwAglaosoma
variegata1212 ThauEpicoma contristis Thau 1216 TThaumetopoea
processionea DeutGazalina chrysolopha233 YunnanPeriergos dispar1047
Nanling 0Gangarides wallacei790 SulawesChadisra bipartita294 Taiwan
0Ellida arcuata244 Akashina 072Hupodonta lignea653 Hokkaido 0Rachia
striata984 Yunnan 0131
100
3799
91
96
99
59100
3598
100
96
100
98
97
100
51
95
38
91
88
64
76
61
95
48
21
30
75
90
55
59
55
76
48
54
69
35
29
32
32
Fig. 1.52. Phylogenetic tree. Basal lineage (28SrRNA, ML).
①Platychasmatinae; ②Scranciinae; ③Pygaerinae; ④Dudusinae;
⑤Thaumetopoeinae; ⑥Periergosinae.
-
Molecular phylogeny of the Notodontidae 13
4. Dudusinae Matsumura, 1929 Dudusinae Matsumura, 1929. Zool.
Mag. 37: 406. Type genus: Dudusa Walker, 1865.Tarsolepidinae
Kiriakoff, 1950. Bil. Jaarb. 17: 104. Type genus: Tarsolepis
Butler, 1872.Dudusinae: Miller, 1991: 181; Kitching & Rawlins,
1999: 363; Lafontaine & Schmidt, 2010: 11; Schintlmeister,
2008: 28; Schintlmeister, 2013: 16.Dudusini: Nakamura, 2007:
318; Kobayashi, 2011: 128.
This subfamily is located next to the Pygaerinae in Fig. 1.1 and
Fig. 1.4. In Fig. 1.3 it branches next to the Thaumetopoeinae. In
Fig. 1.2, it branches from the derived clades. In Fig. 1.51, NJ
tree, the Dudusinae are a sister clade with the Pygaerinae and in
Fig. 1.52, ML, it is a sister clade with the clade composed of the
Thaumetopoeinae and the Periergosinae. The morphological
classification postulates that the Dudusinae are near to
Scranciinae. Miller (1991) broadened the original concept of the
subfamily Dudusinae Matsumura which contained Dudusa and Tarsolepis
and closely related species. He added Scrancia Holland, 1893 and
Gargetta Walker, 1865 to the subfamily Dudusinae and gave them the
tribe name Scranciini. He considered that the most convincing
synapomorphies for his Dudusinae, composed of the Dudusini and
Scranciini, were presence of well-developed ocelli in adults and
absence of seta X on A10 in the larvae. For his Dudusini he counted
the loss of the metascutal bulla. On the other hand, Nakamura
(2007), in studying pupal morphology, placed his Dudusini as one of
tribes of the Notodontinae. The synapomorphy of Notodontinae that
he indicated was the presence of L2 seta on 4th abdominal segment.
However, his sample of the Dudusini was one species only, i.e.
Tarsolepis japonica Wileman & South, 1917, and he did not
examine the Scranciinae species. Considering the close relation of
the Dudusinae to Pygaerinae and Scranciinae, we take its branching
as Fig. 1.1 and Fig. 1.4. The bootstrap value of the Dudusinae is
99 in Fig. 1.51 and 90 in Fig. 1.52. It is possible to divide the
Talsolepis along the cladogram, i.e. Talsolepisoides Nakamura, 1976
for the clade containing T. japonica and T. punicea, Tarsolepis for
the clade of T. taiwana, and Megashachia Matsumura, 1929 for the
clade containing T. brunnea and T. fulgurifera. We could not
examine Stigmatophorina sericeus Rothschild, 1917.
Recognized genera of the Dudusinae
Dudusa Walker, 1865; Talsolepis Butler, 1872.
5. Thaumetopoeinae Aurivillius, 1889 (1861) Thaumetopoeinae
Aurivillius, 1889 (1861) Nordens Fjarilar. Handb. Sveriges, Norges,
Danmarks och Finlands
Macrolepidd.: 68, 71. Type genus: Thaumetopoea Hübner, 1820
[unnecessary replacement name for Cnethocampides but valid by ICZN
art 40.2 (Speidel & Naumann, 2005: 216)].
