149
Ac oss dese and island: Phylogeny o wo hin s ick insec s
om China wi h axonomic insigh s in o he clade “G a idiini II”
(Phasma odea, Cli umninae)
Hao-Ran Gao1, Chen Wang1, Wei-Xian Chen1, Jia-Jin Li1, Hong-Rui Zhang1, Ya-Jin Li1
1 Depa men o En omology, College o Plan P o ec ion, Yunnan Ag icul u al Uni e si y, Kunming, 650201, China
Co esponding au ho : Ya-Jin Li ([email p o ec ed])
Copy igh : © Hao-Ran Gao e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Ex ensi e con e gence led o poo ly unde s ood sys ema ics o s ick insec s (Phas-
ma odea) in he pas . Bo h Scep ophasma bi ube cula um and Macellina souchongia
belong o he ibe G a idiini o he amily Bacillidae, bu hey we e sepa a ed by abou
3500 km and inhabi di e en zoogeog aphic egions. To u he unde s and he Chi-
nese Phasma odea, we sequenced he abo e wo species using nex -gene a ion se-
quencing (NGS) and assembled hei mi ochond ial genomes and s anda d molecula
ma ke s o wo phylogene ic analyses. Mi ochond ial genome (13 PCGs and 2 RNAs)
phylogeny showed ha hese wo species, o iginally belonging o he amily Bacilli-
dae, we e included in he sub amily Cli umninae. Ano he phylogeny including se en
s anda d molecula ma ke s ( ou mi ochond ial genes: COI, COII, 12S, 16S and h ee
nuclea genes: H3, 18S, 28S), co e ing mo e species e ealed ha : S. bi ube cula um
is he sis e g oup o Clona ia spp. om A ica; and genus Macellina o med a complex
clade wi h he O ien al Clona ia and Scep ophasma. Ou esul s co obo a e ha he
amily Bacillidae is no monophyle ic as cu en ly ea ed in he Phasmida Species
File da abase and ha he gene a Clona ia and Scep ophasma, which span di e en
zoogeog aphic egions, a e no monophyle ic and a e in need o u he e ision.
Key wo ds: Cli umninae, Macellina, mi ogenome, nex -gene a ion sequencing, nuclea
genes, Phasmida, phylogeny, Scep ophasma
In oduc ion
Phasma odea, including s ick and lea insec s, ha e long pe plexed axono-
mis s due o hei independen e olu ion o simila mo phological ai s and
beha iou s, which helps hem adap o simila habi a s ac oss di e en egions
(Buckley e al. 2009, 2010; B adle e al. 2015; Glaw e al. 2019; Boisseau e al.
2025). As a esul , molecula phylogeny plays a c ucial ole in unde s anding
hei classi ica ion (B adle and Buckley 2018, 2020).
The amily Cli umnidae wi hin he supe amily Phasma oidea was p oposed
by Cliquennois (2020), o include he sub amilies Cli umninae and Pha naci-
inae. The sub amily Cli umninae comp ises he ibes Cli umnini and Medau ini,
and he clade “G a idiini II” iden i ied h ough molecula biology (B adle e al.
Academic edi o : Ma co Go a do
Recei ed:
29 No embe 2024
Accep ed:
20 Oc obe 2025
Published:
19 No embe 2025
ZooBank: h ps://zoobank.o g/
D742EE1D-93AF-4F82-ABAD-
1186BD87716A
Ci a ion: Gao H-R, Wang C, Chen
W-X, Li J-J, Zhang H-R, Li Y-J (2025)
Ac oss dese and island: Phylogeny
o wo hin s ick insec s om China
wi h axonomic insigh s in o he
clade “G a idiini II” (Phasma odea,
Cli umninae). ZooKeys 1260:
149–160. h ps://doi.o g/10.3897/
zookeys.1260.143038
ZooKeys 1260: 149–160 (2025)
DOI: 10.3897/zookeys.1260.143038
150
ZooKeys 1260: 149–160 (2025), DOI: 10.3897/zookeys.1260.143038
Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
2015). Cu en ly, he majo species online da abase, Phasmida Species File (PSF,
h ps://phasmida.species ile.o g), s ill places he “G a idiini II” clade wi hin he
ibe G a idiini o he amily Bacillidae unde he supe amily Bacilloidea. The s a-
us o o he sub amilies wi hin Bacillidae in he PSF is also unde going e ision
(Bank and B adle 2022; Fo ni e al. 2022): ecen phylogene ic s udies e eal
ha An ongiliinae Zomp o is closely ela ed o Ach iop e ini Gün he and u he
Madagascan axa, which a e now conside ed as Anisacan hoidea (Cliquennois
and B adle , 2022), o ming wi h some A ican G a idiini wha is known as he
“A ican/Malagasy g oup”; Macyniinae Zomp o and Bacillinae B unne on Wa -
enwyl may cons i u e a signi ican componen o Bacillidae.
