355
Fi s eco d o he sub amily Ge acinae B indle, 1971 and genus
Pseudo os ox Bo elli, 1926 (De map e a, Spongipho idae) in
China, wi h a desc ip ion o a new species
Zhi-Teng Chen1, Chao Jiang2
1 School o G ain Science and Technology, Jiangsu Uni e si y o Science and Technology, Zhenjiang 212004, Jiangsu P o ince, China
2 S a e Key Labo a o y o Quali y Ensu ance and Sus ainable Use o Dao-di He bs, Na ional Resou ce Cen e o Chinese Ma e ia Medica, China Academy o Chinese
Medical Sciences, Beijing 100700, China
Co esponding au ho : Chao Jiang ([email p o ec ed])
Copy igh : © Zhi-Teng Chen & Chao Jiang.
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
The sub amily Ge acinae B indle, 1971, and he genus Pseudo os ox Bo elli, 1926, a e
eco ded o he i s ime om China. These new eco ds we e con i med by collec ions
om Guiyang Ci y in Guizhou P o ince, loca ed in sou hwes e n China. Pseudo os ox
guizhouensis sp. no . is desc ibed and illus a ed based on specimens om his e-
gion. Al hough he new species closely esembles he O ien al species Pseudo os ox
ascia us (de Bo mans, 1894), i can be dis inguished by a combina ion o unique mo -
phological cha ac e s. An upda ed key and dis ibu ion map o Pseudo os ox species
a e p o ided o aid u u e esea ch and species iden i ica ion.
Key wo ds: Biodi e si y, ea wig, Ge acinae, iden i ica ion key, mo phology, new eco d,
axonomy
In oduc ion
The amily Spongipho idae is a di e se and ecologically impo an g oup o ea -
wigs (De map e a) mainly ound in opical and sub opical egions. Cu en ly,
he amily is classi ied in o 13 sub amilies, comp ising 42 gene a and 527 spe-
cies (Hopkins e al. 2025). Among hese, he sub amily Ge acinae B indle, 1971
is ela i ely di e se, con aining six gene a and 29 species dis ibu ed ac oss
Sou h Asia and Eas A ica (Hopkins e al. 2025). Species o Ge acinae a e gen-
e ally small o e y small (body leng h sho e han 10 mm) and possess dis-
inc i e mo phological ai s such as a sal a olia and o ceps o simila shape
in bo h males and emales (B indle 1971).
Pseudo os ox Bo elli, 1926 is he mos species- ich genus wi hin Ge acinae,
con aining 13 species ound in bo h he O ien al and A ican egions (S einmann
1990; S i as a a 2003). Species wi hin his genus exhibi conside able mo pho-
logical di e si y, wi h a ia ions in body colo a ion, penul ima e s e ni e s uc-
u e, and geni alia shape, all o which a e essen ial o species iden i ica ion and
axonomic classi ica ion. Despi e he di e si y, he dis ibu ion o Pseudo os ox
has been unde explo ed, pa icula ly in Eas and Sou heas Asia.
Academic edi o : Fabian Haas
Recei ed:
18 June 2025
Accep ed:
8 Sep embe 2025
Published:
8 Oc obe 2025
ZooBank: h ps://zoobank.
