An he aea cas anea Jo dan, 1910, he neglec ed "pseudomuga" silkmo h
(Lepidop e a, Sa u niidae)
Zhengyang Liu1, Changxiong Hu2, Muyang He3
1 Zhangdian Dis ic , Zibo, Shandong P o ince 255000, China
2 Se icul u al and Apicul u al Resea ch Ins i u e, Yunnan Academy o Ag icul u al Sciences, Mengzi, Yunnan P o ince 661101, China
3 Xiangximeilin, Anyuan Dis ic , Pingxiang, Jiangxi P o ince 337000, China
h ps://zoobank.o g/75BD9E13-71DD-480E-922E-D41E021A9DD2
Co esponding au ho : Zhengyang Liu ([email p o ec ed])
Academic edi o : Ma hias Seidel ♦
Recei ed
4 Ma ch 2025 ♦
Accep ed
9 Oc obe 2025 ♦
Published
4 No embe 2025
Abs ac
This s udy epo s he i s comple e li e his o y o An he aea cas anea Jo dan, 1910. Samples collec ed om he sou heas e n bo -
de s o Yunnan and Tibe we e ea ed on speci ic species wi hin he amily Lau aceae, con i ming ha A. cas anea is uni ol ine in
cap i i y. The comple e mi ochond ial genome sequences o A. cas anea and An he aea youngi Wa son, 1915, bo h belonging o he
subgenus An he aeopsis Wood-Mason, 1886, a e p o ided. Genomic and mo phological e idence demons a es ha A. cas anea is
he sis e g oup o all o he An he aeopsis spp. This inding s ongly suppo s he hypo hesis ha his subgenus o igina ed in he sou h-
eas e n egions o he Himalayas, making A. cas anea a key axon o unde s anding he e olu iona y his o y o he genus An he aea
Hübne , 1819. Gi en i s long-s anding neglec in se icul u e, we ecommend inc eased a en ion o A. cas anea in u u e s udies—
pa icula ly o p e en misiden i ica ion wi h he muga silkmo h, An he aea assamensis (Hel e , 1837), which sha es simila habi a s.
Key Wo ds
An he aeopsis, comple e mi ochond ial genome, India, Lau aceae, li e his o y, mo phology, muga, phylogeny, se icul u e, Tibe ,
wild silkmo h, Yunnan, zoogeog aphy
In oduc ion
The amous muga silkmo h, An he aea assamensis
(Hel e , 1837), is one o he mos economically impo an
species in No heas India (e.g., Peigle 1993, 2020). The
se icul u e o his species in Assam was epo ed in ea -
ly li e a u e (e.g., Hugon 1837), and i emains a cu en
ho opic in sa u niid s udies. As one o he h ee clades
cons i u ing he genus An he aea Hübne , 1819 (Nässig
1991), he muga subgenus An he aeopsis Wood-Mason,
1886, is es ic ed o sub opical and opical Asia. Ki ch-
ing e al. (2018) lis ed 12 cu en ly accep ed species.
Based on hei ype locali ies, hese include Sichuanese
An he aea cheng uana Wa son, 1923; Taiwanese An he -
aea o mosana Sonan, 1937; Andamanese An he aea
udlo i B echlin, 2002; Suma an An he aea b unnea an
Eecke, 1921; Bo nean An he aea youngi Wa son, 1915;
Ja an An he aea ubiginea Toxopeus, 1940; Palawan An-
he aea sahi Nässig & T eadaway, 1998; and Philippine
An he aea paniki Nässig & T eadaway, 1998. Howe e ,
An he aea in hanonensis Pauks ad & Pauks ad , 2013,
and An he aea enasse imensis Pauks ad & Pauks ad ,
2013, om Thailand a e e y close o A. assamensis in
e ms o mo phology and ba coding—we belie e hey a e
be e ea ed as di e en subspecies o e en synonyms.
Ano he species, o iginally collec ed in he Megha-
layan Khasi Hills, was named An he aea cas anea Jo -
dan, 1910, bu i has no been discussed by Indian au-
ho s excep o Gup a (2000). Naumann e al. (2008)
con i med ha An he aea mezops B yk, 1944, which is
ypically om Bu mese Kambai i, is a junio synonym
o A. cas anea. All Chinese specimens in ou hands we e
cap u ed only in he ain- ich b oadlea o es s a ele-
a ions o ca. 1200–2200 m in Mêdog, Tibe (Fig. 1B),
D sch. En omol. Z. 72 (2) 2025, 303–315|DOI 10.3897/dez.72.151952
Copy igh Zhengyang Liu e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use,
dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
dez.penso .ne
Zhengyang Liu e al.: Li e his o y and phylogeny o An he aea cas anea304
as well as in he Nujiang, Baoshan (Figs 1A, 4G), De-
hong, Pu’e , and Honghe egions o Yunnan. I has no
been ound in Xishuangbanna, which bo de s Laos and
has ela i ely low al i udes. This aligns wi h he opinion
o Pauks ad and Pauks ad (2008): he samples om
Laos iden i ied as A. cas anea in B osch e al. (1999)
we e de e mined inco ec ly. The gian mo hs o A. cas-
anea a e beau i ully cas aneous, as sugges ed by he e y-
mology o i s speci ic name. Ou obse a ions show ha
hese adul s eme ge om May o Augus in he wilds o
Tibe and Yunnan, which co esponds wi h he Bu mese
eco ds p o ided by Racheli (2008), and a e possibly uni-
ol ine in na u e (see also Sec ions 3 and 4 o Resul s).