Cnethocampides Wallengren, 1861. K. svenska Fregatten Eugenies
Resa omkring Jorden (Zool.) 1 (10, Lepidoptera): 368. Invalid. Type
genus: Cnethocampa Stephens (1828). Cnethocampa is a junior synonym
of Thaumetopoea. Then Aurivillius (1889) made an unnecessary
replacement name, Thaumetopoeinae Aurivillius, 1889, but the latter
name is valid today by ICZN art 40.2.
Cnethocampinae Turner, 1922. Proc. Linn. Soc. N. S. W. 47: 362.
Anaphinae Kiriakoff, 1970. Ent. Z. 80: 66. Nomen nudum.
Unavailable. Type genus: Anaphe Walker, 1855.Epicominae Kiriakoff,
1970. Ent. Z. 80: 66. Nomen nudum. Unavailable. Type genus: Epicoma
Hübner, 1819. Thaumetopoeidae: Common, 1990: 421. Thaumetopoeinae:
Miller, 1991: 175; Kitching & Rawlins, 1999:
362.Thaumetopoeidae: Schintlmeister, 2008: 7.Thaumetopoeinae:
Schintlmeister, 2013: 17.
The rank of the Thaumetopoeinae, whether it is a subfamily or it
receives a family status, had been a long controversy till Miller
(1911) concluded to give it the subfamily rank. He also excluded
Oenosandra from the Thaumetopoeinae and from Notodontidae and
erected the Oenosandridae. And he accepted the family rank of
Doidae as a sister clade of the Notodontidae in Noctuoidea. He
showed presence of symplesiomorphy shared by Doa and the quadrifid
Noctuoidea and lack of important synapomphy of Notodontidae.
Schintlmeister (2008: 7) stated “In the present work the
Thaumetopoeidae are recognized as a distinct family, which is
linked probably to the Lymantriidae and excluded from the
Notodontidae.” He did not give the basis on morphological
cladistics or
-
14 Hideki KOBAYASHI and Masaru NONAKA
molecular phylogeny to his notion. Nakamura (2008, pers. com.)
observed the pupal pattern of Thaumetopoea species and could not
detect any differences from that of Pygaerinae species. He,
therefore, thought his Thaumetopoeini were a tribe of the
Pygaerinae. We think the resemblance of the pupae in them
represents a phenetic situation of affinity of the two primitive
subfamilies. The Thaumetopoeinae branch next to the Dudusinae, in
Fig. 1.1, but before this and next to the Pygaerinae in Figs 1.2
and 1.3. In Fig. 1.4, it contains Platychasma virgo and makes a
Periergosinae-Heterocampinae complex. In Fig. 1.5, however, the
Thaumetopoeinae are the sister clade with the Periergosinae, hence
we take the Thaumetopoeinae branching next to the Dudusinae. The
bootstrap value for the Thaumetopoeinae is 36 in Fig. 1.1, 47 in
Fig. 1.51 and 88 in Fig. 1.52. The diverging order is different in
every tree. The Palearctic group and the Australian group separate
in Fig. 1.51, but the bootstrap values are 41 and 22 each,
consequently the affinity is not strong. Kiriakoff (1970) split his
Thaumetopoeidae into three subfamilies based on the distribution,
that is, African, Palearctic and Australian. Our molecular analysis
results that the stability of separating the Palearctic genera and
Australian genera is only weak.
Recognized genera of the Thaumetopoeinae
Thaumetopoea Hübner, 1820; Gazalina Walker, 1865; Aglaosoma
Scott, 1864; Epicoma Hübner, 1819.
Twenty seven genera are recorded in the world including above
examined (by Schintlmeister, 2013): 12 in Australia (Aglaosoma
Scott, 1864, Axiocleta Turner, 1911, Cynosarga Walker, 1865,
Epicoma Hübner, 1819, Epicomana Kiriakoff, 1970, Mesodrepta Turner,
1924, Ochrogaster Herrich-Schäffer, 1855, Oxymetopa Turner, 1943,
Sthenadelpha Turner, 1922, Tanystola Turner, 1922, Themerastis
Turner, 1903, Trichiocercus Stephens, 1835); 2 in W-Papua
(Paramarane Bethune-Baker, 1910, Tearina Kiriakoff, 1970); 1 in
Asia (Gazalina Walker, 1865); 3 in Europe (Traumatocampa
Wallengren, 1871, Thaumetopoea Hübner, 1820, Helianthocampa de
Freina & Witt, 1985); 6 in Africa (Zastonia Herrich-Schäffer,
1858, Paraphlebs Aurivillius, 1921, Paradrallia Bethune-Baker,
1908, Epanaphe Aurivillius, 1904, Anaphe Walker, 1855, Adrallia
Walker, 1865); 3 in Madagascar (Pseudohypsoides Viette, 1960,
Nesanaphe Kiriakoff, 1969, Hypsoides Butler, 1882).