Ex ensi e mo phological con e gence has led axonomis s o empo a ily
classi y gene a o species o unce ain axonomic s a us in o he ibe, con ib-
u ing signi ican ly o cu en classi ica ion challenges (Cliquennois 2004). The
non-monophyle ic ibe G a idiini Cliquennois, in PSF, is a di e se and complex
g oup wi h 16 gene a and 184 species, making up abou 78% o he species
wi hin he amily (Cliquennois 2005; B ock e al. 2024), dis ibu ed ac oss A -
ica, Eu ope, and Asia. Female G a idiini usually bend hei abdomen o e he
ho ax and head wi h he abdominal ip posi ioned be ween he sho an en-
nae, which hen assis wi h placing he elonga ed adhesi e eggs on a sui able
su ace (B ock and Shlagman 1994; B adle 2009). Molecula phylogeny indi-
ca es his ibe is di ided in o wo lineages: “G a idiini I” p edominan ly ound
in he A ican sa anna and Madagasca (B adle and Buckley 2018; B ock e al.
2024); and “G a idiini II” dis ibu ed o e O ien al egions, and which appea s o
be ela ed o a limi ed numbe o A ican Clona ia spp., wi h nume ous phyloge-
ne ic s udies indica ing a close ela ionship o he Asian Cli umninae B unne
(Robe son e al. 2018; Bank and B adle 2022; Fo ni e al. 2022).
In China, he ibe G a idiini comp ises 5 gene a and 15 species (Wes wood
1859; Chen and He 2008; Hennemann e al. 2008; B ock e al. 2024): Linoce us
G ay (1 sp.), Macellina U a o (4 spp.), Pa agongylopus Chen & He (8 spp.),
Scep ophasma B ock & Seow-Choen (1 sp.), and Zangphasma Chen & He (1
sp.). Recen s udies ha e p ima ily concen a ed on species desc ip ions and
biological aspec s, wi h limi ed esea ch on hei phylogeny (Wu and Fan 2010;
Ho 2014, 2017, 2019; Chen e al. 2015; Gao e al. 2022). No able excep ions
include Scep ophasma bi ube cula um (Red enbache , 1889) om Xinjiang,
China (Cen al Asia), Linoce us g acilis G ay om No h China, and Zangphas-
ma nyingchiense Chen & He om Xizang (Tibe ), which a e dis ibu ed ou side
sou he n egions o China. The Chinese G a idiini was undoub edly puzzling:
he ex eme simila i y o mo phological cha ac e s, he close ela ionship wi h
Cli umninae and he dis ibu ion ac oss zoogeog aphical egions all indica e
he need o a sys ema ic s udy and e ision.
S anda d molecula ma ke s o ma e nally inhe i ed mi ochond ial genomes
do no lead o s ongly suppo ed opology (e.g., Lonchodinae and Nec osci-
inae, which had epea edly been shown o be s ongly ela ed, we e no well
clus e ed in phylogene ic analyses o mi ochond ial genomes) (Kômo o e al.
2011; Robe son e al. 2018; Bank and B adle 2022; Fo ni e al. 2022; Yuan
e al. 2023). Howe e , wi h he educed cos o nex -gene a ion sequencing,
mi ochond ial genome phylogenies ha e succeeded a he genus le el, helping
esea che s quickly cla i y phylogene ic ela ionships wi hin amilies and apid-
ly iden i y species (Li e al. 2022). This s udy in ol ed sequencing wo Chinese
151
ZooKeys 1260: 149–160 (2025), DOI: 10.3897/zookeys.1260.143038
Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
G a idiini species: S. bi ube cula um om U umqi, Xinjiang Uygu Au onomous
Region, and Macellina souchongia om M . Wuzhishan, Hainan Island. Us-
ing nex -gene a ion sequencing (NGS) da a, we assembled he mi ochond ial
genomes and h ee nuclea genes (H3, 18S, 28S) o hese wo species. We
conduc ed wo molecula phylogene ic analyses using: 1) mi ochond ial ge-
nome da a o e i y he gene al classi ica ion o he wo species; and 2) se en
s anda d molecula ma ke s ac oss a b oade ange o species o de e mine
hei phylogene ic posi ion wi hin he amily o ibe. Ou goal is o enhance he
unde s anding o Chinese Phasma odea by s udying hese wo species om
di e en biogeog aphic egions and o o e aluable insigh s o he e ision
o he complex Cli umnidae and G a idiini.