o g/6F3423E4-72E6-4D3D-825C-
896FBC9F6373
Ci a ion: Chen Z-T, Jiang C (2025)
Fi s eco d o he sub amily
Ge acinae B indle, 1971 and
genus Pseudo os ox Bo elli, 1926
(De map e a, Spongipho idae)
in China, wi h a desc ip ion o
a new species. ZooKeys 1254:
355–366. h ps://doi.o g/10.3897/
zookeys.1254.162412
ZooKeys 1254: 355–366 (2025)
DOI: 10.3897/zookeys.1254.162412
356
ZooKeys 1254: 355–366 (2025), DOI: 10.3897/zookeys.1254.162412
Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
P io o his s udy, nei he membe s o Ge acinae no Pseudo os ox had
been epo ed om China. This gap in he axonomic eco d p omp ed an
in es iga ion in o he De map e a auna o Guiyang Ci y, loca ed in Guizhou
P o ince, sou hwes e n China. The egion is cha ac e ized by i s di e se eco-
logical condi ions, which likely suppo a a ie y o ea wig species. In June
o bo h 2024 and 2025, a se ies o ea wig specimens we e collec ed om
Guiyang Ci y. Ini ial mo phological assessmen s indica ed signi ican simila -
i ies wi h Pseudo os ox ascia us (de Bo mans, 1894), a species p e iously
eco ded om Myanma , Thailand, and Vie nam (S einmann 1990; S i as-
a a 2003). Howe e , u he examina ion o he specimens e ealed se e al
key mo phological di e ences, sugges ing he p esence o an undesc ibed
species. In his pape , we desc ibe and illus a e his new species, Pseudo-
os ox guizhouensis sp. no ., which ep esen s he i s o mal eco d o bo h
Ge acinae and Pseudo os ox in China. An upda ed key o he iden i ica ion
o Pseudo os ox species is also p o ided o suppo u u e esea ch on he
genus and i s dis ibu ion.
Ma e ial and me hods
The specimens we e collec ed om di e en a eas o Guiyang Ci y, Guizhou
P o ince, China in June o bo h 2024 and 2025 and p ese ed in 75% alcohol.
The ype specimens o his new species a e deposi ed in he Insec Collec ion
o Jiangsu Uni e si y o Science and Technology (ICJUST), Jiangsu P o ince,
China. Mo phological examina ions and iden i ica ions we e conduc ed using
an SDPTOP SZM45 s e eomic oscope. Images we e cap u ed wi h a Canon
EOS 5DSR came a wi h a Canon MP-E 2.8/65 mm mac o lens. Pla es we e p e-
pa ed using he so wa e Adobe Pho oshop 2024. The mo phological e minol-
ogy ollows S einmann (1990).
Resul s
Pseudo os ox guizhouensis sp. no .
h ps://zoobank.o g/97DAC96D-D9BE-4083-BFD7-EFD0AD4CBDEF
Figs 1–6
Type ma e ial. Holo ype: China • ♂; Guizhou P o ince, Guiyang Ci y, Xiangzhigou;
26.7775°N, 106.9346°E; 1,151 m; 17. i.2024; Zhi-Teng Chen leg. Pa a ypes:
China • 7♂♂16♀♀; same da a as holo ype; 7♂♂2♀♀, Guizhou P o ince,
Guiyang Ci y, He ang; 26.7270°N, 106.8766°E; 1,010 m; 18. i.2024; Zhi-Teng
Chen leg. • 1♂, Guizhou P o ince, Guiyang Ci y, Xiaba Town, nea Baishui Ri e ;
26.7254°N, 106.9313°E; 970 m; 18. i.2024; Zhi-Teng Chen leg; 11♂♂15♀♀,
Guizhou P o ince, Guiyang Ci y, Xiangzhigou; 26.7768°N, 106.9347°E; 1,130 m;
26. i.2025; Zhi-Teng Chen, Qing Li leg.
Di e en ial diagnosis. The new species sha es a simila pale ans e se
s ipe on he egmina wi h P. ascia us (S einmann 1990; S i as a a 2003).
Howe e , i can be dis inguished by se e al mo phological ai s (S einmann
1990; S i as a a 2003): he male head is longe han wide ( s. wide han
long in P. ascia us), he p ono um is long wi h a ounded an e io ma gin
( s. ans e se wi h a unca e an e io ma gin), and he pale s ipe on he
357
ZooKeys 1254: 355–366 (2025), DOI: 10.3897/zookeys.1254.162412
Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
egmina is nea one- ou h o i s leng h ( s. nea one-eigh h in P. ascia us).