Recen ly, ma e ial om he bo de be ween Vie nam
and Yunnan has been co ec ly iden i ied by local en o-
mologis s (e.g., Lien e al. 2014; Wu 2017: 150–151). In
summa y, A. cas anea should be seen as a mon ane and
summe - lying species endemic om no heas e n India
(and sou heas e n Tibe ) o no he n Myanma , h ough
wes e n and sou he n Yunnan, ex ending o no hwes e n
Vie nam bu seemingly excluding Laos. I ungbam and
I ungbam (2019) men ioned a Bhu anese dis ibu ion o
A. cas anea bu p o ided no suppo ing specimens.
Howe e , due o he simila i y be ween adul s and pa -
ial habi a o e lap, A. cas anea was p e iously misiden-
i ied as a subspecies o synonym o A. assamensis (e.g.,
B yk 1944; Holloway 1987; d’Ab e a 2012: 126–127;
Kuma e al. 2016). Al e na i ely, specimens o he o -
me we e e oneously a ibu ed o he la e (e.g., Sei z
1926–1928: 55B; Villia d 1969: 78; Yang e al. 2015:
Figu e 1. Li ing adul s o An he aea cas anea; da es indica e pho og aphy da es. A. ♂, Gaoligong Moun ains, Baoshan, Yunnan,
2214 m, 22 June 2025 ( he mo h was pho og aphed locally); B. ♀ No. 6, Mêdog Coun y, Nyingchi, Tibe , 2182 m, 06 July 2025 ( he
mo h was cap u ed locally and sen o Kunming, Yunnan, o pho og aphy). Scale ba s: 3 cm.
D sch. En omol. Z. 72 (2) 2025, 303–315
dez.penso .ne
305
533; Yi e al. 2015: 35). Fu he mo e, Pauks ad e al.
(1998) co ec ed he e oneous combina ion “A. cas a-
nea youngi,” which had p e iously been misapplied o
he Thai A. assamensis (e.g., Nässig e al. 1996; Lampe
2010: 274), as s a ed by Peigle (2010).
Compa ed o o he An he aeopsis popula ions, se e al
cha ac e is ics o A. cas anea male geni alia a e au apo-
mo phic (Pauks ad e al. 1999), in addi ion o i s uniquely
poin ed o ewing apex (Bou ie 1936: 161). I is possible
ha A. cas anea is he sis e g oup o all o he species wi h-
in An he aeopsis. Since pas s udies based on adul spec-
imens and sho e COI sequences ha e no add essed his
issue, dedica ed esea ch on he p e iously unknown p e-
imaginals o A. cas anea would help o cla i y i s phyloge-
ne ic posi ion. Addi ionally, his would allow highe -quali-
y genomes o be inco po a ed in o phylogene ic analyses.
Ma e ials and me hods
Collec ion and ea ing
This pape u ilizes he eggs om a o al o six wild-caugh
♀♀ A. cas anea o subsequen li e his o y s udies. Da a
o he mo hs a e as ollows: No. 1, Lüchun Coun y, Hong-
he, Yunnan, 1288 m, 18 July 2023; No. 2, same loca ion,
19 July 2023; No. 3, Jinping Coun y, Honghe, Yunnan,
1387 m, 19 June 2024; No. 4, same loca ion, 29 June 2024;
No. 5, same loca ion, 2 July 2024; No. 6, Mêdog Coun-
y, Nyingchi, Tibe , 2182 m, 27 June 2025. Among hem,
he eggs o ♀♀ Nos. 1–4 and 6 we e ea ed by Liu ZY in
Wuhua Dis ic , Kunming, Yunnan (1920 m); ♀♀ Nos. 1–5
we e ea ed by Hu CX in Mengzi, Honghe, Yunnan (1285
m); and ♀ No. 3 was ea ed by He MY in Anyuan Dis ic ,
Pingxiang, Jiangxi (112 m). The samples used o mo pho-
logical desc ip ion a e hose o Liu ZY, wi h he ollowing
e minology abb e ia ions: L1–5 = 1s –5 h la al ins a s;
T1–3 = 1s –3 d ho acic segmen s; A1–10 = 1s –10 h abdom-
inal segmen s. The dimensional da a desc ibed in he Re-
sul s a e based on measu emen s o single samples (n = 1).
The gene al equipmen used includes a e nie calipe
INSIZE 1108-150C 0–150 mm/0.01 mm (± 0.02 mm) (gen-
e al leng hs); a he mohyg ome e BENETECH GM1365
(± 2 RH%; ± 0.3 °C) (humidi y and empe a u es); a
NIKON D5500 wi h a LAOWA 60 mm /2.8–22 lens (Figs
1, 3, 4A–F, 5); and a SIGMA 10–20 mm /4–5.6 lens (Fig.