Ma e ial and me hods
Sampling and sequencing
A sample o S. bi ube cula um was collec ed om U umqi, Xinjiang Uygu Au on-
omous Region, China (43°48'46"N, 87°44'6"E), and a sample o M. souchongia
was collec ed om M . Wuzhishan, Hainan Island, China (Fig. 1). All samples
we e p ese ed in 100% e hanol. The igh midleg and igh hindleg o each sam-
ple we e used o genome ex ac ion. We con ac ed No ogene L d. (Beijing,
China) o ex ac DNA using he TIANamp Genomic DNA Ki (TIANGEN, Beijing,
China) and hen conduc ed 150 bp pai ed-end (PE) sequencing on he Illumina
No aseq pla o m, ob aining 6 Gb o da a pe sample. Vouche specimens we e
deposi ed a he College o Plan P o ec ion, Yunnan Ag icul u al Uni e si y.
Assembly and anno a ion
We assembled he il e ed aw da a using No oplas y . 4.3.1 (Die ckxsens
e al. 2017) and Geneious P ime . 2024.0.5 (Kea se e al. 2012) o he mi-
ochond ial genome and hen submi ed he mi ochond ial con ig o he MI-
TOS2 Web se ice (Be n e al. 2013) in he Galaxy pla o m (h ps://usegal-
axy.o g/) (The Galaxy Communi y 2024) o anno a ion. Fo he h ee nuclea
genes (H3, 18S RNA, and 28S RNA), we used Geneious (Kea se e al. 2012)
o map hese genes wi h e e ence sequences (H3: AY125250, 18S: AY121167,
28S: AY125307). All newly sequenced molecula da a had been deposi ed in
GenBank (h ps://www.ncbi.nlm.nih.go /genbank/).
Phylogene ic analysis
To ob ain a ee in e ence ha is close o he ue ela ionship, ou mi ochond i-
al genome phylogeny will use a cons ained ee sea ch by ollowing he esul s
o ansc ip omic s udies [i.e., Phasma odea, Euphasma odea, O iophasma a
and Occidophasma a, Lonchodidae (Lonchodinae + Nec osciinae)] (Simon e
al. 2019; Tihelka e al. 2020). Embiop e a was no conside ed an ou g oup be-
cause long-b anch a ac ion a e ac s simila o hose p e iously published
appea ed in ou es s (Fo ni e al. 2022).
Thi een p o ein-coding genes and wo RNAs om mi ogenomes we e used
o he mi ogenomic phylogeny. Mi ogenome da a o o he species we e ob-
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Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
Figu e 1. A. Female o Scep ophasma bi ube cula um (©Shao-Shan Wang); B. Female o Macellina souchongia; C. Male
o M. souchongia; D. Sampling map showing he loca ion o he abo e wo newly sequenced species.
153
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Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
ained om p e ious s udies (Suppl. ma e ial 1: able S1). The phylogene ic
ma ix o se en molecula ma ke s was de i ed om Bank and B adle (2022).
We selec ed he clades “Bacillinae,” “G a idiidae,” and “Cli umninae”— ep esen -
ing Bacillidae and Cli umninae wi hin he PSF sys em— ha o med a monophy-
le ic b anch in a p e ious phylogene ic s udy (Bank and B adle 2022) o de e -
mine he posi ion o he newly sequenced species (Suppl. ma e ial 1: able S2).
The alignmen o p o ein-coding genes was compiled in Macse . 2.03 (Ran-
wez e al. 2011). The RNAs we e aligned using MAFFT . 7.0 (Ka oh e al. 2002)
wi h G-INS-i. All aligned genes we e immed using imAI (Capella-Gu ié ez
e al. 2009) wi h he au oma ed1 op ion, hen conca ena ed using PhyloSui e
(Zhang e al. 2020). We es ima ed he bes pa i ioning scheme and model o
ou da ase wi h Pa i ionFinde 2 (Lan ea e al. 2017) (Suppl. ma e ial 1: able
S3, S4). The conca ena ed da ase was pa i ioned in o 13 subse s o mi og-
enome phylogeny and 12 subse s o s anda d molecula ma ke s phylogeny;
he bes - i ing model o each pa i ion is shown in Suppl. ma e ial 1: able S3,
S4. Bayesian in e ence (BI) and maximum likelihood (ML) me hods we e used.