Addi ionally, he male’s penul ima e s e ni e o he new species has a mos ly
unca e pos e io ma gin ( s. conca e in P. ascia us), and he ex e nal
pa ame e is widened subapically and cons ic ed apically wi h an ob use
apex ( s. g adually ape ing o an acu e apex in P. ascia us). The emale
o he new species also di e s om ha o P. ascia us in ha ing pos e io
p ocesses on he ul ima e e gi e and s aigh inne o ceps ma gins, whe e-
as he emale o P. ascia us lacks such p ocesses, and he inne ma gin o
Figu e 1. Pseudo os ox guizhouensis sp. no ., male holo ype. A. Habi us, do sal iew; B. Head and p ono um, do sal iew;
C. Head and ho ax, en al iew. Scale ba s: 1 mm (A); 0.5 mm (B, C).
358
ZooKeys 1254: 355–366 (2025), DOI: 10.3897/zookeys.1254.162412
Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
he o ceps is sinua e (B indle 1973a; S einmann 1990). An upda ed key o
species o Pseudo os ox based on S einmann (1990) is p o ided below o
species delimi a ion.
Desc ip ion. Male. Leng h. Body leng h (excluding o ceps) 4.5–5 mm, o -
ceps leng h 0.5–0.8 mm.
Colo a ion. Head and p ono um da k b own, la e wi h whi e pos e ola e -
al sides (Fig. 1A–C); an ennae yellow a base, pale b own apically. Tegmina
da k b own, wi h a pale ans e se s ipe nea shoulde , he s ipe g adually
na owed mediad (Fig. 1A). Scales o hind wings en i ely whi e. Abdomen da k
b own; legs and o ceps b own (Fig. 1A).
Head. Head longe han wide, umid; pos on al and co onal su u es indis-
inc ; hind ma gin unca e (Fig. 1A–C). Eyes sligh ly sho e han pos -ocula
a ea. An enna 12-segmen ed; basal segmen abou hal he leng h o dis ance
be ween an ennal bases; second segmen as long as wide; hi d segmen h ee
Figu e 2. Pseudo os ox guizhouensis sp. no ., male holo ype. A. Abdomen, do sal iew; B. Abdomen, en al iew;
C. Penul ima e s e ni e, en al iew. Scale ba s: 0.1 mm.
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ZooKeys 1254: 355–366 (2025), DOI: 10.3897/zookeys.1254.162412
Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
imes as long as wide; ou h segmen wo imes as long as wide; i h and sub-
sequen segmen s sligh ly longe han hi d.
Tho ax. P ono um as b oad as long, sligh ly widened pos e io ly; an e io
ma gin sligh ly ounded, la e al and pos e io ma gins ounded; p ozona con-
ex and me azona dep essed; median sulcus dis inc (Fig. 1A–C). Tegmina
well-de eloped, wice as long as p ono um, pubescen and impunc a e; scales
o hind wings nea as long as one- hi d leng h o p ono um (Fig. 1A). Legs
wi h s ong emo a, hin ibiae and a si, and de eloped a sal a olia be ween
claws; hind a si wi h i s segmen 1.5 imes longe han combined leng h o
segmen s 2–3, second segmen nea as long as wide, hi d segmen wice as
long as second (Fig. 1A).
Abdomen. Abdomen usi o m, con ex, wides a segmen s 6–7, cu icle pu-
bescen and impunc a e (Fig. 1A). Ul ima e e gi e ans e se, sloping and
na owed pos e iad; pos e io ma gin conca e medially, wi h sub iangula
p ocesses abo e base o each b anch o o ceps (Fig. 2A). Fo ceps sho ,
con iguous, pubescen , mos ly s aigh , upcu ed, apically ape ing and gen-
ly hooked mediad; inne ma gins den icula e (Fig. 2A, B). Penul ima e s e -
ni e ans e se, 1.5 imes wide han long, pos e io ma gin mos ly unca e;
basola e al p ojec ions sho , nea one- i h o s e nal leng h (Fig. 2C).
Geni alia. Geni alia slende (Fig. 3A–D); pa ame e na owed basally, wid-
ened medially, wi h ounded an e io ma gin; ex e nal pa ame e slende ,
Figu e 3. Pseudo os ox guizhouensis sp. no ., male holo ype. A. Geni alia unde e lec ed ligh , do sal iew; B. Geni alia
unde ansmi ed ligh , do sal iew; C. Geni alia unde ansmi ed ligh , en al iew; D. Geni alia unde e lec ed ligh ,
en al iew. Abb e ia ions: ep – ex e nal pa ame e; pm – pa ame e; – i ga. Scale ba s: 0.5 mm.