4G). One unha ched A. cas anea o um (su ace-d ied and
spu e -coa ed) was used o scanning elec on mic oscope
(SEM) imaging wi h a ZEISS GEMINISEM 360 (Fig. 2).
Molecula sequencing and phylogene ic analysis
One con ibu ion o his s udy is he p o ision o wo new
comple e mi ochond ial genome (CMG) sequences. The
issue was g ound in liquid ni ogen, ollowed by DNA
ex ac ion using 1 ml o p ehea ed (65 °C) CTAB bu -
e wi h 2% β-me cap oe hanol and 50 µl o p o einase
K (20 mg/ml). DNA was pu i ied by chlo o o m–isoamyl
alcohol (24:1) ex ac ion, isop opanol p ecipi a ion, and
RNase A ea men . Nucleic acid concen a ion was mea-
su ed using INVITROGEN QUBIT 3.0 wi h he QUBIT
dsDNA HS Assay Ki , and in eg i y was assessed by 1%
aga ose gel elec opho esis. Fo lib a y p epa a ion (using
he VAHTS Uni e sal Plus DNA Lib a y P ep Ki o Il-
lumina-ND617), we ollowed he ILLUMINA DNA P ep
P o ocol (#1000000025416), which included enzyma ic
agmen a ion, end epai wi h 3′ adenyla ion, adap e li-
ga ion, PCR ampli ica ion, and p oduc pu i ica ion using
VAZYME DNA Clean Magne ic Beads. Pai ed-end se-
quencing was ca ied ou on an ILLUMINA NOVASEQ
6000 pla o m using lib a ies ha passed quali y con ol.
The samples used a e as ollows:
• An he aea cas anea: An L1 la a om ♀ No. 3 (see
abo e), ha ched om o um on 2 July 2024. I was
killed on 5 July 2024 and p ese ed in 95% (± 5%) e h-
anol a −18 °C un il 13 Janua y 2025 o sequencing.
• An he aea youngi: A ♂ adul om T us Madi En o-
mology Camp, Keningau, Sabah, Malaysia, 1183 m,
cap u ed and killed on 6 Feb ua y 2023, hen s o ed
d y in a iangula en elope a oom empe a u e.
Tho acic muscles we e ex ac ed o sequencing on
13 Janua y 2025.
The CMG assemblies we e pe o med using SPADES
3.15.4 (P jibelski e al. 2020), hen anno a ed wi h MITOS2
(Dona h e al. 2019) and manually cu a ed o accu acy. The
co esponding high- h oughpu iles a e accessible in NCBI
BioP ojec PRJNA905660. OGDRAW (G eine e al. 2019)
was used o d aw he linea map, ollowed by manual isual
op imiza ion wi h AFFINITY PHOTO 2 (Fig. 6).
We ollowed he sugges ion o Liu (2024) o use
CMGs a he han he 13 p o ein-coding genes (PCGs)
o cons uc he i s phylogene ic ee (Suppl. ma e ial
1: ee S1). In addi ion o ou new sequences, his da ase
includes 39 o he species o Sa u niidae, wi h wo axa
om Bombycidae and one om Lasiocampidae selec -
ed as ou g oups (Suppl. ma e ial 1: able S1). Sequence
alignmen was pe o med using MAFFT 7 (Ka oh e al.
2019) wi h de aul se ings, and phylogene ic in e ence
was conduc ed using IQ-TREE 2 (Minh e al. 2020) un-
de he bes - i model GTR + F + I + R5, chosen acco d-
ing o BIC (Kalyaanamoo hy e al. 2017), wi h ul a as
boo s ap (1,000 eplica ions) (Hoang e al. 2018).
Fo he second ee (Suppl. ma e ial 1: ee S2), we se-
lec ed COI ba codes o 10 An he aeopsis spp. (based on
he comple e lis o 12 species p o ided by Ki ching e al.
2018, excluding he unce ain A. in hanonensis and A. enas-
se imensis). Th ee o i e sequences o each species ( om
o close o he ype locali y) we e downloaded om BOLD
Sys ems 4 ( 4.boldsys ems.o g) (Suppl. ma e ial 1: able S2)
and added o he abo e CMG da ase . The alignmen and ee
gene a ion we e pe o med using he same so wa e and pa-
ame e s. The esul ing comple e opologies o he wo max-
imum likelihood (ML) ees we e p elimina ily isualized
wi h ITOL 6 (Le unic and Bo k 2021), ollowed by manual
colo and layou design using AFFINITY PHOTO 2 (Fig. 7).
dez.penso .ne
Zhengyang Liu e al.: Li e his o y and phylogeny o An he aea cas anea306
To imp o e cla i y, we c ea ed an app oxima e cu en
dis ibu ion map o An he aeopsis spp. (Fig. 7). I is p i-
ma ily based on logical in e ences om ou phylogene ic
analysis, supplemen ed by specimens om China, Vie -
nam, and Bo neo in ou collec ion. Addi ionally, eco ds
om all a ailable li e a u e lis ed in he Re e ences we e
consul ed, wi h p e e ence gi en o hose om dedica -
ed axonomic s udies (e.g., B echlin 2001; Pauks ad and
Pauks ad 2008). In his wo k, we do no commen on
he axonomic handling o o he An he aeopsis axa by
Ki ching e al. (2018) aside om A. cas anea.