6. Periergosinae Kobayashi, nom. nov. (Fig. 6) Type genus:
Periergos Kiriakoff, 1959: 321.Pydninae Matsumura, 1925. Zool. Mag.
37: 408. Type genus Pydna Walker, 1856.
The genus Pydna Walker, 1856 is a junior homonym of Pydna
Herrich-Schäffer, 1855 (Geometridae). Therefore the subfamily name
Pydninae Matsumura is invalid. Hence the new replacement name for
Pydninae is proposed here according to ICZN art. 39. Since there
exist no available family-group name to replace the Pydninae, and
the current valid name for Pydna is Periergos Kiriakoff, 1959,
accordingly the replacement name for Pydninae is Periergosinae.
Fletcher (in Watson, Fletcher & Nye, 1980:72) proposed the
replacement name Eupydna for Pydna (type species Pydna testacea
Walker, 1856). However Pydna testacea belongs to Periergos
Kiriakoff, 1959 (type species: Periergos obsoleta Kiriakoff, 1958).
Thereby Eupydna is invalid as junior synonym of Periergos. Fletcher
did not establish a family-group name for the Pydninae.
This subfamily comprises the genera that make a stable
monophyletic clade, though the bootstrap values of the clade are
31, 16 and 15 in Figs 1.1, 1.2 and 1.3. In Fig. 6, the bootstrap
value is 93. This clade is very stable in the cladograms presented
here and many other phylogenetic trees which we conducted but are
not presented here. In Figs 1.51 and 1.52, the bootstrap value of
the Periergosinae is 92 and 54, and it takes the position at the
last diverged clade among the basal lineage. In Fig. 1.4, however,
it is divided in two clades and positioned in a
Thaumetopoeinae-Periergosinae-Heterocampinae clade. Of which, the
Gangarides-Periergos clade is located at the sister clade of the
rest, but the bootstrap value is 4, and
Chadisra-Ellida-Rachia-Hupodonta clade is positioned at the sister
clade of the Thaumetopoeinae, the bootstrap value is 10. Though the
divergent order is different, it is depicted that the two clades
are not contained in the Thaumetopoeinae or in the
Heterocampinae.
-
Molecular phylogeny of the Notodontidae 15
In Fig. 1.1 and Fig. 6, the bootstrap value for the clade
comprising Ellida and Chadisra is 100, and for the clade comprised
Hupodonta and Rachia it is 100, and the values are alike in Figs
1.2, 1.3, 1.4 and 1.5. Nakamura (2007) placed the clade configured
of Ellida and Hupodonta in the tribe Notodontini. Though their
belonging tribe is different from ours, his classification
indicates that Ellida and Hupodonta share some morphological
affinity of pupae, and this affinity of the two separated branch in
the Periergosinae can corroborates the validity of this subfamily
clade.
The larvae of Chadisra and Ellida much resemble each other in
appearance: cylindrical body green or yellow with reddish dorsal
beaded line. When resting, it bends its head and thorax laterally
backwards to make U-shaped form (Figs 6.81, 6.82).
The male genitalia of the Gangarides have a small uncus and
large valvae (Fig. 6.32). Gangarides roseus (Walker, 1865) has a
very small trapezoidal uncus with extensions on the caudal corners,
and the valva with long spines on the ampula and the harpe.
Gangarides puerariae Mell, 1922 has very large valvae with big
costal protrusions and strong teeth on the sacculi and harpes. The
uncus is small, roundish square with two small protrusions from the
caudal corners. Those of the Periergos magna are also very curious
(Fig. 6.31). The valva has a spinous costa and a staut sacculus
with a acute process at the distal end. The uncus has two
processes.