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Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
widened subapically, apex na owed, ob use; i ga hick; inne s uc u e
complica ed, wi h wo semici cula basal scle i es and a oo hed apical
scle i e. Geni alia o male pa a ype sligh ly a ied in shape o ex e nal
pa ame al apex and inne scle i e (Fig. 4A, B).
Figu e 4. Pseudo os ox guizhouensis sp. no ., male pa a ype. A. Geni alia unde e lec ed ligh , do sal iew; B. Geni alia
unde e lec ed ligh , en al iew. Abb e ia ions: ep – ex e nal pa ame e; pm – pa ame e; – i ga. Scale ba s: 0.5 mm.
361
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Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
Female. Body leng h wi hou o ceps 4.5–6 mm, leng h o o ceps
0.5–0.8 mm. Body s uc u es and colo a ion iden ical o male holo ype; e g-
i es and s e ni es longe han male (Fig. 5A, B). Penul ima e s e ni e subquad-
a e, pos e io ma gin ounded; basola e al p ojec ions e y sho , abou 1/11
o s e nal leng h (Fig. 5C). Geni alia wi h wo da k, o al scle i es (Fig. 5B).
E ymology. The new species is named a e Guizhou P o ince, he ype locali y.
Dis ibu ion. The species is cu en ly known only om Guizhou P o ince o
sou hwes China (Fig. 6).
Biology. Specimens we e collec ed om ipa ian ege a ion using a sweeping
ne (Fig. 6). This species exhibi ed no speci ic ege a ion p e e ence and was
collec ed om lea es o bamboo, g amineous weeds, ea ees, ege ables and
a ious o he plan s. I occu ed in high densi ies nea he wa e ’s edge. When
dis u bed, indi iduals displayed a ypical de ensi e pos u e by aising hei o -
ceps, a beha iou simila o ha o o he ea wig species (Fig. 7A, B). In addi ion,
hey exhibi ed leg-cleaning beha io (Fig. 7C, D). Du ing a second collec ing ip
o Guizhou P o ince in 2025, specimens we e encoun e ed unde ainy condi-
ions. Indi iduals immedia ely became immobile when hey con ac ed he wa e
d ople s on he inne su ace o he collec ing bo le. A e se e al hou s, once he
wa e had e apo a ed, he specimens became e y ac i e again. This esponse
sugges s ha he new species may exhibi hana osis-like beha io .
Figu e 5. Pseudo os ox guizhouensis sp. no ., emale pa a ype A. Abdomen, do sal iew; B. Abdomen, en al iew;
C. Penul ima e s e ni e, en al iew. Scale ba s: 0.1 mm.
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Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
Upda ed key o species o Pseudo os ox based on males
1 O ien al dis ibu ion .......................................................................................2
– A ican dis ibu ion ........................................................................................7
2 Eyes sho e han leng h o head behind eyes ............................................3
– Eyes longe han leng h o head behind eye ................................................4
3 Head and p ono um longe han wide; pale s ipe o egmina nea one-
ou h o egmina leng h .........................................P. guizhouensis sp. no .
– Head and p ono um wide han long; pale s ipe o egmina nea 1/8 o
egmina leng h ...........................................P. ascia us (de Bo mans, 1894)
4 Tegmina and hind wings unicolo ous ...................... P. bicolo Bo elli, 1926
– Tegmina wi h con as ing colo s ..................................................................5
5 Hind wings absen ................................................ P. my mecus (Bu , 1908)
– Hind wings p esen ........................................................................................6
6 P ono um wi h yellow la e al and pos e io ma gins; egmina wi h yellow
ans e se s ipe nea shoulde and pos e io ma gin ..................................
............................................................................ P. b indlei S i as a a, 2003
– P ono um b oadly yellow pos e io ly; egmina wi h an an e omedial yellow
pa ch .............................................................. P. la o ascialis B indle, 1973
7 Tegmina en i ely o basally yellow ...............................................................8
– Tegmina unicolo ous, da k b own o blackish ...........................................10
Figu e 6. Gene al dis ibu ion o Pseudo os ox species. Inse : collec ing en i onmen in Guizhou P o ince, China.