Resul s
1. Imma u e s ages o A. cas anea
om sou heas e n Yunnan
(based on o sp ing o ♀ No. 3)
1.1. O um (Figs 2, 3A): The o e all shape is i-axial
ellipsoid (2.65*2.22*1.68 mm), wi h a g ayish cho-
ion co e ed by da k b own glue. The mic opyla
ose e is composed o polygonal imp in s le by
ollicle cells. C owned ae opyles a e dispe sed
ac oss he ex e nal su ace, excep in and immedi-
a ely a ound he mic opyla zone.
1.2. L1 (Fig. 3B, C): The shiny ed-b own head capsule
is 1.28 mm wide, wi h long whi e b is les an e io ly.
The ce aco ia is g ay. The g ound colo om T1 o
A10 is yellow, i id do sally and pale en ally. Each
o T2–A8 has wo black ans e se s ipes, one an e io
and one pos e io , o he segmen al ans e se ow o
subdo sal and do sal scoli. Addi ionally, a black s ipe
deco a es each segmen al junc ion om T1 o A8. The
do sal midline T2–A8 is da k in colo . The p o ho acic
shield is black, wi h o wa d-poin ing and cu ed p i-
ma y se ae a he an e oma ginal posi ion (bo ne on he
ac ile do sal and subdo sal scoli). Mos o he scolus
bases a e he same yellowish colo as he su ounding
in egumen ; hey bea g ay, sligh ly cu ed p ima y
se ae. Howe e , he la e al and subdo sal scoli o T1–
A1, he subdo sal scoli o A9, and he do sal scoli o
A10 show mo e melanin a hei bases and hai s, wi h
he melanin ex ending in o he su ounding a eas.
A pai o do sal scoli a e used medially on A8. The
scle i es o he legs T1–3 a e smoo h and da k b own,
while he p olegs A3–6 a e pale yellow, he same colo
as he en e . The la e al pla e o p oleg A10 and he
pos cen al a ea o he anal shield a e black.
1.3. L2 (Fig. 3D, E): The wid h o he head capsule is
2.12 mm, and i s colo is he same as in L1, bu he
ce aco ia has changed o ligh och e. The in egumen
is gene ally yellowish-g een, and he black ans e se
s ipes p esen in he p e ious ins a a e now absen .
The do sal midline om T2 o A8 is da k g een. In he
la e al iew, he subdo sal scoli o A1–9 a e connec ed
by a ho izon al oli e-g een s ipe. In some indi idu-
als, a simila bu e y ain s ipe connec s he do sal
scoli om T2 o A8, hough i is absen in o he s. The
black p o ho acic shield is di ided by a ligh yellow
midline. All p ima y se ae o non-(sub) en al scoli
a e black, wi h he longes ones appea ing on he ac-
ile do sal and subdo sal scoli o T1. Addi ionally, all
la e al, subdo sal, and do sal scoli ha e a bluish-pu -
ple base. The la e op oximal a ea o each do sal
scolus om T2 o A9 displays a me allic sil e y zone,
hough i has no ully de eloped ye . All spi acles
a e b own in colo bu pale on A8. The b own legs
o T1–3 a e ligh e han in L1, and a subci cula black
spo has de eloped on each p oleg A3–6. The plan a
la e al su aces a e ligh b own. Fo he la e pe iod,
i s g eenish hue om T1 o A10 becomes mo e p o-
nounced. The bluish-pu ple wa s appea mo e i id,
wi h enhanced me allic sil e zones. A sup aspi acu-
la yellow band begins o de elop, ex ending om A1
o A10. Each segmen om A2 o A7 is ado ned wi h an
oblique da k g een subspi acula s ipe.
1.4. L3 (Fig. 3F, G): The head capsule measu es 3.69 mm
in wid h, wi h colo a ion simila o L2. The p e iously
dis inc da k do sal midline obse ed du ing L1–2 has
nea ly disappea ed. The sup aspi acula yellow band
p esen in L2 becomes mo e conspicuous, while mos
o he in egumen a y a eas ansi ion o g een. All non-
(sub) en al scolus bases u n pinkish-pu ple excep
o he pe sis en ly blue do sal scoli o A10. The do sal
sil e y spo s become mo e lus ous. The p oximal a -
eas o he coxae T1–3 and he la e al in egumen o T1
exhibi b igh e yellowish ones. Se e al black do s de-
elop on he la e al a eas o p olegs A3–6, each bea ing
a se a. The la e al pla e o p oleg A10 appea s sub ian-
gula , yellow supe io ly and black in e io ly, su ound-
ed by a ligh b own leshy “ ing”. The pos cen al
black pa ch on he anal shield sh inks in o a “V-shape”,
lanked by b own leshy ma gins. A ans e se ow o
yellow hai s appea ing on he an e io egion o each
A1–8. Sho and c ossed whi e se ae eme ge a he do -
sal junc ions o A2/A3, A3/A4, A4/A5, A5/A6, and
A6/A7. All spi acles excep he whi e one on A8 u n
black, and hei in e io oblique s ipes ( isible only on
A2–7) become mo e dis inc compa ed o L2.