Recognized genera of the Periergosinae
Periergos Kiriakoff, 1959; Gangarides Moore, 1866; Rachia Moore,
1879; Hupodonta Butler, 1877; Ellida Grote, 1876; Chadisra Walker,
1862.
We affiliate the following genera based on the resemblance in
formation of the male genitalia. Chalepa Kiriakoff, 1959;
Franzdaniela Sugi, 1992.
①⑤
⑥
⑦
Platychasma virgo798 ArimayamaEpicoma contristis Thau 1226
VHeterocapma astarte741 Louisia
Chadisra bipartita294 Taiwan 0Chadisra vittata Sulawesi
779Ellida branickii404 Hikawa 032
Ellida viridimixta610 Miyagi 0Ellida arcuata244 Akashina 072
Periergos dispar1047 Nanling 0Periergos magna1055 Nanling 09
Hupodonta lignea653 Hokkaido 0Hupodonta lignea Taiwan
Hupodonta corticalis396 AkashiHupodonta corticalis977 Yunnan
Rachia lineata1167 Taiwan 0131Rachia striata984 Yunnan 0131
Gangarides wallacei790 SulawesGangarides vardena Laos268
Gangarides dharma Yunnan 539Gangarides puerariae Laos
284Gangarides puerariae Yunnan685Gangarides puerariae
Nanling916
100
100
99
84
100
100
99
100
71
100
100
53
93
97
77
84
100
94
0.005
Fig. 6. Phylogenetic tree. Periergosinae (28SrRNA, NJ).
①Platychasmatinae; ⑤Thaumetopoeinae; ⑥Periergosinae;
⑦Heterocampinae.
-
16 Hideki KOBAYASHI and Masaru NONAKA
7. Heterocampinae Neumoegen & Dyar, 1894 (Fig.
7)Heterocampinae Neumoegen & Dyar, 1894. Trans. Am. ent. Soc.
21: 183 (key), 200. Type genus: Heterocampa
Doubleday, 1841.Heterocampinae: Packard, 1895; Miller, 1911:
185; Kitching & Rawlins, 1999: 364; Nakamura, 2007: 318;
Lafontaine & Schmidt, 2010: 11; Kobayashi & Chen, 2011:
173; Kobayashi, 2011: 128; Schintlmeister, 2013: 16.
Fentoniinae Matsumura, 1929. Insecta matsum. 4 (1/29): 78. Type
genus: Fentonia Butler, 1881.Disfraginae Thiaucourt, 1995: 455.
Nomen nudum. Unavailable [art. 13.1. ICZN]Disfraginae:
Schintlmeister, 2013: 16.Rifarginae Thiaucourt, 1995: 458. Nomen
nudum. Unavailable [art. 13.1. ICZN] Rifarginae: Schintlmeister,
2013: 17.
In our study, the divergence of the Heterocampinae, Phalerinae,
Spataliinae and Notodontinae is shallow. There are some genera
which unite with the different clade in the different tree;
nevertheless, the core genera of each clade are stable, we hence
accept those four subfamilies.
Miller (1991) placed Heterocampini, Stauropini, Fentonia and
Spatalia in his Heterocampinae. But he stated the unclarity of the
Heterocampinae, “I was unable to identify a single reliable
synapomorphy from adult morphology”, and “The Heterocampinae will
likely prove to be a large and extremely complex subfamily”.
His Heterocampini contain Lochmaeus bilineata (Packard),
Schizura biedermani Barnes and McDunnough, Heterocampa oblique
Packard and Macrurocampa marthesia (Cramer). His Stauropini
contains Schizura unicornis (J. E. Smith), Cnethodonta griscescens
Staudinger, Harpyia microsticta (Hampson) and Stauropus fagi
(Linnaeus).
We have obtained the following affiliations: Fentonia in the
Heterocampinae, Spatalia in the Spataliinae. Concerning Miller’s
Stauropini and the comprising genera, we have acquired the
following result. The tribe Stauropini belongs to the Notodontinae;
Schizura unicornis remains in the Heterocampinae; Harpyia
microsticta belongs to the Dicranurini in the Notodontinae;
Cnethodonta griscescens and Stauropus fagi belong to the Stauropini
in Notodontinae.
