111
A new species o Pseudopoda (A aneae, Spa assidae) om
China, wi h i s desc ip ion o he emale o P. spi alis Zhang,
Jäge & Liu, 2023
He Zhang1,2,3,4* , Chang-Hao Hu4,5,6* , Lu-Yu Wang7, Jie Liu4,5 , Yang Zhong1
1 School o Nuclea Technology and Chemis y & Biology, Hubei Uni e si y o Science and Technology, Xianning, Hubei, China
2 College o Physics and Elec onic Enginee ing, Xing ai Uni e si y, Xing ai, Hebei, China
3 Hebei P o incial Swee Po a o B eeding and Applica ion Technology Inno a ion Cen e , Xing ai, Hebei, China
4 A achnid Resou ce Cen e o Hubei P o ince & Hubei Key Labo a o y o Regional De elopmen and En i onmen al Response, Facul y o Resou ces and En i onmen al
Science, Hubei Uni e si y, Wuhan, Hubei, China
5 Cen e o Beha io al Ecology and E olu ion, School o Li e Sciences, Hubei Uni e si y, Wuhan, Hubei, China
6 College o Li e Sciences, Hunan No mal Uni e si y, Changsha, Hunan, China
7 Key Labo a o y o Eco-en i onmen s in Th ee Go ges Rese oi Region (Minis y o Educa ion), School o Li e Sciences, Sou hwes Uni e si y, Chongqing,
China
Co esponding au ho : Yang Zhong ([email p o ec ed])
Copy igh : © He Zhang 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
One new species o Pseudopoda Jäge , 2000 is desc ibed om Guangxi, sou he n China: P.
ongxianensis sp. no . Addi ional ma e ials o P. de o mis Gong & Zhong, 2023 and P. spi alis
Zhang, Jäge & Liu, 2023 a e also p esen ed, wi h he emale o P. spi alis desc ibed o he
i s ime. Figu es and dis ibu ion da a o hese h ee species a e p o ided.
Key wo ds: Biodi e si y, He e opodinae, hun sman spide s, mo phology, axonomy
In oduc ion
The genus Pseudopoda Jäge , 2000 is cu en ly he mos species- ich genus
wi hin he amily Spa assidae Be kau, 1872, comp ising 270 alid species om
Eas , Sou h, and Sou heas Asia (Wo ld Spide Ca alog 2025). These spide s
a e ypically ound in o es ed and mon ane ecosys ems, o en inhabi ing lea
li e , ege a ion and unde ocks in humid en i onmen s (Jäge 2015; Zhang e
al. 2023). Ac oss i s dis ibu ion, China ep esen s he mos signi ican cen e
o di e si y o he genus, wi h o e 160 species eco ded o da e. This high
numbe o species, along wi h he equen disco e y o new axa, sugges s
conside able endemism and an unde es ima ion o he ue species ichness
(Zhang e al. 2023).
Despi e ongoing axonomic e o s, he classi ica ion and unde s anding o
Pseudopoda emain incomple e. O he known species, only 68 ha e been as-
signed o de ined species g oups based on mo phological o in eg a i e e i-
dence, including he daliensis-g oup, di e sipunc a a-g oup, in e posi a-g oup,
la embola-g oup, ma ensi-g oup, pa ipunc a a-g oup, p omp a-g oup,
Academic edi o : Miquel A. A nedo
Recei ed:
13 May 2025
Accep ed:
23 Sep embe 2025
Published:
18 No embe 2025
ZooBank: h ps://zoobank.o g/
BEB16EAA-5636-46C6-BCC8-
8700D596D37F
Ci a ion: Zhang H, Hu C-H, Wang L-Y,
Liu J, Zhong Y (2025) A new species
o Pseudopoda (A aneae, Spa assidae)
om China, wi h i s desc ip ion
o he emale o P. spi alis Zhang,
Jäge & Liu, 2023. ZooKeys 1260:
111–122. h ps://doi.o g/10.3897/
zookeys.1260.158762
ZooKeys 1260: 111–122 (2025)
DOI: 10.3897/zookeys.1260.158762
* These au ho s con ibu ed equally o his wo k.