1.5. L4 (Fig. 3H): Despi e li le change in colo a ion om
he p e ious ins a , he head capsule wid h inc eased
o 5.23 mm. The ce aco ia is pale wine-colo ed. The
pai ed black spo s on he p o ho acic shield du ing
L2–3 comple ely disappea , eplaced by b igh yel-
low-g een pigmen a ion ha in ensi ies along he
an e io ma gin o T1 and on he coxae T1–3. The su-
p aspi acula yellow band becomes s ikingly i id,
edged wi h ligh b own om A3 o A9 (inc easingly
p ominen pos e io ly, bu no isible on A3–6 in some
indi iduals). The subspi acula oblique s ipes p es-
en in L2–3 a e e y inconspicuous o e en disappea .
All la e al, subdo sal, and do sal scoli de elop mul-
be y-colo ed bases excep hose on T1, which ap-
pea ligh och e. The sho and c ossed whi e se ae
D sch. En omol. Z. 72 (2) 2025, 303–315
dez.penso .ne
307
obse ed in L3 now addi ionally appea a he A1/A2
junc ion, while he ans e se yellow se al ows on
he an e io zones o A1–8 become mo e de eloped,
wi h clubbed ips. The pa e n on A10 esembles ha
o L4, bu he la e al pla e u ns comple ely g een,
wi h a da ke pe iphe al “ ing”. The “V-shaped”
pos cen al black spo on he anal shield spli s cen-
ally in o wo pa s, ading om black o b own,
wi h i s cen al a ea eplaced by b igh g een.
1.6. L5 (Fig. 3I–R): This is he ma u e ins a , wi h a head
capsule 7.57 mm wide. The o e all colo a ion o he
head capsule is pale b ownish- ed, wi h da k b own
p ocesses on he la e al a eas. While supe icially
simila o L4, se e al dis inc mo phological changes
a e e iden . The o e all g een in egumen has become
ligh e in he do sal a ea while simul aneously da ken-
ing en ally, esul ing in a mo e p onounced coun e -
shading e ec . All scoli ha possessed a mulbe y-col-
o ed base in he p e ious ins a ha e now shi ed o
a i id pu ple. Fu he mo e, he wa s T1, p e iously
ligh och e du ing L4, a e now a dis inc lemon-yel-
low, and his yellow pigmen a ion is in ensi ied ac oss
he en i e an e io egion o his segmen —including
he coxae bu excluding he ce aco ia—gi ing i a
b igh e and mo e i id appea ance. No ably, he
majo i y o he yellow se ae loca ed on he an e io
zones o A1–8, which we e desc ibed in L4 as “ha ing
clubbed ips,” ha e now e e ed o poin ed ips.
1.7. Pupa (Fig. 4B–F): The pupal case is da k b own,
excep o a anslucen ‘window’ be ween he com-
pound eyes. The medial ma gins o he an ennae do
no mee along he en al midline, and bo h he max-
illae and he legs T1–2 a e isible. Spi acle T1 appea s
as a na ow gap a he bounda y be ween T1/T2; he
spi acles A2–7 a e well de eloped, bu he spi acle A8
is likely a emnan . A pai o o ewing ube cles is lo-
ca ed on he do sola e al egions o T2. The male geni-
al po e appea s as a na el-like sca on he mid en al
A9. The e minal A10 bea s poin ed c emas e hooks.
1.8. Cocoon (Fig. 4A): The eshly spun cocoons appea
whi e in a ela i ely d y en i onmen (ca. 50–70%
RH), bu a e ull oxida ion (100% RH, 48 h), hey
u n ligh b own. The uppe e minal end o he co-
coon ea u es a peduncle ha w aps a ound he hos
plan , wi h he pupal head o ien ed upwa d and he
exi side comple ely sealed. Cocoons a e enclosed by
su ounding lea es o p o ec ion.
2. Imma u e s ages o A.
cas anea om sou heas e n Tibe
(based on o sp ing o ♀ No. 6)
Al hough SEM obse a ion was no conduc ed, he ex-
e nal mo phology, size, and colo a ion o he eggs a e
nea ly iden ical o hose o he specimens om Yun-
nan (see also Sec ion 1.1). The L1–3 ca e pilla s a e also
gene ally consis en wi h he desc ip ions p o ided in
Sec ions 1.2–1.4. Howe e , se e al no able di e ences
we e obse ed: in he Tibe an popula ion, all bluish-pu -
ple scolus bases—which a e p esen in Yunnan L2 la -
ae—a e b igh aquama ine. Excep on T1, hese wa s
exhibi a dis inc i e yellow uppe po ion and a pinkish
lowe po ion h oughou L3–5. Ano he conspicuous dis-
inc ion is he p esence o mo e o less shiny, clubbed
se ae on all majo scoli in L4–5 (Fig. 5A, B), a ea u e
no obse ed in Yunnan la ae. Fu he mo e, he do sal
yellow hai s wi h clubbed ips, which we e documen ed
in Yunnan L4 la ae, a e consis en ly p esen in bo h L4
and L5 o he Tibe an lineage.