Di e en colo and symbols ep esen di e en a eas: P. a icanus (Angola, Nige ia, Congo, Sudan, Tanzania), P. a um
(Congo, Gabon, Ghana, Mozambique), P. bicolo (Bo neo, Philippine Islands, Vie nam), P. bispinosus (Uganda), P. b indlei
(Vie nam), P. ascia us (Myanma , Thailand, Vie nam), P. la o ascialis (Suma a), P. guizhouensis sp. no . (Guizhou
P o ince o China), P. hincksi (Zimbabwe), P. mame i (Mau i ius), P. my mecus (Ja a), P. udebecki (Angola, Congo,
Gabon), P. unca us (Kenya), P. unicolo (Congo).
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Zhi-Teng Chen & Chao Jiang: Reco d o Pseudo os ox in China
8 P ono um, egmina, and hind wings unicolou ous, yellowish-b own ...........
.................................................................................P. unicolo B indle, 1970
– P ono um o egmina a leas pa ially da kened ........................................9
9 Hind wings blackish; male penul ima e s e ni e wi h a small iangula
pos e io excision .....................................................P. hincksi B indle, 1970
– Hind wings yellow; male penul ima e s e ni e wi h b oadly conca e pos e-
io ma gin.......................................................... P. a icanus (B indle, 1968)
10 Male penul ima e s e ni e wi h almos unca e pos e io ma gin ..............
...............................................................................P. unca us B indle, 1970
– Male penul ima e s e ni e wi h pos e io ma gin no unca e .................11
11 Male penul ima e s e ni e wi h pos e io ma gin p oduced wi h wo dis al
spines ................................................................. P. bispinosus B indle, 1970
– Male penul ima e s e ni e wi hou pos e io spines .................................12
12 Hind wings absen .................................................P. mame i (Hincks, 1950)
– Hind wings p esen ......................................................................................13
13 P ono um as long as wide; male penul ima e s e ni e wi h a na ow pos e-
io excision .........................................................P. udebecki B indle, 1969
– P ono um longe han wide; male penul ima e s e ni e wi h a wide pos e-
io excision ............................................................. P. a um Menozzi, 1935
Discussion
The species o Pseudo os ox a e p ima ily dis inguished by a combina ion o
mo phological ea u es, including eye leng h, head and p ono um shape, colo
pa e n on p ono um and egmina, hind wing con igu a ion, penul ima e s e n-
i e shape, and geni alia s uc u e (B indle 1973b; S einmann 1990; S i as a a
2003). Species o his genus ha e adi ionally been di ided in o wo dis inc
geog aphical g oups: he O ien al g oup and he A ican g oup (S einmann
1990). The O ien al g oup includes species om Sou heas Asia, while he A -
ican g oup comp ises species ound h oughou he A ican con inen (Fig. 6).
Al hough he e is some o e lap in species dis ibu ions wi hin each g oup,
he dis inc ion emains use ul o unde s anding hei b oade biogeog aphy.
S einmann (1990) placed Pseudo os ox mame i (Hincks, 1950) om Mau i ius
in he O ien al g oup; howe e , i is mo e sui able o pu i in he A ican g oup.
This highligh s he complexi y o species bounda ies wi hin Pseudo os ox and
he need o con inued axonomic e isions as new specimens and dis ibu ion
da a become a ailable.
The disco e y o Pseudo os ox in Guizhou P o ince signi ican ly ex ends he
known dis ibu ion o he genus in o sou hwes e n China, a egion ha has e-
mained la gely unexplo ed in e ms o i s De map e a auna. The mo phologi-
cal di e ences be ween P. guizhouensis sp. no . and i s closely ela ed species,
pa icula ly in he head, p ono um, penul ima e s e ni e, and geni alia, a e con-
side able. These di e ences no only jus i y he desc ip ion o a new species
bu also highligh he mo phological di e si y wi hin Pseudo os ox. This new
species adds o ou unde s anding o how Pseudo os ox can adap o di e se
ecological condi ions ac oss i s dis ibu ion. Fu he mo e, his disco e y un-
de sco es he impo ance o explo ing unde s udied egions, such as sou h-
wes e n China, which a e likely o yield addi ional new species, en iching ou
knowledge o Pseudo os ox and he sub amily Ge acinae.