The BI analysis was conduc ed using M Bayes . 3.2.6 (Ronquis e al. 2012)
and pe o med wo Ma ko chain Mon e Ca lo (MCMC) uns o 5,000,000 gen-
e a ions wi h sampling e e y 1000 gene a ions. ML analyses we e pe o med
by IQ-TREE2 (Minh e al. 2020) wi h 1000 Ul a as boo s ap eplica es.
Resul s
Basic ea u es o mi ogenomes
The mi ochond ial genome o S. bi ube cula um is 17,663 bp in leng h (Gen-
Bank accession numbe : PQ469774), while ha o M. souchongia is 16,316 bp
(GenBank accession numbe : PQ469775). Bo h mi ochond ial genomes a e in
he same a angemen , con aining 13 p o ein-coding genes, 22 ans e RNAs,
2 RNAs, and a con ol egion (D-loop) (Fig. 2) (Suppl. ma e ial 1: able S5).
Figu e 2. Ci cula maps o mi ogenomes o he wo newly sequenced species, S. bi ube cula um; A. M. souchongia; B.
The J-s and is isualized on he ou e ci cle and he N-s and on he inne ci cle.
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Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
Phylogene ic analysis
In he mi ogenome phylogene ic analysis, a cons ained ee sea ch p o-
duced a opology ha mo e closely aligned wi h he ansc ip ome esul s
(Simon e al. 2019; Tihelka e al. 2020) (Fig. 3). The newly sequenced spe-
cies o med a s able clade and we e iden i ied as a sis e g oup o Cli umni-
ni (which includes Ramulus Saussu e and En o ia S ål). This clade was
well-suppo ed wi hin Cli umninae. Ou mi ogenome phylogene ic esul s
ea i med ha Eas Asian G a idiini membe s we e included in Cli umninae.
No ably, S. bi ube cula um, despi e i s geog aphical p oximi y o Eu ope,
does no belong o he Eu opean-Bacillidae clade.
The phylogene ic analysis using se en s anda d molecula ma ke s,
which included a b oade ange o species, yielded esul s consis en wi h
hose o Bank and B adle (2022) (Fig. 4). Th ee well-suppo ed clades we e
iden i ied: he “Eu opean-Bacillidae clade,” including Bacillus Be hold, Clo-
nopsis Pan el, Lep ynia Pan el, and Pijnacke ia Scali; he “A ican-G a idiini
clade,” comp ising Macynia S ål, Phalces S ål, and Zehn ne ia B unne on
Wa enwyl; and he complex “Cli umninae clade.” The wo newly sequenced
species we e placed wi hin one o he clades o he “Cli umninae clade,”
which was dis inc om Cli umnini and Medau ini. This clade o med a
well-suppo ed b anch ha spans mul iple biogeog aphic egions, including
Clona ia om he A ican sa annah and Scep ophasma and Macellina om
Eas e n Asia. Scep ophasma bi ube cula um was shown o be s ongly e-
la ed o species o he genus Clona ia om he A ican sa anna, sugges ing
Figu e 3. Consensus phylogeny ee based on he da ase o mi ogenomes (13PCGs + 2 RNA), econs uc ed using ML
and BI. Nume als a nodes a e he Bayesian pos e io p obabili ies and ML boo s ap alues, espec i ely. Suppo o
cons ained nodes is omi ed.
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Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
ha i may no ha e o igina ed in he clades o he majo Asian Cli umninae.
The posi ion o M. souchongia was o be expec ed; i o med a s able b anch
wi h o he hin s ick insec s om he O ien al egion and ma ched well wi h
i s ela i e species.
Discussion
Mi ogenomic phylogeny in Phasma odea
Al hough he mi ochond ial genome p o ides mo e gene ic in o ma ion han
a single DNA ba code o sequence agmen , cu en di ec phylogene ic anal-
yses using mi ochond ial genomes do no accu a ely econs uc he s able
opology o Phasma odea. The mi ochond ial genome has been e ec i ely
used o explo e phylogene ic ela ionships wi hin speci ic amilies, ibes, o
gene a, o e ing aluable insigh s in o a ious complex axon g oups (Li e
al. 2022). In conclusion, using mi ochond ial genome phylogene ic in e ence
wi h a opological-cons ain s ee p o ides a apid me hod o classi ying
species wi h unce ain axonomic posi ions h ough Illumina sequencing.