Miller (1991) stated, “Interestingly, the two Schizura species
fall out separately. I could recognize these two clade by using the
tribal names Heterocampini and Stauropini for each” In our study,
also, Schizura is not a monophyletic genus. Schizura unicornis
unites with Oligocentria semirufescens; Schizura badia unites with
Hyparpax aurora; Schizura leptinoides unites with Heterocampa
umbrata. Oligocentria and Heterocampa should also be reexamined on
their monophyly, although all of them belong to the Heterocampinae
(Fig. 7).
The bootstrap value for the clade composed with Destolmia,
Formofentonia and Cascera is 72 in Fig. 1.1 and 70 in Fig. 1.3, 79
and 74 in Fig. 1.2 and Fig. 1.4; however their belonging subfamily
is not stable. In Fig. 1.2 and Fig. 1.4 the clade is in the
Stauropini of the Notodontinae. We consider this clade belonging to
the Stauropini according to Fig. 10.2, though the bootstrap value
is low 50 (q. v. section Stauropini).
As regard to the Disfraginae Thiaucourt and Rifarginae
Thiaucourt, they are nomen nudum, therefore not available [art.
13.1., ICZN]. Weller (1992) placed Disphragis notabilis (Schaus) in
the Heterocampinae and she placed Rifargia in the Heterocampinae as
unknown tribe. In our cladogram, Disphragis sabaria makes a clade
with Heterocampa, the bootstrap value is 87 (Fig. 7). We think
those genera do not need new subfamily for now.
Revived genus
Hemifentonia Kiriakoff, 1967Parafentonia Kiriakoff, 1964: 277.
Homonym of Parafentonia Roepke, 1944.Hemifentonia Kiriakoff, 1967:
152. Replacement name for Parafentonia Kiriakoff, 1964. Neopheosia
Matsumura, 1920: Schintlmeister & Pinratana, 2007: 159.
Type species: Parafentonia inconspicua Kiriakoff, 1963 (synonym
of Drymonia mandschurica Oberthür, 1911).
-
Molecular phylogeny of the Notodontidae 17
Schintlmeister & Pinratana (2007) treated Hemifentonia as a
junior synonym of Neopheosia, on the basis of the shape of the
uncus being Y-shaped. But from the viewpoint of the molecular
analysis, Neopheosia is a paraphyletic clade from Hemifentonia and
a sister clade with Fentonia (Fig. 7). Neopheosia fasciata (Moore,
1888), the type species of the Neopheosia, has indeed the Y-shaped
uncus, however it has a ventral process at its base (Fig. 7.26b,
c), a very unique formation, while in Hemifentonia mandschurica,
the uncus has no ventral process on its base and the uncus itself
forms gutter-shaped in entire length and has an U-shaped incisura
at distal end (Fig. 7.25b, c). In addition, the socius, valva,
aedeagus and 8th sternite have different patterns between the two,
thus mandschurica does not belong to the genus Neopheosia. Thereby
we think the Hemifentonia should be treated as a good genus on
phenetic and phyletic classification.
Revived combination
Hemifentonia mandschurica (Oberthür, 1911) Drymonia mandschurica
Oberthür, 1911: 323. Fentonia fasciculate Filipjev, 1927: 224
(synonymized by Tshistjakov, 1977).Microphalera styxana Shaus,
1928: 89 (synonymized by Schintlmeister, 1989).Parafentonia
inconspicua Kiriakoff, 1963: 278 (synonymized by Schintlmeister,
1989).Hemifentonia mandschurica (Oberthür, 1911): Kiriakoff, 1967:
153; Schintlmeister, 1992: 84; Kobayashi & Chen,
2011: 183.Neopheosia mandschurica: Schintlmeister &
Pinratana, 2007: 160; Schintlmeister, 2008: 197; Schintlmeister,
2013:
278.
Recognized genera of the Heterocampinae
Lochmaeus Doubleday, 1841; Hyparpax Hübner, 1825; Schizura
Doubleday, 1841; Oligocentria Herrich-Schäffer, 1855; Wilemanus
Nagano, 1916; Hemifentonia Kiriakoff, 1967; Neopheosia Matsumura,
1920; Fentonia Butler, 1881; Betashachia Matsumura, 1925;
Disphragis Hübner, 1820;
①②
③
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