The la ae om Yunnan exhibi a da ke appea ance
in he head capsule and legs T1–3 du ing L2–5 compa ed
o hose om Tibe . Fu he mo e, he b ownish a eas on
A10 in L4–5 a e also da ke in he Yunnan popula ion han
in he Tibe an one (Fig. 5C, D) (see also Sec ions 1.5 and
1.6). The pupae and cocoons bo h con o m o he desc ip-
ions p o ided in Sec ions 1.7 and 1.8.
3. Rea ing epo o A. cas anea
om sou heas e n Yunnan
(based on o sp ing o ♀ No. 3)
Du ing he summe ea ing o 2023, all L1 la ae o Liu
ZY pe ished wi hin one week a e eeding on Cam-
pho a o icina um (Lau aceae) and Cinnamomum bu -
mannii (Lau aceae). In con as , Hu CX success ully
ea ed a ew indi iduals o L2 using he la e plan bu
did no con inue obse a ions. In he summe o 2024,
he h ee au ho s e-e alua ed hese wo plan s, and all
es ed la ae once again died du ing L1.
Liu ZY subsequen ly o e ed Que cus yunnanensis
(Fagaceae), Alnus nepalensis (Be ulaceae), and Liquid-
amba o mosana (Al ingiaceae) o 21 newly ha ched
L1 indi iduals, all o which comple ely ejec ed hese
Figu e 2. Cho ion o An he aea cas anea, SEM. A. The an e io pole, whose cen al a ea is he mic opyla zone; B. Mic opyla
ose e; C. Ae opyle c owns. Scale ba s: 100 μm (A); 20 μm (B); 10 μm (C).
dez.penso .ne
Zhengyang Liu e al.: Li e his o y and phylogeny o An he aea cas anea308
Figu e 3. O a and L1–5 o An he aea cas anea om sou heas e n Yunnan. A. O a; B, C, E. On hos plan Li sea populi olia; D, F–R,
R. On hos plan Campho a sep en ionale; B. L1, la e al iew; C. L1, do sal iew; D. L2, la e al iew; E. L2, do sal iew; F. L3,
la e al iew; G. L3, head and T1–3, an e ola e al iew; H. L4, la e al iew; I. L5, la e al iew; J. L5, do sal iew; K. L5, subdo sal
scolus o A2, apicola e al iew; L. L5, do sal scolus o A2, apicola e al iew; M. L5, do sal scoli o A8 (medially used), apical iew;
N. L5, leg T3, la e al iew; O. L5, head and T1–2, an e ola e al iew; P. L5, A10, do sal iew; Q. L5, A10, la e al iew; R. L5, spinning
he ex e nal pa [ loss] o cocoon, do sal iew. Scale ba s: 1 mm (K); 2 mm (L, M); 3 mm (A, N); 1 cm (D, E); 5 mm (B, C, G,
O–Q); 1.5 cm (F, H, I, J, R).
D sch. En omol. Z. 72 (2) 2025, 303–315
dez.penso .ne
309
op ions. Ul ima ely, a e accep ing Li sea populi olia
(Lau aceae) and Campho a sep en ionale (Lau aceae)
( ea ing unde condi ions o 16.5–25.8 °C wi h ca. 70.6–
90.4% RH), nine indi iduals success ully spun cocoons.
Howe e , h ee o hese died du ing he p e-pupa ion,
while he emaining six pupae a e all male.
L1 ca e pilla s li e independen ly on he unde sides
o lea es wi hou in egumen a y con ac wi h o he s,
hough 2–4 indi iduals may occasionally be ound on
he same lea . When a es , hey ypically assume a “J”-
shaped pos u e. Du ing L2–5, la ae a e s ill gene ally
soli a y, esiding ei he on he unde sides o lea es o
a he b anches. As wi h mos sa u niids, supplemen a y
wa e is essen ial o la al de elopmen . The la ae o
L4–5 usually de ach hei legs T1–3 (some imes includ-
ing p olegs A3–4) om he ege a ion, and du ing L2–5
hey we e consis en ly obse ed o ien ing hei heads
owa d he side wi h highe illuminance. A e eeding
ended [liquid de eca ion], la ae all spun cocoons on
hos plan s. The scoli o he whole la al s age ha e no
isible exoc ine sec e ion, wi hou i i a ion o human
skin. When he ma u e la ae a e a acked, hey will
Figu e 4. Cocoon and pupa o An he aea cas anea om sou heas e n Yunnan. A. Cocoon, spun and s o ed in a d y en i onmen ,
la e al iew; a: peduncle; b: he exi side; B–F. Pupa ♂; B. a: la e al iew; b: en al iew; c: do sal iew; C. Head and T1–2, en o-
la e al iew; D. T1–2, do sola e al iew; a: spi acle T1; b: o ewing ube cle; E. en al iew; a–c: A8–10 ( he geni al po e is loca ed a
he cen e o A9); F. C emas e on he ip o A10, do sal iew; G. The habi a in Gaoligong Moun ains, Baoshan, Yunnan P o ince,
China, 2214 m, 24 ♂♂ we e collec ed he e on 21 June 2024 and 21–23 June 2025 by Liu ZY. Scale ba s: 1 mm (F); 1.5 mm (E);
3 mm (C. D); 8 mm (B); 1 cm (A).