112
ZooKeys 1260: 111–122 (2025), DOI: 10.3897/zookeys.1260.158762
He Zhang e al.: Pseudopoda species om Sou h China
schwendinge i-g oup, and signa a-g oup (Jäge 2001; Zhang e al. 2017, 2019;
Li e al. 2019; Zhang e al. 2025). The emaining 202 species a e s ill un-
g ouped, highligh ing he complexi y and un esol ed ela ionships wi hin he
genus. Mo eo e , 138 species a e cu en ly known only om a single sex, which
poses ongoing challenges o accu a e species iden i ica ion and compa a i e
mo phological analyses (Wu e al. 2024).
In his s udy, we desc ibe one new species o Pseudopoda collec ed om
Guangxi Zhuang Au onomous Region, China. We also p esen new mo pho-
logical in o ma ion and a dis ibu ional eco d o Pseudopoda de o mis Gong
& Zhong, 2023 (Gong e al. 2023), based on a emale specimen om Chongq-
ing Municipali y, which documen s no able in aspeci ic a ia ion in he in e -
nal duc sys em. In addi ion, we desc ibe he emale o Pseudopoda spi alis
Zhang, Jäge & Liu, 2023 (Zhang e al. 2023) o he i s ime. These indings
con ibu e o a mo e comple e unde s anding o he genus’s di e si y and
dis ibu ion in China.
Ma e ial and me hods
All specimens we e collec ed by hand and examined using an Olympus SZX16
s e eomic oscope; u he de ails we e in es iga ed using an Olympus BX51
compound mic oscope. Colou a ion is desc ibed in all species om specimens
in e hanol. Copula o y o gans we e examined and illus a ed a e dissec ion
om he spide bodies; epigynes we e clea ed wi h P o einase K. Habi us pho-
os we e ob ained using a Leica M205C digi al mic oscope a ached wi h a
Leica DMC4500 digi al came a.
Leg measu emen s a e shown as: o al leng h ( emu , pa ella, ibia, me a-
a sus, a sus). The numbe o spines is lis ed o each segmen in he ol-
lowing o de : p ola e al, do sal, e ola e al, en al (in emo a and pa ellae
en al spines a e absen , and he ou h digi is omi ed in he spina ion
o mula). The e minology used in ex and igu e legends ollows Jäge
(2000). All measu emen s a e gi en in millime es. The classes o soma ic
size ollow he c i e ia o Jäge (2001). A ising poin s o he egula append-
ages o male palps a e gi en acco ding o clock posi ions o he le palp in
en al iew (Rheims 2013). The map was p oduced using A cMap e sion
10.8.1.
Abb e ia ions used in ex and igu es: ALE, an e io la e al eyes; AME,
an e io median eyes; C, conduc o ; CH, clypeus heigh ; CO, copula o y
opening; CP, cymbial p ocess; dRTA, do sal b anch o RTA; DS, do sal shield
o p osoma; E, embolus; EP, embolic p ojec ion; FD, e iliza ion duc ; Fe,
emu ; FW, i s winding; IDS, in e nal duc sys em; LL, la e al lobes; M ,
me a a sus; OS, opis hosoma; Pa, pa ella; PLE, pos e io la e al eyes; PME,
pos e io median eyes; Pp, palp; RTA, e ola e al ibial apophysis; Sp, spe -
mopho ; S , sub egulum; T, egulum; Ti, ibia; RTA, en al b anch o RTA;
VTA, en al ibial apophysis; I, II, III, IV, legs I o IV.
The s udied specimens ha e been deposi ed in he Cen e o Beha io al
Ecology and E olu ion, College o Li e Sciences, Hubei Uni e si y, Wuhan, Chi-
na (CBEE), and he School o Nuclea Technology and Chemis y and Biology,
Hubei Uni e si y o Science and Technology, Xianning, China (HUST).
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He Zhang e al.: Pseudopoda species om Sou h China
Taxonomy
Family Spa assidae Be kau, 1872
Sub amily He e opodinae Tho ell, 1873
Genus Pseudopoda Jäge , 2000
Type species. Pseudopoda p omp a (O. Picka d-Camb idge, 1885).
Diagnosis. See Jäge (2000).
Dis ibu ion. Eas , Sou h and Sou heas Asia.