This app oach also enables u he analyses, including ances y econs uc-
ion and di e gence ime es ima ion. Thus, mi ochond ial genome phyloge-
ne ic in e ence using high-suppo ee opological cons ain s, despi e some
limi a ions, emains an e ec i e me hod o apid species classi ica ion
h ough Illumina sequencing and o e s po en ial o u he analyses, such
as ances y econs uc ion and di e gence ime es ima ion.
Figu e 4. ML ee based on he da ase o se en s anda d molecula ma ke s (18S, 28S, H3, COI, COII, 12S and 16S).
Nume als a nodes a e he ML boo s ap alues.
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Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
Ad ances in nex -gene a ion sequencing echnologies, along wi h hei
dec easing cos s, ha e made mi ochond ial genomes a apid, a o dable
and accessible way o phylogene ic s udies. Howe e , a common issue in
many s udies was he po en ial misiden i ica ion o species o axa wi h
uzzy iden i ica ion. Fo ins ance, an ea lie s udy misiden i ied a winged
Lanceoce ca a species as Phobae icus se a ipes (GenBank accession:
AB477467) (Kômo o e al. 2011). Addi ionally, some in alid species names
ha e been encoun e ed, making i di icul o e i y he accu a e species in-
o ma ion (Zhang and Guo 2022). We s ongly ecommend ha u u e s ud-
ies include comp ehensi e e idence o he iden i ica ion o Phasma odea,
such as pho og aphs and de ailed specimen in o ma ion.
Taxonomic s a us o wo newly sequenced species
B ock and Seow-Choen e ec ed he genus Scep ophasma o di e en ia e
species om Sou heas and Sou h Asia om hose o he genus G a id-
ia ound in A ica and he Middle Eas (Seow-Choen 2000). Subsequen ly,
Hennemann e al. (2008) mo ed S. bi ube cula um om he genus G a id-
ia o Scep ophasma. Scep ophasma bi ube cula um signi ican ly di e s
om o he species in he same genus in eggs: he ope culum lacks dis inc
aised la e al im- o ming ee h (Wes wood 1859; Seow-Choen 2021, 2023).
This ans e is deba able based on ou new phylogene ic s udies, as molec-
ula e idence indica es a close ela ionship o Clona ia o G a idia species
ound in A ica and he Middle Eas . Un o una ely, in he absence o addi-
ional specimens and molecula samples, ou s udy could no go u he o
explain he ela ionship o S. bi ube cula um wi h i s ue “ ela i es” and o
ob ain a mo e highly suppo ed ee opology.
Acknowledgemen s
We wish o hank Shao-Shan Wang (Shihezi, Xinjiang) o p o iding pho os. We ex-
p ess ou g a i ude o he edi o Ma co Go a do (I ay), and e iewe S en B adle
(Ge many) o hei e o s and sugges ions in enhancing his manusc ip .
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
This esea ch is suppo ed by Yunnan Ag icul u al Uni e si y 17 h S uden Science and
Technology Inno a ion and En ep eneu ship Ac ion Funds (No. 2024N031) and Scien i ic
Resea ch Funds P ojec o Yunnan P o incial Depa men o Educa ion (No. 2024Y269).
157
ZooKeys 1260: 149–160 (2025), DOI: 10.3897/zookeys.1260.143038
Hao-Ran Gao e al.: Mi ogenomic phylogeny o wo hin s ick insec s om China
Au ho con ibu ions
Concep ualiza ion: HRG. Da a cu a ion: HRG. Fo mal analysis: HRG. Funding acquisi-
ion: HRG. In es iga ion: CW, WXC. Me hodology: CW. P ojec adminis a ion: HRZ.
Resou ces: WXC. So wa e: JJL. Supe ision: HRZ, YJL. Valida ion: YJL. Visualiza ion:
HRG. W i ing – o iginal d a : HRG. W i ing – e iew and edi ing: YJL, HRZ.
Au ho ORCIDs
Hao-Ran Gao h ps://o cid.o g/0000-0001-5093-0474
Chen Wang h ps://o cid.o g/0000-0002-2711-2358
Wei-Xian Chen h ps://o cid.o g/0009-0003-0595-0523
Jia-Jin Li h ps://o cid.o g/0000-0002-9914-464X
Hong-Rui Zhang h ps://o cid.o g/0000-0002-0089-1099
Ya-Jin Li h ps://o cid.o g/0000-0003-3815-3021
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex o
Supplemen a y In o ma ion.
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