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Zhengyang Liu e al.: Li e his o y and phylogeny o An he aea cas anea310
shake T1–A3 om side o side, some imes clicking he
mou hpa s o make noise.
The de elopmen al calenda is as ollows (1 ♂, ed on
C. sep en ionale in 2024): 20 June, o iposi ed; 01 July,
ha ched in o L1; 06 July, 1s p e-ecdysis; 07 July, mol ed
in o L2; 12 July, 2nd p e-ecdysis; 13 July, mol ed in o L3;
17 July, 3 d p e-ecdysis; 18 July, mol ed in o L4; 25 July,
4 h p e-ecdysis; 27 July, mol ed in o L5; 12 Augus , eed-
ing ended and spun cocoon. On 28 May 2025, he mo h
eme ged om i s cocoon.
4. Rea ing epo o A. cas anea
om sou heas e n Tibe
(based on o sp ing o ♀ No. 6)
Based on he ea ing expe ience in 2023–2024 (see also
Sec ion 3), du ing he summe o 2025, a o al o 99 L1
la ae ( om 137 eggs) we e ea ed on C. sep en ionale
and Li sea ubescens ( he only a ailable L. populi olia
had died and was he e o e unusable), unde condi ions
o 17.8–26.4 °C wi h ca. 66.3–87.4% RH. The la e hos
was mo e eadily accep ed by newly ha ched ca e pilla s
han he o me . Compa ed wi h he L1 g oup ea ed on
L. ubescens, hose p o ided wi h C. sep en ionale ex-
hibi ed highe mo ali y and slowe de elopmen al p o-
g ession, in a iably ailing o each he L2. The e o e, all
su i ing la ae we e ul ima ely ea ed on L. ubescens.
Ul ima ely, se en ma u e la ae spun cocoons di ec ly on
his hos plan , whe eas he emainde le he ege a ion,
yielding a o al o 35 cocoons. The pe iod om ini ial
o iposi ion o he spinning o he las cocoon ex ended
om 27 June o 24 Augus . As o 19 Sep embe , no adul
eme gence had been eco ded. O he biological ai s ob-
se ed du ing he la al s age we e consis en wi h he
co esponding desc ip ions p o ided in Sec ion 3.
5. Phylogene ic Analysis on A. cas anea and
An he aeopsis
The CMG sequence leng hs a e 15,325 bp o A. cas anea
(GenBank accession numbe : PV136007) and 15,307 bp o
A. youngi (GenBank accession numbe : PV136008) (Fig. 6).
In he CMG-based phylogeny, all An he aeopsis nodes
ecei ed maximal suppo , whe eas ce ain nodes in he
CMG + COI opology showed lowe suppo (Fig. 7). The
wo esul s o m a single opology, indica ing ha he sub-
genus An he aeopsis is composed o h ee majo species
g oups. The CMG + COI ee shows a monophyle ic g oup
o ou species [((A. assamensis + A. udlo i) + A. cheng u-
ana) + A. o mosana] is dis ibu ed ac oss he mainland
o Sou heas Asia and i s no he n opical and sub opical
e i o ies ( he “cool colo s” in Fig. 7), i.e., he assamen-
sis-g oup. Meanwhile, ano he lineage occupies Sundaland
and he Philippines ( he “wa m colo s” in Fig. 7), known as
Figu e 5. Ma u e la ae o An he aea cas anea om sou heas e n Tibe , on hos plan Li sea ubescens. A. L5, la e al iew; B. L5,
do sal scolus o A4, la e al iew; C. L5, head and T1–3, an e ola e al iew; D. L5, A10, do sola e al iew; Scale ba s: 2 mm (B); 5 mm
(C, D); 1.5 cm (A).
Figu e 6. Linea map o he CMGs o An he aea cas anea ( op) and An he aea youngi (bo om).
D sch. En omol. Z. 72 (2) 2025, 303–315
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311
he youngi-g oup, which con ains i e species [((A. paniki
+ A. sahi) + (A. youngi + A. b unnea)) + A. ubiginea].