Pseudopoda ongxianensis sp. no .
h ps://zoobank.o g/C55D09FD-D7BA-41F8-B3AF-60ACEF0E7027
Figs 1, 2, 8
Type ma e ial. Holo ype male: China • Guangxi Zhuang Au onomous Region:
Yulin Ci y, Rong Coun y, Duqiaoshan Fo es Pa k, 22.8216°N, 110.6429°E,
187 m, 5 Oc obe 2023, Qian-Le Lu leg. (CBEE, GXRX-23-1001).
E ymology. The speci ic name is de i ed om he ype locali y, Rong Coun y.
Diagnosis. Male o P. ongxianensis sp. no . is simila o ha o P. s ombu-
li o mis Zhang, Jäge & Liu, 2023 (c . Fig. 1 and Zhang e al. 2023: ig. 231) in
ha ing la ge al eolus, and an ex emely elonga ed and ili o m embolus, bu
can be dis inguished om P. s ombuli o mis by: 1) RTA longe han he con-
nec ing pa be ween RTA and dRTA; 2) VTA hinne , almos 1/4 o he wid h o
ibia; 3) p ola e al pa o egulum wi hou a dep ession; 4) embolic p ojec ion
long, almos as long as he diame e o he i s loop o embolus; and 5) em-
bolus long, o ming 3 loops ( s. RTA sho e han he connec ing pa be ween
RTA and dRTA, VTA almos 1/3 o he wid h o ibia, p ola e al egulum wi h
dep ession, embolic p ojec ion sho e , almos hal he leng h o he diame e
o he i s loop o embolus, embolus o ming 7 loops in P. s ombuli o mis).
Desc ip ion. Male (holo ype): Small-sized. Body leng h 8.9, DS leng h 4.0,
wid h 3.5, OS leng h 4.3, wid h 2.9. Eye measu emen s: AME 0.18; ALE 0.32;
PME 0.25; PLE 0.31; AME–AME 0.11; AME–ALE 0.04; PME–PME 0.16; PME–
PLE 0.28; AME–PME 0.24; ALE–PLE 0.13; CH AME 0.28; CH ALE 0.28. Mea-
su emen s o palp and legs: Pp: 6.5 (2.0, 0.7, 1.1, 2.7); I 19.0 (5.3, 1.5, 5.8, 4.8,
1.6); II 20.7 (5.9, 1.6, 6.2, 5.2, 1.8); III 15.7 (4.6, 1.4, 4.4, 3.9, 1.4); IV 18.8 (5.6, 1.3,
5.0, 5.2, 1.7). Spina ion: Pp: 131, 101, 2111; legs: Fe I–II 323, III 322, IV 321; Pa
I–IV 001; Ti I –II 2026, III–IV 2126; M I–II 2024, III 3036, IV 3236. Leg o mula: II-
I-IV-III. Chelice ae wi h 3 p oma ginal, 4 e oma ginal ee h, and c. 16 den icles.
Palp (Fig. 1A–F): As in diagnosis. Tibia almos 1/3 o he leng h o cymbium,
RTA a ising medially om ibia, RTA wi h ounded dis al pa in en al iew,
dRTA quad angula in en al iew, en al dis al ibia wi h a sho blun apophy-
sis. Re op oximal cymbium wi h a sho blun p ocess (pa acymbium in Ramí ez
2014); al eolus ex ending in o dis al cymbial hal . Conduc o ili o m, a ising
om egulum a 11 o’clock posi ion. Embolic p ojec ion sligh ly cu ed, almos
2/5 o he leng h o cymbium. Embolus a ising sub-cen ally om egulum.
Colou a ion (Fig. 2A, B): Ca apace ligh o ange, wi h black spo s excep pos-
e io ma gin. Chelice ae ligh o ange, wi h black spo s. Legs yellow, wi h black
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He Zhang e al.: Pseudopoda species om Sou h China
spo s. S e num yellow, wi h o ange ma gins. Opis hosoma yellowish-g ey, do -
sum medially wi h ou ligh b own do s, en e la e ally wi h ligh b own spo s,
in on o spinne e s wi h a ligh b own V-shaped ma king.
Female: unknown.