A. cas anea is he sis e axon o hese wo clades.
Discussion
Al hough he ca e pilla s om Honghe, Yunnan, and
Mêdog, Tibe , exhibi some mo phological di e ences,
we cu en ly do no p opose o ea hem as sepa a e
species—speci ically, we e ain om esu ec ing he
name A. mezops o he o me popula ion o naming
he la e as a new species. This decision is based on
se e al ac o s: no li e-his o y da a a e a ailable om
Meghalaya, wes e n Yunnan, o no he n Myanma ,
and he geni alia o he Tibe an mo hs equi e u he
s udy, as no males ha e ye been ob ained om ield
collec ion o ea ing expe imen s. Mo eo e , e en i
he wo we e o be classi ied as dis inc subspecies o
species, hey would s ill ep esen he closes known
sis e lineages wi hin he subgenus An he aeopsis, and
hese di e ences a e no suppo ed by eliable diagnos-
ic cha ac e s in he imaginal ex e nal mo phology, a
leas in emales. Technically, any po en ial u u e ax-
onomic e isions would no a ec he cu en discus-
sion, which ea s ma e ial om Yunnan and Tibe as a
monophyle ic g oup unde he name A. cas anea.
Regie e al. (2005) desc ibed he eggshell o A. ubigi-
nea (o iginally as “A. youngi om Ja a,” hen co ec ed by
Pauks ad and Pauks ad (2010)) as “ he ae opyle c owns
we e uni o mly dis ibu ed ac oss he eggshell excep im-
media ely su ounding he egion o he mic opyle a he
an e io pole.” Kuma and Kamble (2008) s a ed ha “all
he ae opyles o A. assamensis we e co e ed wi h a cup-
shaped s uc u e, and he e minals o his s uc u e ha e
many inge -like p ojec ions.” Ou obse a ions on A. cas-
anea con i m ha i s ae opyle c own dis ibu ion pa e n
aligns wi h hose o he wo a o emen ioned species.
Indian au ho s ha e published many mo phological s ud-
ies on he li e his o y o A. assamensis (e.g., Jolly e al. 1979:
122–126; Goswami and Singh 2012; Kaka i e al. 2012),
while Lampe (2010: 274) and Wu (2017: 135–144) ha e
illus a ed he imma u e s ages o he Thai and Chinese pop-
ula ions, espec i ely. Recen ly, Veenakuma i e al. (2011)
ound some wild la ae o A. udlo i om he sou he n An-
daman. We ha e ea ed ma e ials om Yunnan and Hain-
an, which we empo a ily e e o as A. assamensis, as hei
la ae appea nea ly indis inguishable om he ca e pilla s
o A. cheng uana om Ya’an, Sichuan, and A. assamensis
om Mêdog, Tibe ( ea ed by Liu ZY in 2022 and 2025,
espec i ely). The p eimaginal A. o mosana om Taiwan
a e also documen ed in se e al wo ks (e.g., Lin e al. 1993;
Wang 1994: 51–57; Wang and Heppne 1994; Peigle and
Wang 1996: 240–245). Pauks ad and Pauks ad (2001)
p o ided a comple e li e cycle o he Ja anese A. ubiginea
(o iginally as “A. youngi,” co ec ed by Pauks ad and Pauk-
s ad (2010)). Addi ionally, an Eecke (1929) igu ed a ma-
u e la a o A. b unnea om Suma a and p o ided a b ie
desc ip ion. In all he compa ed specimens, L1 consis en ly
exhibi ed a shiny black o blackish-b own head capsule,
yellow do sal colo a ion wi h black ans e se s ipes on T1,
and a la e al black spo a he base o each p oleg on A3–6.
Despi e a ia ions in do sal scolus colo a ion ( anging om
black o ligh g ay), hese la ae a e mac oscopically indis-
inguishable. In compa ison, A. cas anea has a ed-b own
head capsule wi h an almos en i ely black do sum T1 and no
black ma kings on he p olegs A3–6. Mo eo e , in L2–3 o An-
he aeopsis, as known in he pas , a con inuous da k g een
s ipe uns along each side (di ided along he do sal and
en al midlines), connec ing he do sal scoli om T2 o A8.
Nässig (1991) iden i ied his as a subgene ic synapomo phy;
howe e , in A. cas anea, i appea s indis inc . Fo L4–5, mo -
phological ea u es ha could iden i y speci ic popula ions
a e no longe disce nible excep o A. cas anea om Tibe ,
which has cha ac e is ic clubbed se ae. Howe e , we cu -
en ly know ha he “yellow-g een la e al pla e o p oleg A10
is inged by a black o da k b own zone (ma u e la a)” is
exhibi ed by all membe s o An he aeopsis—i may ac ually
unc ion as an eyespo playing a ole in de ensi e wa ning.
This ea u e is no p esen in any o he known An he aea
Figu e 7. In asubgene ic phylogeny o he subgenus An he aeopsis and cu en dis ibu ion. The s ipes on he map (a he le mos )
indica e “unce ain anges.” The phylogene ic ee on he le is he CMG esul (Suppl. ma e ial 1: ee S1), and he one on he igh
is based on CMG + COI (Suppl. ma e ial 1: ee S2); he ee scale is only o he CMG esul . The colo s o he map co espond o
he colo s o he axa on he phylogene ic ees.