Rema ks. This species shows simila palpal mo phology o Pseudopoda
s ombuli o mis (e.g., la ge al eolus, ex emely elonga ed and ili o m em-
Figu e 1. Pseudopoda ongxianensis sp. no . om Guangxi, le palp o male holo ype (A–C). A. Ven al; B. Do sal;
C. Re ola e al; D. Schema ic cou se o embolus, p ola e al iew; E. Schema ic cou se o embolus, en al iew; F. Le
male palpal ibia, e ola e al. Abb e ia ions: C, conduc o ; CP, cymbial p ocess; dRTA, do sal b anch o RTA; E, embolus;
EP, embolic p ojec ion; Sp, spe mopho ; S , sub egulum; T, egulum; RTA, en al b anch o RTA; VTA en al ibial apoph-
ysis. Scale ba s: 0.5 mm.
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He Zhang e al.: Pseudopoda species om Sou h China
bolus, and sho and quad angula dRTA), di e ing om o he congene s.
These wo species p obably belong o an unde ined species g oup. The
s ick-shaped embolus p ojec ion, and long and ili o m embolus a e simila
o Ma ensopoda Jäge , 2006, a genus endemic o he sou he n moun ain
anges o India (Jäge 2006). The e o e, we conside ed ha hese wo Pseu-
dopoda species may ep esen a ansi ional g oup be ween Pseudopoda
and Ma ensopoda, o ha he mo phological simila i ies may esul om
con e gen e olu ion.
Dis ibu ion. China (Guangxi Zhuang Au onomous Region, Fig. 8).
Pseudopoda de o mis Gong & Zhong, 2023
Figs 3, 4, 8
Pseudopoda de o mis Gong & Zhong, in Gong e al. 2023: 191, igs 1A–F, 2A–F,
3A–J (male holo ype, male and emale pa a ypes om Guanmenshan Sce-
nic A ea, Muyu Town, Shennongjia Fo es y Dis ic , Hubei P o ince, China,
deposi ed in HUST, examined).
O he ma e ial examined. China • Chongqing Municipali y: 1 emale, Wuxi Coun-
y, Shuangyang Township, Yin iaoling Na u e Rese e, Linkouzi, 31.4746°N,
109.8778°E, 1224 m, 22 June 2024, Luyu Wang leg. (CBEE, LJ20240053).
Diagnosis. See Gong e al. (2023). The addi ional emale examined has mo e
compac and medially shi ed copula o y duc s, consis en wi h in aspeci ic
a ia ion.
Desc ip ion. Female (Figs 3, 4): In e nal duc sys em mo e compac ly coiled,
FD mo e wis ed, medially shi ed (Fig. 3A, B). Fo u he desc ip ion, see Gong
e al. (2023).
Figu e 2. Pseudopoda ongxianensis sp. no . om Guangxi, male habi us. A. Do sal;
B. Ven al. Scale ba s: 2 mm.
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He Zhang e al.: Pseudopoda species om Sou h China
Figu e 3. Pseudopoda de o mis Gong & Zhong, 2023 om Chongqing, emale. A. Epigyne, en al; B. Schema ic cou se
o IDS, do sal; C. Vul a, do sal. Abb e ia ions: CO, copula o y opening; FD, e iliza ion duc ; FW, i s winding; IDS, In e nal
duc sys em; LL, la e al lobes. Scale ba s: 0.5 mm.
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He Zhang e al.: Pseudopoda species om Sou h China
Figu e 4. Pseudopoda de o mis Gong & Zhong, 2023 om Chongqing, emale habi us. A. Do sal; B. Ven al. Scale ba s: 2 mm.
Rema ks. All examined specimens a e conside ed conspeci ic based on he
gene al s uc u e and con igu a ion o he epigynal ield and ul al o gans. The
in e nal duc sys em is highly con olu ed and exhibi s no consis en o ien a ion
o symme y, e lec ing a high deg ee o s uc u al complexi y and no able in-
aspeci ic a ia ion.
Dis ibu ion. China (Hubei P o ince and Chongqing Municipali y, Fig. 8).
Pseudopoda spi alis Zhang, Jäge & Liu, 2023
Figs 5–8
Pseudopoda spi alis Zhang, Jäge & Liu, 2023 in Zhang e al. 2023: 247, igs
229A–C, 230A–B (male holo ype om Maoxian Moun ain Ecosys em Posi-
ioning S a ion, CAS, Sichuan, China, deposi ed in CBEE, examined).
O he ma e ial examined. China • Sichuan P o ince: 4 males, 4 emales, wi h
same da a as o holo ype, excep o : 25–26 May 2015, Gui-Qiang Huang, Lu-
Yu Wang & Yan-Chen Zhou leg. (CBEE, LJ20240001–LJ20240003, LJ20240007,
LJ20240015, LJ20240096–LJ20240098).
Diagnosis. The emale o his species can be dis inguished om congene s
by i s in e nal duc sys em, which is elonga e, na ow, and o ms a compac ,
longi udinally aligned looping pa e n, clea ly isible in do sal iew.
Desc ip ion. Male (Figs 5, 7A, B): o de ails see Zhang e al. (2023).
Female (LJ20240007): Measu emen s: Small-sized. Body leng h 8.2, DS
leng h 3.3, wid h 3.1, OS leng h 4.9, wid h 3.7. Eyes: AME 0.13, ALE 0.20, PME
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He Zhang e al.: Pseudopoda species om Sou h China
0.16, PLE 0.18, AME–AME 0.17, AME–ALE 0.07, PME–PME 0.21, PME–PLE
0.32, AME–PME 0.24, ALE–PLE 0.22, CH AME 0.36, CH ALE 0.30. Spina ion: Pp
131, 101, 2121, 1014; Fe I–II 323, III 322, IV 321; Pa I–III 101, IV 000; Ti I–II 2228,
III–IV 2126; M I–II 3034, III–IV 3036. Measu emen o palps and legs: Pp 3.9
(1.0, 0.6, 0.8, –, 1.5); I 10.3 (3.1, 1.2, 2.6, 2.4, 1.0), II 13.3 (3.6, 1.5, 3.9, 2.9, 1.4),
III 9.2 (2.9, 1.1, 2.0, 2.3, 0.9), IV 10.7 (3.4, 1.0, 2.4, 2.7, 1.2). Leg o mula: II-IV-I-III.
Chelice ae wi h h ee p oma ginal, ou e oma ginal ee h, and c. 22 den icles.
Epigyne (Fig. 6A–C): As in diagnosis. Epigynal ield as wide as long, wi hou
an e io bands. An e io ma gins o la e al lobes o ming a dis inc V-shape,
median ma gins o la e al lobes ouching each o he along he middle line. In-
e nal duc sys em ex ends longi udinally and clea ly isible in do sal iew, an-
e io ma gins sligh ly co e ed by FW. FD a ising pos e o-la e ally.
Colou a ion (Fig. 7C, D): Ca apace o ange o eddish b own, wi h dis inc a-
dial s eaks ex ending om he o ea o he ma gins. Eye egion da k b own.
Chelice ae o ange o eddish b own; endi es and labium o ange-b own. Legs
ligh o ange, wi h nume ous da k eddish-b own spo s. OS do sally yellowish
b own wi h a da ke median a ea and a ans e sal whi e line in pos e io pa .
OS en ally ligh yellow wi h i egula b own spo s, dense la e ally.
Dis ibu ion. China (Sichuan P o ince, Fig. 8).
Figu e 5. Pseudopoda spi alis Zhang, Jäge & Liu, 2023 om Sichuan, le male palp. A. Ven al; B. Do sal; C. Re ola e al.
Abb e ia ions: C, conduc o ; dRTA, do sal b anch o RTA; E, embolus; EP, embolic p ojec ion; Sp, spe mopho ; S , sub eg-
ulum; T, egulum; RTA, en al b anch o RTA. Scale ba s: 1 mm.
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He Zhang e al.: Pseudopoda species om Sou h China
Figu e 6. Pseudopoda spi alis Zhang, Jäge & Liu, 2023 om Sichuan, emale. A. Epigyne, en al; B. Schema ic cou se o
IDS, do sal; C. Vul a, do sal. Abb e ia ions: CO, copula o y opening; FD, e iliza ion duc ; FW, i s winding; IDS, In e nal
duc sys em; LL, la e al lobes. Scale ba s: 0.5 mm.
